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Soo/h cr> /'/^i 




28, e ?e 






Printed from the 
W. W. Whitney Publication Endowment 



Printed for the Society 



f^r Zoo'ogv »*; 

NOV 29 1938 ' 


U. S. Grant, IV, Chairman 

L. M. Klauber Clinton G. Abbott, Editor 


1. A New Subspecies of Pocket Gopher from Sonora, Mexico. By 

Laurence M. Huey. Published August 10, 1934 1-2 

2. A New Puff-Bird from El Salvador. By A. J. van Rossem. Pub- 

lished August 10, 1934 3-4 

3. Notes on the Races of Claravis mondetoura. By A. J. van Rossem. 

Published August 10, 1934 5-8 

4. Notes on Some Races of Ceophloeus lineatus (Linnaeus) . By A. J. 

van Rossem. Published August 10, 1934 9-12 

5. Three New Species of Pinnixa from the Gulf of California. By 

Steve A. Glassell. Published March 21, 1935 13-14 

6. New Marine Mollusca from West Mexico, Together with a List of 

Shells Collected at Punta Penasco, Sonora, Mexico. By Herbert 

N. Lowe. Published March 21, 1935 15-34 

7. New Species of Mollusks of the Genus Triphora. By Fred Baker 

and V. D. P. Spicer. Published March 21, 1935 35-46 

8. New Trilobite Species from the Anthracolithic of Northern Cali- 

fornia, and Griffithides conwayensis, a New Name for a Trilo- 
bite Species from the Atoka Formation of Arkansas. By Harry 
E. Wheeler. Published March 21, 1935 47-58 

9. Revision of Some California Species of Astrodapsis. By George L. 

Richards, Jr. Published March 21, 1935 59-66 

10. The Mangrove Warbler of Northwestern Mexico. By A. J. van 

Rossem. Published August 24, 1935 67-68 

11. A New Race of Brown Towhee from the Inyo Region of California. 

By A. J. van Rossem. Published August 24, 1935 69-72 

12. A New Silky Pocket Mouse from Sonora, Mexico. By Laurence 

M. Huey. Published August 24, 1935 73-74 

13. A New Subspecies of Crotalus confluentus, the Prairie Rattlesnake. 

By Laurence M. Klauber. Published August 24, 1935 75-90 

14. New or Little Known Crabs from the Pacific Coast of Northern 

Mexico. By Steve A. Glassell. Published August 24, 1935 91-106 

15. A New Genus and Species of Pigmy Goose from the McKittrick 

Pleistocene. By Roland Case Ross. Published August 24, 1935.107-114 

16. Description of a Race of Myiarchus cinerascens from El Salvador. 

By A. J. van Rossem. Published March 12, 1936 115-118 

17. A New Pelecypod Genus of the Family Cardiidae. By A. Myra 

Keen. Published March 12, 1936 119-120 


San Diego Society of Natural History 

18. Notes on Birds in Relation to the Faunal Areas of South-Central 

Arizona. By A. J. van Rossem. Published May 29, 1936 121-148 

19. Crotalus mitchellii, the Speckled Rattlesnake. By Laurence M. 

Klauber. Published May 29, 1936 149-184 

20. A Key to the Rattlesnakes With Summary of Characteristics. By 

Laurence M. Klauber. Published December 7, 1936 185-276 

21. New Porcellanids and Pinnotherids from Tropical North Ameri- 

can Waters. By Steve A. Glassell. Published December 7, 1936..277-304 

22. West American Species of the Genus Phos. By A. M. Strong and 

H. N. Lowe. Published December 1, 1936 305-320 

23. A Further Report on Birds of Sonora, Mexico, with Descriptions 

of Two New Races. By A. J. van Rossem and The Marquess 
Hachisuka. Published June 15, 1937 321-336 

24. Paleontology of the Pleistocene of Point Loma, San Diego County, 

California. By Robert W. Webb. Published June 15, 1937 337-348 

25. Descriptions of New Mammals from Arizona and Sonora, Mexico. 

By Laurence M. Huey. Published June 15, 1937 349-360 

26. A Northwestern Race of the Mexican Black Hawk. By A. J. van 

Rossem and The Marquess Hachisuka. Published June 15, 1937-361-362 

27. A New Snake of the Genus Sonora from Mexico. By Laurence M. 

Klauber. Published December 15, 1937 363-366 

28. A New Sea-Urchin from the "Oligocene" of Oregon. By Hubert 

Lyman Clark. Published December 15, 1937 367-374 

29. An Extinct Puffin from the Pliocene of San Diego, California. 

By Loye Miller. Published December 15, 1937 375-378 

30. An Upper Pleistocene Fauna from the Baldwin Hills, Los Angeles 

County, California. By George Will ett. Published December 15, 

1937 379-406 

31. A New Hummingbird of the Genus Saucerottia from Sonora, Mex- 

ico. By A. J. van Rossem and The Marquess Hachisuka. Pub- 
lished January 18, 1938 407-408 

32. A New Muskrat from Utah. By Laurence M. Huey. Published 

January 18. 1938 409-410 

33. New and Obscure Decapod Crustacea from the West American 

Coasts. By Steve A. Glassell. Published May 31, 1938 411-454 

34. A Study of the Skull of the Pleistocene Stork, Ciconia maltha, 

Miller. By Loye Miller. Published May 31, 1938 455-462 





Vol. VIII, No. 1, pp. 1-2 August 10, 1934 



Laurence M. Huey 

Curator of Birds and Mammals, San Diego Society of Natural History 

During February, 1934, a party representing the San Diego Society 
of Natural History made collections in the vicinity of Punta Penascosa, 
on the northeast coast of the Gulf of California, Sonora, Mexico. Among 
the limited number of mammals obtained is a series of 12 Thomomys 
which represents an uradescribed form of the wide-ranging bottae group. 
This race may be know« as : 

Thomomys bottae vanrossemi subsp. nov. 

Punta Penascosa Pocket Gopher 

Type. — From Punta Penascosa, Sonora, Mexico; no. 10922, collection of 
the San Diego Society of Natural History; adult male; collected by Laurence 
M. Huey, February 15, 1934. 

Characters. — In color vanrossemi is almost identical with its nearest relative, 
Thomomys bottae phasma from southwestern Arizona, but cranially some 
characters showing constant difference are present. The skull is relatively nar- 
rower than that of phasma with less widely spreading zygomatic arches, narrower 
rostrum and more fully inflated bullae, that bulge farther below the basioccipital. 

Measurements. — Type: Total length, 215; tail, 72; hind foot, 28; ear, 4. 
Skull (type) : Greatest length, 38.5; spread of maxillary arches, 24.0; length of 
nasals, 13.5; interorbital constriction, 6.5; alveolar length of upper molar series, 

Range. — So far as known, the vicinity of Punta Penascosa, Sonora, Mexico, 

2 San Diego Society of Natural History 

where it was found living at almost tide level on the land side of a series of large 
sand dunes that bordered the sea beach. 

Remarks-— Since Nelson and Goldman have recently described a number 
of forms of Thomomys from Sonora (Journ. Mam., Vol. 15, no. 2, May, 1934, 
pp. 105-124), the writer forwarded this series of specimens to Washington, 
where it was compared with topotypical material by Major Goldman and the 
differences confirmed. For this aid the writer's thanks are here expressed. 

Specimens examined by the writer. — Thomomys bottae phasma: 5 from 
Tinajas Altas and 13 from 4 miles south of Gadsden, Yuma County, Arizona; 
Thomomys bottae vanrossemi: 12 from Punta Pehascosa, Sonora, Mexico [type 
locality] . 

This race is dedicated to Mr. A. J. van Rossem of the California 
Institute of Technology, a life-long friend of the author and co-worker in 
the field of natural science. 



Vol. VIII, No. 2, pp. 3-4 August 10, 1934 



California Institute of Technology 

In view of Ridgway's remarks (Birds of North and Middle America, 
6, 1914, 377, footnote) concerning the individual variation found in 
Dyson's Puff-Bird, the eight specimens of this species which I collected 
in El Salvador in 1925, 1926, and 1927 had been referred to Notharchns 
hyperrhyncbus dysoni, purely on assumption. Last fall when at the Mu- 
seum of Comparative Zoology, I chanced to see a large series of dysoni 
which were so very different from my recollection of the Salvador birds 
that I asked Mr. Peters to forward a good selection in order to make 
possible a direct comparison. Further specimens were borrowed from the 
Bureau of Biological Survey and the Smithsonian Institution. The total 
series of dysoni examined at this time numbers 28 specimens from prac- 
tically the entire Central American range save for the Pacific coast of 
Nicaragua and Guatemala. 

The Salvador birds represent a strongly characterized race which 
evidently occupies a limited territory on the Pacific coast of Central 
America. This race is here named 

Notharchus hyperrhynchus cryptoleucus subsp. nov. 

Type. — Male adult, no. 18714 Dickey collection; Barra de Santiago, Dept. 
of Ahuachapan, El Salvador, April 3, 1927, altitude sea level; collected by A. J. 
van Rossem, original number 11580. 

Subspecific characters.— Compared with Notharchus hyperrhynchus dysoni: 
feathers of lower rump and upper tail -coverts with one or two concealed grayish- 
white bars, instead of being immaculate proximal to the terminal tipping; dark 

4 San Diego Society of Natural History 

portions of the plumage everywhere, save on the crown, paler, duller, and more 
slaty, and with the white margins notably wider; rump and upper tail-coverts 
"mouse gray" instead of "dark mouse gray" or "blackish mouse gray"; flanks 
with dr.rk area much restricted and narrowly barred with brownish slate and 
white in about equal widths instead of being black with narrow white bars. 

Range. — Coastal plain of El Salvador and, probably, closely adjacent areas 
in western Guatemala and western Nicaragua. 

Remarks. — The concealed barring is a feature which is apparently totally 
absent in dysoni and always present in cryptoleucus. Aside from this, cryptoleucus 
is easily distinguished by general pallor of coloration and wide white tipping and 

Sclater's type of Bucco dysoni came from "Honduras" and was collected 
by Dyson. This necessarily restricts the type locality to northwestern Honduras, 
since the chief purpose of Dyson's visit was to secure live specimens of the 
Ocellated Turkey for the aviary of Lord Derby. 

Specimens of dysoni have been examined from Honduras (Santa Ana); 
British Honduras (Toledo District) ; Guatemala (Gualan) ; Costa Rica (Pigres; 
Boruca) ; Panama (Loma del Leon; Panama City; Almirante; Fruitdale) ; Chia- 
pas (San Benito). I can make out no variation in this extensive range, that is 
other than individual or seasonal in character. The margins to the dark parts of 
the plumage are naturally wider when the feathers are fresh and gradually disap- 
pear with wear. 

The fact that dysoni occurs in typical form at the Chiapas-Guatemala boun- 
dary on the Pacific coast, and as far north as Pigres on the Gulf of Nicoya in 
Costa Rica, argues a relatively limited range for cryptoleucus. However, since 
cryptoleucus was found to inhabit the full length of the coastal plain of El Salva- 
dor, it would be remarkable if it did not extend for some slight distance into the 
adjoining portions of Guatemala and Nicaragua. 



Vol. VIII, No. 3, pp. 5-8 August 10, 1934 




California Institute of Technology 

Ludlow Griscom (Occ. Pap. Bost. Soc. Nat. Hist., 5, June 14, 
1930, pp. 287-292) has recently recognized four races of this little dove, 
though he was able to assemble little more than a dozen specimens from 
American collections. In the interval since Griscom's paper appeared I 
have had the opportunity to examine most of his material and also some 
twenty additional skins in Europe and England. As might be expected 
the foreign material modifies, somewhat, the characters of the four races 
defined by Griscom. This comment is given below and, in addition, two 
more races are described, respectively from the southern and northern 
extremes of the range of the species. 

Claravis mondetoura mondetoura (Bonaparte) 

As first noted by Count Salvadori (Cat. Birds Brit. Mus., 21, 1893, 495) 
and later by Ridgway and Griscom, South American specimens are sharply dis- 
tinguished from those of Central America and Mexico by the reddish intermix- 
ture in the slate color of the under wing-coverts and axillars of the males. 

The typical race, mondetoura, is distinguished from all the others, both north 
and south, by the longer wing. Seven fully adult males measure: wing, 117-121; 
tail, 85-91 mm. Five fully adult females measure: wing, 115-118; tail, 74-82 mm. 
Bonaparte's types (male and female) from Caracas, Venezuela, were examined 
in the Paris Museum. They are typical of the northern South America sub- 

Sixteen specimens of both sexes and representing adult and immature indi- 
viduals have been examined from Venezuela, Colombia and Ecuador. Age for 

6 San Diego Society of Natural History 

age, I am unable to see any differences between birds from these three countries 
and believe that the characters noted by Griscom for some Colombia females 
were due to age. Old females are very much grayer than immatures. 

Claravis mondetoura inca subsp. nov. 

Type. — Male adult, no. British Museum; Huasampilla, Peru, 
March 18, 1872; collected by H. Whitley. 

Subspecific characters. — Adult male with under wing-coverts and axillars 
much more extensively rufous than in typical mondetoura, in fact with rufous in 
excess of slate; flanks paler gray, and posterior median underparts more exten- 
sively white than in any other race. Wing shorter than in mondetoura. The type, 
which is apparently the only known specimen, measures: wing, 111; tail, 76; 
exposed culmen, 13.5; tarsus, 24.0; middle toe minus claw, 21.5 mm. 

Range. — Known only from the type locality. 

Remarks. — Colors of soft parts as recorded by the collector are: "Bill black; 
eye pink; legs and toes lead colour." This is not at all in agreement with the 
colors given by Salvin and Godman in the 'Biologia' for an adult male of salvini 
from Guatemala. 

Claravis mondetoura pulchra Griscom 
Claravis mondetoura umbrina Griscom 

Western Panama and Costa Rica specimens appear to be non-existent in 
European collections. Therefore, I have nothing to add to the diagnosis of either 
of these races. I have examined the same specimens used by Griscom, however, 
and believe that they are both valid subspecies. 

Claravis mondetoura salvini Griscom 

This race, though described on the basis of a single adult male, is verified 
through two males and a female from Volcan de Fuego in the British Museum. 
Males are indistinguishable from mondetoura dorsally. They show more white 
ventrally, but this may be due to the "make" of the skins. The supposed 
character of a more extensively black tail does not prove constant. In other 
words the males of saWnu may be distinguished from mondetoura by the posses- 
sion of uniform slaty (instead of mixed slaty and rufous) under wing-coverts 
and axillars; and shorter wing with proportionally longer tail. 

A single female (also from Volcan de Fuego), is very close to mondetoura, 
but differs in having a slightly darker and less reddish rump and upper tail-coverts, 
more slaty (less reddish) under wing-coverts, and shorter wing. 

Two adult males measure: wing, 110-114; tail, 79-83 mm. One adult female 
measures: wing, 110; tail, 79 mm. 

Though Volume 21 of the 'Catalogue of Birds' cites one male and two 
females from Volcan de Fuego, this is in error for there are two males and one 
female from that locality in the British Museum collection at this time. One of 
the males was recorded by Salvin and Godman as having the colors of the soft 
parts as follows: "Iris reddish-orange; bill black; tarsi and toes dull red; claws 

van Rossem — Races of Claravis Mondetura 

Claravis mondetoura ochoterena subsp. nov. 

Type. — Male adult no. British Museum; Jalapa, Vera Cruz, 
Mexico, 1872; collected by [Rafael Montes] de Oca. 

Subspecific characters. — Adult males similar to Claravis mondetoura salvini 
of Guatemala, but dorsal coloration darker and more fuscous (less grayish) 
slate; underparts darker everywhere and with the red of the pectoral region ex- 
tending back laterally to tinge the slate color of the flanks. Female unknown. 

Range. — Mountains of the State of Vera Cruz, Mexico. 

Remarks.— Ridgway (Birds of No. and Mid. Amer., Pt. 7, 1916, 436, 
footnote) , has remarked on the characters shown by the two subadult males in 
the United States National Museum, but he suspected that the peculiarities 
might be due to age. The fully adult male in the British Museum shows that such 
is not the case. The Mexican race is easily distinguished by being the darkest 
of the known subspecies. 

Three males, two of them subadult, but which possess the primaries and 
rectrices of maturity, measure: wing, 107-113; tail, 72-83 mm. 

I take pleasure in naming this dove for Professor Isaac Ochoterena, 
Director of the Biological Institute of Mexico, not only in recognition of 
his own work but in appreciation of his assistance in furthering my own 
activities in northwestern Mexico. 



Vol. VIII, No. 4, pp. 9-12 August 10, 1934 




California Institute of Technology 

A few years ago J. L. Peters briefly reviewed 1 the races of this species, 
but in certain instances he was handicapped either by lack of material or 
because type specimens were not available for critical examination. It has 
been my good fortune to be able to study, during the summer of 1933, 
types of some of the races described by European ornithologists, and it is 
now possible to settle some, at least, of the questions concerning which 
Peters was obliged to make arbitrary decisions. Notes on these types and 
changes in nomenclature (where necessary) are given below. 
Ceophloeus mesorhynchus Cabanis and Heine 

The three specimens on which this name 2 is based are still in the Zoological 
Museum in Berlin, where they are numbered 16,292-3 and 4, respectively. They 
are all mounted in fair condition though all show some degree of moult. 

This name comes dangerously close to being a synonym of Picus similis 
Lesson. One of the three cotypes ( 8 no. 16,292) is a white-billed bird from the 
Aguacate Mountains, collected by Hoffmann; another, ( $ no. 16,293), col- 
lected by von Frantzius in "Costa Rica," has a pale yellowish brown bill and is 
also intermediate in other particulars. Only one specimen, ( $ no. 16,294), ful- 
fills all requirements and is typical of the race which has so long borne the name 
mesorhynchus. Although the original description expressly calls for "Bill bright 
bluish-horncolor," the name "mesorhynchus" would seem to indicate recognition 

1 Occ. Pap. Bost. Soc. Nat. Hist., 5, Sept. 2, 1930, pp. 317-22. 

2 Mus. Heir.eanum, 4, heft 2, 1863, 86. See also critical comment on these specimens 
in Journ. fur Orn., 1862, 176. 

10 San Diego Society of Natural History 

of the intermediate character of the bill when the three birds as a whole were 

Since the Cabanis and Heine description is a composite, based on two races, 
it is proper to designate one of the three cotypes as the type. Therefore I name 
number 16,294, as the only course which will permit (for the time at least) reten- 
tion of this long-established name for the dark-billed race of the Pileated Wood- 
pecker in eastern and southern Costa Rica. 

This specimen is a mounted bird, a female, presumably adult, which has 
nearly completed the annual moult. It has the following measurements: wing, 
181; tail, 122; culmen from base, 35.0; tarsus, 26.7; outer anterior toe minus 
claw, 20.5. The bill is dark horn-color on the maxilla and basal two-thirds of the 
mandible. The tip of the mandible is dull brown. Though the type locality as 
published was simply "Costa Rica," Dr. von Frantzius has given a good indica- 
tion of the locality in which his two specimens were collected. In his paper on the 
distribution of birds in Costa Rica he states, under the name of Dryocopus 
scapularis? that the species is relatively rare and is known only from the Aguacate 
Mountains and on the Sarapiqui. The Aguacate record was made by Hoffmann, 
and therefore the locality from whence came Dr. von Frantzius' two specimens 
must necessarily be the Sarapiqui River. This is a region of intergradation between 
mesorhyncbus and similis. It may be that the male, which is an intermediate 
(though nearer similis) , was taken at a point further down stream and closer to 
the Honduras Boundary than was the female, which is typical of the southern 

There is still the possibility that a good series of specimens from even the 
headwaters of the Sarapiqui would show a mass average considerably closer to 
similis, and in that case the name mesorhyncbus will have to become a synonym 
of the former race. It is significant that Ridgway 4 has considered a specimen 
from the Rio Revantazon, a point considerably to the south of the Sarapiqui, as 
being an intermediate, which is closer to similis than to mesorhyncbus. I have 
tried to make out a case which will permit the retention of an old name rather 
than to scrap it for a new one, though until adequate material from the type 
locality is collected there is no certainty that even the arbitrary selection of a type 
finally disposes of the matter. 

Campephilus leucorhamphus "Licht." Reichenbach 

Peters is entirely correct in considering this name a synonym of Picus siniilis 
Lesson, for two of the three specimens which were the basis of Lichtenstein's 
manuscript name 5 are still present in the collection of the Zoological Museum in 
Berlin. Both are females (the male possibly went to Reichenbach, though I could 
not find it at Dresden), and are normal representatives of similis. They were 
collected by Ferdinand Deppe; no. 10375 at "Cosmalvapan" | =Cosamaloapam], 
and no. 10376 at Alvarado. Both of these localities are in southern Vera Cruz. 

3 Joum. fur Orn., 1869, 364. 

4 Birds of No. 6C Mid. Amer., pt. 6, 1914, 151, footnote. 

5 Nom. Aves. Mus. Berol., 1854, 75. 

van Rossem — Races of Ceophloeus Lineatus 1 1 

The wing measurement of these two cotypes, which are mounted and in fair 
condition, are 174 and 167 mm., respectively. 

Campephilus lineatus Var.? C. leucopterylus Reichenbach 

Reichenbach's type of leucopterylus 6 is in the Dresden Museum. It is an 
adult male, a skin in good condition and bears the following label "Geophloeus 
[sic] lineatus (L.) /7487/ Siid-Amerika." This relatively new label has the old, 
badly torn, original pasted on the reverse. All that is left is the single word 
"'Leucopterylus" and the terminal letter (n) of a word which once was written 
on the now missing lower left hand corner of the label. Dr. Wilhelm Meise, the 
curator of birds at the Dresden Museum, told me that this skin (formerly 
mounted) is a part of the old collection of Reichenbach's regime. So far as Dr. 
Meise could determine there never was a female leucopterylus in the collection 
and he thinks it possible that the figure of the female was pictured by the analogy 
of the color characters of allied species. 

This specimen belongs to none of the northern races but is one of those rare 
individuals of Ceophloeus erythrops which show traces of a white scapular 
stripe! It fits the colored plate and accompanying description perfectly, even to 
the light spots on the tips of the alula and primaries, and my own measurement 
of the wing, 191 mm., is exactly the length given by Reichenbach (7" 4"' using 
the Rhineland foot) . 

The missing corner of the label, which presumably bore the locality, may 
have been torn off even prior to the time leucopterylus was described, and Reich- 
enbach evidently guessed at the source when he ascribed his bird to "Mexiko." In 
the Berlin Museum there is another example of erythrops which shows the same 
type of coloration. It was collected by Euler at Cantagallo, Province of Rio de 
Janeiro, Brazil, and it is not impossible that Reichenbach's bird has the same 
origin. Dr. Meise could give me no definite information as to why the "Siid- 
Amerika" had been put on the newer label. 

Whether traces of white scapular stripes in occasional specimens of erythrops 
indicate intergradation with lineatus or whether the character is a sporadic one 
which appears in a limited number of individuals, regardless of locality, I do not 
know. At any rate it is doubtful if erythrops is entitled to more than subspecific 
rank as a race of lineatus. 

The large race of lineatus in north-eastern Mexico, Ceophloeus lineatus 
leucopterylus Peters (nee. Reichenbach), is here named as 

Ceophloeus lineatus petersi subsp. nov. 

with the type an adult male, no. 31,833 Dickey collection at the California Insti- 
tute of Technology; Cuidad Victoria, Tamaulipas, Mexico, March 3, 1908; col- 
lected by F. B. Armstrong. Measurements of the type are: wing, 190; tail, 116; 
exposed culmen, 38.0; tarsus, 32.6; outer anterior toe, 26.5. 
Picus scapularis Vigors 
Peters had only two birds which he considered to be referable to scapularis, 
a male and female taken at Alamos, Sonora. 

6 Handb. Scans. Picinae, 1854, 392, pi. 647, figs. 4319-4320. 

12 San Diego Society of Natural History 

The type locality of Picus scapularis is San Bias, in what is now the state 
of Nayarit, western Mexico. Although Vigors' types of this race are stated to 
have gone to the Zoological Society, they are not now in the collection at the 
British Museum and were probably long ago destroyed. However, I have ex- 
amined two specimens of this race from the type locality (in the collections of the 
British Museum and U. S. Biological Survey) as well as eleven others from 
various points in Guerrero, Jalisco and southern Sinaloa north to Mazatlan. The 
race scapularis as represented by specimens from the above states is not markedly 
different in color from similis, the race which occurs over most of Central America 
north to central Vera Cruz on the Atlantic coast and to central Oaxaca on the 
Pacific. The only color differences which I can observe are that scapularis is 
slightly less buffy below as an average character, and the white sub-ocular streak 
is narrower and is often broken up by the intrusion of black streaks from the 
surrounding areas. These characters are fairly uniform over the entire area out- 
lined above, though Guerrero specimens are, naturally, very close to similis. In 
northern Sinaloa and southern Sonora there is" an abrupt color change, in that 
the sub-ocular streak becomes practically obsolete, the ground color of the under- 
pays is whitish rather than buffy, and the wing lining is cream-colored instead 
of distinctly yellow. On a color basis, therefore, scapularis, though uniform over 
a large area, is an intermediate between similis and birds of the Alamos district 
of Sonora and northern Sinaloa. 

In size, however, scapularis is small, decidedly smaller than similis and 
slightly smaller than the Alamos birds. The largest male out of five from Ma- 
zatlan, the northernmost point from which I have seen specimens of scapularis, 
has a wing and tail length of 167 and 105 mm. respectively. Ridgway gives essen- 
tially the same figures as the average for the race, but I suspect that central 
Sinaloa specimens are included. 

Specimens from the Alamos Faunal Area represent the pale extreme reached 
by Ceophloeus lineatus in North America. The race is here described as 

Ceophloeus lineatus obsoletus subsp. nov. 

Type.— Adult male, no. 224,294, Mus. Comp. Zobl.; Alamos, southern 
Sonora, Mexico; March 16, 1888; collected by M. A. Frazar. 

Subspecific characters. — Nearest to Ceophloeus lineatus scapularis (Vigors) 
of central western Mexico, but ground color of underparts pale buffy white or 
grayish white instead of pale buff; wing lining cream-color instead of light yellow; 
sub-ocular and sub-auricular streak nearly obsolete; wing and tail slightly longer. 

Range. — The Alamos District in southern Sonora, northern Sinaloa and 
probably the adjacent portions of Chihuahua. 

Remarks. — Seven specimens of this race have been examined from Alamos, 
Sonora, and from Rosario and La Guasimas in extreme north-eastern Sinaloa. 
Two, from Rosario and La Guasimas, respectively, are young birds. Five adults 
of obsoletus average (sex ignored): wing, 172.3; tail, 110.5. The characters as 
given by Peters for "scapularis" apply, of course, to obsoletus. 

In addition to the institutions mentioned above I am indebted to 
Robert T. Moore for the loan of specimens from northern Sinaloa. 

2* Y?7 



Vol. VIII, No. 5, pp. 13-14 March 21, 1935 



Steve A. Glassell 

Research Associate in Crustacea, San Diego Society of Natural History 

The crabs here described were collected at the upper end of the 
Gulf of California, near the village of San Felipe, in Lower California, 
Mexico, by the author. Fuller descriptions and plates of the following 
species will appear in a forthcoming partial revision of the genus Pjnnixa 
by the writer. 

Pinnixa abbotti, sp. nov. 

Type. — Female, San Diego Society of Natural History, Cat. No. 208; San 
Felipe, Lower California, June 10, 1933. Allied to P. floridana. 1 Carapace a little 
wider than twice the length. Antero-lateral margin rounded, marked with a dis- 
tinct punctate ridge, extending from the cervical groove and meeting the concave 
postero-lateral margin at an acute angle. Entire carapace coarsely punctate; 
pubescent near outer angles; a transverse groove between gastro-cardiac regions; 
a median sulcus. Front truncate, slightly concave, entire. Mesogastric region 
depressed. Chelipeds weak, hairy, pubescent; carpus punctate; chela tapering dis- 
tally; inside of hand smooth; fingers feebly denticulate, nearly horizontal, not 
gaping, tips hooked. Length of carapace 2.95, width 6.5 mm. 

Pinnixa fusca, sp. nov. 

Type. — Female, San Diego Society of Natural History, Cat. No. 209; San 
Felipe, Lower California, May 29, 1934. Allied to P. longipes. 2 Carapace slightly 

1 Bull. 97, U. S. Nat. Mus., 1918, pi. 30, figs. 4-7. 

2 Proc. Calif. Acad. Sci., ser. 2, vol. 4, 1894, p. 573, pi. 20, figs. 19-20. 

14 San Diego Society of Natural History 

more than three times the length, and rounding down to margins, smooth; regions 
poorly denned. Front truncate, slightly convex, not extending beyond carapace; 
posterior margin concave. Chelipeds equal; upper distal end of merus extends 
over carpus; carpus and hands smooth; hands rounded, tapering distally; fingers 
slightly gaping, one large central tooth; pollex slightly deflexed, tip hooked. 
First and second ambulatory legs, slight, third and fourth, heavy; fourth reaches 
nearly to middle of propodus of third; dactyls of first and second legs long and 
slim, curved third and fourth, long and heavy, curved at tip, upper crest of tip 
of third dactyl, tri-dentate, fourth smooth. Abdomen covers sternum. Length of 
carapace 3.7, width 11.5 mm. 

Pinnixa felipensis, sp. nov. 

Type.— Female, San Diego Society of Natural History, Cat. No. 210; San 
Felipe, Lower California, June 1, 1934. Carapace slightly more than 21/ 2 times 
the length, and rounding down to margins in front, smooth, depressed in central 
portions; regions defined; an unbroken, straight, transverse cardiac ridge, extend- 
ing across the carapace; front very broadly triangular, concave toward orbits; 
posterior margin straight. Chelipeds unequal, dissimilar; merus long, increasing 
in breadth with length, carpus narrow, long, overhanging the manus, smooth; 
hands crested, wide, smooth; pollex of larger hand, short, horizontal, thick; of 
smaller hand, longer, lighter and pointed; fingers, greatly curved, gaping, a tuft 
of dense pubescence in the gape. First and second ambulatory legs, slight, dactyls 
long, tapering, bi-curved; dactyls of third and fourth legs, long, lanceolate, 
straight; dactyl of fourth leg reaches carpus of third leg. Abdomen nearly covers 
sternum. Length of carapace 3.2, width 8.3 mm. 



Vol. VIII, No. 6, pp. 15-34, plates 1-4 March 21, 1935 



Herbert N. Lowe 

San Diego Society of Natural History 

The following undescribed species of mollusks were taken by the 
author on the west mainland coast of Mexico in 1930 and 1931, at various 
points in the Gulf of California in 1932, at San Felipe, Lower California, 
in May, 1933, and at Punta Penasco, Sonora, in February, 1934. Descrip- 
tive accounts of the various collecting trips will be found in "The Nau- 
tilus" as follows: Mazatlan-Guaymas-Topolobampo, Vol. XLIII, pp. 
135-138; Tres Marias-Manzanillo-Acapulco, Vol. XLIV, pp. 24-27; 
Gulf of California and its islands, Vol. XLVI, pp. 73-76, 109-115; San 
Felipe, at the head of the Gulf, Vol. XLVII, pp. 45-47; Punta Penasco, 
on the northern Sonora coast, Vol. XLVIII, pp. 1-4, 43-46. 

I am indebted to Mr. J. R. Pemberton of Los Angeles for the privilege 
of being included in the party of scientists which cruised in the Gulf 
of California during the early part of 1932. I also wish to thank the offi- 
cials of the San Diego Society of Natural History for facilities provided 
on the co-operative expedition to Punta Penasco, Sonora, in February, 
1934. The splendid research library of Dr. U. S. Grant at the University 
of California at Los Angeles was of great assistance in the preparation 

1 Also known as Punta Penascosa. 

16 San Diego Society of Natural History 

of the paper. Finally, the fullest credit is due to Mr. Ernest H. Quayle 
of the Department of Geology, University of California at Los Angeles, 
for the excellent pen and ink drawings here reproduced. 

Area gordita, new species. Plate 1, fig. 1. 

Acapulco. 20 fathoms (1931). Type 11387, Lowe collection; paratype, 
Lowe collection, Guaymas, 20 fathoms, (1932). 

Shell irregularly ovoid, solid; anterior end roundly sloping downward and 
backward, posterior end roundly, obliquely produced; color flesh-white; about 
27 radiating ribs with somewhat wider interspaces; dorsal margin nearly straight, 
ventral margin much rounded; greatest altitude of shell in almost vertical line 
with umbones; anterior wing more sharply produced than the posterior. 

Diameter 30 mm., altitude 19.1 mm. 

This shell has about nine less ribs than A. aviadoides Rve., and is a thicker 
and heavier shell for the same size. 

This species and the two following, A. delgada and A. reinharti, have 
subsequently turned up in the dredgings of the Templeton Crocker Expedition 
off West Mexico, (August, 1933) . 

Area delgada, new species. Plate 1, fig. 2. 

Manzanillo, 20 fathoms (1930). Type 11388, Lowe collection. 

Shell very obliquely ovoid, rather solid; anterior end roundly sloping down- 
ward and backward; posterior end roundly obliquely produced; color gray white; 
about thirty delicately nodulous ribs, growing smaller and closer together toward 
the anterior end. Between the longest four ribs are delicate riblets. The shell is 
rather flat s"ided and the ligamental area about normal. 

Diameter 12.3 mm., altitude 8 mm. 

Area (Anadara) reinharti, new species. Plate 1, figs. 3a, 3b, 3c. 

Guaymas, 20 fathoms (1932). Type 11389, Lowe collection. 

Shell ivory white, covered with a brown, horny epidermis; obliquely rhom- 
boid, solid, equivalve; edges of valves thick; anterior end rounded, posterior end 
angularly extended downward; about 25 radiating ribs, with narrow inter- 
spaces; the ribs toward the anterior end are strongly nodulous. The shell some- 
what resembles a miniature specimen of A. multicostata Sby., but is much more 
oblique, has about ten less ribs, and has a ligamental area (in specimens of equal 
size) of three times the diameter. In young A. multicostata the left valve over- 
laps the right. 

In young A. multicostata the edges of the valves are quite thin, while those 
of the species under discussion are abnormally thickened; the grooves on the 
inner margin of the valves extend almost four times as far within the shell. 

To compare the new species with one of almost the same size of A. multi- 
costata the following measurements are given: 
Area reinharti — diameter 27.7 mm., altitude 22.1 mm., thickness 24.5 mm. 

Lowe — Mollusc a from West Mexico 17 

A. multicostata (young) — diameter 29 mm., altitude 27.5 mm., thickness 20 mm. 
The species is named in honor of Philip W. Reinhart, of Stanford Uni- 
versity, who has done most excellent work in West Coast Paleontology, especially 
in the Arcidae. 

Phacoides (Pleurolucina) leucocymoides, new species. Plate 1, fig. 4. 

Tres Marias (1930). Type 11386, Lowe collection; paratypes, San Diego 
Society of Natural History, Carmen Island, Gulf of California, 20 fathoms 
(1932) and Lowe collection, Angel de la Guardia Island, Gulf of California, 
20 fathoms (1932). 

Shell convex, thin, white; entire surface covered with sharply reflexed con- 
centric lirae, which are much stronger and further apart than mPhacoides undatus 
Cpr. Instead of three radiating costae with four narrow interstices on each valve, 
as in Carpenter's species, there is but a single wide costa with a channeled groove 
on either side. The shell is higher and narrower than P. undatus and somewhat 
resembles P. leucocyma Dall (Proc. U. S. Nat. Muse., vol. 12, p. 263, pi. 14, 
figs. 6-7, 1889) from the Atlantic coast. The interior marginal crenations and 
cardinal teeth are more prominent than in P. undatus Cpr., while the subumbonal 
pit is not so deep as in that species. 

Diameter 10.7 mm., altitude 11.1 mm. 

Lithophaga abbotti, new species. Plate 1, fig. 5. 

Kino Bay, Sonora, tidal zone (1932). Type 11390, Lowe collection; para- 
types, San Diego Society of Natural History and Academy of Natural Sciences 
of Philadelphia. 

Shell cylindrical, thin, posteriorly obtusely rounded, anteriorly tending to 
subangulation above, evenly rounded below; growth lines are plainly visible 
under the shining light brown epidermis. The whole shell is covered with a lime 
incrustation somewhat ruffled in the central portion of both valves. The anterior 
end is less attenuated than either L. attenuata Desh. or L. aristata Hanley. 

Diameter 62.5 mm., altitude 19.5 mm. 

The type and several additional specimens were obtained in a mass of worm 
tubes, coralline growths and lime incrustations on a tidal bar a mile or more 
back in the estuary at Kino Bay, January, 1932. A single specimen was taken in 
1933 at San Felipe on the western side of the Gulf of California. 

In the U. S. National Museum is a single specimen (#381411) of the 
above species marked from San Lucas Island, Costa Rica, which measures as 
follows: length 40.4 mm., width 12.5 mm. 

This largest of our West Coast Lithophaga is named in honor of Clinton 
G. Abbott, Director of the Natural History Museum, San Diego, California. 

Solen pazensis, new species. Plate 1, fig. 6. 

La Paz, Lower California, tidal zone (1929). Type 11391, Lowe collec- 
tion; paratypes, San Diego Society of Natural History and Academy of Natural 
Sciences of Philadelphia. 

Shell transversely oblong, with anterior terminal beaks; anterior extremity 

18 San Diego Society of Natural History 

obliquely truncated; posterior extremity rather squarish; dorsal and ventral edges 
very slightly curved; hinge and ligament similar to S. sicarius Gld. Epidermis 
shining horn color, with a darker blotch on the anterior ends and a darker 
triangle formed by a line from anterior dorsal end to posterior ventral end, in- 
stead of the rosy suffusion as in S. rosaceus Cpr. Where the epidermis is removed 
near the beaks, a somewhat darker color is seen in parallel lines corresponding 
to the lines of growth. 

Diameter 57.5 mm., altitude 11.5 mm. 

Comparative dimensions are as follows: 
Solen sicarius Gld.: diameter 55.5 mm., altitude 14.5 mm. 
Solen rosaceus Cpr.: diameter 57.5 mm., altitude 13.5 mm. 
So/en mexicanus Dall: diameter 60 mm., altitude 8.5 mm. 

Psammosolen guaymasensis, new species. Plate 1, fig. 7. 

Guaymas, 20 fathoms (1932). Type 11392, Lowe collection; paratype, 
Lowe collection, off Angel de la Guardia Island, Gulf of California, 20 fathoms. 

Shell oblong-oval, rather thin, convex; extremities equally rounded; dorsal 
and ventral markings nearly parallel. Beaks not prominent, much nearer the 
anterior end. Color white; unequal striae of growth crossed by numerous diagonal 
incised lines. Pallia! sinus wide and three-fourths length of shell. 

Type: diameter 48.5 mm., altitude 20.3 mm. 

Paratype: diameter 18 mm., altitude 8.5 mm. 

Both type and paratype are right valves. 

Leda (Adrana) penascoensis, new species. Plate 1, fig. 8. 

Punta Penasco, Sonora, dredged 10 fathoms (1934). Type 11393, Lowe 

Shell white, with a straw-colored glossy periostracum; strongly compressed 
beaks much nearer the anterior end. Dorsal line nearly straight, ventral margin 
curved, anterior and posterior ends about equally angular. Dorsal edges of both 
valves slightly crenate the entire length. Sculpture of fine concentric lines of 
growth over the entire surface of both valves, except a narrow portion bordering 
the posterior dorsal margin, which is entirely smooth. 

Diameter 37.5 mm., altitude 9.4 mm. 

This shell differs considerably in sculpture and shape from the three other 
forms described in this group from West America. 

Venus kellettii, Fbs. Plate 2, fig. 1. 

Carmen Island, Gulf of California, 20 fathoms (1932) . 

Venus mariae Orb. Plate 2, fig. 2. 

Santa Maria Bay, Lower California, 20 fathoms (1931). 

Plate 2, figure 1, shows the young stage of Venus kellettii Fbs. At this 
period of its growth, it more resembles Venus mariae Orb., figured in Plate 2, 
figure 2, than the adult form, which is well illustrated in Reeve, Conch. Icon., 
vol. 14, pi. 18, fig. 82, but which shows none of the exquisite earlier sculpture. 
Had I not an adult specimen of this species, I should have unhesitatingly con- 

Lowe — Mollusca from West Mexico 19 

sidered it a new species in a group with Venus mariae Orb., which it resembles 
both in form, size, and sculpture. For this reason it seems well to figure the two 
species for comparison. 

Measurements of the Venus kellettii Fbs. figured are diameter 16.7 mm., 
altitude 1 1.3 mm.; of the Venus mariae Orb., diameter 15.4 mm., altitude 12 mm. 


Calliostoma marshalli, new species. Plate 2, fig. 3 

San Felipe, Gulf of California (1933). Type 11380, Lowe collection; para- 
types, San Diego Society of Natural History, U. S. National Museum and 
Academy of Natural Sciences of Philadelphia. 

Shell conic, elevated, rather thin; imperforate, light cinnamon brown, with, 
on the body whorl, about 14 sagittate flames of darker brown, bordered by a 
white anterior zone, running from suture to umbilicus. Five rounded whorls 
besides 2 nuclear; sutures distinct; on the penultimate whorl thirteen fine but 
sharply crenate spiral threads, on the preceding whorl seven, and on the others 
three. Base slightly convex with about 20 flat spiral threads with narrower inter- 
spaces, the fourth, eighth, and twelfth showing regular square dots of dark 
brown. Columella excurved, pearly white; umbilical callus slightly depressed. 
Aperture rounded, irridescent within; outer lip thin, with faint lirations within. 

Diameter 13.5 mm., altitude 14.1 mm. 

I take pleasure in naming this species for Mr. W .H. Marshall, who has 
given so many years of valuable service to the U. S. National Museum in the 
Department of Mollusks. 

Calliostoma gemmuloides, new species. Plate 2, fig. 4. 

Tepopa Bay, Sonora (1932). Type 11382, Lowe collection. 

The description of this beautiful species may most clearly be given by com- 
parison with the well known C. gemmulatum Cpr. It is a narrower shell; being 
a full millimeter less in diameter than a gemmulatum of equal height. The sutures 
are well defined but lack the deep channeling of gemmulatum, whorls more slop- 
ing, less angular. The beaded spiral lines are not so prominent as in gemmulatum 
and more in number. On the base are two extra spirals; above the periphery of 
body whorl are three extra spirals; on the penultimate one extra. Color dark 
reddish brown with nine radial flames of a lighter color. Six whorls exclusive 
of the two nuclear. 

Diameter 12 mm., altitude 13.5 mm. 

Calliostoma angelenum, new species. Plate 2, fig. 5. 

Angeles Bay, Lower California (1932). Type 11381. Lowe collection. 

Shell conic elevated, thin, imperforate, color reddish brown, with a few 
white dots around periphery of body whorl; whorls rather flat, six in number 
besides the two smooth nuclear; sutures distinct; base rounded; columella ex- 
curved, pearly white; umbilical callus slightly depressed. Aperture rounded, 
irridescent within; outer lip thin, with faint lirations within. There are on the 
base 16 strap-like spirals with equal interspaces, three nearest the umbilical callus 

20 San Diego Society of Natural History 

stronger; on the body whorl above the periphery are 13 regularly beaded spirals 
and on the penultimate whorl seven, antepenultimate five. 
Diameter 13 mm., altitude 14.5 mm. 

Tritonalia carmen, new species. Plate 2, fig. 6. 

Angel de la Guardia Island, Gulf of California, 20 fathoms (1932). Type 
11378, Lowe collection; paratype, San Diego Society of Natural History. 

Shell solid turreted, of four angular whorls and three rounded nuclear and 
post-nuclear whorls, suture distinct, each whorl with sloping shoulders, the lower 
four prominently angular at the periphery. Below the periphery on the body 
whorl is a lesser spiral angulosity and a smaller spiral cord below that. Outer lip 
thin, inner lip covered with a white callous, canal short, moderately wide, and 
slightly bent to the left. Faint incremental lines are visible over the entire surface. 
Color of shell a light cream with a few light brown flecks on upper portion of 
each whorl. 

Diameter 5 mm., altitude 9 mm. 

The paratype specimen was dredged off Carmen Island in 20 fathoms. 
Under catalogue number 96326, the U. S. National Museum has three examples 
of this species dredged in 9 fathoms off La Paz. They had been tentatively 
identified as young of Murex squamulatus Cpr. 

Mitrella granti, new species. Plate 2, fig. 7. 

San Felipe, Gulf of California (1933). Type 11383, Lowe collection; 
paratypes, San Diego Society of Natural History and Academy of Natural 
Sciences of Philadelphia. 

Shell smooth, solid, with seven slightly convex whorls; sutures distinct; 
axial sculpture entirely absent except inconspicuous lines of growth; the body 
whorl and the two preceding whorls are covered with regularly spaced spiral 
grooves with wider interspaces. Color dark brown, somewhat suffused with pale 
yellow. Aperture rather wide; outer lip slightly undulate; a well marked callus 
on the straight columella. 

Diameter 3.4 mm., altitude 9.4 mm. 

This interesting species has been dedicated to Dr. U. S. Grant, of the 
Department of Geology, University of California at Los Angeles. 

Anachis sanfelipensis, new species. Plate 2, fig. 8. 

San Felipe, Gulf of California, lower tidal zone (1933). Type 11384, 
Lowe collection; paratypes, San Diego Society of Natural History and Academy 
of Natural Sciences of Philadelphia. 

Shell solid, turreted, of eight rather flat whorls exclusive of the lost nucleus. 
Sutures distinct, about thirteen strong axial ribs to the whorl. Entire shell covered 
with fine microscopic spiral threads. On the back of the columella and base are 
about eleven strong spiral cords with equal interspaces; above these on the base 
are about the same number of lighter spiral threads. Body of shell of warm flesh 
color, with longitudinal blotches of light brown between the axial ribs, and, on 
the base, wavy longitudinal lines of same color. 

Lowe — Mollusca from West Mexico 21 

Diameter 6.5 mm., altitude 17 mm. 

This shell belongs in the same group as Anachis vexilhim Sby. and A. julva 
Sby. The former species, which comes from Mazatlan, is a somewhat stouter 
shell, of much darker color and fewer axial ribs and spiral cords. A. julva Sby., 
which comes from the Panamic region, is also a broader shell with fewer axial 
ribs and is of an even light brown color. 

Strombina carmencita, new species. Plate 3, fig. 1. 

Carmen Island, Gulf of California, dredged 20 fathoms (1932). Type 
11375, Lowe collection. 

The shell has ten rounded, rapidly enlarging whorls, including two smooth 
nuclear; the four early whorls almost smooth; on the fifth whorl four spiral cords 
appear just below the suture, which grow stronger on the body of the last whorl; 
on the last three whorls are fourteen axial ribs which are obsolete below the 
periphery of body whorl; entire body whorl covered with wavy spiral threads 
with about equal interspaces. Color white, slightly mottled with brown, a little 
darker on the ribs of body whorl. Aperture rather narrowly oblique, with heavily 
calloused inner and outer lips; canal short and recurved. 

Diameter 1 1 mm., altitude 29.7 mm. 

Strombina subangularis, new species. Plate 3, fig. 2. 

Carmen Island, Gulf of California, dredged 20 fathoms (1932). Type 
11374, Lowe collection. 

Shell with acuminate spire, oblong, pyramidal; pale, variegated with brown; 
eight flattish whorls exclusive of the lost nucleus, ten rather sharp axial ribs, with 
much wider interspaces, to the whorl; middle of the last whorl gibbously angled, 
reflected at base; aperture somewhat square, canal long, slightly recurved, lip 
much thickened, slightly ribbed inside. 

Diameter 11.7 mm., altitude 32.2 mm. 

This species was subsequently taken in two locations off the Mexican West 
Coast by the 1933 Crocker Expedition. 

The most nearly comparable representative in the group is S. angularis 
Rve. (Conch. Icon., vol. 11, pi. 1, figs, la, lb, 1859), which has four more ribs 
to the whorl and a much shorter canal. 

Turbonilla (Ptycheulimella) penascoensis, new species. Plate 3, fig. 3. 

Punta Penasco, Sonora, dredged 10 fathoms (1934). Type 11588, Lowe 

Shell elongate conic, of a warm flesh color, with two yellowish brown spiral 
bands, the one a little below the suture over twice as wide as the one on the 
periphery. Nuclear and all post nuclear whorls lacking all axial or spiral sculp- 
ture, except the last three, which show very faint microscopic spiral threads visible 
under a high power lens. There are seventeen rather flat whorls including the 
nucleus; sutures well appressed, base and aperture well rounded. 

Diameter 1.5 mm., altitude 10.4 mm. 

22 San Diego Society of Natural History 

Pyramidella (Triptychus) hermosa, new species. Plate 3, fig. 4. 

San Felipe, Gulf of California (1933). Type 11376, Lowe collection; 
cotype, California Academy of Sciences. 

Shell small, semiopaque, ivory white. Eight moderately rounded whorls, 
including the smooth nuclear. Rather strongly tabulated at the shoulders. 
Sculptured by three strong rounded spiral cords, of which the second and third 
are stronger than the one just below the suture. In addition to the spiral cords, 
the whorls are marked by axial ribs which are of about equal strength over the 
entire shell. Their junction with the spiral cords forms prominent tubercles, which 
are the outstanding part of the sculpture pattern. There are about thirty-two of 
these axial ribs on the body whorl. Base moderately rounded, marked with a 
single spiral cord. Outer lip a little thickened and slightly reflexed. Columella 
covered with a heavy white callus. 

Diameter 2.4 mm., altitude 6.7 mm. 

This very interesting species differs considerably in sculpture from Tripty- 
chus olssoni Bartsch from Santa Elena Bay, Ecuador (Proc. U. S. Nat. Muse., 
vol. 69, pi. 1, fig. 11, 1926), which seems to be the only other species in this 
group described from this coast except Odostomia pedroana Dall and Bartsch, 
which was provisionally placed in their new subgenus Ividella. 

Simnia quaylei, new species. Plate 3, fig. 5. 

San Felipe, Gulf of California (1933). Type 11379, Lowe collection; 
paratypes, San Diego Society of Natural History, University of California at 
Los Angeles and Academy of Natural Sciences of Philadelphia. 

Shell thin, fusiform, swollen at the middle; color a bright shrimp pink; sur- 
face polished and glossy; under a lens are seen many fine longitudinal striations; 
the low spiral cords at either end of the shell appear wavy where crossed by 
these striations. The callus on the outer lip is not very heavy; aperture rather 
wide, especially toward the base. There is no trace of an angulated callus on 
the body whorl side of the aperture, as in S. aequalis Sby. and other species. 

Diameter 7.8 mm., altitude 23.2 mm. 

I have named this finest of all West Coast species of Simnia in honor of 
Mr. E. H. Quayle, who accompanied me on my trip to San Felipe in May, 1933, 
and who has executed the very excellent drawings for this paper. 

Clavus pembertoni, new species. Plate 3, fig. 6. 

Angeles Bay, Lower California (1932). Type 11377, Lowe collection; 
paratypes, San Diego Society of Natural History and Academy of Natural 
Sciences of Philadelphia. 

Shell heavy, turreted, with eleven rounded, strongly nodulous whorls, ex- 
clusive of two smooth nuclear whorls. About thirteen nodes on the penultimate 
whorl, a heavy callosity on back of body whorl. A few strong spiral incised lines 
are below the periphery of each whorl and on the body whorl extend to the canal. 
Anal fasciole large, marked with numerous fine incremental lines. Anal sinus 
very deep; siphonal sinus short and wide. Outer lip slightly thickened and undu- 

Lowe— Mollusca from West Mexico 23 

lated. Columella pillar rather straight, covered with a strong glistening callus. 
Shell of a deep cream color with a light brown blotch on each node. 

Diameter 17.3 mm., altitude 49 mm. 

In Dr. R. E. C. Stearns' paper on the shells of the Gulf of California (Proc. 
U. S. Nat. Muse., vol. 17, p. 172, 1894), he lists a specimen (No. 55239 U. S. 
N. M.) under the name Pleurotoma unimaculata Sby. and compares it with 
P. echinata Lam. and P. gibbosa Kiener. I have examined this specimen in the 
U. S. National Museum and find it to be identical with my specimens of Clavus 
pembertoni. It is quite different from the glistening porcelain white shell of 
P. unimaculata Sby. in color, size and texture. 

I take pleasure in dedicating this species to Mr. J. R. Pemberton, owner of 
the yacht "Petrel" and sponsor of the cruise in 1932 in the Gulf of California. 

Elaeocyma acapulcana, new species. Plate 4, fig. 1. 

Acapulco, dredged 20 fathoms (1930). Type 11587, Lowe collection. 

Shell turreted, acute, smooth, of a delicate flesh color with a pinkish spot 
on each axial rib at the periphery. There are ten whorls including the smooth 
nucleus. Suture distinct, slightly undulated by the ribs of preceding whorl; spiral 
sculpture of sharp, narrow grooves, with much wider, flat, smooth interspaces; 
there are about twenty of the grooves on the body whorl anterior to the siphonal 
fasciole; the wide anal fasciole is faintly spirally striate under a high power lens; 
axial sculpture of about ten straight sharp-edged ribs, with wider interspaces, on 
the body whorl. Aperture rather wide and short, with a deep, rounded anal 
sulcus and prominent subsutural callosity; outer lip subvaricose, sharp-edged, 
smooth within; inner lip with thick layer of enamel; pillar short, straight; canal 
deep, short, wide, slightly recurved. 

Diameter 7 mm., altitude 17 mm. 

This shell differs from Elaeocyma aerope Dall (Proc. U. S. Nat. Muse., 
vol. 56, pi. 1, fig. 3, 1920), in having about twice as many spiral grooves on the 
body whorl, and in the prominently colored peripheral spots on the axial ribs. 

Clathrodrillia pilsbryi, new species. Plate 4, fig. 2. 

Punta Pefiasco, Sonora, dredged 10 fathoms (1934). Type 11587, Lowe 
collection; paratype, San Diego Society of Natural History. 

Shell pale horn-color, with sienna brown blotches between the whitish ribs. 
There are three smooth nuclear whorls, with nine succeeding whorls; axial sculp- 
ture of seven prominent ribs to the whorl, which undulate the well-defined suture. 
The ribs are obsolete on the base and the wide anal fasciole. The spiral sculpture 
consists of four or five flat strap-like cords, with wider interspaces, which pass 
over the periphery and continue over the base. Anal sulcus deep, and prominent 
anal fasciole smooth, except for strong growth striae; a strong subsutural callus. 
Outer lip moderately thickened, crenulated by spiral sculpture of the body 
whorl. Siphonal sinus short, of medium width; columella pillar straight. 

Diameter 7.3 mm., altitude 23.5 mm. 

This fine species is one of the most interestingly colored and sculptured in 
the genus, and so far as is now known seems to be confined to the upper end of 

24 San Diego Society of Natural History 

the Gulf of California. It has not turned up in any of the numerous dredgings 
south to Panama. 

It is named for Dr. H. A. Pilsbry, of the Academy of Natural Sciences of 
Philadelphia, in appreciation of his kindly assistance through many years. 

(?) Homalopoma concepcionensis, new species. Plate 4, fig. 3. 

Concepcion Bay, Lower California, 15 fathoms (1932). Type 11593, Lowe 

Shell small, pure white, solid, globose, suture strongly appressed; five whorls, 
including the smooth nucleus, strongly tabulated by a peripheral keel. On the 
penultimate whorl is a strong sutural keel and two almost equally strong just 
below it; on the flat shoulder just anterior to the major keel are three secondary 
flat spiral cords with wide interspaces. On the body whorl, just below the suture, 
are two major spiral cords with a secondary spiral thread between; next three of 
the strong cords with four spiral threads anterior to each; posterior to the last 
are thirteen flattened spiral cords of about equal strength and equal interspaces. . 
The entire surface between the spiral sculpture is covered with microscopic, 
diagonally radial striae. Aperture circular, outer lip thin; heavy callus on the 
columella, back of which is a large, flattened chink with four radial threads on 
its flat surface. 

Diameter 5.6 mm., altitude 5.6 mm. 

As there was no sign of an operculum attached to the animal, I am in doubt 
whether to place the species in Homalopoma or Liotia. In DalPs paper on the 
Florida Fossils (Trans. Wagner Free Inst. Sci., vol. 3, Aug., 1890) is a species 
which is certainly congeneric — very similar in form and sculpture, and even in 
having the same umbilical chink with four radial threads. Dall has placed the 
shell with a question in Gibbula, as G. americana Dall (Plate 22, fig. 32) . 

Hemitonia hermosa, new species. Plate 4, fig. 4. 

Carmen Island, Gulf of California, 20 fathoms (1932). Type 11385, Lowe 

Shell small, thin, oblong oval, much elevated, narrowest anteriorly; apex 
posterior, prominent and somewhat recurved; outline in front of apex slightly 
convex, from apex to the posterior margin slightly excavated, sides descending 
nearly straight; sinus moderate, situated at the extremity of a prominent, strongly 
nodulous rib. Three slightly less prominent, but nodulous ribs on either side, 
with weaker ribs between, giving the margin of the shell a crenulated appearance. 
Inside of the shell is a glossy horn color, outside chalky of a lighter shade. 

Diameter 7.3 mm., breadth 5 mm., altitude 4.7 mm. 

Fusinus fredbakeri, new species. Plate 4, fig. 5. 

San Felipe, Gulf of California (1933). Type 11590, Lowe collection; 
paratypes, San Diego Society of Natural History and California Academy of 

Shell with six well-rounded, strongly sculptured whorls exclusive of the 
nuclear whorls. Axial sculpture, on the penultimate whorl, of twelve rounded 


Plate 1 

1. Area gordita n. sp. 

2. Area delgada n. sp. 

3a, b, c. Area (Anadara) reinharti n. sp. 
4. Phacoides (Pleurolucina) 

leucocymoides n. sp. 

5. Lithophaga abbotti n. sp. 

6. Solen pazensis n. sp. 

7. Psammosolen guaymasensis n. sp. 

8. Leda (Adrana) penascoensis n. sp. 

Lowe — Mollusca from West Mexico 

Plate 2 

1. Venus kcllettii Fbs. 

2. Venus mariae Orb. 

3. Calliostoma marshalli n. sp. 

4. Calliostoma gemmuloides n. sp. 

5. Calliostoma angelenum n. sp. 

6. Tritonalia carmen n. sp. 

7. Mitrella granti n. sp. 

8. Anachis sanfelipensis n. sp. 

Lowe — Mollusca from West Mexico 

Plate 3 

1. Strombina carmencita n. sp. 

2. Strombina subangularis n. sp. 

3. Turbonilla (Ptycheulimella) 

penascoensis n. sp. 

4. Pyramidella (Triptychus) hermosa n. sp. 

5. Si mn 1. 1 quaylei n. sp. 

6. Clavus pembertoni n. sp. 

Lowe — Mollusca from West Mexico 

Plate 4 

1. Elaeocyma acapulcana n. sp. 4. Hemitonia hermosa n. sp. 

2. Clathrodrillia pilsbryi n. sp. 5. Fusinus fredbakeri n. sp. 

3. (?) Homalopoma concepcionensis n. sp. 6. Fusinus felipensis n. sp. 

7. Fusinus hertleini n. sp. 

Lowe — Mollusca from West Mexico 25 

ribs, with about equal interspaces, most prominent on the periphery. There are 
eight or nine spiral cords of unequal strength on the penultimate whorl. The 
type specimen is of a deep cream color, on other specimens shaded to a warm 
sienna brown. Canal straight, narrow and of medium length, aperture broadly 
rounded; outer lip thin, crenulated by the spiral sculpture, which shows through 
on the inside. 

Diameter 15.5 mm., altitude 38 mm., 7 whorls, 12 varices. 

In all stages of growth this shell is much broader than F. ambustus Gld., 
measurements of which are diameter 13.3 mm., altitude 38 mm., 8 whorls, 10 
varices. It also has more, although less prominent, axial ribs. It is named in 
honor of my good friend Dr. Fred Baker, of San Diego, who has done so much 
valuable work in West Coast Conchology and whose kindly assistance and advice 
to me have been of great help. 

Fusinus felipensis, new species. Plate 4, fig. 6. 

San Felipe, Gulf of California (1933). Type 11589, Lowe collection; 
paratypes, San Diego Society of Natural History and California Academy of 

Shell small, purplish brown, nearly the same size as the average Fusinus 
luteopictus Dall of the upper California coast. There are seven rounded whorls, 
including the smooth white nucleus. There are eleven axial ribs with somewhat 
wider interspaces on the penultimate and ten on the antepenultimate whorl, 
which are continuous from suture to suture; they gradually fade out on the body 
whorl. There are four or five strong, spiral cords to the whorl, with a weaker 
spiral thread between, which render the axial ribs nodulous. Aperture oval, of a 
purplish color; outer lip thin, slightly crenulated by the spiral sculpture; inside 
smooth; canal straight, of medium length and width. 

Diameter 7.7 mm., altitude 19.2 mm. 

Fusinus hertleini, new species. Plate 4, fig. 7. 

Concepcion Bay, Lower California (1932). Type 11592, Lowe collection; 
paratypes, San Diego Society of Natural History and California Academy of 

Shell elegantly and regularly fusiform, of six or seven well rounded whorls. 
On the body whorl are eleven or twelve rounded axial costae, which become 
obsolete below the periphery, crossed by three strong, spiral cords and several 
lesser spiral threads; canal straight and narrow; aperture suboval; outer lip 
slightly crenate at the margin. Color sienna brown with cream-colored costae. 

Diameter 15.1 mm., altitude 41.1 mm. 

At Sargent's Point on the Sonora coast, off the north end of Tiburon Island, 
I collected a form entirely cream-colored, except two or three post-nuclear whorls 
which show the brown blotches between the costae. This may take the name of 
variety albescens. 

At the same locality I collected another form with wide white subperipheral 
band on body whorl and a narrow dark brown band just below. This may be 
known as variety bruneocincta. 

26 San Diego Society of Natural History 

The new species has more prominent axial ribs than Fusinus ambus tus Gld., 
which has sharper spiral sculpture. It also has two more axial ribs to the whorl 
than F. ambus tus. 

The shell is named in honor of Dr. L. G. Hertlein, of the California 
Academy of sciences, who has been studying the West Mexican molluscan 
faunas for a number of years. 

Fusinus cinereus (Reeve) and varieties 

Specimens of a Fusinus collected by me at La Paz and also at Guaymas 
match the figure given by Reeve for his Turbinella cinerea 2 so closely that I do 
not hesitate to identify them as typical examples of his species. Since Reeve's 
cinerea was described under the genus Turbinella and Say's earlier Fusus cinereus 3 
under Fusus (a group generally known as Fusinus, though Say's cinereus is really 
a Urosalpinx) , it does not seem advisable to consider Reeve's specific name a 
homonym. The two species bear the same specific name, but were described under 
different genera and actually are not congeneric or even members of the same 

The present species, Fusinus cinereus Reeve (olim Turbinella id.) is prob- 
ably the species which Dall 4 once identified as F. taylorianus Reeve, 5 but in all 
my collecting I have never encountered a west coast shell which I could identify 
unquestionably as taylorianus, and I believe that Dall must have overlooked 
Reeve's cinereus because it was included in Turbinella. 

On the Coronado Island in the Gulf of California I collected a smaller, 
lighter colored form of Fusinus cinereus Reeve with white axial ribs. This may 
take the varietal name of coronadoensis. 

On the Sonora coast, north from Guaymas to Sargent's Point (opposite the 
north end of Tiburon Island) , I collected in several localities an almost black 
form, with only the first three whorls showing white on the axial ribs. This color 
form may be known as variety sonoraensis. 

2 Conch. Icon., vol. 4, Turbinella, pi. 13, fig. 68, 1847. 

3 Acad. Nat. Sci. Philadelphia, Journ., vol. 2, p. 236, 1822. 

4 Nautilus, vol. 29, no. 5, p. 55, 1915. 

5 Conch. Icon., vol. 4, Fusus, pi. 20, fig. 85, 1848, (unknown habitat). 

Lowe — Mollusca from West Mexico 27 





Herbert N. Lowe 

Sim Diego Society of Natural History 


Solemya panamensis Dall — 10 fathoms, dredged. 

valvulus Cpr. — a single example of each dredged in 10 fathoms; gray mud. 
Nucula declivis Hds. — many valves taken at 10 fathoms. 
Leda impar Pils. and Lowe — a few pairs and many valves at 10 fathoms. 
leviradius Pils. and Lowe — four pairs only at 10 fathoms. 

(Adrana) penascoensis Lowe — the type and a damaged paratype at 10 fathoms; 
Glycimeris maculata Brod. — a large colony at low tide near Punta La Cholla in 
coarse gravely sand. Many valves found in Indian kitchen middens near by. 
gigantea Rve. — a single beach valve only. 

multicostata Sby. — many young valves brought up in dredge at 10 fathoms. 
Area alternata Sby. — valves only taken in dredge at 10 fathoms. 
gradata B. and S. — valves only taken with preceding species. 
illota Sby. — living examples not rare under rocks. 
pacifica Sby. — one pair only, although plentiful in kitchen middens. 
reeveana Orb. — three pairs taken under rocks. 
reinharti Lowe — valves only at 10 fathoms. 
solida B. and S. — common living under rocks. 
Pinna rugosa Sby. — several very young pairs on beach. 
maura Sby. — several very young pairs on beach. 
Pteria peruviana Rve. — three young pairs washed in attached to sea fans. Many large 

valves in kitchen middens. 
Melina (Pedalion) chemnitziana Orb. — common under rocks. 

(Pedalion) anomioides Rve. (=janus Cpr.) — not rare under rocks. 
Ostrea chilensis Phil. — a few attached to rocks at half tide. 
palmula Cpr. — fairly plentiful with preceding species. 

dalli Lamy (=serra Dall) — a few valves brought up in dredge at 10 fathoms. 
Pecten circularis Sby. — beach valves and a few very young brought up in dredge. 
Lima pacifica Orb. — a few living pairs under rocks, extreme tide. 

orbignyi Lamy — a few beach valves only. 
Anomia peruviana Gray — two beach valves only. 
Mytilus adamsianus Dkr. — not common. 

multiformis Cpr. — abundant in rock crevices at half tide, though not in such 
profusion as at San Felipe. 
Modiolus capax Conr. — a few good pairs washed up. 

guyanensis Lam. (=braziliensis Chem.) — plentiful living in sandy mud flats. 

Gregariella denticulata Dall — a few good pairs. 

Lithophagus attenuata Desh. — boring in ledges of fossiliferous sandstone. 

aristata Dill. — with preceding species, but more abundant. 
Crenella divaricata Orb. — a few valves in dredge. 
Thracia curta Conr. — a few perfect pairs. 

squamosa Cpr. — one young pair dredged at 10 fathoms. 

28 San Diego Society of Natural History 

Pandora claviculata Cpr. — several fragments dredged. 
Lyonsia inflata Conr. — on reefs with ascidians. 

sp. ? — dredged at 10 fathoms. 
Cuspidaria didyma Hds. — a few pairs dredged. 

dulcis Pils. and Lowe — six valves only dredged. 
Crassatellites gibbosus Rve. — odd valves and a few very young examples dredged. 
Cardita aihnis var. californica Desh. — very large and abundant under rocks at half 

Chama buddiana C. B. Ads. — one pair only; common in the Indian kitchen middens. 

echinata Brod. — beach valves; abundant in kitchen middens. 
Diplodonta subquadrata Cpr. — a few valves. 
Felaniella serricata Rve. — not common. 
Divaricella eburnea Rve. — valves brought up in dredge. 
Codakia distinguenda Tryon — beach specimens not rare. 

mexicana Dall — dredged at 10 fathoms. 

chiquita Dall — odd valves plentiful with foregoing species. 
Phacoides cancellaris Phil. — odd valves in dredge. 

mazatlanicus Cpr. — odd valves in dredge. 

nuttallii var. centrifugus Dall — odd valves in dredge. 

(Cavilucina) lamprus Dall — a few beach valves. 
Cardium (Papyridea) aspersum Sby. — beach valves only. 

(Fragum) biangulatum Sby. — a few pairs brought up in dredge. 

(Laevicardium) elatum Sby. — - young shells in dredge and a few full grown 
valves on beach. 

(Laevicardium) elenensis Sby. — a few in dredge. 

(Trigonicardia) graniferum B. and S. — many valves in dredge. 

(Bingicardium) procerum B. and S. — good pairs on tide flats. 
Dosinia dunkeri Phil. — a few fresh pairs. 

ponderosa Gray — single valves abundant on beach. 
Tivela delesserti Desh. — rare living on sand flats at low tide. 
Chione fluctifraga Val. — living on mud flats. 

succincta Val. — with preceding species. 

purpurascens Dall — one beach valve only. 

mariae Orb. — odd valves plentiful in dredge. 
Macrocallista squalida Sby. — a few beach pairs. 
Pitar concinna Sby. — a few valves in dredge. 

newcombiana Gabb — valves only in dredge. 
Paphia grata Sby. — plentiful in sand between small stones near mouth of estuary. 
Cyclinella singleyi Dall — one pair and a few valves in dredge. 
Petricola denticulata Sby. — not rare in fossiliferous limestone reefs. 

robusta Sby. — seemingly a rare species; only two pairs taken with preceding 
Metis excavata Sby. ■ — one beach valve only. 
Tellidora burneti B. and S. — valves only in dredge. 
Tellina crystallina Chem. — valves only in dredge. 
Macoma panamensis Dall — valves in dredge. 

(Cymatoica) undulata Hanlcy (=occidentalis Dall) — many valves at 10 

indentata Conr. ■ — many pairs on mud flats. 
Tellina simulans C. B. Ads. — odd valves on beach. 

(Moerella) meropsis Dall — dredged at 10 fathoms. 

(Moerella) reclusa Dall — dredged at 10 fathoms. 

(Angulus) amianta Dall — dredged at 10 fathoms. 

Lowe — Mollusca from West Mexico 29 

Semele flavescens Gld. — three beach specimens. 

guaymasensis Pils. and Lowe — a few pairs in dredge. 

pacifica Dall — odd valves only in dredge. 

sp. ? — one valve only in dredgings. 
Donax gracilis Hanley — living on sand flats. 

navicula Hanley — living on sand flats. 
Heterodonax bimaculatus Orb. — beach valves. 
Tagelus affinis C. B. Ads. — plentiful on mud flats. 

Psammosolen guaymasensis Lowe — two valves dredged at 10 fathoms in mud. 
Solen rosaceus Cpr. — two pairs only on sand flats. 
Mactra dolabriformis Conr. — a single beach valve. 

californica Conr. — valves in dredge. 
Sphenia fragilis Cpr. — two pairs only. 
Corbula marmorata Hds. — a few in dredge. 

nasuta Sby. — plentiful in dredgings. _ 

bicarinata Sby. — a single pair under a rock. 

sp. ? — odd valves in dredge. 
Solecardia eburnea Conr. — one valve in dredge. 
Crassinella varians Cpr. — valves plentiful in dredge. 


Dentalium inversum Desh. — dredged at 10 fathoms. 

fisheri Stearns — dredged at 10 fathoms. 

splendidum Sby. — dredged at 10 fathoms. 

numerosum Dall — dredged at 10 fathoms. 
Cadulus panamensis Sby. — dredged at 10 fathoms. 
Retusa paziana Dall dredged at 10 fathoms. 

gonzagensis Baker and Hanna — dredged at 10 fathoms. 
Volvulella californica Dall — dredged at 10 fathoms. 
Acteocina infrequens C. B. Ads. — dredged at 10 fathoms. 
Bulla gouldiana Pils. — several taken living in sand pockets in reefs. 
Haminea virescens Sby. — one specimen. 
Terebra bridgesi Dall — a few in dredge at 10 fathoms. 

larvaeformis Hds. — dredged at 10 fathoms. 

sp. ? — dredged at 10 fathoms. 

sp. ? — dredged at 10 fathoms. 

sp. ? — dredged at 10 fathoms. 

sp. ? — dredged at 10 fathoms. 
Turritella goniostoma Val. — dredged at 10 fathoms. 

tigrina Kiener — dredged at 10 fathoms. 
Conus interruptus Brod. — a fine colony of extra large specimens taken in gravely 
sand with Glycimeris metadata. 

puncticulatus Hws. — a few in dredge. 

regularis Sby. — a few live ones on mud flats. 
Turris olivacea Sby. — a number taken living on reef. 

tuberculifera Brod. and Sby. — two beach specimens only taken of this very 
rare form. 
Crassispira bottae Val. — two living specimens taken on reef, in sand pockets; an 
exceedingly rare species. 

nymphia Pils. and Lowe — four taken on reef. 

nigerrima Sby. — a few in the 10 fathom dredgings. 

pluto Pils. and Lowe — abundant living on moss-covered rocks of reef. 

30 San Diego Society of Natural History 

Clathrodrillia halis Dall — dredged at 10 fathoms; not rare. 

alcestis Dall — dredged at 10 fathoms. 

thestia Dall — dredged at 10 fathoms. 

callianira Dall — dredged at 10 fathoms. 

rosea Sby. — one fine specimen in dredgings. 

pilsbryi Lowe — a few in dredgings. 
Elaeocyma unimaculata Sby. — dredged at 10 fathoms. 

aeolia Dall — dredged at 10 fathoms. 

ianthe Dall — dredged at 10 fathoms. 

palmeri Dall — dredged at 10 fathoms. 

sp. ? — dredged at 10 fathoms. 
Glyphostoma adria Dall — a few choice specimens. 
Cytharella phaethusa Dall — a single shell dredged. 
Mangelia arteaga roperi Dall — a few dredged at 10 fathoms. 

antipyrgus Pils. and Lowe — a few dredged at 10 fathoms. 

cymatias Pils. and Lowe — a few dredged at 10 fathoms. 
Cancellaria cassidiformis Sby. — a few beach specimens. 

obesa Sby. — two beach specimens. 

funiculata Hds. — one dredged living at 10 fathoms. 
Oliva incrassata Sol. (=angulata Lam.) — fine large ones living with Conns inferrup- 
fits at low tide. 

polpasta Duclos — dredged; this species seems to live only in deep water. 
Olivella dama Gray — abundant in sand pockets in reefs. 

zonata Duclos — very rare on beach; living. 

gracilis B. and S. — taken in dredge. 
Agaronia testacea Lam. — many fine specimens taken living on sand beach at half tide. 
Marginella californica Tomlin — not rare, under stones. 
Mitra attenuata Rve. — a few fine specimens dredged. 

dolorosa Dall — a single example taken on reef. 
Latirus lugubris C. B. Ads. — three specimens from reef. 
Galeodes patula Brod. — beach specimens. 
Hanetia pallida Brod. and Sby. — abundant on reef. 
Fusinus dupetithouarsi Petit — a number of young specimens in dredge. 

felipensis Lowe — several live specimens under rocks. 

Nassa iodes Dall — many living in sand flats. 

leucops Pils. and Lowe — abundant in sandy mud. 

tiarula Kiener — a few taken on sand flats. 

pagoda Rve. — dredged at 10 fathoms. 

versicolor C. B. Ads. — taken alive in sand pockets in reef. 

versicolor striatula C. B. Ads. — with preceding species. 

angulicostis Pils. and Lowe — dredged at 10 fathoms. 

Anachis coronata Sby. — living specimens under rocks. 

hilli Pils. and Lowe — four living specimens under rocks. 

vexillum Rve. — four living specimens under rocks. 

varia Sby. — four living specimens under rocks. 
Columbella fuscata Sby. — ■ common under rocks. 

major Sby. — not common. 
Mitrella diminuta C. B. Ads. — a few of this tiny species. 

ocellata var. guttata Sby. — common under rocks. 
Strombina dorsata Sby. — dredged at 10 fathoms. 

gibberula Sby. — ■ dredged at 10 fathoms. 

maculosa Sby. — dredged at 10 fathoms. 
Parametaria dupontii Kiener — a few living under rocks. 

Lowe — Mollusca from West Mexico 3 1 

Cosmioconcha palmeri Dall — two specimens in dredge. 
Phos veraguensis Hds. — two young in dredge. 

mexicanus Dall — a number of fine specimens in dredge. 
Murex elenensis Dall (=plicatus Sby.) — beach shells only. 

Phyllonotus bicolor Val. — many fine specimens feeding on bivalves on sand beach at 
very low tide. 

nigritus Meusch. ■ — abundant on reefs feeding on Ceritbium stercus-muscarum. 
Acanthina angelica Oldroyd — very abundant living on exposed wave-beaten rocks. 

muricata Brod. — very good examples taken on reefs. 
Thais triserialis Blv. — a few taken on reef. 
Muricopsis erynaceoides Val. — a few taken in dredge. 
Eupleura muriciformis Brod. — some good specimens taken with dredge. 

triquetra Rve. — not rare on reefs feeding on Ceritbium; a few were yellow 
and some almost white. 
Epitonium crenimarginata Dall — three beach specimens. 

crenatoides Cpr. — one dredged. 

(Asperoscala) canna Dall — two dredged. 

bialatum Dall — two dredged. 

sp. ? - — two dredged. 
Melanella mexicana Dall — dredged at 10 fathoms. 

rutila Cpr. — one dredged at 10 fathoms. 
Strombiformis lapazana Bartsch — four dredged at 10 fathoms. 

townsendi Bartsch — ■ one dredged at 10 fathoms. 
Niso excolpa Bartsch — a few fine examples dredged. 
Turbonilla ceralva Dall and Bartsch — dredged at 10 fathoms; 3 specimens. 

may ana Baker, Hanna and Strong — dredged at 10 fathoms. 

calvini Dall and Bartsch — dredged at 10 fathoms. 

sanctorum Dall and Bartsch — dredged at 10 fathoms. 

pazana Dall and Bartsch — dredged at 10 fathoms. 

penascoensis Lowe — dredged at 10 fathoms. 

azteca Baker, Hanna and Strong — several dredged at 10 fathoms. 

subangulata Cpr. — three specimens dredged. 

macbridei Dall and Bartsch — dredged at 10 fathoms. 
Pyramidella mazatlanica Dall and Bartsch — a few dredged at 10 fathoms. 

bicolor Dall and Bartsch — a few dredged at 10 fathoms. 
Odostomia telescopium Cpr. — dredged at 10 fathoms. 

convexa Cpr. — six specimens dredged. 

gabrielensis Baker, Hanna and Strong — two specimens dredged. 

effusa Cpr. — several dredged. 
Cypraea annettae Dall (=sowerbyi Kiener) — some fine living specimens under rocks. 
Trivia solandri Gray — many specimens taken feeding on upper side of moss-covered 

californica Gray — a few taken with preceding species. 
Cymatium adairensis Dall — a few taken alive in crevices of rocks; a rare form. 
Nearly topotypes, as Adair Bay is only a few miles north of Punta Penasco. 
Cerithiopsis sp. ? — ■ three specimens in dredge.. 
Alabina diomedeae Bartsch — common in dredgings. 
Seila assimillata C. B. Ads. — several taken living on under side of old valves of 

Dosinia pondcrosa on reef. 
Cerithium maculosum Kiener — living in sand pockets in reefs. 

incisum Sby. — common living under rocks. 

stercus-muscarum Val. — thousands living on reefs at half tide. 
Cerithidea mazatlanica Cpr. — abundant on mud flats. 

32 San Diego Society of Natural History 

Caecum firmatum Cpr. — common in dredgings. 

liratocinctum Cpr. — common in dredgings. 
Vermetus pellucidus Brod. — a few under rocks. 

tripsycha Pils. and Lowe — one beach specimen. 
Rissoina barthelowi Bartsch — four crab specimens dredged at 10 fathoms. 

mexicana Bartsch — four specimens dredged at 10 fathoms. 
Hipponyx barbatus Sby. — extra fine specimens with lower plate developed into a deep 
concave valve; taken on outer reefs. 
serratus Cpr. — under rocks at low tide. 
Calyptraea mamillaris Brod. — dredged at 10 fathoms. 

conica Brod. — dredged at 10 fathoms. 
Crucibulum spinosum Sby. — a few in dredge. 
Crepidula arenosa Brod. — half grown specimens in dredge. 
onyx Sby. — one beach specimen. 
nivea Gld. — one beach specimen. 
Natica marochiensis Gmel. — two living specimens on mud flats. 

Polinices bifasciatus Gray — extra large specimens taken in sandy gravel with Gly- 
cimeris maculata. 
uber Val. — living specimens taken on sand flats. 
recluzianus" Petit — young specimens in dredge. 
Lamellaria diegensis Dall — a number taken alive with ascidians on beach after storm. 
Acmaea mesoleuca Menke — abundant on rocks at half tide. 

mitella Menke — not common; almost at high tide; a very tiny species. 
Turbo fluctuosus Wood — plentiful under rocks. 
Leptothyra concepcionensis Lowe — one specimen dredged. 
Liotia carinata Cpr. — several in dredgings at 10 fathoms. 
Tegula globulus Cpr. — abundant under stones in same zone as A. mesoleuca. 
rugosa A. Ads. — extra large specimens taken on upper side of rocks. 
mariana Dall — a few good living specimens taken under rocks at low tide. 
Calliostoma palmeri Dall — a few in dredgings. 

marshalli Lowe — a single example dredged. 
Circulus annulatus Cpr. — dredged at 10 fathoms. 

tricarinatus C. B. Ads. — dredged at 10 fathoms. 
Neritina picta Sby. — plentiful on rocks at mouth of estuary. 
Nerita scabricosta Lam. — on rocks near high tide. 
bernhardi Reel. — on rocks near high tide. 
Strombus galeatus Sby. — half grown specimens on mud flats. 
Diadora alta C. B. Ads. — four specimens taken under rocks at low tide. 

inaequalis Sby. — not rare under rocks. 
Ficus decussatus Wood — three fair beach specimens. 
Cassis abbreviatus Lam. — several beach specimens. 
Heliacus radiatus Mke. — two crab specimens under rocks. 
Aplysia sp. ? — the animal looks much like our californicus. 
Chiton virgulatus Sby. — abundant under rocks. 
Ischnochiton acrior Cpr. — plentiful under rocks. 
clathratus Rve. — plentiful under rocks. 
(Stenoplax) limaciformis Sby. — five specimens on reef. 
Callistochiton infortunatus Pils. — not common under rocks, 
sp. ? — a beautiful color series taken on reef. 

S p_ } — a f ew taken on reef; both this and preceding species were taken in 193 3 
at San Felipe. 
Acanthochites diegensis Pils. — three specimens on reef. 

Dendrochiton sp. ? — three specimens on reef; similar to D. thamnophora Berry. 
Nuttallina sp. ? — a very small species; taken on outer rocks. 






Vol. VIII, No. 7, pp. 35-46, plate 5 March 21, 1935 



Fred Baker and V. D. P. Spicer 

San Diego Society of Natural History 

This report covers seven species of the genus Triphora which seem 
to be new. Five species were collected in 1927 by Mr. Wray Harris, of the 
U. S. Navy, on the outer edge of the coral reefs surrounding the Island 
of Ofu of the American Samoan Group at low tide levels and where 
they were subject to the full force of the wave movement. The other two 
species were collected in 1896 by the late Capt. Geo. D. Porter for the 
late Miss Jeannette M. Cooke of San Diego, who maintained a small 
collecting vessel on the coast of Lower California and in the Gulf of 
California under command of Capt. Porter for many years. These shells 
were taken during Capt. Porter's final disastrous trip, when he and his 
companion were murdered on Tiburon Island by the Seri Indians. The 
specimens were labeled as from the Gulf of California, but, as Capt. 
Porter made most of his collections of minute shells from Espiritu Santo 
Island near the southern end of the Gulf, it is extremely probable that 
they came from that locality. 

Our sincere thanks are due to Mr. Wray Harris for kindly allowing 
us to describe and figure the Samoan species, and to Mr. E. M. Thorp 
of the Scripps Institution of Oceanography at La Jolla for making the 
microphotographs illustrating the paper. 

TRIPHORA (Deshayes) Blainville, 1828 

In the discussion of this genus, Tryon (Man. Conch., IX, 121 ) 
says 'An anomaly of the shell is the occasional preservation of a second 

36 San Diego Society of Natural History 

canal upon the back of the body whorl, showing the termination of a for- 
mer aperture. This is present in the fossil species upon which Deshayes 
established his genus, 1 and which is named from this character. * * * 
Having examined several species with three apertures I incline to the 
opinion that the posterior canal is only accidentally preserved in some 
cases by reason of its deflection, which causes its tube to be surrounded 
with shelly matter during the growth of the shell, and that it bears no 
relation to the economy of the animal. This view is fortified by the con- 
sideration that neither in this group nor in any other group of the 
mollusca are we acquainted with any organ which might occupy or occa- 
sion this tube for the purpose of its economy. Moreover, in one of the 
species before me two individuals possess the third opening, whilst two 
others have it not." 

In a large suite of Triphora violacea Quoy taken on Ofu by Mr. 
Harris, most of the specimens show the third opening or meatus noted 
above. In a few specimens it is missing and all of these are immature. The 
series shows the posterior canal in the usual position within the aperture, 
open as is common in most species of univalves; partially enclosed and, 
finally, as a completed tube. It is evident that when the shell is within 
half a turn of maturity, the animal proceeds to change the open canal to 
the tubular form which, with the advancement of the outer lip through 
additions, is left further and further from the outer lip until it reaches 
a position opposite the aperture, where it remains permanently. Such 
specimens show no sign of the open posterior canal within the aperture. 
The series shows every stage of this process up to the point where, having 
taken its permanent place, the animal proceeds to build the outer lip and 
aperture into final shape. Some species show this process to have taken 
place in nearly all, or all, mature specimens, while in other species of 
which we have large suites, the process does not take place at all. No 
specimen of T. abbotti or of T. granti of this paper shows any sign of the 
tubulation of the posterior canal. 

While the question cannot be settled finally until careful examina- 
tions of living animals can be made, the writers differ radically from 
Tryon's dictum and believe that the third opening, when present, has a 
very decided "relation to the economy of the animal." We believe it will 
be shown that, as the posterior canal is thus left behind, the excretory 

1 A reference to a description in 1825 under a vernacular name without validity under 
the Code. 

Baker & Spicer — New Triphora 37 

siphon also remains stationary within the canal while a half turn is added 
to the shell, so that the third opening does service for the rest of the life 
of the animal as an excretory meatus. We wonder that Tryon forgot 
that a similar process takes place in Haliotis, in which the notch on the 
edge of the shell is finally sealed as a round hole. This hole and the en- 
closed excretory siphon remain stationary as additions are made to the 
edge of the shell and new holes are added. Finally, from being the newest 
hole it is left to the oldest and, later, is closed by the deposition of new 
shell material and the corresponding excretory siphon atrophies. 

As to the same species showing specimens with and without the third 
opening as noted by Tryon, it seems most probable that two specimens 
were immature, not having begun the formation of the body whorl, in 
which case the posterior canal would be open and within the aperture. 
The other two specimens were evidently mature, showing the posterior 
canal in its normal position opposite the aperture, in which case there 
would be no sign of a posterior canal within the aperture. 

Triphora harrisi Baker and Spicer, new species 

Plate 5, figs. 1-2 

Shell sinistral, of medium size, spindle-shaped; nuclear whorls four, straw- 
yellow, the first nearly smooth, rounded and shining, rhe others bearing a single 
prominent, spiral subcarinate cord, slightly concave below the carina, the carina 
doubled on the final nuclear whorl, crossed by numerous, prominent vertical 
ribs forming deep pits at the sides of the carinae and in the space formed by the 
doubling of the carina of the fourth whorl; postnuclear whorls eleven, white, 
polished and shining; transition of color and sculpturing abrupt to two low 
spiral cords, each bearing a row of nearly round tubercles, the anterior cord more 
prominent up to the last turn, where the posterior cord becomes more prominent; 
sutural channel narrow, slightly sinuous; median channel smooth except for in- 
cremental lines, not crossed by vertical riblets, rendered sinuous by the place- 
ment of the upper tubercles opposite the lower interspaces; tubercles appearing 
as two parallel rows of shining beads immersed in the shell structure rather than 
as if formed by the intersections of axial ribs and spiral cords; median channels of 
the lower whorls slightly tinted with brown; a weak, nodular peripheral keel 
margining the aperture; base irregularly convex, faintly brown, bearing two 
tuberculate cords, the posterior about equalling the peripheral cord, the anterior 
weaker and smoother, continuing for about half a turn and ending abruptly 
behind the outer lip; aperture round, tubular, extending radially beyond the 
periphery of the shell, smooth and pearly within, wrinkled and lined externally; 
anterior canal short, tubular, nearly straight, projecting in the axis of the shell; a 
third meatus opposite the aperture projecting as a round tube about 0.25 mm. 
beyond the periphery of the shell; no posterior canal showing within the aperture. 

38 San Diego Society of Natural History 

Length, 5.5 mm.; diameter, 1.75 mm. 

Holotoype: No. 23761, collection of the San Diego Society of Natural 
History, and ten paratypes; collected by Mr. Wray Harris on the coral reefs 
of Ofu, Samoa. Paratypes in the U. S. National Museum and in the collections 
of Mr. Harris, for whom the species is named, and of the authors. The 
species seems to be new and distinct from any other species of the region except 
the succeeding one. 

Triphora ofuensis Baker and Spicer, new species 
Plate 5, fig. 3 

Shell sinistral, minute, spindle-shaped; nuclear whorls five, light brown, 
postnuclear whorls seven, white; first nuclear whorl papilliform, nearly smooth 
above, exserted; all nuclear whorls bearing a sharp, well-developed carina, nearer 
the anterior than the posterior suture, crossed by numerous, very fine, sharply- 
defined, retractive vertical ribs, 20 appearing on the lower part of the first, 24 
entirely crossing the second, 26 on the third, fourth and fifth; postnuclear whorls 
increasing rapidly in diameter, the last decreasing slightly through the diminish- 
ing size of the last anterior spiral cord; transition from the nuclear portion very 
abrupt, the color becoming a clear, shining white, and the sculpture changing to 
two well marked spiral cords, the posterior smaller on the first two whorls, larger 
on the last and nearly equal on all the others; spiral cords continuing to the 
border of the aperture and bearing throughout prominent, roundish tubercles; 
axial ribs distinctly, but irregularly, protractive; sutural channels not sinuous, 
very sharply incised, deep and much narrower than the median channel, truncat- 
ing the tubercles of the spiral cords; median channels rendered sinuous by the 
placement of the upper tubercles opposite the interspaces of the lower cord; 
peripheral keel double, arising at the juncture of the suture with the margin of 
the aperture, the two parts slowly diverging, nodose, the posterior portion being 
slightly stronger; basal keels two, nodose, beginning beneath the rather heavy 
parietal callus; base convex, stained with yellow; aperture circular, outer lip 
thin; a third meatus or posterior canal opposite the aperture on the last whorl; 
no sign of a posterior canal at the margin of the aperture; anterior canal long, 
completely enclosed, opening nearly round, the extremity squarely truncate. 

Length, 3 mm.; diameter, 1 mm. 

Holotoype: No. 23762, collection of the San Diego Society of Natural 
History, and sixteen paratypes; collected by Mr. Harris on Ofu, Samoa. Para- 
types in the U. S. National Museum and in the collections of Mr. Harris and 
the authors. 

The species is similar to T. harrisi Baker and Spicer of this paper in color 
and general appearance, but the number of nuclear whorls and their sculpturing 
differ widely, the postnuclear whorls are fewer on the mature shell, and the 
tubercles of the spiral cords are sharply truncated by the sutures, a characteristic 
not present in T. harrisi. All the specimens taken show the third meatus charac- 
teristic of the genus, but it is far less prominent than in T. harrisi. 

Baker & Spicer — New Triphora 39 

Triphora abbotti Baker and Spicer, new species 
Plate 5, fig. 4 

Shell sinistral, elongate-conic, rather stout, large for the genus, dove-gray; 
early nuclear whorls decollated, the one remaining bearing two prominent, shin- 
ing, obsoletely tuberculate spiral cords separated by a deep, pitted channel nearly 
equalling the cords in width; postnuclear whorls eleven; transition to postnuclear 
sculpture rather abrupt, the nearly equal spiral cords quickly losing their indis- 
tinct tuberculation and becoming polished and shining, the intercostal channel 
showing numerous very fine, sinuous, incised spiral lines, deepest in the middle 
and rising nearly to the edges of the spiral cords; sutures channeled and showing 
the same sculpture as the intercostal channel, the one above the initial postnu- 
clear whorl nearly equalling the intercostal channel in width; intercostal channel 
widening much more rapidly on the later whorls; a fine spiral cord beginning on 
the sixth whorl almost midway between the other two and gradually increasing 
until nearly, but not quite equalling the other two on the penultimate turn; 
median cord dividing the median channel into nearly equal parts, each part 
about equal to the suture in width; last whorl and base showing a fine, transverse 
wrinkling, becoming close-set, rugose, and nearly equalling the spiral sculpture on 
the last quarter turn; aperture broadly pear-shaped, showing the external sculp- 
ture within; color the same as the exterior but darker beneath the spiral cords; 
posterior canal a deep, narrow notch; outer lip effuse and abruptly expanding at 
the margin, wrinkled, thin; parietal wall with a thin callus, free at the outer edge 
and continuous with the outer lip above; anterior canal long, reflexed, tubular 
for one-half its length; periphery subangulated by an obscurely tuberculate peri- 
pheral keel nearly as large as the anterior cord and slightly nearer to it than the 
middle cord; base moderately convex, marked by two smaller keels beginning 
under the parietal callus, the first obsoletely tuberculate and placed wholly on 
the base, the anterior smaller, smooth, and circling the tubular anterior canal; 
columella not well marked, showing a distinct callus. 

Length, 8.5 mm.; diameter, 2.5 mm. 

Holotype: No. 23763, collection of the San Diego Society of Natural 
History, and fifty paratypes; collected by Mr. Harris on Ofu, Samoa. Para- 
types are in the U. S. National Museum and in the collections of Mr. Harris and 
the authors. 

The shell is like a sinistral Seila. It somewhat resembles T. incisa Pease, as 
far as can be judged from the inadequate description and poor figure of Tryon, 
(Man. Conch., IX, 190) and the fuller description of Hedley (Shells of 
Funafuti), but the coloration is quite distinct, the anterior and posterior post- 
nuclear cords are nearly equal in size and prominence and nowhere show a ten- 
dency "to divide into beads." Hedley's specimen evidently represented a smaller 
species. Of the large number of specimens taken, most of which seem mature, 
all lack the third meatus and show the posterior canal in its ordinary position 
within the aperture. 

The species is named for Mr. Clinton G. Abbott, Director of the San 
Diego Society of Natural History. 

40 San Diego Society of Natural History 

Triphora granti Baker and Spicer, new species 
Plate 5, fig. 5 

Shell sinistral, rather large for the genus, elongate-conic, white, profusely 
flamed with chestnut except on the spiral cords and tubercles; nuclear whorls 
decollated; postnuclear whorls fourteen, with a superior, sinuous, non-tuberculate 
spiral cord near the suture, a wide median channel filled with fine, sharply in- 
cised spiral striae and numerous minute, slightly protractive incremental lines, 
about equal to the spiral striae, not crossing the spiral cords, followed by a 
similar, weaker median cord and a second median channel of the same width, 
and sculptured the same as the preceding one, followed by a very prominent, 
strongly tuberculate inferior cord; tubercles large, rather distant, spirally elon- 
gate, four or five times as long as broad, polished white on the summits, twelve 
to fourteen appearing on each whorl; a narrow, sinuous spiral cord between the 
row of tubercles and the suture, white, articulated with brown, producing a false 
appearance of tuberculation, and sculptured with fine spiral striae; sculpture 
and coloration of all the whorls very similar, but becoming less distinct and less 
distinctly marked on the upper whorls; periphery marked by a narrow channel 
separating the last spiral cord from a nearly equal, smooth basal keel, followed 
by a second basal keel bearing small, close-set tubercles; balance of base smooth 
except for fine, incised spiral striae and incremental lines; suture narrow, sinuous, 
sharply incised, but not as deep as the peripheral channel; aperture roughly 
quadrilateral; anterior canal short, moderately retracted, closed for about one- 
fourth its length; posterior canal a shallow groove within the aperture; parietal 
callus heavy, with a lobe descending and partially occluding the anterior canal; 
outer lip thin, translucent, smooth inside, showing the external colors and 
sculpture within. 

Length, 10 mm.; diameter, 3 mm. 

Holotype: No. 23764, collection of the San Diego Society of Natural 
History, and forty-three paratypes; collected by Mr. Harris on Ofu, Samoa. 
Paratypes in the U. S. National Museum and in the collections of Mr. Harris 
and the authors. 

The shell resembles T. crenulata Deshayes, but differs in the extreme size 
of the tubercles of the anterior cord, while the secondary cords are of much 
smaller size. 

The species is named for Dr. U. S. Grant, IV, professor of Paleontology 
in the University of California at Los Angeles. 

Triphora peleae Baker and Spicer, new species 

Plate 5, fig. 6 

Shell sinistral, dull white, minute, spindle-shaped, the upper part of the 
spire slightly concave; nuclear whorls two, the first the larger, rounded and 
exserted, giving a club-shaped appearance to the nucleus; smaller nuclear whorl 
spirally carinated, smooth; postnuclear whorls nine, bearing two equal tubercu- 
late spiral cords, about fourteen tubercles appearing on each whorl; the tubercles 

Baker & Spicer — New Triphora 41 

slightly elongated spirally and increasing in size on each succeeding whorl; median 
channel sinuous, wider and shallower than the suture; suture fine, not well- 
defined, lying at the bottom of a channel separating the two rows of tubercles; 
base bearing four equal, wavy keels starting from the parietal callus and diverg- 
ing gradually, the lower two extending on the tube of the anterior canal; parietal 
callus prominent, semilunar, extending onto the anterior canal; aperture nearly 
circular, interior white, smooth, outer lip projecting; anterior canal tubular, 
tapering, projecting from the base at an angle of about thirty degrees from the 
line of the axis of the shell; third meatus round, enclosed, immediately adjacent 
to the posterior angle of the aperture. 

Length, 5.5 mm.; diameter, 1.75 mm. 

Holotype: No. 23765, collection of the San Diego Society of Natural 
History, and four paratypes; collected by Mr. Harris on Ofu, Samoa. Paratypes 
in the U. S. National Museum and in the collections of Mr. Harris and the 

The species seems to be new and distinct from all the Triphoras of the 
region. It is named for Miss Pele Spicer who was born in Samoa. 

Triphora cookeana Baker and Spicer, new species 

Plate 5, fig. 7 

Shell sinistral, minute, elongate-conic; nuclear whorls four, smooth, shining, 
dingy-white, the first papilliform, the others moderately convex, increasing in 
size very gradually, with well-defined sutures marked vertically by regular, fine, 
incised lines, about thirty appearing between the third and fourth whorl; postnu- 
clear whorls eight and a half, very slightly convex, ashen-brown, increasing in 
diameter rather more rapidly than the nuclear whorls to about the sixth, the 
remaining whorls being about equal; postnuclear sculpture consisting of nearly 
vertical ribs, about 12 appearing on the first turn; 14 on the second, 16 on the 
third, 20 on the fourth and fifth and 22 on the remaining turns, crossed by three 
spiral cords, the middle slightly nearer the posterior than the anterior and 
slightly stronger on the first five turns, all becoming nearly equal on the remain- 
ing whorls; intersections of the ribs and cords marked by nearly round tubercles, 
sloping a bit more abruptly posteriorly than anteriorly; sutures well-defined but 
not channeled; periphery marked by a rather strong keel commencing at the 
upper curve of the outer lip, definitely tuberculate for about a third of a turn, 
then becoming narrow, sharp and distinctly wavy, the waves correlated with the 
tubercles of the vertical cords, this keel continuing in all the sutures and defining 
the sutures anteriorly; base showing no continuation of the vertical ribs but 
marked by a single wavy, spiral keel separating from the peripheral keel above 
the aperture and diverging widely to a termination on the columella; aperture 
rounded, anterior canal open, straight, very short; posterior canal a broad notch 
within the aperture, not well-defined; outer lip fractured, thin, showing the ex- 
ternal sculpture within; columella darker than any other portion of the shell, 
short, stout, obliquely truncated anteriorly. 
Length, 3.5 mm.; diameter, 1 mm. 

42 San Diego Society of Natural History 

Holotype: No. 23766, collection of the San Diego Society of Natural 
History, a unique specimen; collected by Capt. Geo. D. Porter in the Gulf of 
California, Mexico. 

The shell is very distinct from all other species from this Coast, but falls 
into the very small class of T. callipyrga Bartsch in that the three spiral cords 
are continuous on all the postnuclear whorls. The species is named for Miss 
Jeannette M. Cooke. 

Triphora stephensi Baker and Spicer, new species 

Plate 5, figs. 8-9 

Shell sinistral, of medium size, stout, spindle-shaped; nuclear whorls decol- 
lated; postnuclear whorls seven and a half, scarcely convex, tuberculate at the 
intersections of the vertical ribs and spiral cords; the first and second bearing 
two nearly equal tuberculate spiral cords, quite close together but separating on 
the third turn; a faint median cord beginning on the fourth turn, increasing 
very gradually and equalling the other two on the last whorl; first whorl having 
about 14 axial ribs, the second 16, the third 17, the fourth 18 and the last three 
about 20; peripheral keel about two-thirds the width of the others, sinuous and 
scarcely tuberculate; sutures moderately channeled and showing an extension 
of the peripheral cord about six turns; axial ribs slightly, but irregularly protec- 
tive, the upper ribs on each whorl ending opposite the interspaces of the succeed- 
ing whorl, producing a sinuous suture; base rather evenly rounded, carrying two 
fairly developed, sinuous keels, beginning close together beneath the parietal 
callus, but spreading until about equally separated from each other and from 
the peripheral keel; spiral cords of the first four whorls pale ashen-brown, the 
posterior slightly lighter, the intervening channels and sutures darker; median 
cord of the same color as the sutures, becoming pale ashen-brown on the last 
turn; axial ribs weaker and more widely spaced than the spiral cords, producing 
a slight spiral elongation of the enclosed pits; parietal callus well-developed, 
extending on the columella; aperture irregularly subquadrangular; anterior canal 
short, open, very moderately twisted; posterior canal a broad notch within the 
aperture; columella rather stout, obliquely truncate below. 

Length, 4 mm.; diameter, 1.75 mm. 

Holotype: No. 23767, collection of the San Diego Society of Natural 
History, and a less mature paratype, also in the same collection; collected by 
Capt. Porter in the Gulf of California, Mexico. Three worn specimens in the 
collections of the authors are probably of this species. 

The paratype retains all but a fraction of the nucleus consisting of nearly 
three nuclear whorls which can be described as follows: Nuclear whorls nearly 
three, changing very gradually to postnuclear sculpture, the first slightly frac- 
tured above, consisting of a very convex, light yellow spiral cord, separated from 
a similar cord on the second nuclear whorl by a broad, dark brown, flat-bottomed 
suture, cord and suture being everywhere marked by very minute tubercles; 
second nuclear whorl similar to the first in color but beginning to show the start 
of postnuclear sculpture; remaining nuclear whorl similar in color but with 

Baker & Spicer — New Triphora 43 

rather distinct axial ribs, while the nuclear spiral cord becomes tubercular and 
gradually merges into the anterior spiral cord of the first postnuclear whorl. 

The species somewhat resembles T. oweni Fred Baker, but the color is dif- 
ferent, the shell is stouter, with smaller tubercles, and the broken lines of the 
axial riblets, with earlier incidence of the median spiral cord, constitute a very 
marked distinction. 

The species is named for Mr. Frank Stephens, Curator Emeritus, San 
Diego Society of Natural History. 

44 San Diego Society of Natural History 


Fig. 1. Triphora harrisi Baker and Spicer, sp. nov. Holotype. Alt. 5.5 mm. 

Fig. 2. Same, lateral view, showing aperture and both canals. 

Fig. 3. Triphora ofuensis Baker and Spicer, sp. nov. Holotype. Alt. 3 mm. 

Fig. 4. Triphora abbotti Baker and Spicer, sp. nov. Holotype. Alt. 8.5 mm. 

Fig. 5. Triphora granti Baker and Spicer, sp. nov. Holotype. Alt. 10 mm. 

Fig. 6. Triphora peleae Baker and Spicer, sp. nov. Holotype. Alt. 5.5 mm. 

Fig. 7. Triphora cookeana Baker and Spicer, sp. nov. Holotype. Alt 3.5 mm. 

Fig. 8. Triphora stephensi Baker and Spicer, sp. nov. Holotype. Alt. 4 mm. 

Fig. 9. Triphora stephensi Baker and Spicer, sp. nov. Paratype, immature 
specimen showing nucleus. 

Baker 8c Spicer — New Triphora 

Plate 5 



Volume VIII, No. 8, pp. 47-53, plate 6 







Harry E. Wheeler 

Stanford University, California 

Printed for the Society 
March 21, 1935 


U. S. Grant, IV, Chairman 

Fred Baker Clinton G. Abbott, Editor 



Harry E. Wheeler 

Stanford University, California 

The first recorded trilobites from the late Paleozoic of California 
were collected in 1892 by H. W. Fairbanks 2 from the Baird formation 
near the U. S. Government Fish Hatchery on the McCloud River. His 
two fragmentary specimens were referred by A. W. Vogdes to Proetus 
ellipticus Meek and Worthen, a species from the Kinderhook group of 

Unfortunately, Fairbanks' specimens are no longer available for 
study. Nevertheless, several pygidia collected by J. P. Smith 4 the follow- 
ing year, and referred by him to the same species, are still in the Stanford 
collection. These, together with a specimen which Professor Muller and 
I collected in 1931, form the basis for the present description of the 
Baird species. 

The Nosoni formation (Permian) has yielded the other trilobite 
species described in the following paper. 

Genus PROETUS Steininger, 1830 

Genotype: Proetus curieri Steininger, 1830 

Proetus bairdensis Wheeler, new species 

Plate 6, figs. 1-3 

1892. ?Proetus ellipticus Meek and Worthen, Vogdes, A. W., "Proceedings 

1 Read before the Paleontological Society of America, Pacific Coast Branch, Berkeley 
meeting, April 14, 1934. 

2 Fairbanks, H. W., "Geology and Mineralogy of Shasta County," 1 1th Rep. Calif. 
State Mineral., p. 39, 1893. 

"Notes on some Localities of Mesozoic and Paleozoic in Shasta County, California," 
Amer. Geol., vol. 14, no. 1, p. 29, 1894. 

3 Zoe, vol. 3, p. 274, 1892. Under the heading of "Proceedings of the Societies, Cali- 
fornia Academy of Science, Oct. 17, 1892," appears the following quotation: "The secretary 
read an announcement of the discovery by H. W. Fairbanks of Proetus ellipticus Meek, a 
trilobite from the Waverly Group, in Shasta County, California, identified by Captain 
A. W. Vogdes." 

4 Smith, J. P., "The Metamorphic Series of Shasta County, California," Jour. Geol., 
vol. 2, no. 6,' p. 595, 1894. 

50 San Diego Society of Natural History 

of the Societies, California Academy of Science [meeting] , Oct. 17, 
1892," Zoe, vol. 3, p. 274. 

1894. ?Proetus ellipticus Meek and Worthen, Fairbanks, H. W., "Notes on 
some Localities of Mesozoic and Paleozic in Shasta County, Califor- 
nia," Amer. Geol., vol. 14, no. 1, p. 29. 

1894. Proetus ellipticus Meek and Worthen, Smith, J. P., "The Metamor- 
phic Series of Shasta County, California," Jour. Geol., vol. 2, no. 6, 
p. 595. 
Not Proetus ellipticus Meek and Worthen, Geol. Surv. Illinois, vol. 
3, p. 460, pi. 14, fig. 3, 1868. 

Description. — The specimens of this species now available for study are 
imperfectly preserved fragments of pygidia, an external mold of one individual 
showing the pygidium, thorax and part of the glabella, and a free cheek (prob- 
ably of the same individual) . 

Although the anterior border of the cephalon is unknown, the remaining 
outline suggests the general form of an elongate ellipse. 

The cephalon is straight at the sides, the genal angles being drawn out into 
slender spines which extend backward to about the fifth thoracic segment. The 
lateral borders (and probably the anterior as well) are folded upward to form a 
raised border which is separated from the cheeks by a deep furrow. The occipital 
ring is about twice as wide as a thoracic segment, and is raised above the level of 
the highest part of the glabella. These two structures are separated by a distinct 
groove which extends laterally across the cheeks to the marginal furrow, which it 
intersects perpendicularly. The posterior-lateral lobes of the glabella stand in low 
relief, and are bordered anteriorly by shallow lateral furrows which extend 
obliquely backward to intersect the neck furrow. 

The thorax is about one-third wider than long, and consists of nine seg- 
ments. The moderately arched axis is about equal in width to the lateral lobes, 
from which it is separated by well-defined dorsal furrows. The lateral lobes are 
depressed below the axis, are somewhat flattened near the dorsal groove, bend 
downward at the fulcral point, and are flattened again from there to the margin. 
Each of the pleurae is marked by a median groove inside the fulcral point. 

The pygidium is sub-semicircular, is nearly twice as wide as long, and is of 
moderately high convexity. It bears about thirteen axial, and eight or nine pleural 
segments. The axial lobe is prominent, its anterior width being a little greater 
than that of the lateral lobes. The top of the axis is flattened, and the sides slope 
steeply to the furrows. The pygidium is entirely surrounded by a smooth mar- 
ginal border of approximately the same width at all points. 

Comparisons and affinities. — This species is in many respects similar to 
Proetus ellipticus Meek and Worthen from the Kinderhook of Illinois, to which 
it was referred by both Vogdes and Smith. However the Baird species possesses 
a greater number of segments in the thorax and in both the axial and pleural lobes 
of the pygidium. Furthermore, both the pygidial axis and the entire pygidium 
are relatively wider in Proetus bairdensis than in the Kinderhook species. 

Wheeler — Trilobite Species 5 1 

Holotype. — Stanford Univ. Paleo. Type Coll., catalogue no. 777-a. 

Paratype.— Stanford Univ. Paleo. Type Coll., catalogue no. 777-b. 

Plastotype. — San Diego Society of Natural History Trilobite Coll., cata- 
logue no. 272. 

Type locality.— L. S. J. U. loc. 1041, Redding Quadrangle, Shasta County, 
California. Highly indurated buff colored shale on crest of spur in the S. W. ]/ 4 
of the S. E. 14, sec. 14, T. 34 N., R. 4 W. Elevation 1000 feet. 

Formation and age. — Baird formation. The age of the strata at this locality 
has not as yet been precisely determined. I have shown elsewhere 5 that the 
Gigantella-bearing strata of the Baird formation (which apparently lie strati- 
graphically below the beds at the Proetus bairdensis locality) are of latest 
Dinantian age. On the basis of stratigraphic position, therefore, it is probable 
that the strata at the type locality of P. bairdensis belong to the Lower Moscovian 

Collectors.— S. W. Muller and H. E. Wheeler, 1931. 

Genus GRIFFITHIDES Portlock, 1843 

Genotype: Griff ithides longiceps Portlock, 1843 

Griffithides nosoniensis Wheeler, new species 

Plate 6, figs. 6 and 7 

Description. — Although the posterior portion of the pygidium is unknown, 
the outline of the remainder of the specimen suggests an elongate ellipse as the 
general form. The greatest width is probably about eight-thirteenths of the length. 
Measured along the axis, the length of the cephalon is nearly equal to the length 
of the thorax. 

The outline of the cephalon (including the spines) forms slightly more than 
half of an ellipse. The spines extend backward to about the sixth thoracic seg- 
ment. The glabella, which is of low convexity, is pyriform, and is especially ex- 
panded anteriorly. It is marked by two pairs of obsolete lateral furrows which 
extend inward from the forward portion of the broad and deep grooves which 
lie in front of the basal lobes. The occipital ring, whose width is about one- 
third greater than that of a thoracic segment, is marked anteriorly by a broad fur- 
row. The surface of the eyes has been removed, and, in consequence, their exact 
form is unknown. 

The thorax consists of nine segments. The axis, which is moderately arched, 
is slightly wider than the lateral lobes, and is separated from them by deep dorsal 
furrows. The pleurae are also moderately arched, their crests being at the fulcral 
points, where they bend rather abruptly downward and slightly backward. 

The pygidium, which is known only in part, is of fairly low convexity, bears 

5 Wheeler, H. E., "The Carboniferous-Permian Dilemma," Jour. Geol., vol. 42, no. 1, 
p. 68, 1934. 

52 San Diego Society of Natural History 

a greater number of axial than lateral segments (as judged from the forward 
portion) , and possesses, at least anteriorly, a narrow and smooth border. 

Comparisons and affinities. — Among the known trilobites, Griffithides 
nosoniensis most closely resembles G. acanthiceps Woodward from the Carboni- 
ferous limestone of England. The Nosoni species differs from G. acanthiceps 
in the greater anterior expansion of its glabella, its proportionally shorter cepha- 
lon, its wider and more anteriorly arched occipital ring, and its obsolete lateral 
grooves on the glabella. 

Holotype. — Stanford Univ. Paleo. Type Coll., catalogue no. 778. 

Plastotype. — San Diego Society of Natural History Trilobite Coll., cata- 
logue no. 272. 

Type locality.- — L. S. J. U. loc. 1034, Redding Quadrangle, Shasta County, 
California. Dark shale on the south side of the ridge south of Potter Creek, 
about 250 feet stratigraphically above the McCloud-Nosoni contact. Elevation 
1800 feet. N. E. ]/ 4 of the S. W. l/ 4 , sec. 24, T. 34 N., R. 4 W. 

Formation and age. — Nosoni formation, Permian (Kungurian?) . 

Collectors.— S. W. Muller and H. E. Wheeler, 1931. 





Harry E. Wheeler 

Stanford University, California 

An endeavor to locate comparative material representing the genera 
Griffithides and Phillipsia has brought to my attention a specimen from 
the Stanford University Paleontological Collection, which carries the 
label, "Phillipsia (Griffithides) ornatus Vogdes, Lower Coal Measures, 
Conway County, Arkansas. Original Type Specimen." A comparison 
of this specimen with Vogdes' original description 1 and J. P. Smith's 
republication of that description reveals that both the description and 
figure of this species are inadequate. Furthermore, a nomenclatural study 
shows that the specific name is a homonym, and must accordingly be re- 
jected. Thus, the purpose of this paper is to rename, redescribe and re- 
figure this Anthracolithic trilobite species. 

Genus GRIFFITHIDES Portlock, 1843 
Genotype : Griffithides longiceps Portlock, 1843 
Griffithides conwayensis Wheeler, new name 

Plate 6, figs. 4 and 5 

1895. Griffithides ornata Vogdes, "Notes on Paleozoic Crustacea No. 4. — 
On a New Trilobite from Arkansas Lower Coal Measures," Proc. 
Cal. Acad. Sci., ser. 2, vol. 4, pp. 589-591, text fig. 

1895. Griffithides ornata Vogdes, "A Supplement to the Bibliography of 
Paleozoic Crustacea," Proc. Cal. Acad. Sci., ser. 2, vol. 5, p. 73, 

1897. Phillipsia (Griffithides) ornata (Vogdes), Smith, J. P., "Marine Fos- 
sils from the Coal Measures of Arkansas," Proc. Amer. Phil. Soc, 
vol. 35, no. 152, pp. 61-63, pi. 22, fig. 6. 
Not Phillipsia ornata Portlock, "Report on the Geology of the County 
of Londonderry and Parts of Tyrone and Fermanaugh," p. 307, fig. 
2, Dublin, 1843. 
Not Phillipsia? (Brachymetopus?) ornata Hall, "Illustrations of De- 
vonian Fossils," New York Geol. Surv., pi. 21, fig. 1, 1876. 

1 See synonymy for citation of references not footnoted. 

54 San Diego Society of Natural History 

1898. Griffithides ornatus Vogdes, Weller, S., "A Bibliographic Index of 

North American Carboniferous Invertebrates," U. S. Geol. Surv., 

Bull. 153, p. 302. 
1933. Griffithides ornatus Vogdes, Williams, J. S., "A New Pennsylvanian 

Trilobite from Missouri," Jour. Washington Acad. Sci., vol. 23, no. 9, 

pp. 431-432. 

Description. — The general form is sub-ovate, with the greatest width being 
about two-thirds that of the length. Measured along the axis, the cephalon, thorax 
and pygidium are all of about equal length. 

The outline of the cephalon (excluding the spines) forms about half of a 
near circle. The head-shield is bounded by a marginal border which extends 
backward from the posterior-lateral angles, in the form of genal spines, to at least 
the fourth or fifth thoracic segment. The cephalic border and spines bear about 
eight very fine parallel costae. The glabella is pyriform and gibbous in front. 
Its basal lobes are small, but well marked by deep furrows, whose anterior ends 
are connected by a shallow groove across the glabella. From the center of this 
groove, a faint furrow extends backward, thus defining two small rounded nodes 
near the posterior margin of the glabella. The occipital ring is broad and well 
defined, its width being nearly twice that of a thoracic segment. The eyes are 
large, smooth and quarto-spherical. 

The thorax consists of nine segments. Its axis is strongly arched, and bears 
a series of about ten small tubercles across the center of each segment. The pos- 
terior border of each pleural segment, beyond the fulcral point, bears a row of 
extremely fine tubercles. 

The pygidium is very convex, and is surrounded by a marginal border 
which widens anteriorly. Its strongly arched axis consists of eleven segments, 
each bearing, along its flattened crest, a row of six nodes which are slightly larger 
than those of the thorax. Segmentation on the sides of the axis is but faintly 
defined. The lateral lobes of the pygidium consist of seven segments with a ves- 
tige of an eighth. These segments are divided by deep furrows, and each bears a 
node at its fulcral point, from whence it is abruptly truncated. 

Comparisons and affinities. — Vogdes compared this trilobite with Griffi- 
thides scitula Meek and Worthen from the "Illinois Coal Measures." However, 
as pointed out by Williams, 2 any comparison with G. scihda is of little value, 
since the type is apparently no longer available for study, and since Meek and 
Worthen were the only authors known to have examined the type. 

G. conwayensis compares favorably in many respects to G. olsoni Williams 3 
from the Cherokee shale (Lower Pennsylvanian) of Missouri. G. conwayensis 
differs from G. olsoni, however, in its more conspicuous basal glabellar lobes, its 
greater number of pleural segments of the pygidium, and in the noded character 
of its axis. 

2 Williams, J. S., Jour. Washington Acad. Sci., vol. 23, no. 9, p. 431, 1933. 

3 Ibid., pp. 429-435. 

Wheeler — Trilobite Species 55 

G. conwayensis appears to be more closely allied to G. parvulus Girty 4 
from the Wewoka formation of Oklahoma. The Arkansas species differs from 
G. parvulus in its much smaller width-length ratio, its narrower neck ring, its less 
distinctly defined transverse glabellar furrow, the absence of any indication of a 
second such furrow, and the presence of a longitudinal groove dividing the basal 
glabellar lobe. 

Discussion.— In 1895 Vogdes described and figured this species under the 
heading "Griffithides ornata sp. nov." At the time of its proposal, 
the validity of that name depended entirely upon the taxonomic rank of 
Griffithides, as interpreted from Vogdes' paper. If it be construed that he re- 
garded Griffithides as a subgenus of Phillipsia, his specific name becomes a 
homonym of Phillipsia ornata Portlock, 1843. In his description, Vogdes makes 
no mention of the genus Phillipsia, from which we may interpret that he treated 
Griffithides as being of full generic rank. On the other hand, under the sub- 
heading of "Affinities and differences," he cites alternately Phillipsia {Griffi- 
thides) scitula and Griffithides scitula, thus, apparently sanctioning the sub- 
generic usage of Griffithides. Furthermore, it should be noted that the name 
Griffithides is masculine, while the specific name ornata is feminine. This dis- 
cordance might be cited as additional evidence for Vogdes' intention to employ 
Griffithides as a subgenus of Phillipsia (feminine) ; otherwise, the name should 
have been ornatus. One might construe, therefore, either that Vogdes did or 
did not regard Griffithides as a subgenus of Phillipsia. J. P. Smith, however, in 
his republication ot Vogdes' description, leaves no doubt as to the status of the 
name in question. Under the heading of "Genus Phillipsia, Portlock," he 
cites the species as Phillipsia (Griffithides) ornata A. W. Vogdes. Through this 
action, Vogdes species unquestionably becomes a homonym of Phillipsia ornata 
Portlock; and it must accordingly be rejected and supplanted by a new name. I 
therefore propose the name Griffithides conwayensis for this species. 

Holotype.— Stanford Univ. Paleo. Type Coll., catalogue no. 5077. The 
holotype of Griffithides conwayensis is the same specimen upon which Vogdes 
based his species "ornata." 

Plastotype. — San Diego Society of Natural History Trilobite Coll., cata- 
logue no. 274. 

Type locality.— L. S. J. U. loc. 1040, Morrillton Quadrangle, Conway 
County, Arkansas. Near the center of the N. W. T/ A , sec. 17, T. 5 N., R. 16 W. 

Formation and age. — Atoka 5 formation, Lower Moscovian (Lower Penn- 
sylvanian) . 

Collectors. — Arkansas Geological Survey. 

4 Girty, G. H., New York Acad. Sci. Annals, vol. 21, p. 154, 1911; Bull. U. S. Geol. 
Surv., no. 544, pp. 268-270, pi. 18, figs. 14-15, 1915. 

5 Carey Croneis shows ("Geology of the Arkansas Paleozoic Area," Ark. Geol. Surv., 
Bull. 3, 1930, structural map in fold) that the locality of Griffithides conwayensis lies very 
close to the axis of the Redemption anticline; and on page 265 of the same paper, he 
states that the surface rocks of that structure belong to the Atoka formation. A personal 
communication from Dr. Croneis (Feb., 1934) gives reassurance that this locality is under- 
lain by the Atoka formation. 

56 San Diego Society of Natural History 


Fig. 1. Proetus bairdensis Wheeler, new species. Rock surface containing the 
holotype (incomplete external mold of cephalon, thorax and pygi- 
dium), and paratype (free cheek). Holotype, Stanford Univ. Paleo. 
Type Coll., no. 777-a, paratype, no. 777-b. L. S. J. U. loc. 1041, 
Baird formation, Redding Quadrangle, Shasta County, California, 
x 1.9. 

Fig. 2. Proetus bairdensis Wheeler, new species. Paratype; same specimen as 
in upper left corner of fig. 1. x 1.9. 

Fig. 3. Proetus bairdensis Wheeler, new species. A clay cast impressed from 
the external mold (holotype) shown in fig. 1. x 1.9. 

Fig. 4. Griffithides conwayensis Wheeler, new name. Holotoype, Stanford 
Univ. Paleo. Type Coll., no. 5077. L. S. J. U. loc. 1040, Atoka 
formation, Lower Moscovian, Morrillton Quadrangle, Conway 
County, Arkansas, x 2.1. 

Fig. 5. Griffithides conwayensis Wheeler, new name. A line drawing traced 
from a photograph of the specimen shown in fig. 4. 

Fig. 6. Griffithides nosoniensis Wheeler, new species. Holotype, Stanford 
Univ. Paleo. Type Coll., no. 778. L. S. J. U. loc. 1034, lower No- 
soni formation, Permian, Redding Quadrangle, Shasta County, Cali- 
fornia, x 2.1. The dark areas on either side of the anterior half of the 
specimen represent the cavities remaining after the dissolution of the 
marginal border and genal spines of the cephalon. 

Fig. 7. Griffithides nosoniensis Wheeler, new species. A line drawing of the 
cephalon traced from a photograph of the specimen shown in fig. 6. 
Gl, glabella; L, basal glabellar lobe; OR, occipital ring; S, suture; 
GS, genal spine. 

Wheeler — Trilobite Species 

Plate 6 



Volume VIII, No. 9, pp. 59-66, plate 7 



George L. Richards, Jr.. 

Stanford University, California 

Printed for the Society 
March 21, 1935 


U. S. Grant, IV, Chairman 

Fred Baker Clinton G. Abbott, Editor 



George L. Richards, Jr. 

Stanford University, California 

Extensive field work throughout the central Coast Ranges of Cali- 
fornia during the past two years has shown that various species of the 
genus Astrodapsis are of value in establishing the stratigraphic correla- 
tion of Upper Miocene and Lower Pliocene formations. 

During the paleontological study of these late Tertiary marine 
sediments, the valuable works of Clark and Twitchell, 1 and Kew, 2 were 
found to be almost indispensable in the determination and identification 
of the various genera and species of the Pacific Coast Echinoidea. How- 
ever, a study of the original description and figure of the type of Astro- 
dapsis antiselli Conrad suggested that the workers had misidentified 
this species, which is the type of the genus Astrodapsis. 

Through correspondence with the Curator, Division of Mollusks, 
U. S. National Museum, photographs of specimens were obtained which 
confirmed this opinion, and indicated that the specimen figured by 
Clark and Twitchell (U. S. National Mus. Cat. No. 165466a) cannot 
be regarded by Kew as a type specimen of A. antiselli Conrad. Further- 
more, this specimen is not from the original lot, nor even from the type 
locality. A photograph of the above mentioned specimen shows it is not 
a true Astrodapsis antiselli as originally defined and figured by Conrad. 
Unfortunately, Kew followed Clark and Twitchell and recognized this 
figured specimen as the holotype. The problem is therefore two-fold: 
(l) biologic identification of species, and (2) nomenclatorial. In order 
to correct this confusion it is necessary to make several corrections and, 
in addition, propose one new name, to wit : 

Astrodapsis salinasensis, new name 

"Astrodapsis antiselli Conrad," of Clark and Twitchell, 1915, also of Kew, 1920, 
but not of Conrad, 1856. 

1 Clark, W. B. and Twitchell, M. W. The Mesozoic and Cenozoic Echinodermata 
of the United States. U. S. Geological Survey, Monogr. 54, 1915. 

2 Kew, W. S. W. Cretaceous and Cenozoic Echinoidea of the Pacific Coast of North 
America. Univ. Cal. Pub. Geol., Vol. 12, no. 2, pp. 23-236, 1920. 

62 San Diego Society of Natural History 

Determinative characters. — Clark and Twitchell, verbatim: 3 "Test medium 
in size; regularly oval in marginal outline, longer than broad, slightly truncated 
at anterior end, slightly pointed at posterior end, with faint notches opposite 
ends of petals; margin rounded and very thick, almost as thick as rest of test. 
The whole form is considerably depressed, almost equally so from edge to edge, 
and therefore subdiscoidal; the upper surface with broad flattened ambulacral 
ridges alternating with narrow interambulacral depressions; apex eccentric an- 
teriorly, in front of depressed apical system; lower surface slightly concave. Am- 
bulacral petals large, broad, tumid, especially near apical system; poriferous 
zones narrow, at first diverging, then converging slightly from one-fourth to one- 
third the way to margin and again diverging to the wide open ends which are 
nearly to the margin. Peristome central; the main ambulacral grooves straight, 
well denned, and rather deep from peristome to margin and continuing as faint 
lines over margin to near apex, two faint lines are given off about half way to 
margin, which continue over margin to near apical system. Periproct small, infra 
marginal, almost marginal." 

Dimensions. — Specimen B, type, (U. S. National Mus. Cat. No. 
165466a) : length, 57 mm.; width, 50 mm.; height, 14 mm. 

Localities. — Specimen B (No. 165466a) : "2 miles south of San Lucas, 
Monterey County, Cal." (Clark and Twitchell). 

Collection.— U. S. National Mus. Cat. No. 165466, Specimen A, cotype. 
U. S. National Mus. Cat. No. 165466a, Specimen B. Both the type and the 
specimens collected by Ralph Arnold, which include specimens A and B. 4 

Remarks. — Astrodapsis salinasensis (No. 165466a) misidentified as "Astro- 
dapsis antiselli Conrad" by Clark and Twitchell, and by Kew, is herein figured 
(Plate 7, figs. 2a, 2b, and 2c) for comparison with the original type of true 
Astrodapsis antiselli Conrad (U. S. National Mus. Cat. No. 13337), which is 
lebelhjd "Conrad's type," from Estrella, Monterey County, California. (See 
Plate 7, figs, la, and lb). A. salinasensis differs from A. antiselli Conrad in the 
following characters: 

Astrodapsis salinasensis Astrodapsis antiselli Conrad 

Test: Discoidal, oval; slightly notched. Pentagonal; markedly notched oppo- 
site ends of petals. 

Margin: Broadly rounded and thick; almost Sightly rounded; greatest elevation 

as thick as rest of test— biscuit adjacent to the depressed apical 

sha P ed - system. 

Apical system: 

Moderately depressed. Deeply depressed. 

Tubercles: Very prominent. N ot prominent. 

Petals: Low, broad, and tumid. Elevated, narrow, angular. 
Interambulacral areas: 

Rounded, shallow grooves. Angular, deep grooves. 

3 Op. cit. p. 198. 

4 This information from Clark and Twitchell, 1915, p. 199. 

Richards — California Species of Astrodapsis 63 

Distribution. — Geographically Astrodapsis salinasensis occurs abundantly 
in the fine, medium to coarse, white, littoral marine sandstones at the top of the 
Santa Margarita formation, or sandstone facies of the Upper Miocene Monterey 
Shale, throughout the entire Salinas Valley, Monterey County, California, as 
well as in similar standstones of the Santa Margarita formation as exposed along 
Bean Creek, Santa Cruz County, California, and the uppermost Santa Margarita 
sandstones exposed along Saucelito Creek, Nipomo Quadrangle, San Luis 
Obispo County, California. 

Stratigraphically Astrodapsis salinasensis occurs in a monoclinal, upper 
Miocene section, approximately 250 feet above organic and siliceous shales con- 
taining a N onion schencki foraminiferal assemblage, which in turn overlies sand- 
stones containing Astrodapsis tumidus, Astrodapsis whitneyi, Ostrea titan cor- 
rugata, and associated faunal assemblage. It occurs below sandstones containing 
Astrodapsis cf. jacalitosensis and lower Pliocene mollusks belonging to the Jaca- 
litos faunal assemblage. 

Associated faunal assemblage. — Astrodapsis salinasensis occurs with the 
following forms: Pecten estrellanus Conrad (18-20 rib var.), Tritonalia sp., 
Balanus concavus Bronn, "Tamioso?na" gregaria Conrad, Astrodapsis spatiosus 

Additional revision. — In the monographs by Clark and Twitchell, and Kew, 
the true Astrodapsis antiselli Conrad was named Astrodapsis arnoldi through 
the unfortunate misidentification mentioned above. In order to correct this 
nomenclatorial problem, it is necessary to consider those forms originally de- 
scribed by Kew (1921) as subspecies of Astrodapsis antiselli Conrad, or to 
regard them as of full specific rank. The revised nomenclature of all the forms 
involved in this problem is as follows: 

Old arrangement (Kew) New arrangement 

Astrodapsis arnoldi arnoldi Astrodapsis antiselli Conrad 












odapsis antiselli Conrad" of Kev.'. Astrodapsis salinasenis, new name 
not Conrad 

Acknowledgments. — The writer is indebted to Dr. Alexander Wetmore, 
Assistant Secretary of the Smithsonian Institution, and Mr. Wm. B. Marshall 
of the Division of Mollusks, U. S. National Museum, for permission to repro- 
duce photographs of the type of Astrodapsis antiselli Conrad (U. S. National 
Mus. Cat. No. 13337); photographs of Astrodapsis salinasensis new name (U. 
S. National Mus. Cat. No. 165466a) were supplied through the courtesy of the 
U. S. Geological Survey. He is also grateful to Dr. Hubert G. Schenck of 
Stanford University, and Dr. U. S. Grant of University of California at Los 
Angeles, for suggestions concerning nomenclatorial problems and the prepara- 
tion of the manuscript. 

64 San Diego Society of Natural History 


All figures approximately natural size. 

Fig. la. Astrodapsis antiselli Conrad. 

Genotype, U. S. National Mus. Cat. No. 13337. Upper surface of 
test. Estrella, Monterey Co., California. 

Fig. lb. Astrodapsis antiselli Conrad. 

Same specimen. Lateral view of test. 

Fig. 2a. Astrodapsis salinasensis, new name. 

Holotype, U. S. National Mus. Cat. No. 165466a, Specimen B. 
Lower side of test. Two miles South of San Lucas, Monterey Co., 

Fig. 2b. Astrodapsis salinasensis, new name. 

Same specimen. Upper surface of test. 

Fig. 2c. Astrodapsis salinasensis, new name. 

Same specimen. Lateral view of test. 

Richards — California Species of Astrodapsis 

Plate 7 

PgK* r 

2 a. 


2 a 

2 b. 



Vol. VIII, No. 10, pp. 67-68 August 24, 1935 



San Diego Society of Natural History 

In the spring of 1930 while at Tobari Bay on the coast of southern 
Sonora I collected a series of breeding mangrove warblers. These, on a 
more critical examination than I was able previously to give them, prove 
not to be of the subspecies castaneiceps, but to belong to an undescribed 
race which is apparently more closely related to Dendroica erithachorides 
xanthotera of the Pacific coast of Central America. At present the com- 
bined material in the San Diego Society of Natural History and the 
Dickey collection (for the privilege of using which I am under obligations 
to Mrs. Dickey) totals 64 specimens. These consist of 15 xanthotera, 30 
castaneiceps, and 19 of the Sonora race. A description of the new race, 
together with comment on the others is herewith offered. The new race 
may be called 

Dendroica erithachorides rhizophorae subsp. nov. 

Type. — Male adult, no. 17090, collection of the San Diego Society of 
Natural history; Tobari Bay, Sonora, Mexico, April 30, 1930; collected by A. J. 
van Rossem. 

Subspecific characters. — General size very similar to Dendroica erithacho- 
rides xanthotera Todd of the Pacific coast of Central America, though averaging 
even smaller in wing length and size of bill; definitely smaller in all dimensions 
than Dendroica erithachorides castaneiceps Ridgway of southern Lower Cali- 
fornia. The tail has less yellow than in xanthotera; more than in castaneiceps. 

68 San Diego Society of Natural History 

Adult males have the chestnut of the throat more restricted and the under parts 
usually more heavily streaked than in either xanthotera or castaneiceps; colora- 
tion of both sexes otherwise similar to castaneiceps, that is to say less richly col- 
ored than xanthotera. 

Range. — Coast of southern Sonora from the northern limit of mangroves 
at Tepopa Bav south (Kino Bay; Guaymas; Tobari Bay) to Agiabampo on the 
Sonora-Sinaloa boundary, and probably for some distance further south. 

Remarks. — In considering the most characteristic features of the three races 
under comparison there are immediately noticeable the large size of castaneiceps, 
the restriction of chestnut and heavy streaking of rhizophorae, and the rich gen- 
eral coloration of xanthotera. 

There are two very distinct color phases in the young of this species, and 
these persist at least through the post-juvenal moult. Whether or not they nor- 
mally persist beyond that stage I do not know. There is evidence both ways in 
the combined series of 64 skins. However, both the gray and the yellowish olive 
green phases are present in unquestionably young females of castaneiceps and in 
spring females of uncertain age of rhizophorae. In xanthotera the pale phase is 
almost indistinguishable from the olive green phase of the northern races and 
the bright phase is, of course, infinitely richer and yellower. Evidence that the 
pale phase persists at times into the adult stages is shown by two fully adult males 
of xanthotera from Costa Rica. 

It is certain that a large amount of carefully collected specimens will be 
necessary in order to determine not only the duration of the pale phase, but its 
relative abundance in different geographic areas. The material before me indi- 
cates that it is much more prevalent southerly. 

Measurements of Adult Males in Millimeters 

Extremes and Averages 





Middle Toe 
minus Claw 




























Vol. VIII, No. 1 1, pp. 69-72 August 24, 1935 




San Diego Society of Natural History 

In the latter part of May of the present year I was invited by Mr. 
Christopher Henne of Pasadena to accompany him on a short trip to the 
Argus Mountains, a small, excessively arid range in southern Inyo and 
extreme northern San Bernardino Counties. This range forms, in part, the 
western rim of Panamint valley. The region is adequately mapped by the 
U. S. Geological Survey and is covered by the Searles Lake and Ballarat 
Quadrangles, and Plate X of Water Supply Paper No. 490. 

Our three day reconnaissance of Mountain Springs Canyon on the 
west slope of the range in Inyo County was made more than casually in- 
teresting by the discovery that the thickets of willows, which in interrupted 
fashion follow the canyon bed from 4200 to 5500 feet altitude, harbored 
a breeding colony of brown towhees. This colony is of course effectually 
isolated from the geographically nearest race of brown towhee, Pipilo fus- 
cus carolae, by the Sierra Nevada as well as by intervening deserts. 

The six specimens collected cannot satisfactorily be placed with any 
of the known races of this rather "plastic" species, a not surprising circum- 
stance in view of the fact that the isolated habitat lies in a region far re- 
moved faunally from the coastal and interior valley habitats of other 
brown towhees of the crissalis group. This region, a part of the Inyo 
Division of the Great Basin Faunal Area, is already characterized by 
numerous subspecies of birds and mammals. The new towhee is named as 

70 San Diego Society of Natural History 

Pipilo fuscus eremophilus subsp. nov. 

Type.— Breeding male adult, no. 17083, collection of the San Diego 
Society of Natural History; Lang Spring, 5500 feet altitude, Mountain Springs 
Canyon, Argus Mountains, Inyo County, California, May 22, 1935; collected 
by A. J. van Rossem. 

Subspecifc characters.— Most closely resembles Pipilo fuscus carolae Mc- 
Gregor of the Sacramento-San Joaquin Valley of California, but bill smaller, 
tarsi and toes decidedly shorter, and coloration slightly darker and grayer. 
Resembles Pipilo fuscus crissalis of the Pacific slope of southern California, but 
wing and tail longer, bill much more slender, both in lateral and vertical profile, 
and coloration grayer. 

Range. — Argus Mountains of Inyo and San Bernardino Counties, south- 
eastern California. 

Remarks. — The gray, dark coloration of the desert race is more pronounced 
in the single juvenile than in the worn adults. Since wear tends to obscure com- 
parative color values between races of this species it seems likely that fresh- 
plumaged adult specimens will show even more definite differences than are ap- 
parent in worn series. 

There are several matters of interest connected with the characters displayed 
by this desert race of brown towhee. First, there is no approach in any particular 
toward mesoleucus, indeed the tendencies are away from that race. Second, there 
would appear to be every reason to suspect, a prion, that a race resident in the 
Argus range would show relative pallor, compared with other crissalis subspecies, 
for not only is the region one of extreme aridity and high temperatures, but the 
soil color is definitely pale — a light colored granite which weathers reddish rather 
than gray. Large size and pallid coloration rather generally characterize Inyo 
subspecies, but the trend in this instance is an exception. 

In going over the published literature I was surprised to find that Frank 
Stephens had taken a brown towhee in the Argus Mountains on April 25, 1891, 
at which time he was a member of the Death Valley Expedition. The record 
seems to have been generally overlooked by reviewers, though Ridgway (in Pt. 
1 of Birds of No. & Mid. Amer., p. 435), in the bibliography of "Pipilo crissalis 
senicula," repeats the record which was first published by Fisher on page 105 of 
North American Fauna No. 7. 

Through the courtesy of the Bureau of Biological Survey I am able to ex- 
amine this specimen. It has become reddened by post-mortem color change and 
is now indistinguishable in color from recently collected examples of carolae. 
However, the mensural characters accord strictly with those of eremophilus. The 
precise locality on the label is given as "Searles Garden," presumably at, or near, 
Searles Borax Works at the south end of the range. 

In addition to this specimen borrowed from the Biological Survey, I must 
acknowledge the use of specimens in the Dickey collection, and the courtesy of 
Dr. Joseph Grinnell in sending me a comprehensive series of carolae from the 
west slope of the Sierra Nevada. 

van Rossem — New Brown Towhee 


Measurements of Males in Millimeters 1 
Extremes and Averages 


Depth at 

Middle Toe 



Cut »i i n 



mi ii n s Clau 

1 crissalis 

from Los Angeles 







and Ventura Cos. 







10 carolac from 







the range 







4 eremophilus 













1 Wing and tail measurements are all from worn specimens, so far as possible in com- 
parable stages of abrasion. Fresh plumaged birds measure somewhat longer. In this connec- 
tion, as well as for modern systematic treatment of various California races, see the measure- 
ments recorded by Swarth, Condor, 1918, pp. 117-121, and Grinnell and Swarth, Univ. 
Calif. Pub. Zool., 21, no. 18, 1926, pp. 427-433. 



Vol. VIII, No. 12, pp. 73-74 August 24, 1935 



Laurence M. Huey 

Curator of Birds and Mammals, San Diego Society of Natural History 

In a collection of mammals secured in early 1935 by the writer for 
the San Diego Society of Natural History at Bahia Kino, Sonora, Mex- 
ico, are four specimens of Perognathus longimembris that differ in several 
characters from the hitherto described forms of this species. This race 
may be known as : 

Perognathus longimembris kinoensis subsp. nov. 
Kino Silky Pocket Mouse 

Type. — From Bahia Kino, Sonora, Mexico (more precisely — from the 
northern end of the sand dune peninsula that borders the bay and forms the north- 
ern arm of the estuary); no. 11300, collection of the San Diego Society of 
Natural History; adult male; collected by Laurence M. Huey, February 26, 1935. 

Characters. — In color, kinoensis is darker than Perognathus longimembris 
bombycinus. its nearest relative. The most prominent characters of this form are 
cranial, and compared with bombycinus the skull of kinoensis is more rounded 
and narrower across the bullae. The interparietal is almost square in shape, and 
the nasals are longer and more attenuated. In some respects it approaches P. I. 
pacifcus from the coastal region of San Diego County, California; for example, 
in its very small size and compressed, arched skull with rounded bullae. How- 
ever, in color these two races are widely different. 

Measurements. — Type: Total length, 135; tail, 80; hind foot, 17; ear, 4. 
Skull (type): Greatest length, 20.7; width across bullae, 11.4; interorbital con- 
striction, 4.6; nasals, 7.2; tooth row, 2.6. 

74 San Diego Society of Natural History 

Range. — So far as known, only the type locality. 

Remarks. — P. I. kinoensis provides an interesting illustration of color devel- 
opment, as compared with the other two races of Perognathns longimembris 
mentioned in this paper. It is a well established fact that desert and coastal forms 
are respectively light and dark. This is true of P. I. bombyanus and P. I. pacificus. 
In P. I. kinoensis, however, we have a coastal race whose range borders the humid 
tropical desert of central Sonora. Here it has developed a grayish cast, as con- 
trasted with the rich black tendencies found along the coast in southern California. 

Specimens examined. — Perognathus longimembris bombycinus: 2 from 6 
miles east of Yuma, Arizona (type locality) ; 2 from 3 miles west of Pik>t Knob, 
Imperial County, California; 3 from San Felipe, Lower California, Mexico. 
Perognathus longimembris pacificus: 60 from Tia Juana Valley, San Diego 
County, California (type locality) ; 6 from 4 miles north of Oceanside, San Diego 
County, California; 1 from San Onofre, San Diego County, California. Perogna- 
thus longimembris kinoensis: 4 from Bahia Kino, Sonora, Mexico (type local- 




Volume VIII, No. 13, pp. 75-90, plate 8, map 





Laurence M. Klauber 

Curator of Reptiles and Amphibians, San Diego Society of Natural History 


Printed for the Society 

August 24, 1935 





Laurence M. Klauber 

Curator of Reptiles and Amphibians, San Diego Society of Natural History 

It is with some hesitation that the writer proposes the differentiation 
of the Prairie Rattlesnake of the Little Colorado Basin and the surround- 
ing territory in Arizona as a new subspecies of Cro talus conjluentus. The 
latter is a wide-spread and rather variable snake ; it has already been divided 
into five territorial races, e. g., conjluentus, abyssus, concolor, lutosus, and 
oreganus. However, surveying the type subspecies conjluentus conjluen- 
tus, as now recognized, we find that the Arizona specimens differ from 
those inhabiting the rest of the range from Canada to Mexico in consis- 
tent and conspicuous characters. 

I do not favor the indiscriminate splitting of reptile species in each 
instance where significant differences in some character can be found to 
exist between two or more geographical groups. Thus, specimens of 
Crotalus conjluentus oreganus from Washington can be quite readily dis- 
tinguished from southern California specimens by color and pattern ; and 
it can be shown mathematically that Texas specimens of Crotalus atrox 
have a significant difference in such important characters as dorsal scale 
rows, ventrals, and labials from Arizona individuals. But, after all, these 
differences are largely technical; the snakes themselves are essentially the 
same, and it will serve no practical purpose to recognize each of such dif- 
ferences, with the multiplicity of subspecies that would result in plastic 

But the prairie rattlesnakes of Arizona, and particularly those found 
in the drainage area of the Little Colorado River, between Canyon Padre 
on the west and Bibo on the east, are so conspicuously different in size and 
color, and so significantly different in scale counts from the snakes found 
beyond the Continental Divide, that the divergence will appeal to the ncn- 
herpetologist as well as to the specialist. Venom data will be clarified by 
the recognition of this form, owing to the differences from the typical sub- 
species in yield, and possibly in quality as well. Thus it would seem that 
it is desirable to make this segregation. 

78 San Diego Society of Natural History 

It is true that there is some lack of uniformity in the specimens 
found in this area, so that the relationship pattern is not as clear as 
might be desired. Much of this may be attributed to a few inaccurate 
locality records, or to such effect as the religious rites of the Indians may 
have had upon distribution; but in any case, even if we consider all of 
the Arizona specimens (rather than only the stunted specimens from the 
Winslow area), we will still find a significant divergence, when compari- 
sons are made with the type subspecies. 

While the differences herein mentioned were first recognized in 
1927, initially leading to some confusion with C. tigris, a discussion of 
the problem has been postponed until adequate material has become 
available. Studies have now been made of 200 Arizona specimens of this 
form, and scale counts of 1900 specimens of confluentus confluentus 
from other states are at hand for purposes of comparison. 

Crotalus confluentus nuntius 1 subsp. nov. 
Arizona Prairie Rattlesnake 

Type. — No. 3105 in the collection of L. M. K. Collected at Canyon 
Diablo, Coconino County, Arizona, by R. L. Borden, August 9, 1930. 

Diagnosis. — A stunted subspecies of Crotalus confluentus, predominantly 
reddish-brown in coloration and with low dorsal and ventral scale counts. 

Description of Type. — Adult male. Length (live measurements) 468 mm. 
to rattles, tail length 38 mm., ratio 0.081. Length of head 24 mm., times con- 
tained in body length 19.5. Width of head 17 mm. Width across the supraocu- 
lars 1 1 mm., distance between supraoculars 4.5 mm., ratio 2.44. 

The head is subtriangular, depressed, and, except for the supraoculars, cov- 
ered with small scales. These are raised and unkeeled, excepting those in the tem- 
poral area and toward the neck. 

The dorsal scale rows are 23-23-19; the first row dropped is the 6th, the 
second the 5th. At mid-body all scale rows are keeled, excepting the first two on 
either side. The central dorsal rows are smaller than the lateral; they are the 
more strongly keeled and have moderate posterior bosses. The ventrals number 
166, and the caudals 26, in a single series. The anal is entire. The supralabials 
number 16-16; the infralabials 15-14. The rostral is higher than wide; eight scales 
contact it posteriorly, a first supralabial and prenasal on each side, and 4 interna- 
sals. Between the internasals and the supraoculars there are two canthals on each 
side. The scales on the top of the head, anterior to the supraoculars, number 18. 
The anterior intersupraoculars are 4+6; the anterior boundary of these scales is 

1 Nuntius, the messenger. In the Hopi Snake Ceremonial, these snakes are used as 
messengers to the gods of the underworld. 

Klauber — New Prairie Rattlesnake 79 

indefinite. The nasals are 2-2, the anterior larger; there are 0-1 Ioreals. The 
upper preocular contacts the prenasal on the left; on the right the contact is pre- 
vented by the juxtaposition of loreal and posterior canthal. There are two pre- 
oculars on either side, the upper larger, the lower crescent-shaped and bordering 
the pit above. The postoculars are 3-3, the total scales in the orbit, 8-9; scales 
from the labials to the orbit, 2+3, 2+4. The first and eighth supralabials are 
the largest. The small scales anterior to the pit number 4-3. 

The first infralabials are undivided and are in contact on the median line; 
there are no intergenials. The mental is subtriangular, contacting only the first 
infralabials and a small submental. The genials are in a single pair, short and 
obtuse, and contact 4-5 infralabials. 

The head above is light red-brown, irregularly spotted with darker. The 
supraoculars are crossed with light marks, widening inwardly. On the side of the 
head there is a light preocular stripe passing backward to the angle of the mouth, 
and a second narrower postocular light stripe, about \\ scales wide, the upper 
edge of which is rather indefinite. Between the two light stripes there is a dark 
ocular stripe about 2h scales wide ending above the commissure. The infrala- 
bials are punctated. 

The ground color of the body is light reddish-brown, upon which there are 
superimposed 43 blotches of darker red-brown. On the sides there are secondary 
and tertiary series of ill-defined spots; posteriorly these are confluent with the 
dorsal series so that the last ten blotches become transverse rings, of a somewhat 
lighter color than the anterior blotches. The dorsal blotches are irregular both in 
shape and outline; at mid-body they are ellipses with the major axes transverse 
to the snake. They are about 11 scale rows wide, and longitudinally are 2 to 3 
scales (end to end) long. The blotches are wider (along the body of the snake) 
than the interspaces. The blotches internally are somewhat darker at the borders 
than centrally; exteriorly there is a white edge, but this is neither regular nor 
always present. The blotch borders are independent of scale edges, which is typi- 
cal of confluentus as opposed to scutulatus. The tail is crossed with 10 rings, all 
being brown except the last two, which are black, thus being in strong contrast 
with the rest of the body. The ventrals are straw colored, and somewhat punctated, 
particularly adjacent to the dorsals. 

The rattles, of which 3 remain, measure about 7.3 mm. across. Studies of 
the rattles of this form indicate that there were not less than ten rattles in the 
complete string. The base of the rattles is black. 

The hemipenis is completely bifurcate with divided sulcus. The base on the 
outer shoulders is covered with short, heavy spines, there being about 24 major 
spines on each shoulder, and some 55 smaller points. There are no spines in the 
crotch. The branches are covered with laminate fringes, there being about 27 on 
each lobe. The boundary between spines and fringes is sharply defined. The ratio 
between lobe length and diameter is 2.1, which is approximately the proportion 
usually found in confluentus confluentus. 

General Description and Remarks. — The following is a summary of scale 
counts and measurements of 108 specimens from the area between Canyon Padre, 

80 San Diego Society of Natural History 

Coconino County, Arizona, on the west and Bibo, Apache County, Arizona, on 
the east, and will serve to indicate character variations in specimens from that 
area wherein this form adheres most closely to the type. Other Arizona speci- 
mens are subsequently discussed, but in this summary of the new subspecies it is 
deemed advisable to omit specimens which might be considered intergrades with 
other subspecies. 

Size, small. Scale rows at midbody usually 23 (48 per cent) or 25 (50 per 
cent); rarely 21, 22, or 27 (less than 1 per cent of each). The scales are keeled, 
except the first two on the sides. Posterior scale bosses are not conspicuous. Ven- 
trals: males, max. 181, min. 166, av. 172.29±0.26, interquartile range 170.1 — 
174.5 (68 specimens); females, max. 182, min. 169, av. 177.51±0.32, interquar- 
tile range 175.5 — 179.5 (39 specimens). Anal entire. Caudals: males, 21 to 28, 
average of 68 specimens 24.8; females, 14 to 21, average of 39 specimens 18.5. 
These extremes are seldom attained; the males usually have from 23 to 26 and the 
females from 17 to 20. The caudals, while generally entire, may have a few at 
either end of the series divided. 

The supralabials average 14.8; they usually number 15 (43 per cent), or 14 
(30 per cent); occasionally 16 (18 per cent) or 13 (6 per cent); rarely 17 (less 
than 3 per cent). The infralabials average 15.3; they generally number 15 (38 
per cent), 16 (35 per cent); occasionally 14 (18 per cent) or 17 (7 per cent); 
rarely 12 or 13 (less than 1 per cent of each) . 

The rostral is higher than wide, and in contact with the prenasals. The pre- 
nasals are always in contact with the supralabials. The internasals (scales in con- 
tact with the rostral between nasals, regardless of size or relative position) usually 
number 3, 4, or 5; rarely 2 or 6, the average being exactly 4. The scales on the 
crown, anterior to the supraoculars, vary from 12 to 31; the average is 20.2, with 
an interquartile range of 17.6 to 22.7. The minimum scale rows between supra- 
oculars are usually 3 or 4, rarely 2 or 5, averaging 3.52. Supraocular sutures or in- 
dentations are not present. 

The nasals are 2-2. About 90 per cent of the specimens have one loreal, the 
rest two or none; the upper is always the smaller when present. The scales along 
the canthus rostralis from internasals to supraoculars usually number two, rarely 
1 or 3; the posterior is the largest of the series. 

The upper preocular, which is the larger, is usually not in contact with the 
postnasal. In 82 per cent such contact is prevented by the contact of the post-can- 
thai with the loreal, in 5 per cent by the presence of a small upper loreal. 

The upper preocular is usually undivided; only in one instance is an upper 
corner cut off at the eye. The lower preocular is crescent shaped and constitutes 
the upper border of the pit. The small scales anterior to the pit usually number 3 
to 5; they are not carried forward to the rostral. 

The scale rows from labials to orbit usually number 2+3 or 2+2. 
Generally the 5th and 6th supralabials are the largest; however they do not con- 
spicuously exceed the others in size. Usually the third and fourth are in contact 
with the lower pit border. 

The first infralabials are usually undivided (only 5 per cent divided) . Nor- 
mally 4 are in contact with the genials on each side. 

Klauber— New Prairie Rattlesnake 81 

The mental is subtriangular. The genials are in a single pair, relatively short 
and obtuse, intergenials being present in 13 per cent. Also 13 per cent of the 
specimens have submentals. 

The equation for the head length of nuntius approximates H = 0.0318L+ 
7.7, where the head and body length are given in millimeters. Thus a 500 mm. 
snake would have a head length of about 23.6 mm. L /H is, of course, not a 
constant, but closely approximates 21.2 in adults. The ratio of the distance across 
the supraoculars to the space between averages 2.64 (range 2.25 to 3.13) in 86 

The ratio of the length of tail to total length, exclusive of rattle, varies from 
about 0.065 to 0.089 in the males (average 0.077), and 0.045 to 0.074 in the 
females (average 0.056) . 

The largest preserved specimen examined measured 732 mm. (29 in.). The 
average size at birth is probably 165 mm. (6| in.) . Specimens exceeding 650 
mm. (26 in.) are not common. The smallest gravid female measured 395 mm. 
(15 J in.). 

In color the typical specimens from the Little Colorado Basin are pink, 
red-brown, brown, or gray-brown. Pinks predominate about Adamana and Hol- 
brook; west of Dennison dark-brown is the typical color. Those from the vicinity 
of Moqui have an orange tinge. An occasional olive-brown specimen may be 
found in the vicinity of Winslow. Dark gray-brown specimens are found at 
Meteor Crater. The reddish hues tend to fade in preservative so that preserved 
specimens show less of this color than live material. 

The head is rather brightly marked. Supraocular light cross-dashes are always 
in evidence in well preserved material; usually these are inwardly divergent. The 
postocular light line is l| to 2 scales wide, thus being intermediate between 
typical conjluentus and oreganus. The infralabials are punctated, otherwise the 
underside of the head is immaculate. 

The body blotches number from 35 to 52, interquartile range 39.8 to 44.6, 
mean 42.2 ±0.23. The blotches are of the conjluentus confluentus type, that is, 
the edges do not follow scale outlines. Longitudinally, they are wider than the 
interspaces. In shape they are highly irregular but are usually cross-ovals, rectan- 
gles, or figure-eights. The internal edges are darker than the blotch centers and 
are sometimes black; the external edges are lighter than the ground color and are 
sometimes almost white. Secondary and tertiary blotches, while usually present, 
are ill-defined. Caudad the blotches become transverse rings and are lighter than 
the anterior blotches. 

The ventral surfaces are straw-colored. Usually the ends of the ventral 
scales are punctated, but they may be immaculate. 

The tail rings vary from 5 to 12 (usually 8 to 11) in the males (average 
9.4), and 5 to 10 (usually 6 to 8) in the females (average 7.3). The anterior 
rings are not in strong contrast to the ground color and are often ill-defined. The 
posterior rings (1 to 3) are black, in strong color contrast with the rest of the 
body, but are so poorly outlined as not to be conspicuous. The rattle matrix is 

82 San Diego Society of Natural History 

The rattles are small and delicate. The average widths of the first seven 
rattles in mm./ 10 are 43-50-60-69-73-80-82. 

The hemipenis is completely bifurcate with divided sulcus. There are about 
67 short spines on the shoulders; some are quite small so that the counting is not 
always accurate. There are no spines in the crotch. The lobes are covered with 
fringes which are laminate in front and reticulate in back, as is usual in confluen- 
tus. The fringes vary from 20 to 31; most specimens have from 25 to 29, the 
average being 27. This is distinctly lower than the confluentus average. The ratio 
of the lobe length to diameter is 2.2. The border between spines and fringes is 
sharply defined. 

The venom yield is about 38 mg. of dry purified venom per fresh adult snake. 
The fang length, measured from upper lumen to tip, of a 500 mm. snake (head 
length 23.6 mm.) will closely approximate 4.1 mm. 

Range. — Specimens of the typical stunted nuntius have been collected at 
the following points located along the line of the Santa Fe Railway or adjacent 
to U. S. Highway 66 between Canyon Padre, Coconino County, Arizona, on the 
west and Bibo, Apache County, Arizona, on the east: 

Coconino County: Navajo County: 
Canyon Padre (at U. S. 66) Winslow 

Babbitt Tank 3, 6, and 22 mi. N. of Holbrook 

7 mi. and 6 mi. W. of Two Guns ( Road to Keams Canyon) 

(on U. S. 66) Apache County: 

5 mi. E. of Canyon Padre (on Adamana 

U S - 66 > 6 mi. N. of Adamana 

Canyon Diablo (Type locality; gji 

station on Santa Fe Railway) 
Two Guns 

4 mi. NW. of Meteor Crater 

6 mi. W. of Winslow 

This territory is an arid prairie about 4800 to 5200 feet in altitude; it is 
cut by deep arroyos, of which Canyon Diablo is the most conspicuous, and buttes 
are scattered about. 

Discussion. — Having described and summarized the new subspecies, nuntius, 
two interrelated problems remain: First, the disposition of the specimens from 
those areas of Arizona outside the Canyon Padre-Bibo section, hitherto considered 
confluentus confluentus; and secondly, the relationship of the new form with the 
other confluentus subspecies. 

Nuntius is clearly a stunted offshoot of confluentus confluentus; this is 
shown by the pattern on both head and body. The characteristic arrangement of 

Klauber — New Prairie Rattlesnake 83 

the head marks, the nature of the blotch edges, the number and form of the tail 
rings, all show a close affinity to the parent form. The difference between the two 
is found in the reduction in scale counts so often seen in stunted races. This may 
be exemplified by comparing the dorsal scale rows and ventral scales of the two 
forms. For our basic confliientus conjluentus data we may use 875 specimens 
from Colorado, in which State the type specimen was collected. We have the 

Scale Rows — Per Cent Distribution 2 
29 27 25 23 21 

Conjluentus (Colo.) 3 67 30 

Nuntius 1 50 48 1 

The averages of the ventral scale counts are as follows : 

Average Ventral Scales 

Males Females 

Confliientus (Colo.) 178.84±0.10 185.73zt0.ll 

Nuntius 172.29±0.26 177.51 ±0.32 

In addition there are differences in color and size, although it is admitted 
that these characters are of less importance than scale counts, since they are more 
plastic. Co7ifluentus, from its type area, is usually green or olive-green, while 
nuntius is pink or red-brown. A large adult male conjluentus from Colorado will 
have a length of about 1000 mm.; in other parts of the range the size may exceed 
1200 mm.; nuntius seldom exceeds 650 mm. The smallest Colorado female with 
eggs (out of 149 gravid females) was 588 mm. long; the smallest nuntius (out 
of only 6 gravid specimens) was 395 mm. Thus, without doubt, there is a real 
difference in adult size in these forms, a fact further validated by rattle studies. 

Cope's pulverulentus 1 does not anticipate nuntius; the type of the former 
is a large snake with 27 scale rows, although the ventral scale count is low for 
typical conjluentus. The punctations and the number of intersupraoculars, which 
led Cope to describe this as a new subspecies, are found not to differ either in 
the type of pulverulentus, or in other specimens collected in the same area in 
New Mexico, from specimens of conjluentus taken near its type locality in Colo- 

From lutosus we find nuntius to differ in size, pattern, color, number of ven- 
trals, and number of scales on the snout before the supraoculars; and the same is 
true to a less extent in comparing abyssus and nuntius. Just as nuntius is a stunted 
form of conjluentus conjluentus , so concolor seems to be a stunted form of 
lutosus; concolor is superficially more like nuntius than is any other of the con- 
jluentus subspecies, although it is doubted whether the relationship is a direct one. 
In any case, they differ in color, pattern, and head scales, especially the number of 
scales before and between the supraoculars. 

2 Even rows are distributed equally to the next odd number above and below. 

3 Proc. Acad. Nat. Sci. Phila., 1883, p. 11. 


San Diego Society of Natural History 

Nuntius differs from Arizona oreganus in color, pattern, and head scales. 

These differences are discussed somewhat more in detail in considering the 
other specimens hitherto classified as confluentus confluentus from the area in 
northeastern Arizona surrounding the territory in which nuntius reaches its most 
typical development. Of these there are available 82 specimens from the following 

Coconino County: 

*Lee's Ferry Bridge, South Side 

*Base of Echo Cliffs, near Cedar 
Ridge Trading Post 

Havasupai Point, South Rim, 
Grand Canyon 

El Tovar, Grand Canyon 

15 mi. S. of El Tovar 

Red Butte 

22 mi. N. of Williams 
(EI Tovar Road) 


5 mi. N. of Valle 

36 mi. N. of Maine Sta. 

12 mi. N. of Deadman's Flat 

Deadman's Flat 

Medicine Valley, NE. of San 

Francisco Mt. 
Near San Francisco Mt. 
Tanner Tank 
15 and 20 mi. NE. of Flagstaff 

(Tolchaco Road) 
7 mi. NE. of Leupp 
12 mi. E. of Mouth of Moencopie 

East Foot Monument Point 

6 mi. E. of Flagstaff 
1 mi. N. of Winona 
Angel 1 

Navajo County: 

*Marsh Pass 


8, 10 mi. S. of Oraibi 
(Road to Leupp) 

Apache County: 

*Four Corners 
10 mi. NE. of Chin Lee 

8 mi. E. of Sanders 

5 mi. W. of Lupton 
40 mi. S. of Navajo 
10 mi. NE. of St. Johns 

* Of these localities, all should be considered to be within the range of nuntius except 
those marked*; those so marked are to be considered confluentus confluentus. (See map). 

Klauber — New Prairie Rattlesnake 85 

In addition the species has been observed at the Hopi villages of Hotevila, 
Oraibi, Shipaulovi, Mishongnovi, Toreva, Sichomovi, and Walpi. 

The following specimens of conjluentus confluentus or nuntius are contained 
in the U. S. National Museum: No. 5271 from Fort Buchanan, No. 8395 from 
Fort Apache, and No. 11879 from Fort Whipple. These three localities are in 
oreganus territory from which, in the last 25 years, no specimens of confluentus 
have been forthcoming. In the days when these snakes were collected it was the 
custom to label specimens with the name of the military post from which they 
were forwarded to the Smithsonian Institution. Sometimes the actual point of 
collection was some hundreds of miles away. It is therefore deemed advisable to 
omit consideration of these three specimens as being of uncertain locality. 

Summarizing the situation it may be said that we have sufficient material, 
upon which to draw conclusions, from areas to the east and west of the Canyon 
Padre-Bibo area, but not from the north. 

East of Bibo we have undoubted intergradation between nuntius and con- 
jluentus conjluentus. As we pass through Navajo, Chambers, and Houck there 
is a gradual increase in body size, and a shift in color from pink through red- 
brown and olive-brown toward green. Twenty-five dorsal scale rows becomes the 
mode and there is a moderate increase in the dorsal scale count. At Gallup, New 
Mexico, larger, olive-green specimens with 25 scale rows predominate; these are 
to be considered conjluentus conjluentus, although the ventral scale counts are 
lower than in the typical form from Colorado. 

This easterly intergradation is broad and gradual, for the habitat conditions 
change slowly; a sharp line of demarcation is not to be expected since true inter- 
grades occur over a wide territory. Merely for purposes of allocation we may 
consider that the Arizona-New Mexico line (at U. S. 66) is the approximate 
location of the boundary between the forms; thus the Arizona specimens are 
assembled with nuntius rather than coyijluentus conjluentus. 

West of Canyon Padre the situation is not so simple. First, we have, at such 
points as Angell and Winona, small snakes only slightly larger than those from 
Winslow, but distinctly darker and more brightly marked. While the majority 
are dark-brown or red-brown, a few are olive-brown. The same situation exists 
at Deadman's Flat in the area northeast of the San Francisco Peaks. All of these 
snakes have low scale counts; they are clearly nuntius, differing only in color, and 
with a slight increase in size, from the typical specimens. 

Proceeding further west we come to the Coconino Plateau, lying south of the 
Grand Canyon. A good series of specimens is available from Anita, Valle, and a 
few other scattered points round about. Here the snakes are superficially much 
more like conjluentus. They are decidedly larger than typical nuntius. Browns 
predominate, with large dorsal blotches close together and without light edges; 
olive-greens and greens are likewise present. They are much punctated. Yet with 
all these conjluentus tendencies, they are far from typical conjluentus, for the 
dorsal and ventral scale counts are as low or lower than in nuntius. Thus, in these 
all-important characters they more nearly resemble the latter and will be so class- 

86 San Diego Society of Natural History 

A few specimens are available from the south rim of the Grand Canyon; 
these show decided abyssus tendencies, particularly in high number of intersupra- 
oculars and scales on the snout. Even the specimens from as far south as Anita 
show this tendency to a slight degree. Thus, I consider these snakes to be nuntius, 
intergradation with abyssus occurring at the south rim of the Canyon. 

Also, in this Coconino Plateau area, we have the nuntius-oreganus relation- 
ship to determine, and this is the most difficult of all. A number of oreganus are 
available from the vicinity of Gleed; these show undoubted resemblances to the 
Valle nuntius; two of them might almost be considered intergrades. The Valle 
specimens present some interesting tendencies as compared with the main group 
of nuntius, particularly in color and pattern, toward these Gleed oreganus. Un- 
fortunately, no specimens have been taken between Valle and Gleed. Although 
the intervening territory is suitable to either subspecies, I do not affirm that inter- 
gradation occurs, for there are some differences in head scales which rather 
sharply divide the two. For instance, almost all the Gleed specimens have the 
prenasals separated from the supraoculars, which is not the case with the Valle 

Another uncertainty is the following: There are some areas, particularly in 
the vicinity of the San Francisco Mountains, where nuntius and oreganus have 
been taken so near to each other that an actual overlap is indicated. If this be 
the case they could hardly be expected to intergrade in the Valle-Gleed territory. 
Only the receipt of additional material can resolve this doubt. 

It may be of interest ot note that the actual and direct intergradation of ore- 
ganus and conflue?itus conjluentus, although possible in central Idaho or west- 
central New Mexico has not yet been demonstrated. It may occur through lutosus 
in southern Idaho; the lutosus-oreganus intergradation is demonstrated near the 
Oregon border, but the confluentus-lutosus merger is not. Thus, the most certain 
intergradation (as known today) of the two terminal forms, oreganus and con- 
jluentus, is that via the detour conjluentus, nuntius, abyssus, lutosus, oreganus, 
and this is not as certain as is desirable. 

Lastly there arises the question as to the classification of the prairie rattlers 
found to the north of the Little Colorado Basin. Of these unfortunately we have 
insufficient specimens to determine their position definitely. The situation is fur- 
ther complicated by the fact that the Hopi Indians use these rattlesnakes in their 
Snake Dance and sometimes have brought in specimens from distant points. On 
one occasion when I saw the dance there was a mixture of "large greens" and 
typical nuntius. In scale counts the snakes of this north area are more like conjlu- 
entus conjluentus than those from the vicinity of Gallup or the Coconino Pla- 
teau, this being especially true of the specimens from beyond the Hopi Reserva- 
tion to the north. Thus, tentatively, I am disposed to consider the snakes of the 
extreme northeastern corner of Arizona (i. e., the San Juan drainage area) as 
conjluentus conjluentus rather than nuntius. This would also seem to prevent 
direct intergradation between nuntius and concolor. A definite decision on this re- 
lationship cannot be made until more specimens are available from the San Juan 
basin, and especially along the San Juan River, from which two or three peculiar 
specimens have been seen. 

Klauber — New Prairie Rattlesnake 87 

Conclusion. — Crotalus confluentus nuntius is described as a new subspecies. 
It is a stunted form reaching its most typical development in the Winslow-Hol- 
brook area in Arizona. Eastward it intergrades with C. c. confluentus and west- 
ward, at the south rim of the Grand Canyon, with abyssus. It may intergrade with 
oreganus southwest of the Coconino Plateau. The snakes of the San Juan basin 
in Arizona are of uncertain status, but are probably C. c. confluentus. 

San Diego Society of Natural History 


Fig. 1. Comparison of adult Crotalus confluentus nuntius (left) with adult 
C. c. confluentus (right) . The former is from near Winslow, Ari- 
zona; the latter from Kansas. 

Klauber — New Prairie Rattlesnake 

Plate 8 

Fig. 1 




Volume VIII, No. 14, pp. 91-106, plates 9-16 



Steve A. Glassell 

Research Associate in Crustacea, San Diego Society of Natural History 


Printed for the Society 

August 24, 1935 


U. S. Grant, IV, Chairman 

Fred Baker Clinton G. Abbott, Editor 



Steve A. Glassell 

Research Associate in Crustacea, San Diego Society of Natural History 

The marine decapod crustacean fauna of the Gulf of California and 
neighboring Pacific waters has such a rich and varied number of species 
that it is not to be wondered at that so many new forms have been found 
there from time to time. Geographic isolation and lack of roads or facilities 
for observation has kept this territory a terra incognita, except for the 
sporadic efforts of collectors on a few expeditions whose chief interest did 
not lie primarily in the crustacean field. It is not possible for a single ex- 
pedition into these waters to make more than a reconnaissance of the lit- 
toral zones. Extreme low tides do not occur with sufficient frequency dur- 
ing daylight hours to allow an intensive study of many and varied localities 
in any one season. So it is not surprising that diligent effort is rewarded by 
prodigal returns. The following described new species bear testimony to 

During the several times I have collected there I have found a num- 
ber of obscure species in the Gulf of California of which some lacked de- 
scriptions of the opposite sex, and some were of forms the type specimen 
of which had been destroyed. I have thought it best to include a description 
of at least one already named species in this paper, in order to replace the 
lost type by a new one which could be kept in a safe repository for the 
benefit of future workers in this field. 

I am greatly indebted to Dr. Mary J. Rathbun and Dr. Waldo L. 
Schmitt, of the U. S. National Museum, for their unfailing cooperation, 
to Dr. Edith Berkeley, of the Pacific Biological Station, at Nanaimo, 
British Columbia, for her prompt determination of the worm host of the 
new Polyonyx, and to Mr. Anker Petersen, of Beverly Hills, California, 
who has made and donated, for the benefit of science and the embellish- 
ment of these pages, the splendid drawings which accompany the text. 
For their accuracy and fidelity I can vouch. 

Polyonyx quadriungulatus, n. sp. 

Plate 9 
Type. — Female, holotype; Cat. No. 750, San Diego Society of Natural 

94 San Diego Society of Natural History 

History; male, paratype; Cat. No. 751, S. D. S. N. H; one paratype, 
Cat. No. 71336, U. S. National Museum: from Estero de la Punta Banda, south 
of Ensenada, Baja California, Mexico; January 3, 1935; collected by Steve A. 
Glassell, Beverly Hills, California, in whose collection the remaining paratypes 
are located. 

Diagnosis. — Carapace ^ broader than long. Hands unequal. Carpus of cheli- 
peds f as wide as long. Dactyli of ambulatory legs armed with four unguicles on 
the inner edge. 

Description. — Carapace about 3 broader than long, appears smooth and 
shining, but has a fine transparent pubescence growing in a series of transverse 
lines, convex fore and aft, transversely ovate, regions indistinct; two subcrescentic 
pits in center of carapace, at base of gastric region; front truncate, straight, entire, 
in dorsal view. Antero-lateral margin divided into two arcs by a broad sinus 
above the base of the first antennal joint. Posterior margin concave. Chelipeds un- 
equal, dissimilar, microscopically punctate; merus stout, short, produced into a 
prominent semi-oval lamina distally and anteriorly; carpus stout, about | as wide 
as long, with a deep concavity on the under side for the reception of the posterior 
side of the manus; manus, small at juncture with carpus, increasing in breadth and 
thickness to the central part of the hand, which here forms an obtuse angle, the 
distal portion pointing outward; dense, long pubescence fringes the outer lower 
margin of manus; pollex of major hand turning outward, upward at tip, armed 
with a row of low blunt teeth, largest and highest in the center; the prehensile 
finger is curved outward, tip downward, and meshes on the outside of the tip of 
the pollex; it is armed with a cutting edge of small teeth extending from the tip 
to a large tooth in the middle; from this point to the gape, on a different axis, are 
five blunt, low teeth, the last a lobe; the minor hand has a straight finger and 
pollex, curved at the tips; the tip of the dactylus meshes on the inside of the pollex, 
the pollex being the longer; a double row of small sharp teeth on both members, 
the outside row entire, the inner row extending half way from the tip. Ambula- 
tory legs longest in the order 1-2-3; merus wide, crested on 1-2, flattened on 3, a 
row of sharp outward pointing spines on posterior margin of merus of legs 2-3; 
propodi compressed, posterior margin armed with four forward pointing spines, 
a transverse pair at distal end, followed by a single spine set a little back of these, 
also a spine near center and on the same axis as the last; dactyli with four ungui- 
cles, numbers 1-2-3 curving inward, the 4th outward; upper margin of carpus and 
both margins of propodi and dactyli fringed with hair. The telson is composed 
of seven plates. 

Sexual variation. — Males smaller than females, also darker in color. 

Color in life. — Ground color of carapace a dark brown, mottled with green 
and red, chelipeds the same, while the legs are lighter and banded. Abdomen 
mottled and opalescent. 

Measurements. — Female holotype: length of carapace 9.1 mm., width 13.5 
mm. Male paratype: length 7.9 mm., width 10 mm. Largest female (imperfect) : 
length 10.2 mm., width 15.5 mm. 

Range. — Known only from type locality. 

Glassell — Crabs from Mexico 95 

Material examined. — A series of 8 females and 5 males. 

Habitat. — This species is found commensal with the chaetopodous annelid, 
Chetopterus vanopedatus (Renier), which was found in a leathery double-ended 
tube, located at mean low water level, on an eel grass mud flat. Only the larger 
tubes were found to have crabs in them. These tubes are about a yard long by an 
inch in diameter. 

Remarks. — Related to P. nitidus Lockington, 1878, but differs in the fol- 
lowing respects: hands unequal, instead of equal; carpus of chela § as wide as 
long, instead of \ as wide; dactyli of ambulatory legs with four unguicles, instead 
of with various numbers of unguicles. I am unfamiliar with any other species of 
this genus and it seems remarkable to me that the dactyls of the chelipeds are not 
symmetrically disposed, the dactyl of the major hand overlapping on the outside 
of the pollex, while the reverse is true of the dactyl of the minor hand. 


Speloephorus schmitti, n. sp. 

Plate 10 

Type.— Female, holotype; Cat. No. 67728, U. S. National Museum: from 
San Felipe, Baja California, Mexico, low tide; May 10, 1933; collected by E. H. 
Quayle. One paratype male, one paratype female, Cat. No's. 752 and 753, S. D. 
S. N. H.; collected by S. A. Glassell, same locality, June 12, 1933; five female 
paratypes in the collection of Steve A. Glassell, Beverly Hills, California. 

Diagnosis. — Carapace subtriangular, one and a half times as broad as long. 
Hinder edge of carapace sharp, straight. 

Description. — Carapace subtriangular, approximately one and a half times 
as broad as long; surface granulate, formed by a pavement of flat and rounded, 
close-set granules. Subhepatic region prominent in dorsal view, postero-Iateral 
margin a broad arch which terminates posteriorly in a slight tooth at the begin- 
ning of the posterior hollow. Two small excrescences on the anterior branchial 
margin, a short transverse raised line above. Hepatic region high, frontal teeth 
deeply separated, a median groove on the cardiac region. Intestinal region thick, 
forming a large protuberance within the posterior cavity, not visible in dorsal 
view; the border of the cavity roughly pentagonal. Hinder edge of carapace sharp, 
straight, invisible from above. Merus of chelipeds trilobed on outer margin; car- 
pus short; hands dilated at proximal end, a straight crest on upper margin; ringers 
thin, flat, curved, grooved, fitting close together. Ambulatory legs short, merus 
lobed, carpus and propodus cristate, dactyli spinous. Enitire ventral side including 
legs and chelipeds tuberculate. 

Sexual and juvenile variations. — Abdomen of female heavily eroded. Male 
abdomen tuberculate, with a backward pointing spine at the proximal end of 
penult segment. Juvenile carapace with a high tuberculate median crest, a trans- 
verse row of four granulated lobes, two branchial, two epibranchial, a deep trans- 
verse sulcus posterior to these lobes. In the very small specimens the posterior 
border is visible in a dorsal view. 

96 San Diego Society of Natural History 

Color in life. — Carapace a salmon pink, blotched with white, ventral side 
and legs a muddy white. 

Measurements. — Female holotype: length of carapace 27 mm., width 36.8 
mm. Male paratype: length 27.4 mm., width 36.5 mm. 

Range. — Upper end of Gulf of California, Mexico. 

Material examined. — Angeles Bay, Baja California, January 4, 1932, by 
S. A. Glassell: two females and one male, juveniles. San Felipe, Baja California, 
May and June, 1933, by E. H. Quayle and S. A. Glassell: two females (one the 
holotype), and one male. San Felipe, Baja California, May 28, 1934, by S. A. 
Glassell: two females. 

Habitat. — Found at low tide, under dense sea lettuce and weeds among 

Remarks. — This species is dedicated to Dr. Waldo L. Schmitt, of the U. S. 
National Museum, to whom I am indebted for assistance and counsel. 


Panoplax mundata, n. sp. 

Plate 1 1 

Type. — Male, holotype; Cat. No. 754, San Diego Society of Natural His- 
tory; female, paratype; Cat. No. 755, S. D. S. N. H; one paratype, Cat. No. 
71337, U. S. National Museum: from San Felipe, Baja California, Mexico, near 
upper end of Gulf of California; June 2, 1934; collected by Steve A. Glassell, 
Beverly Hills, California, in whose collection the remaining paratypes are located. 

Diagnosis. — Carapace convex anteriorly and posteriorly. Front bilobed, 
truncate, straight. Sixth segment of male abdomen broader than base of seventh. 
Antero-lateral borders converging slightly posteriorly. 

Description. — Carapace slightly depressed at cardiac region, convex anter- 
iorly and posteriorly; transversely flat at fourth marginal tooth; regions fairly 
well marked, surface finely punctate, granulate on outer regions. Front truncate, 
divided into two straight margined lobes by a sharp V shaped median notch. 
Width of orbit and frontal lobe subequal. Five lateral teeth including the orbital 
tooth; the second shallow, separated from the first by a shallow sinus; the third 
large, blunt, right-angled, pointing forward and slightly upward; the fourth 
sharper, right-angled, with anterior margin short and nearly transverse to the 
carapace; the fifth small, not projecting beyond the general outline. Postero- 
lateral margins moderately converging behind. Hinder edge of carapace straight. 
Merus stout, dentate on upper margin. Carpus oblong, with a blunt tooth at inner 
angle, granulate on upper surface; a slight anterior transverse groove. Hands un- 
equal, smooth and rounded; a row of hairs on upper crest of hand and finger; a 
lateral groove from near tip of pollex, paralleling lower outer margin; prehensile 
teeth broad, low, a large one at base of dactylus on major hand; no gape; a dark 
brown patch of color on pollex, extending slightly on palm of male; fingers brown, 
curved at tips. Ambulatory legs long, hairy on margins, merus slightly com- 
pressed; dactyli long, lanceolate, hairy to tip. 

Glassell — Crabs from Mexico 97 

Sexual variation. — Slight. Color on pollex of female not continued on palm. 

Color in life. — Ground color of carapace pinkish cream, overlaid with a few 
reddish orange spots, irregularly broadcast. Chelipeds,merus and carpus same color 
as carapace; fingers a deep dark brown, tips white. Ambulatory legs a yellowish 
cream, with a few reddish-orange spots; tips of dactylus amber. Ventral side same 
as dorsal, but without spots. 

Measurements. — Male holotype: length of carapace 4.6 mm., width 6 mm. 
Female paratype, length 4.9 mm., width 6.5 mm. 

Range. — Known only from the type locality. 

Material examined. — A series of over fifty specimens of both sexes, many 

Habitat.— This species inhabits a soft mud bottom, covered by five to seven 
fathoms of muddy water. 

Remarks. — The fifth tooth of the carapace is hard to distinguish in any but 
the largest specimens, and even then the pubescence will have to be removed to 
show as in the plate. This species is the Pacific analogue of Panoplax depressa 
Stimpson, 1871, but differs from the species in the squarer carapace, straighter 
front, smaller size, and lack of distinct epigastric lobes. 


Pinnotheres clavapedatus, n. sp. 

Plates 12, 13 

Type.— Female, holotype; Cat. No. 756, San Diego Society of Natural 
History: from San Felipe, Baja California, Mexico, low tide; June 1, 1934; col- 
lected by Steve A. Glassell. One paratype, male, Cat. No. 757, S. D. S. N. H.; 
one female and one male, paratypes, Cat. No's. 71338 and 71333, U. S. National 
Museum; three males, sixty females, paratypes, in the collection of Steve A. Glas- 
sell, Beverly Hills, California. 

Diagnosis. — First ambulatory leg stout, propodus greatly dilated at distal 
end, clavate, second leg similar but not so stout, third and fourth legs slender. 
Dactylus of outer maxilliped narrow-spatulate, slightly curved, attached near mid- 
dle of propodus and reaching \ its length past the extremity of the latter. 

Description of female. — Carapace smooth, shining, semi-hard, convex in 
both directions, eyes not visible in dorsal view; regions not denned; posterior mar- 
gin concave. Merus of outer maxillipeds widest at distal f , narrowing at distal 
end; propodus longer than carpus; dactylus narrow-spatulate, slightly curved, 
attached near middle of propodus and reaching \ its length past the latter mem- 
ber. Hands cylindrical, similar, increasing in width to base of prehensile finger, 
smooth with exception of a row of forward-pointing hairs on inner side of pollex, 
extending back to a point under the gape; inner tip of dactylus hairy. First and 
second legs stout, smooth, with propodi cylindrical and distended at distal ends; 
first leg much the larger, dactylus short, stout, straight on under side, arched on 
upper, with needle tip; second dactylus longer and heavier; third and fourth legs 

98 San Diego Society of Natural History 

very slender; dactylus of third leg long, slender, slightly curved; dactylus of fourth 
leg arched on the anterior margin, larger at proximal third than at base, a row of 
hairs on the inner margin, as has the propodus of this leg at the distal inner half. 
Abdomen unusually large, even to covering the mouth parts when non-ovigerous. 

Description of male. — Much smaller, width of carapace less than \ that 
of female. Carapace suboctagonal, lightly sculptured, finely pubescent, posterior 
margin straight. Cardiac region transversely ovate, surrounded by a groove except 
posteriorly; branchial and gastric regions grooved; a light median sulcus. Front in 
dorsal view advanced, arcuate, divided; in front view, the front is deflexed and 
pointed. Orbits oval, eyestalks stout, filling the hiatus. Antennae short. Chelipeds 
stout, pubescent; carpus short; hands thick; palm with upper surface concave, inner 
proximal end distended, lower margin of pollex convex from end to end, tip fal- 
cate, armed with a tooth and a groove immediately behind it, fingers heavy, gap- 
ing when closed, heavily curved at tip, armed with a prominent tooth which 
meshes into the groove of the pollex. Ambulatory legs similar, slender, slightly 
pubescent; merus long, cylindrical; propodus convex on both margins; dactylus 
falcate, slight, long and sharp; carpus and propodus of second and third legs with 
long fringes of hair on posterior surface. Abdomen with first segment wide at 
base, widest at third segment, sides converging to tip, seventh segment semi-oval, 
slightly longer than wide. 

Color in life. — The female carapace is a light cream, with an orange red de- 
sign showing through the cardiac region, and extending on to the branchials. Male, 
a light brown. 

Measurements. — Female holotype: length of carapace 7.6 mm., width 12.4 
mm. Male paratype: length of carapace 2.33 mm., width 2.5 mm. 

Range. — From Magdalena Bay, Baja California, Mexico, to the head of the 
Gulf of California. 

Material examined. — A series of over two hundred females, taken through- 
out its range, and five males taken at San Felipe, Baja California, Mexico. 

Habitat. — Found commensal in the boring mollusk, Lithophaga attenuata 
(Deshayes), which I have taken from a depth of 15 fathoms to just below the 
inter-tidal zone. 

Remarks- — This species closely resembles Pinnotheres lithodomi Smith, 
1870, which was described from a damaged juvenile specimen, but differs from 
that species in the following particulars: Merus of outer maxillipeds convex on 
anterior margin, convex on distal posterior margin, and thence concave to base, 
instead of broadest at distal extremity, sides nearly straight; first and second am- 
bulatory legs stout, clavate at distal end of propodus, instead of slender. The dis- 
tinctive characters of this new species, especially the club-shaped first propodus, 
which is a striking feature even in juveniles, could hardly have missed the critical 
eye of such a carcinologist as S. I. Smith, and since he makes no mention of them 
I am convinced his species was not the one here described as new. 

Glassell — Crabs from Mexico 

Plate 9 


■-■■ --_ ■ ? •-'-> ^ 






,_iiii**»* > ' 

' '" .^...iw**-- 


Polyonyx quadriungulatus, n. sp. 

Fig. 1. Female holotype, dorsal view. Fig. 4. Propodus and dactyl of ambulatory leg. 

Fig. 2. Female third maxilliped. Fig. 5. Major chela. 

Fig. 3. Female telson of abdomen. Fig. 6. Minor chela. 

Glassell — Crabs from Mexico 

Plate 10 


:■ ■ 


6 § j 






lO rrvm. . 


— si »• 




2£ V. 



¥ - f , -^ 

Speloephorus schmitti, n. sp. 

Fig. I. Male paratype, dorsal view. Fig. 2. Male paratype, ventral view. 

Glassell — Crabs from Mexico 

Plate 11 

,y. 1 1 

\ i 

W ... 


Panoplax mundata, n. sp. 

Fig. 1. Male liolotype, dorsal view. Fig. 3. Outer maxilliped. 

Fig. 2. Major chela without hair. Fig. 4. Female abdomen. 

Fig. S. Male abdomen. 

Glassell — Crabs from Mexico 

Plate 12 


2&*E - ^ ■■-V 

-— m; 



V 1 

Pinnotheres clavapedatus, n. sp. 

Fig. 1. Male paracype, dorsal view. Fig. 3. Male right chela, dorsal view. 

Fig. 2. Male right chela. Fig. 4. Male antennal and buccal area. 

Fig. J. Male abdomen. 

Glassell — Crabs from Mexico 

Plate 13 

Pinnotheres clavapedatus, n. sp. 

Fig. 1. Female holotype, dorsal view. Fig. 3. Female, ambulatory legs. 

Fig. 2. Female, right chela. Fig. 4. Female, third maxilliped. 

Fig. 5. Female, antcnnal and buccal area. 

Glassell — Crabs from Mexico 

Plate 14 

Pinnotheres angelicus Lockington, female. 
Fig. 1. Female neotype, dorsal view. Fig. 3. Female antennal and buccal area. 

Fig. 2. Female chela. Fig. 4. Female outer maxilliped. 

Fig. 5. Female frontal-ventral view. 

Glassell — Crabs from Mexico 

Plate 15 

Pinnotheres angelicus Lockington, male. 

Fig. 1. Male allotype, dorsal view. Fig. 4. Male chela, dorsal view. 

Fig. 2. Male chela. Fig. S. Male antenna! and buccal area. 

Fig. 3. Male abdomen. Fig. 6. Male chela, front view of palm 

Glassell— Crabs from Mexico 

Plate 16 

Dissodactylus lockingtoni, n. sp. 

Fig. 4. Female, right chela. 
Fig. S. Male abdomen. 
Fig. 6. Female abdomen. 

Fig. 1 . Female, dorsal view. 

I ig. 2. Female, third maxilliped. 

Fig. 3. Female, first ambulatory leg. 

Glassell — Crabs from Mexico 99 

Pinnotheres angelicus Lockington 
Plates 14, 15 

Pinnotheres angelica Lockington, Proc. California Acad. Sci., vol. 7, 1876 
(1877), p. 155 [10] (type-locality, Angeles Bay, Gulf of California, in 
oysters; type not extant) . Not P. angelicus Miers, Journ. Linn. Soc. London, 
Zool., vol. 15, 1880. (Rathbun). 

Pinnotheres angelicus Lockington, (Rathbun), Bull. U. S. National Museum, 
No. 97, Jan. 25, 1918, pp. 72-73, pi. 16, figs. 5-6, text fig. No. 34. 

This species was found on the Gulf coast of Baja Califorina, Mexico, at 
Angeles Bay, and was described by Lockington in 1877. In 1906 the female holo- 
type and the female paratypes were destroyed in the San Francisco fire. No males 
were in the original collection. 

In January 1932 at Angeles Bay, and in June 1933, at San Felipe, on the 
Gulf coast of Baja California, Mexico, I took a large series of the females and a 
few males. From the Angeles Bay material I am designating one female the neo- 
type and one male the allotype. 

Neotype. — Female; No. 758, collection of the San Diego Society of Natural 
History: from Angeles Bay, Baja California, Mexico; January 4, 1932; collected 
by Steve A. Glassell. 

"Diagnosis. — Female transverse, smooth, shining. Dactylus of second leg 
much the longest. Prominent tubercle on basal joint of antennae. Dactylus of 
endognath attached to end of propodus. 

"Description of female. — Smooth, shining. Carapace thin, easily wrinkled, 
transverse, with anterior margin strongly arcuate, posterior margin long, slightly 
concave, sides rounded; gastric region well defined; a large pit on branchial region 
near inner angle. Front advanced, edge rounded. Orbits and eyes oval, hidden 
from dorsal view. A large prominent tubercle at posterior end of basal joint of an- 
tenna. Propodus of endognath distally rounded, dactylus small, attached on inner 
portion of extremity of propodus. Chelipeds elongate, manus slightly compressed 
and increasing distally; immovable finger slightly deflexed, swollen in basal half: 
fingers fitting together when closed, tips curved and crossing each other. Legs 
slender, second longest, third next, fourth shortest; dactyli nearly straight, except 
that of second leg, which is twice as long as of other legs, and nearly equaling its 
propodus. Abdomen unusually large." (Rathbun). 

Allotype. — Male; No. 759, collection of the San Diego Society of Natural 
History: from Angeles Bay, Baja California, Mexico; January 4, 1932; collected 
by Steve A. Glassell. 

Description of male allotype. — Much smaller than female. Carapace flat, 
suborbicular, with the front advanced in a triangle, tip rounding downward to a 
blunt point, not observable in dorsal view; surface hard, lightly pubescent, punc- 
tate; cardiac region faintly defined. Eye stalks large and stout, diminishing from 
base to tip; pubescent on upper forward surface; cornea large. A small tubercle 
at posterior end of basal joint of antenna. Chelipeds stout, similar, slightly pubes- 

100 San Diego Society of Natural History 

cent, palm smooth, swollen in basal half, decreasing distal ly; exterior of manus 
crossed longitudinally by a granulate ridge. Pollex short, stout, hooked at tip; a 
large ridge-like tooth occupying entire central portion; a deep notch at gape. Dac- 
tyli long and curved at tip, armed with two well-developed teeth at proximal end, 
the distal one the larger; fingers fitting together when closed, curved tips crossing 
each other; a row of hairs at bottom margin of hand. Legs slender, second and 
third subequal, first shorter, fourth shortest. Dactyls long and straight, with tips 
hooked, acuminate. Popodus of last three legs crested with setae. Lower distal 
edges of carpus of second and third legs with setae. Abdomen widest at third seg- 
ment which is almost three-quarters of entire length; gradually narrowing from 
third to seventh segment, which is obtusely rounded. Abdomen and sternum 

Color in life. — Carapace and chelipeds of male a deep chocolate brown; am- 
bulatory legs much lighter. Carapace and chelipeds of female a deep chocolate 
brown; ambulatory legs a light cream color. 

Measurements. — Female neotype: carapace 8 mm. long, by 1 1 mm. wide. 
Male allotype: carapace 2.7 mm. long, by 2.8 mm. wide. 

Range. — Gulf of California, Mexico, from upper to lower end. 

Habitat. — Found in the mantle cavity of a small plicated oyster which is 
attached to rocks or mangrove roots, this oyster may prove to be either Ostrea 
cumingiana Dunker, or O. amara Carpenter. It is also reported in the mussel 
Modiolus capax Conrad, a series of which I examined at San Felipe, Baja Cali- 
fornia, Mexico, but without results. The male is undoubtedly free swimming and 
its capture at any time is highly problematical. Out of several hundred specimens 
examined, a series of only six or eight males was taken. 

Dissodactylus lockingtoni, n. sp. 

Plate 16 

Type. — Female, holotype; Cat. No. 760, San Diego Society of Natural His- 
tory; from Punta Penasco (Rocky Point) , Sonora, Mexico, low tide; May 3, 
1935; collected by Steve A. Glassell. One paratype, male, Cat. No. 761, S. D. S. 
N. H.; one paratype female, juvenile, Cat. No. 71339, U. S. National Museum; 
25 of both sexes, paratypes, in the collection of Steve A. Glassell, Beverly Hills, 

Diagnosis. — Carapace convex. Dorsal ridge slightly oblique to postero- 
lateral border. Upper border of merus-ischium of outer maxilliped serrate; palp 
three-jointed. Last segment of male abdomen semi-oval. 

Description. — Carapace convex in both directions, high in middle, minutely 
pubescent; regions ill denned, almost smooth. Front concave, slightly produced. 
Anterolateral borders arcuate, sinuous, with a milled margin starting at the cervi- 
cal groove, which at the lateral angles, extends obliquely inward on the carapace; 
postero-Iateral borders with a nearly straight rim; posterior margin sinuous. 
Merus of outer maxillipeds suboblong, upper crest serrated; lower distal angles 
rounded; segments of palp long; second segment spatulate, distal end broad and 

Glassell — Crabs from Mexico 101 

squarely truncate; third segment small, located at the distal inner angle of the sec- 
ond segment. Merus of chelipeds short, not extending far beyond carapace; carpus 
short, minutely granulate; hands similar, suboblong, granulate, faint ridges on 
inner and outer sides, a pubescent fringe on inner lower margin of palm, extend- 
ing to pollex. Pollex horizontal, unarmed. Prehensile finger longer than pollex, 
curved, granulate, and armed with three or more small teeth, increasing in size 
towad the gape. Legs slightly hairy, short, stout; dactyli of first three legs smooth, 
naked, divided at proximal third, secondary tip short, bifid; dactylus of fourth 
simple, long, lanceolate, slightly curved at tip, lower margin with long hair to 
distal third. 

Sexual variation. — The female resembles the male. The female abdomen 
covers the sternum, is suboblong, with seven segments; seventh segment broadly 
triangular, height less than £ the width. Male abdomen, first and second segments 
narrower than third, fused; third to sixth fused, expanded at base, contracted at 
distal end; first to sixth coffin-shaped; seventh segment semi-oval, height a little 
more than \ the width. 

Color in life. — Male, bluish-white, with fingers of chelipeds yellowish. Female, 
carapace pigmented on bluish-white ground with minute dark brownish specks, 
yellow of internal organs showing through. Ambulatory legs specked with dark 
spots. Row of black hairs on anterior margin of sternum, in both sexes, partly 
covering the mouth parts. 

Measurements. — Female holotype: length of carapace 7 mm., width 7:5 mm. 
Male paratype: length of carapace 5.5 mm., width 5.8 mm. 

Range. — Found both at San Felipe, Baja California, Mexico, and at Punta 
Penasco (Rocky Point), Sonora, Mexico, but undoubtedly ranges throughout 
the Gulf of California. 

Habitat. — Commensal on the ventral exterior surface of the following 
Echinoids: Mellita longifssa Michelin, Encope micropora Agassiz, E. grandis 
Agassiz and E. californica Verrill. It is usually found, when on the Encope, 
located in the proximal portion of the posterior interambulacral Iunule. From this 
position to a point near the peristome or periproct of the echnoid, the crab clears 
the actinal spines, thus forming for itself a roadway but little wider than its out- 
stretched ambulatory legs. D. nitidus Smith, may also occupy the same echnoid 
with D. lockingtoni, but the former ranges over the entire ventral surface and 
has no fixed place of abode. 

Affinity. — Related to D. nitidus Smith, 1870, which it somewhat resem- 
bles, but differs in the following respects: carapace convex, instead of compressed; 
crest of outer maxillipeds serrate, instead of smooth; terminal segment of male 
abdomen semi-oval, instead of triangular; color bluish-white, instead of purplish 

Remarks. — This species is dedicated to Mr. W. N. Lockington, who did 
such splendid work in West American carcinology during the 1870's and late 
1880's, both as a collector and systematist, while with the California Academy of 
Sciences, at San Francisco, California. 

102 San Diego Society of Natural History 

Pinnixa plectrophoros, n. sp. 

Type.— Male; Cat. No. 762, San Diego Society of Natural History: from 
Punta Pefiasco (Rocky Point), Sonora, Mexico; May 1, 1935; collected by Steve 
A. Glassell. 

Description. — Allied to P. retinens. 1 Carapace three times as wide as long; 
nearly flat except toward margins, where it slopes gradually downward at lateral 
angles, abruptly to posterior margin. Antero-lateral margin feebly indicated by 
minute granules. Orbits of eyes, the eyes and sub-hepatic regions visible in dorsal 
view. Front not projecting, bilobed, a median sulcus. Two transverse lunate pits 
at cardiac region; a pit in line with these on mesobranchial region; other regions 
ill defined. Chelipeds weak, hairy, similar; carpus smooth, with pubescent mar- 
gins; hands weak, margins parallel, compressed; pollex in line with lower margin 
of hand, uncolored, microscopically denticulate; dactyl curved, crested with hair, 
closely fitted to pollex. First and second legs slight, dactyli long, lanceolate; third 
leg very heavy, dactyl triangular, with two spines near base of anterior side, a 
posterior propodal spine, a large outward curving spur at distal posterior end of 
merus, a small blunt spine at its base, two ischial spines; fourth leg very short, 
slight, hairy, not reaching past merus of third leg, with two ischial spines and a 
spine on the proximal lower margin of the merus. Male abdomen, first and sec- 
ond segments free, third to sixth fused, seventh segment as in P. transversalis, 2 
female abdomen with seven segments, covering sternum. 

Measurements. — Length of carapace 2 mm., width 6 mm. 

Habitat. — Commensal in the sand tube of a species of annelid worm 
(Clymenella) . 

Remarks. — Fuller descriptions and plates of this and the two following 
species will be included in a forthcoming partial revision of the genus Pinnixa by 
the writer. 

Pinnixa pembertoni, n. sp. 

Type. — Male; Cat. No. 763, San Diego Society of Natural History: from 
San Felipe, Baja California, Mexico; June 19, 1935; collected by Steve A. Glas- 

Description. — Allied to P. \loridana? Carapace twice as wide as long, con- 
vex posteriorly, punctate, sharply rounding at the angles and to the posterior mar- 
gin. Antero-lateral angle forming a shoulder, the side walls steep and tapering 
outwardly. Antero-lateral border with a fine granulate ridge. Cardiac and gastric 
regions slightly raised, a sulcus between, a shallow depression on each side. Front 
truncate, slightly projecting, pubescent. Eyes large, filling the orbits. Chelipeds 
similar, strong; merus short, pubescent; carpus smooth on upper surface, mar- 
gined with pubescence; hands smooth on inner palm, smooth on outer side, with 
a row of fine hair on upper and lower margins, a longitudinal row from gape to 

i Bull. 97, U. S. Nat. Mus., 1918, p. 139, pi. 41, figs. 1-2, text figs. 83-84. 

2 Bull. 97, U. S. Nat. Mus., 1918. p. 132, text figs. 75-76. 

3 Bull. 97, U. S. Nat. Mus., 1918, p. 138, pi. 30, figs. 4-7, text fig. 82. 

Glassell — Crabs from Mexico 103 

carpus; lower margin sinuous, pollex convex, distal end upturned, armed with three 
or more triangular blunt teeth; dactyl curved at tip, armed with two or more blunt 
teeth in center, crested with fine hair, tip not crossing pollex; gape pubescent, open 
to tip of fingers. First and second legs slight, merus trihedral, dactyli long, slim, 
straight, lanceolate; third leg heavy, dactyl long, heavy, straight, pubescent, pos- 
terior margin of leg heavily pubescent; fourth leg similar but much smaller, dac- 
tyl reaching slightly past merus of third leg, pubescent on both margins. Terminal 
segment of palp of outer maxilliped reaches nearly to base of ischium. Abdomen 
widest at third segment; fourth, fifth and sixth segments fused, sides sinuous, 
narrowest at sixth, seventh broader than sixth, broadly rounded at sides, tip 

Measurements. — Length of carapace 3.8 mm., width 7.6 mm. 

Habitat. — Commensal with a species of the "lug-worm" (Arenicola) . 

Remarks- — Dedicated to Mr. J. R. Pemberton, of Los Angeles, California, 
whose generous constructive criticism has been greatly appreciated by the writer. 

Pinnixa huffmani, n. sp. 

Type.— Female; Cat. No. 764, San Diego Society of Natural History: from 
Punta Penasco (Rocky Point), Sonora Mexico; May 4, 1935; collected by Steve 
A. Glassell. 

Description. — Allied to P. barnharhf' Carapace little wider than long, very 
convex in both directions, thin, not firm, internal organs showing through, punc- 
tate on posterior half, smooth on anterior portion. A faint antero-lateral margin. 
Sub-hepatic region prominent in dorsal view. Front a wide arc, not projecting. A 
transverse, narrow sulcus across entire carapace at cardiac-gastric regions. Regions 
not defined. Eyes not large, cornea small, red pigment. Chelipeds similar, equal; 
merus long, hairy; carpus long, rounded, smooth; hands very long, compressed, 
margins sub-parallel, curving inward at distal end, upper margin highest over dac- 
tyl, crested with hair, outside of palm smooth; pollex slightly depressed, triangu- 
lar, sharp pointed, armed with cutting edge; dactyl very heavy at gape, curved 
downward to a sharp pointed tip, armed with a large tooth at proximal side of 
middle, very slightly gaping; the fingers when closed cross the tips of their respec- 
tive pollices on opposite sides from each other. Legs similar, hairy on margins, 
not compressed, third longest; dactyli long, nearly straight, curved slightly at tip, 
which is very sharp, corneous; fourth leg long, reaching to propodus of third; 
dactyl straight. Abdomen semi-globular, punctate, covering sternum and part of 
maxillipeds. Second and third segments of palp of outer maxillipeds very large, 
third the longer. 

Measurements. — Length of carapace 6.1 mm., width 7.6 mm. 

Habitat. — Commensal in a species of the sea cucumber CThyone) . 

Remarks. — Dedicated to Mr. Earl C. Huffman, of Pasadena, California, 
conchologist, and a companion on many a mile of mud and sand. 

4 Bull. 97, U. S. Nat. Mus., 1918, p. 149, pi. 32, text fig. 91. 

104 San Diego Society of Natural History 

Ozius tenuidactylos (Lockington) , corrected name 

Ozius agassizii A. Milne Edwards, should be known as Ozius tenuidactylos 
(Lockington) . A careful study of Lockington's original description of this species 
(Proc. California Acad. Sci., vol. 7, 1876 [1877], p. 98 [4] : type-locality, La 
Paz, Lower California; type not extant) , under the name Xantho tenuidactylos 
nov. sp., and allowance for the obvious reversing of his length and width measure- 
ments, shows that Milne Edwards' Ozius agassizii (Crust. Reg. Mex., 1880, p. 
279, pi. 55, figs. 1-ld: type-locality, Panama; type in M. C. Z.), must be conspe- 
cific. Hence agassizii being of later date must be suppressed as a synonym. 

Glassell — Crabs from Mexico 


Extension of Range and New Locality Records 

Fabia grand Glassell 


Lophopanopeus bellus (Stimpson) 

San Pedro, California 
Lophopanopeus bellus (Stimpson) variety 

San Pedro, California 

Taken at low tide 
Glyptoxanthus meandricus (Lockington) 

Punta Penasco, Sonora, Mex. 
Pilumnus limosus Smith 

Punta Penasco, Sonora, Mex. 


Pachycheles holosericus Schmitt 

Ensenada, Baja Calif.. Mex. 
Pachycheles pubescens Holmes 

Ensenada, Baja Calif., Mex. 


June 1, '34 2 $ juv. 

10 5, 10 2 

3 5,35 

6 5 juv., 6 9 juv. 

San Felipe, Baja, Calif., Mex. June 1, '34 

Host, Acmaea mesoleuca Menke 

San Felipe, Baja Calif., Mex. June 19, '35 2 9 

Host, Crucibulum spinosum (Sowerby) 

Punta Penasco, Sonora, Mex. May 3, '35 1 9 

Host, Crepidula nieva C. B. Adams 
Opisthopus transversus Rathbun 

San Felipe, Baja Calif., Mex. 
Pinmxa transversalis (M. Edw. & Lucas) 

San Felipe, Baja Calif., Mex. June 1, '34 

Punta Penasco, Sonora, Mex. May 2, '35 

Pinnixa occidentalis Rathbun 

San Felipe, Baja Calif., Mex. June 1, '34 

Pinmxa longipes (Lockington) 

Ensenada, Baja Calif., Mex. Jan. 2, '35 15,15 

Pinnotheres reticulatus Rathbun 

San Felipe, Baja Calif., Mex. June 19, '35 12 9 

Host, Tagelus affinis C. B. Adams 

Host, Paphia grata Sowerby 


Pugettia producta (Randall) 

Ensenada, Baja Calif., Mex. 


Portunus (Portunus) xantus'u (Stimpson) 
Santa Barbara, California 

Jan. 2, '35 1 $ 

Mar. 17, '33 1 5 

Jan. 8, 



Jan. 8, 



May 2, 


10 5,10 9 

May 1, 



10 5 , 10 9 

Jan. 3, 



Jan. 3, 


2 5 juv., 3 9 juv. 




Vol. VIII, No. 15, pp. 107-1 14, text figs. 1-6 August 24, 1935 



Roland Case Ross 

Los A ngeles City Schools 

Among the Pleistocene avian remains found by the California Insti- 
tute of Technology in the asphalts of McKittrick are fourteen (complete 
and partial) tarsometatarsal elements of a goose more slender than any 
of the living Anserinae. Without attempting at this time to establish the 
identity of the form on the basis of other elements, it is here proposed to 
describe and record a new genus and species of goose on tarsal characters 
alone, the tarsus being a characteristic avian element, and in this instance 
well preserved, numerous and clearly generically distinct. 


Thanks are due Dr. Loye Holmes Miller for suggesting the problem 
and for use of material. 

Mr. John L. Ridgway, scientific illustrator of the California Institute 
of Technology, is acknowledged gratefully for the preparation of text 

Dr. Chester Stock, in whose laboratory this study has been made, is 
thanked for continued encouragement and guidance. 

Anabernicula gracilenta, n. gen. and n. sp. 
Type. — No. 1169 Calif. Inst. Tech. Vert. Pale. Coll., left tarsometatarsus of 

1 Contribution No. 171. Balch Graduate School of the Geological Sciences, California 
Institute of Technology. 

108 San Diego Society of Natural History 

mature Anserine bird, complete and unworn, from the brea deposits of McKit- 
trick, Calif., Pleistocene age. 

Paratypes. — No. 1168, right tarsometatarsus, complete except for extended 
portions of hypotarsus, length 60.3 mm.; and No. 1170, left tarsometatarsus, 
proximal three-fifths 

Description. — Goose-like in general structure and proportions. Small and 
slender with delicately tapered shaft, broad spread of distal condyles, and dimin- 
ished tarsal articulation. 

Inner cotyla deeply cupped and of noticeably slight extent an tero posteriorly. 1 

Hypotarsus in lateral aspect rectangular, of relatively slight vertical extent, 
markedly strong in plantar depth. Inferior terminus of inner calcaneal ridge 
abrupt, with slight gradient into shaft. 

Intermuscular line of the posterointernal border of shaft deflected inwardly 
toward the proximal end, effecting thus a disjunction with the inner calcaneal 
ridge. Extensor groove flattens out in its oblique descent across inner surface of 
shaft and weakens intermuscular line at junction. 

Measurements of Type in Millimeters 

Total length over all 61.8 

Minimum shaft width (transverse) 4.4 

Minimum shaft depth (anteroposterior) 3.4 

Minimum shaft area, cross section (sq. mm.) 14.96 

Transverse width proximal end 11.6 

Anteroposterior depth proximal end (inner cotyla) 5.5 

Anteroposterior depth proximal end including hypotarsus 11.0 

Anteroposterior depth first calcaneal ridge 5.5 

Transverse (total) extent of distal end (condyles 2, 3, 4) 10.4 

Transverse extent of condyles 3 and 4 8.4 

Minimum transverse width of condyle 2 2.9 

Maximum transverse width of condyle 3 5.0 

Mean transverse width of condyle 4 3.6 

Anteroposterior depth of condyle 2 5.6 

Anteroposterior depth of condyle 3 7.2 

Anteroposterior depth of condyle 4 7.1 

Minimum transverse width of prong shaft to condyle 4, taken from 

distal foramen to outer border 3.2 

Generic Characters of Anabernicula 
Although assignable only to the subfamily Anserinae or true geese, 
there are characters consistently present in the fossil tarsal specimens that 
relate to ducks and tree-ducks, Anatinae and Dendrocygninae respective- 
ly. Chief among these are the square-cornered outline of the hypotarsus as 
viewed from the inner side, its undercut effect at the distal confluence with 

- Anteroposterior is the rotular-plantar axis. 

Ross — New Pigmy Goose 109 

shaft, and its marked plantar extent. Fig. 1A shows these distinctions in 
contrast to the characters seen in species of living pigmy geese (fig.l, B, 
C, D). Of 24 races of geese studied, none displays the above characters, 
while ducks and tree-ducks both possess them. 

The deflection of the intermuscular line internolateral of the hypo- 
tarsus is a strong dendrocygnine structural feature, as is also the strap-like 
groove for the muscle extensor hallucis breris upon the inner side of the 
shaft. In the Anserinae the groove is fine or cord-like as opposed to ribbon 
or strap-like in tree-ducks. This character in the fossil bird is intermediate, 
being fine-grooved proximally but widened distally on the approach to the 
intermuscular line. An unique feature is the flattening or weakening 
effect this has on the intermuscular line at this junction. This effect has not 
been noted in ducks (excepting the Muscovy Duck) or tree-ducks, and is 
only weakly if at all defined in geese 

Proximal view of the head (fig. 1A upper) shows a more restricted 
articulation area as compared with that in other geese (actual and rela- 
tive), an excessive development of the first calcaneal ridge anteroposteri- 
orly, and a small circular deeply cupped inner cotyla of limited anteropos- 
terior diameter. 

The above structural features deserve generic distinction, while the 
following goose-like characters maintain the status of the new group 
within the Anserinae : The tapering and proportions of the shaft elimin- 
ate Cygninae, Anatinae, Dendrocygninae and are typical of Anserinae. 
The swelling of shaft into condylar prongs, the spacing, and the total 
spread of the distal condyles are separable by detail or proportion from 
all others but the Anserinae. 

Specific Characters of Anabernicula gracilenta 

Anabernkula gracilenta as a species can be "hand picked" from min- 
gled tarsi of other pigmy geese by an appearance of delicacy and dwarf- 
ness. The ranges in size of its component parts as given below are consid- 
ered fair delimitation for the species, inasmuch as the number of fossil 
specimens equaled or surpassed Recent specimens of each species studied. 
Both mature and immature individuals were present. 

The seventeen fossil specimens of tarsometatarsus fall into a very con- 
sistent clustered group when measured in certain ways. Various units of 
study and ratios between measurements give specific and even generic dis- 
tinctness to the fossil group. In the living pigmy geese no such sharpness 

110 San Diego Society of Natural History 

between groups and rarely any tendency to generic separation has 
appeared in the numerous measurements and ratios exacted of their tarsi. 
There are, however, size and proportional figures that yield no dis- 
tinctions between Anabernicula gracilenta and the four pigmy races of 
native geese. Such characters are frequently of subfamily rank, inasmuch 
as they unite the goose group and serve to distinguish it from Anatinae, 
Cygninae, and Dendrocygninae. 

Life Characteristics 

This small goose outnumbers several times over the individuals of 
other geese occurring in the McKittrick collections of the California In- 
stitute. The larger geese are noticeably infrequent in this and other col- 
lections of McKittrick material. The relative abundance of the pigmy 
goose would seem to indicate special attractiveness of the locality as af- 
forded perhaps by shallow ponds and mud flats. 

Anabernicula gracilenta possesses more slender proportions than any 
known goose, and is smaller in bulk and weight than any goose living with 
the exception of Chenonetta of Australia. 

The resemblance of the fossil goose to Branta bernicla in length of 
tarsometatarsus and in some other gross features leads one to picture the 
extinct waterfowl as of similar but slighter build than the black or sea 
brant. Standing in equal height of limb, A. gracilenta weighed much less 
than the brant, probably approaching the tree-duck in weight. 

The spread of the distal condyles for digital articulation, while dis- 
tinctly goose-like and not duck-like, nevertheless falls away from the goose 
type when considered in proportion to length of shaft. In this regard it 
joins in close proportional similarity to the tree-ducks. Chloephaga, the 
upland goose, noted for its terrestrial habits, favors this narrow foot and 
long shaft proportion. It is significant that this proportional study throws 
sharp distinction between the fossil goose and the goose nearest it in size, 
namely Branta bernicla, while at the same time placing it nearer to B. c. 
minima and C. rossii. The latter two geese spend a considerable part of 
their lives upon the land, while Branta bernicla is a marine forager that 
spends little time ashore. It might be further pointed out that Philacte 
canagica, our most maritime goose, shows an extreme disproportion 
toward short shank and wide foot, and that the swan (Cygnus columbia- 
nus) , strongly aquatic, falls well within the same grouping. 

Considering the characters, slender shaft, length of shaft, small 
proximal weight-bearing articulation, reduced spread of digital condyles, 

Ross — New Pigmy Goose 111 

and considering the inference given above as based upon the ratio of foot- 
spread to tarsal length, this Brea Pigmy Goose appears in mind as an 
agile, light-bodied goose of active, walking habits frequenting, in company 
with shorebirds, mud flats and borders of ponds. 

Anabernicula (Anas-bernicla, duck-goose) denotes the relationship 
of the new goose genus, while gracilenta denotes the delicacy and grace 
suggested in many features by the specimens at hand. 

Faunal Range 

In the fossil collections of the University of California at Los Ange- 
les there are six tarsi of pigmy geese : two from Rancho La Brea, three 
from McKittrick, and one from Fossil Lake, Oregon. These were shown 
to me in 1931 by Dr. Loye Holmes Miller, who kindly permitted the 
study of them. 3 One of the two "pigmy geese" cited from Rancho La Brea 
a.s Branta(?) sp. by Miller 4 and two from McKittrick referred to the 
Chen hyperboreas group of the same author 5 classify readily as Anaber- 
nicula gracilenta. The Fossil Lake specimen, however, is not referable to 
the latter form. 

Branta minuscula, described by Alexander Wetmore 6 from a proxi- 
mal half of a humerus presents characteristics which might be expected in 
humeri of A. gracilenta. If the geologic position of Branta minuscula is 
Upper Pliocene, as determined by J. W. Gidley, considerable difference in 
age prevails between these two similar forms from Arizona and Southern 


A total of sixteen tarsometatarsal elements of a small goose from the 
Pleistocene asphalts of McKittrick and one specimen from Rancho La 
Brea furnish the basis for establishing a new genus of goose. In certain 
characters, as displayed in the regoin of the hypotarsus and on the inner 
extensor groove, this form shows resemblance to the tree-ducks. The 
species represented is smaller than Branta bernicla and evidently more 
slender than any living goose. 

? Miller, Loye H. A second avifauna from the McKittrick Pleistocene: The Condor, 
vol. 37, p. 76, 1935. 

4 Miller, Loye H. The birds of Rancho La Brea : Carnegie Inst. Wash. Publ. No. 
349, p. 73, 1925. 

5 Miller, Loye H. Avifauna of the McKittrick Pleistocene: Univ. Calif. Publ., Bull. 
Dept. Geol. Sci., vol. 15, No. 9, pp. 314, 315, 1925. 

6 Wetmore, Alexander. Fossil birds from southeastern Arizona: Proc. U. S. Nat. 
Mus., vol. 64, art. 5, pp. 6-7, 1924. 


San Diego Society of Natural History 

Fig. 1. 










Right tarsometatarsi of pigmy geese. Natural size. 
A 1 . Anabernicula gracilenta, Type, anterior view. 

A. Anabernicula gracilenta, Type, inner lateral and proximal views. 

B. Branta bernicla hrota, inner lateral and proximal views. 

C Branta canadensis minima, inner lateral and proximal views. 
D. Chen rossii, inner lateral and proximal views. 

Key to Figures on Page 113 

Total length of tarsometatarsus. 

Extensions beyond arrows indicate extremes listed in standard texts for tarsal lengths 
in skin specimens. In C (B. c. minima) skin measurements taken by author have 
been added to skeletal series. 
Inner cotyla, proximal end, anteroposterior depth. 

Articulation area end. Transverse width x cotyla depth (anteropos- 
terior) . 

Maximum depth condyle 4 (anteroposterior). 

Ratio of anteroposterior depth of inner proximal cotyla (Fig. 3) to anteropos- 
terior extent of inner calcaneal ridge. 

Ross — New Pigmy Goose 113 

A. Anabernicula gracilenta. C. Branta canadensis minima. 

B. Branta bernicla (2 races). D. Chen rossii. 

Fig- 2. 

1\5 60 6 1 ! 70 1*5 80 MM 

A — ■ 

B „ i '- J 

C • { ■ 

Fig. 3. 


C _ 


Fig. 4. 

.££ 7£ SjO 9^_ ioc) lib 

Square millimeters 

C ' 

Fig. 5. 

6.0 6lS 7.0 





Fig. 6. 

6,0 6,5 7,0 75 80 85 90 95 I00MM 

— I L 1 I I I I i i_. 

A . — 


Explanation of figures will be found on opposite page. 




Vol. VIII, No. 16, pp. 115-118 March 12, 1936 



San Diego Society of Natural History 

Studies in this rather complex genus of flycatchers have received im- 
petus through various papers published since the comprehensive revision- 
ary work of Ridgway (1907) and Hellmayr (1927). Chief, perhaps, of 
the problems which have been discussed are the relationships of Myiar- 
chus nuttingi and Myiarchus inquietus to Myiarchus cinerascens cinera- 
scens. Although the two most recent writers (van Rossem, 1931, and Gris- 
com, 1932 and 1934) are now in substantial agreement that cinerascens, 
inquietus, and nuttingi are conspecific, there remains considerable uncer- 
tainty as to the manner of intergradation. In this regard I can only reiter- 
ate that in Sonora cinerascens intergrades gradually and in perfectly con- 
ventional manner with inquietus. In regard to the behavior of these two 
forms in Guerrero I am not convinced, after an inspection of the same 
series that Griscom has studied from that State, that the collector has not 
mistaken belated migrants or non-breeding individuals of cinerascens for 
residents. It would appear most improbable that the geographic behavior 
of these two races should be so different in two far separated regions unless 
it be that cinerascens extends a great deal further south in the interior high- 
lands than has heretofore been suspected. 

It may be appropriate to note here that a personal examination of the 
type of Salvin and Godman's Myiarchus inquietus in the British Museum 

116 San Diego Society of Natural History 

shows it to be without question a perfectly typical example of the race to 
which the name has currently been applied. It is a female in fresh fall plum- 
age, collected at Acaquizotla, Guerrero (alt. 3500 feet) by Mrs. H. H. 
Smith on October 18, 1888. In addition to the somewhat yellower and 
riche;: coloration, as compared with cinerascens, it possesses the pro- 
nounced dusk)' shaft streak on the inner webs of the outer rectrices which, 
in part, distinguishes inquietus from nuttingi, but this streak does not ex- 
pand terminally as in cinerascens. The measurements of the type (Brit. 
Mus. 99-4-20-1479) are: wing, 85.5; tail, 82.5; culmen from base, 21.5; 
tarsus, 20.1 ; middle toe minus claw, 13.2 mm. 

Mr. Griscom has recently (1932) called attention to certain peculi- 
arities in a series of Myiarchus "nuttingi" from the Pacific slope of Guate- 
mala, and has suggested that possibly they are subspecifically distinct from 
true nuttingi of northwestern Costa Rica and western Nicaragua. For 
several years there has been a series of twelve specimens from El Salvador 
in the Dickey collection which are obviously neither inquietus nor nuttingi, 
but for various reasons it has been deemed advisable to delay the bestowal 
of a formal name until now. Several years ago I compared this Salvador 
series with Griscom's Guatemala material (in part) and concluded that 
the two lots were racially identical, though the Guatemala birds averaged 
slightly paler. A diagnosis of the new race follows. 

Myiarchus cinerascens flavidior subsp. nov. 

Type. — Male adult in fresh fall plumage, no. 15,556 Dickey collection; Lake 
Olomega, Depto. San Miguel, El Salvador, August 26, 1925; collected by A. J. 
van Rossem, original no. 8626. 

Subspecific characters. — Differs from all known forms of Myiarchus cine- 
rascens in richness and brightness of coloration; in this respect it is a striking dupli- 
cate of Myiarchus crinitus, save that the throat and chest of flavidior are slightly 
paler than in crinitus. Differs, further, from nuttingi in possessing a broad, well 
defined dusky stripe along the shafts (on the inner webs) of the lateral rectrices 
and in being very slightly larger. Besides the brighter and richer coloration, flavi- 
dior differs from inquietus in definitely smaller size. 

Range. — Lowlands of El Salvador and the Pacific coast of Guatemala. 

Remarks. — The relatively intense coloration, which in flavidior attains the 
yellowest underparts and brownest upperparts displayed by the species cinera- 
scens, is not dependent on season, for the remarkably uniform series was collected 
at various seasons throughout the year. This race is resident in El Salvador but is 
nowhere common. I did not encounter it above an elevation of 1500 feet. 

van Rossem — Race of Myiarchus cinerascens 117 

Measurements of Myiarchus cinerascens flavidior 
Extremes and Averages in Millimeters 

Culmen Middle Toe 

Wing Tail frcnn Base Tarsus minus Claw 

7 m ales 83-85 79.85 19.6-21.7 20.0-21.5 11.3-12.5 

(84.1) (81.6) (20.9) (21.0) (11.8) 

5 females 78-82 77-81 19.5-20.4 19.7-21.2 10.8-12.2 

(80.5) (78.2) (20.0) (20.4) (11.5) 

For the various conceptions of the relationships of the forms mentioned in 
this paper, reference may be made to the following publications: 

Nelson, E. W. 

1904. A revision of the North American mainland species of Myiarchus. 
Proc. Biol. Soc. Wash., 17, 1904, pp. 21-50. 


1907. The birds of North and Middle America. Bull. U. S. Nat. Mus., No. 
50, Part 4, pp. 625-632. 

Hellmayr, C. 

1927. Catalogue of birds of the Americas [etc.]. Field Mus. Nat. Hist., 
Zool. Ser., Part 5, pp. 160-161. 

van Rossem, A. J. 

1931. Report on a collection of land birds from Sonora, Mexico. Trans. San 
Diego Soc. Nat. Hist., 6, No. 19, pp. 260-261. 

Griscom, L. 

1932. The distribution of bird-life in Guatemala. Bull. Am. Mus. Nat. Hist., 
Vol. 64, pp. 253-254. 

Griscom, L. 

1934. The Ornithology of Guerrero. Bull. Mus. Comp. Zool., 75, No. 10, 
pp. 386-388. 

3L 9 Tf? 




Vol. VIII, No. 17, pp. 119-120 March 12, 1936 




A. Myra Keen 

Stanford University 

An evaluation of the pelecypod family Cardiidae (summarized else- 
where 1 ) , by means of a graphic comparison of the genotypes of the fifteen 
principal named groups, indicates the desirability of erecting a new genus. 
None of the genotypic species examined exhibits the distinctive characters 
which constantly appear in the Northwest American cardiids hitherto, 
though incorrectly, classified as Cerastoderma. The purpose of this pre- 
liminary note is to validate a name used in manuscript ; detailed discussion 
of the species included in the genus is withheld for a monographic study 
whose publication may be considerably delayed. 

CLINOCARDIUM Keen, new genus 
Genotype : Cardium nuttallii Conrad, 1837 

Description. — Shell medium to large, trigonal, oblique, usually ventricose; 
beaks recurved, prosogyrate; position of the umbones varying with age but usually 
at two-thirds the distance between posterior and anterior ends of the shell; dorsal 
margin very broadly arched, sloping downward at an angle of about 25°, ventral 
and anterior margins broadly rounded; epidermis closely adherent, brownish; 
sculpture of 28 to 55 rounded radial ribs and concentric growth lines which may 
cross the ribs as conspicuous loops, never as spines; lunule when present circum- 
scribed, never impressed; escutcheon inconspicuous; ligament in dorsal view long, 
narrow, and oval. Interior porcellaneous, ventral and anterior margins crenulate; 
hinge arched; cardinals in each valve slightly nearer anterior than posterior laterals; 
anterior cardinal of left valve stronger than posterior, recurved, posterior cardinal 

1 Keen, A. Myra, Revision of Cardiid Pelecypods; Proc. Geol. Soc. Amer. for 1935 (1936). 
Preliminary abstracts, Dec, 1935, p. 61. 


San Diego Society of Natural History 

of right valve stronger than anterior, also recurved; ligament not elevated on a 
short, shelly platform; beaks originating at a point slightly anterior to the anterior 
cardinals; muscle scars large; pallial line simple. Specimens range in length up to 
about 120 mm. 

Remcrks.— Clinocardium is distinguished from its nearest relative, Cerasto- 
derma, by the markedly forward-pointing beaks, the long, narrow, low ligament, 
the arched hinge -line, and the greater number of ribs. From Laevicardium it is 
distinguished by the presence of elevated ribs and by the long, depressed ligament. 

Clinocardium nuttallii (Conrad) 
Cardium nuttallii Conrad, Jour. Acad. Nat. Sci. Phila., vol. 7, 1837, p. 229, pi. 

17, fig. 3. 
"Cardium corbis (Martyn)" of West Coast authors; (not Corbis Martyn, Univ. 
Conch., Tab. 2, fig. 80, 1788) . 
Type locality. — A few miles from the estuary of the Columbia River. 
Repository of holotype. — Academy of Natural Sciences, Philadelphia; Cata- 
logue No. 54036. 

The name Cardium corbis (Martyn) is unavailable for two important rea- 
sons: First, as Winckworth 2 has pointed out, Martyn's Universal Conchologist 
is not consistently binomial and hence is to be rejected. Second, the identification 
of the West American species with that figured by Martyn is erroneous, a fact 
recognized by Conrad 3 in 1869. The first available name, therefore, is nuttallii 
Conrad, 1837. 

Other Species of Clinocardium 

The following tabulation lists all of the other species which have been deter- 
mined to be Clinocardium. Several Japanese Tertiary species will probably prove 
to belong here, as well as some two or three unnamed species from the Tertiary 
of California. 


blandum (Gould) 18 50 

bulowi (Rolle) 1896 

calif orniense (Deshayes) 1839 

ciliatum (Fabricius) 1780 
coosense (Dall) 1909 
comoxense (Dall) 1900 
decoratum (Grewingk) 1850 

fucanum (Dall) 1907 
meekianum (Gabb) 1866 

yakatagense (Clark) 1932 

Type Locality Age 

Puget Sound, Washington Recent 

Yokohama, Japan Recent 

(Of lectotype) Kamtschatka; 

(Of description) "Mers de Cali- 
fornia" Recent 

Greenland Recent 

Coos Bay, Oregon; "Miocene" Pliocene 

Vancouver Island, B. C. Pleistocene 

Aleutian Islands, "Jiingsten 

Tertiarzeit" Pleistocene? 

Juan de Fuca Strait Recent 

Eagle Prairie, Humboldt County, 

California, Pliocene Pliocene 

Yakataga formation, southern 

Alaska, "Upper Oligocene" Pliocene? 

2 Winckworth, R., Notes on nomenclature; Proc. Mai. Soc. London, vol. 18, pt. 5, July, 

1929, pp. 228-229. 

3 Conrad, T. A., Notes on Recent Mollusca; Amer. Jour. Conch., vol. 5, pt. 2, 1869, p. 105 



Volume VIII, No. 18, pp. 121-148, plates 17-18, maps 1-3 






San Diego Society of Natural History 


Printed for the Society 

May 29, 1936 



U. S. Grant, IV, Chairman 

Fred Baker Clinton G. Abbott, Editor 

van Rossem— Birds of South-central Arizona 

Plate 17 

Fig. 1. Lower Sonoran Plains along the north-east slope of the Atasco Mountains; the 
Tumacacori Mountains in the distance. 

w "'''^v^ai^ '*'-■ 


/. '4L 


Fig. 2. Pena Blanca Spring in the Pajarito Mountains, Santa Cruz County. Elf owls, 
spotted screech owls, and Mexican screech owls have been found to be common in this locality. 





San Diego Society of Natural History 

In the spring of 1931 I suggested to the late Donald Dickey, at 
that time my chief at the California Institute of Technology, that certain 
investigations be carried on along the Arizona-Sonora border west of 
Nogales. The ends sought were: (l) that the radically differing 
opinions of Mearns and Swarth as to the limits of certain faunal areas 
west of the Santa Rita Mountains might be brought into some sort of 
agreement, and (2) that a better understanding of the faunal areas of 
northern Sonora would be an inevitable result of such work. Mr. Dickey 
at once acquiesced and accordingly Dr. W. H. Burt, then mammalogist 
at the California Institute, and the writer arrived in south-central Arizona 
on April 23. Our first base was at Continental on the Santa Cruz River. 
From this station we worked south (including the west slope of the Santa 
Ritas) to Tumacacori and Nogales and thence west, including the 
Atasco and Pajarito Mountains, to the east slope of the Baboquivaris. 
Later in the season (on May 29) we returned to the Santa Ritas and 
worked for some few days in Madera Canon — from about 4500 feet to 
the summit of the range. 

In June, 1932, I again returned to southern Arizona and in com- 
pany with Dr. Walter P. Taylor of the Biological Survey and Mr. David 
Gorsuch of the University of Arizona worked from Peria Blanca Spring 
in the Pajaritos west around the northern end of the Baboquivaris to 
Fresnal on the west side of that range, and thence west to Ajo and south 
to Bates Well. One side excursion was made to the border village of 
San Miguel on the west side of the Baboquivaris. R. T. Moore of Pasa- 
dena who contributed, in part, the necessary field expenses, accompanied 
us as far as Fresnal. After three days at Bates Well, Gorsuch and I 
returned (on June 25) to the Santa Ritas, where I spent several days in 
Stone Cabin and Madera Canons before returning to Pasadena. 

Briefly, the discrepancy between Mearns' (1907) concept of the 
"Elevated Central" and "Western Desert" tracts, and Swarth's (1929) 

124 San Diego Society of Natural History 

limits of his "Eastern Plains" and "Western Desert" areas involves the 
not inconsiderable distance of some 120 miles! Mearns considered the 
longitude of the Ajo Mountains (specifically, Monument 163 in the 
Sonoyta River Valley) to be the dividing line; Swarth decided on the 
Santa Rita Mountains. Such opposition between two observers as ex- 
perienced as Mearns and Swarth was incomprehensible to me until per- 
sonal investigation showed each to have basis for his belief insofar as the 
territory explored by him was concerned. Mearns, except for side trips 
to Tucson and Fort Lowell, worked close to the border and was never 
more than a few miles north of that line; Swarth had never been along 
the border west of Nogales. 

Our own investigations showed Mearns' account of the border west 
of Nogales to be an excellent picture of actual conditions; in other words, 
from Nogales to the Baboquivaris (see Plates 17 and 18) there is ex- 
ceedingly close resemblance in most particulars to the Sonoran zones of 
his "Elevated Central Tract" and to Swarth's "Eastern Plains Area." 
Topographically the country is, in places, more broken than east of the 
Santa Ritas, but floral and ecological conditions are practically the same. 
On the higher levels are scattered stands of blue oaks, with sycamores 
and other characteristic Upper Sonoran growth in the canons. On levels 
below the oak belt and at intervals between oak areas are grass covered 
plains, more or less rolling and less level than to the east, but virtually 
the same type of country nevertheless. These grass areas, particularly 
those on fairly level ground, frequently support a thin growth of mes- 
quites — small trees spaced well apart and so regularly that, as Swarth has 
so graphically described them, one is forcibly reminded of a "young peach 

Along the border west of the Baboquivaris is a type of desert which 
I have not encountered elsewhere (save very locally) in Arizona. In 
most places the terrain consists of broad, level, grass plains which are 
varied along the usually dry water courses with dense thickets of large 
mesquite. At occasional points one encounters a more typical Colorado 
Desert topography and flora. Separating these plain-like valleys are 
stony ridges or low mountains which support a more familiar desert 
vegetation — giant and other cactus, greasewood, and low thorny scrub. 
This alternation of valleys and ridges extends west nearly to Ajo, at 
about the point where Mearns terminated his "Elevated Central Tract." 
Presumably, Mearns supposed that conditions similar to those in the im- 
mediate vicinity of the border persisted to the northward, and since the 

van Rossem — Birds of South-central Arizona 





pKjarito mts 




Map 1. Southern border of Arizona and surrounding territory. A — Western boundary of the "Elevated 
Central Tract" as determined by Mearns. Two subspecies of Lower Sonoran Zone birds (Pipilo fuscus meso- 
leucus and Phalaenoptilus nuttallii nuttallii) here reach their western limits. B — Western boundary of the 
"Eastern Plains Area" as determined by Swarth. This point marks the western limits of two Lower Sonoran sub- 
species (Toxostoma curvirostre curvirostre and Agelaius phoeniceus nevadensis). C — Here, at the Baboquivari 
Mountains, terminate, westwardly, the ranges of seven species and subspecies characteristic of the Lower Sonoran 
Zone of the "Apache Faunal Area" and also many of that Area's Upper Sonoran forms. The seven Lower 
Sonoran birds are: Colinus virginianus ridgwayi (formerly), Callipepla squamata pallida, Otocoris alpestris 
adusta, Corvus cryptoleucus, Geothlypis trichas chryseola, Sturnella magna lilianae, and Spizella carpalis 

general features of this part of the border more closely resemble the 
Lower Sonoran Zone of the "Elevated Central Tract" than they do the 
excessively barren desert to the westward one surely cannot criticize him 
for linking it with the former. Concerning the faunal affinities of this 
stretch from the Baboquivaris to Ajo, I cannot be certain at this time. 
At least two of the Lower Sonoran subspecies which are characteristic 
of southeastern Arizona range over this stretch, and possibly there are 
others. On the other hand "Western Desert" birds appear to be vastly 
in the majority. The faunal affinities, as a whole, are more probably with 
the Colorado Desert than otherwise. I have not been along the border 
between Ajo and Yuma, save for a very short distance east of Yuma. 
Mearns' "Western Desert Tract" and Swarth's "Western Desert 

126 San Diego Society of Natural History 

Area" are, in effect, practically synonymous (although not entirely so) 
with Grinnell's (1928) "Colorado Desert Differentiation Area." Mearns' 
"Elevated Central Tract" and Swarth's "Eastern Plains Area," on the 
other hand, are only in part comparable. Mearns' concept (properly, I 
believe) included the Sonoran and Boreal Zones; Swarth, specifically, 
considered in his studies only the Lower Sonoran. There is every reason 
to believe, in fact intensive studies of the geographic behavior of the 
birds of Sonora and Chihuahua make it almost certain, that southeastern 
Arizona is, on the basis of its Lower Sonoran, Upper Sonoran, and 
Boreal Zone birds, the northwestern portion of a large and well charac- 
terized area of differentiation which has as its geographical hub the 
mountain mass of the northern Sierra Madre which lies north of the 
east-west course of the Yaqui River. This may appropriately be desig- 
nated as the Apache Faunal Area. 

In the following pages I have confined observations to "new" data, 
that is such as change or modify current ideas of faunal areas, range ex- 
tensions, or systematic status. Maps have been added when thought 
necessary or desirable in order to make more graphic the problems in- 
volved. Carefully avoided have been data which would simply be 
repetition of the work of former observers. Needless to say, the Arizona- 
Sonora border is still a relatively unexplored territory and I have no 
illusions as to the finality of any ideas or statements expressed. Future 
field work may well, and probably will, modify any or all of them. 

Accipiter atricapillus striatulus (Ridgway) 

The American Goshawk seems not to have been previously recorded from 
the Santa Ritas. On May 1, 1931, and on two subsequent occasions Dr. Loye 
Miller and I saw an adult in the pines at about 7500 feet altitude in Madera 
Canon. Considering the date and the actions of this bird, which obviously resented 
our presence, it seems reasonable to suppose the probability of a nest somewhere 

Specific records of the goshawk in extreme southern Arizona are still suffi- 
ciently rare to make permissible the recording of some occurrences in the Chirica- 
huas. The Dickey collection contains an adult male taken by H. H. Kimball on 
July 10, 1918, at Paradise, an adult female taken by Kimball at an unknown 
locality (in the Chiricahuas) on December 5, 1918, and a juvenal female collected 
by Frank Hand in Pinery Canon on February 9, 1928. None of these three indi- 
viduals can be assigned to the eastern race, for they all possess the dark dorsal 
coloration and, in the juvenile, the broad blackish-brown ventral streaking, which 
characterize the race striatulus as now understood. In fact they are slightly but 
appreciably darker than average goshawks from the northwest coast. In the British 
Museum is a similarly dark-colored adult taken by W. Lloyd at Yecora in the 

van Rossem — Birds of South-central Arizona 127 

high mountains of east central Sonora on April 13, 1888, and also two nearly full- 
grown juveniles (which may well have been collected directly from the nest) 
taken by W. B. Richardson in the Sierra Nayarit, Jalisco. This last record is ap- 
parently far south of any previously known breeding station. 

There is the possibility that a definable race, distinct from striatulus, exists 
in the Mexican highlands and that its range includes the mountains of extreme 
southern Arizona. The appearance of the Jalisco juveniles, which are the darkest 
I have ever seen, is further indication that such is the case. 

I most emphatically follow Peters (1931), and most European ornitholo- 
gists, in considering Astur congeneric with Accipiter. 

Accipiter cooperii mexicanus Swainson 

Several pairs of the Western Cooper's Hawk were found in the Baboqui- 
varis, a point which marks the extreme western limit of territory suitable for 
breeding purposes. An adult male was collected with a set of four eggs in a small 
canon at 5500 feet altitude at the east base of Baboquivari Peak on May 27, 1931. 
Other breeding pairs were noted at intervals in the oak regions along the border 
between Nogales and the Baboquivaris and, in common with former observers, 
we found several breeding pairs in the Santa Ritas. 

Buteo albonotatus Kaup 

A single adult Zone-tailed Hawk was seen on the ground beside the road 
near Arivaca on May 23, 1931. In the Baboquivaris, a pair was found nesting 
in the oaks in Thomas Canon on the east side of the range. One parent (the 
female) was collected on May 24; the other, the male, who was then incubating 
the set of two eggs, on the 25th. 

Save for the several records from the lower Colorado River, some of which 
are dubious and more probably pertain to Harris's Hawk, I believe the specimens 
taken in the Baboquivaris to be the westernmost known occurrences of the Zone- 
tailed Hawk in southern Arizona. Since this hawk is distinctly partial to oak 
and sycamore timber (chiefly Upper Sonoran Zone) during the breeding season, 
the Baboquivaris in this, as in many other cases, probably constitute the western 
range limit along the border. 

Callipepla squamata pallida Brewster 

We found the Scaled Quail to range west along the border foothills and 
uplands to the east slope of the Baboquivari Mountains. In this territory we 
found it common, although usually outnumbered by GambeFs Quail. We noted 
it almost continuously along grassy, broken ground north almost to Picacho, but 
at no point west of the Baboquivaris. 

While I agree with Swarth that the Scaled Quail (or in a more properly 
restricted sense the subspecies pallida) is one of the characteristic birds of the 
"Eastern Plains" fauna, I cannot for a moment subscribe to a belief that Gambel's 
Quail is an indicator of his "Western Desert Area." Gambel's Quail is common, 
locally, clear across the "Eastern Plains" to the Chiricahua Mountains at least, 
where it is abundant. In fart in certain washes along the western foothills of that 
range I found it, in 1914 and 1915, to outnumber the Scaled Quail. Similarly, 


San Diego Society of Natural History 

A' 113" II 











• ; 







[m ~^~ — 



■ 1 



•) — 













\ i 




Map 2. Distribution of two species of quail, Callipepla squamata pallida (squares), and Lophortyx gam- 
belii gambelii (dots) in southern Arizona and northern Sonora. The broken line which encloses the range of 
pallida also indicates in a general way the western and northern section of the "Apache Faunal Area." 

south of the boundary, the Scaled Quail is an excellent faunal indicator, but 
gambelii ranges east across the state of Sonora to the foothills of the Sierra Madre. 

Cyrtonyx montezumae mearnsi Nelson 

In common with many other Upper Sonoran birds, Mearn's Quail ranges 
west of the Baboquivaris (both slopes; see Bruner, 1926), which mountains 
necessarily mark the extreme western limit of the species along the border. We 
also found it to be fairly common in the Atasco Mountains and about Pena 
Blanca Spring in the Pajaritos. 

Otus asio cineraceus (Ridgway) 

The distribution of the common screech owls in extreme southern Arizona 
west of the Santa Ritas cannot be finally outlined until more specimens have been 
collected from critical localities. One thing is certain, however, and that is that 
cineraceus occurs west along the border considerably beyond the range now ac- 
corded it. Dr. Miller and Berry Campbell (1934) took several specimens (ex- 
amined by me) at Pena Blanca Spring, Pajarito Mountains, in June, August, 
and September. I found a fully grown juvenile dead in the road between Oro 
Blanco and Arivaca on June 19, and Dr. Taylor collected a juvenile and an adult. 

van Rossem — Birds of South-central Arizona 129 

(also examined in the present connection), at Arivaca (a Lower Sonoran locality, 
incidentally) , on June 20, 1932. In the Baboquivaris a screech owl, probably of 
this subspecies, was heard but not collected. 

The subspecies gilmani has been recorded from the Santa Ritas (Bailey, 
1923), but Dr. Oberholser informs me that the specimen which was the basis 
of the record cannot now be found in the Biological Survey collection. In view 
of the fact that Swarth (1929) took a family of cineraceus there, the gilmani 
record is decidedly open to question. 

In considering the western limits of the range of cineraceus, it has been ad- 
visable to re-examine the type series of Otus asio cardonensis Huey from Lower 
California, since cardonensis has been synonymized with cineraceus by Grinnell 
(1928) . Entirely aside from the incongruity of such a "split" range — that is two 
colonies of cineraceus separated from each other by gilmani — I can see no valid 
reason for not recognizing cardonensis on its own character merits. Although 
cardonensis is very close to cineraceus and has the same gray cast of plumage 
(that is with almost complete absence of brown tones), cardonensis is definitely 
darker, and is more leaden (less ashy) than cineraceus, both above and below. 

Otus trichopsis trichopsis (Wagler) 

The occurrence of this rare screech owl in the mountains of south-central 
Arizona was to be expected, but that it should prove to be common in a locality 
which had prevoiusly been worked extensively was surprising. 

About 9 o'clock on the evening of June 4, 1931, the notes of a small owl 
were heard in the sycamore and oak timber along the stream which runs past the 
resort in Madera Canon in the Santa Ritas. These notes, while unmistakably 
"screech-owl" in character, were very different in cadence from those of any 
member of the asio group covered by my experience. They consisted of a repeti- 
tion of three short notes, a slight pause, and a fourth, terminal note, " -, 

, f -, - - - -•" A whistled imitation soon decoyed the caller, which proved 

to be a Spotted Screech Owl, within range. On June 5, a night trip of several 
hours' duration pretty well prospected Madera Canon between the resort at 5500 
feet altitude and Littleshot Cabin at 7000 feet. Among other nocturnal rarities 
encountered, Spotted Screech Owls were located, by the unmistakable call notes, 
at five different points and by calling the birds from a central location I had 
four individuals close to me at one time. On this occasion a mated pair was col- 
lected. Two nights later, two pairs were located in the south fork of the canon 
at altitudes of 6000 and 6500 feet respectively. One of these pairs was collected 
in an oak grove near the stream. 

At no time were asio call notes heard in Madera Canon, and I was tempted 
to conclude that trichopsis and asio were not likely to be found in the immediate 
vicinity of each other. However, I subsequently learned that Dr. Miller and his 
party took several examples of each species at Pena Blanca Spring in the Pajari- 

Stomachs of the pair collected on June 7 were submitted to the Biological 
Survey and their report is herewith appended. 

130 San Diego Society of Natural History 

Male — Condition of stomach: full 

Percentage on animal matter, 100; of vegetable, — 
Contents: 1 Diplotaxis sp. = 2%; 13 Noctuidae larvae = 87%; 
1 adult Lepidoptera — 2% ; 2 Formicidae = trace; 
1 Spider = 7 % ; fur of a small rodent = 2 % . 
Female — Condition of stomach: nearly full 

Percentage of animal matter, 100; of vegetable, — 
Contents: 1 Acrididae=5%; 6 Gryllus assimilis = 60%; 2 Noctui- 
dae larvae = 25%; 1 other Lepidopterous larva = trace; 1 
adult Noctuidae = 5%- 1 spider = trace; 1 scorpion = 5 % . 

Otus flammeolus (Kaup) 

Flammulated Screech Owls were found only in the Santa Ritas, from which 
range they have not previously been reported. At Littleshot Cabin, in the mixed 
oaks and pines at 7000 feet, a male was collected at dusk on June 6, 1931, as he 
was flying about through the trees. This bird was not at all shy and decoyed 
readily to a squeak. Three days later, on June 9, I noticed a feather clinging to 
the entrance of an old flicker hole some ten feet from the ground in a dead pine 
stub a quarter of a mile above the spot where the male had been shot on the 6th. 
The stub was pushed over without much difficulty, and the hole was found to 
contain a female Flammulated Screech Owl and two newly-hatched young. This 
nest probably did not belong to the male taken on the 6th, for another Flammu- 
lated Screech Owl was seen to visit the site on several occasions during the night 
after the female and young had been collected. 

I could perceive no definite eye shine from either of the two individuals 
seen at night, though the visitor to the nest site occasionally gave out a greenish 
white glint as the bird was viewed in profile. No notes, which could be certainly 
attributed to this species, were heard, although on one occasion a small owl, 
which I concluded because of its size to be a Flammulated Screech Owl, was 
heard calling in an oak grove at 6000 feet in Madera Canon. The call notes, 
which were given persistently, might be described as a poor-will call reversed — 
that is to say with the higher of the two notes given first. This bird was so shy that 
it flew rapidly from tree to tree in order to avoid the beam of the flashlight, and 
never remained stationary long enough to be collected, even by a snap shot. 

The two newly-hatched young collected on June 9, 1931, are thickly cov- 
ered with snowy white down, with, in life, the bills and feet flesh color. The irides, 
both of adults and young, were very dark, nearly blackish, brown — very different 
from the yellow irides of the common and spotted screech owls. In plumage, 
neither adult is in an extreme color phase; the female, though, is definitely more 
rufescent than the male. 

I am indebted to the Bureau of Biological Survey for the following stomach 
analyses of the two adults collected. 

Male — Condition of stomach: full 

Percentage of animal matter, 100; of vegetable, — 
Contents: 3 Diplotaxis sp. = 25%; 4 Noctuidae larvae = 35%; 2 
other Noctuidae larvae=5%; 6 adult Melipotis sp., 
probably indomita, =35%. 

van Rossem — Birds of South-central Arizona 131 

Female — Condition of stomach: full 

Percentage of animal matter, 100; of vegetable, — 
Contents: 1 Gryllus assitnilis = trace; 5 Diplotaxis sp. = 35%; 
1 1 Noctuidae larvae = 65 % . 

Glaucidium minutissimum gnoma Wagler 

Three specimens of the Pigmy Owl were collected, a breeding pair in the 
Atasco Mountains and a male in the Santa Ritas. In view of the rarity of the 
species, and particularly this subspecies which is here recorded for the first time 
from the United States, the circumstances of their capture seem worth recording. 

The breeding pair was encountered on May 19, 1931, as Dr. Burt and I 
were coming down through a grove of blue oaks at 4500 feet in Piskiorski Canon. 
As we passed under a small oak, a Pigmy Owl flew out and perched at the very 
tip of another oak nearby. It proved to be an adult male. Investigation of the tree 
from which he had flown showed three old nest holes of the Arizona Woodpecker 
in the main trunk, at heights of from 12 to 15 feet above the ground. Since the 
holes were drilled in the hard live wood they could not be opened at the time, 
but I returned next day and chopped out the most favorable looking one. The 
female refused to leave the hole during chopping operations, but flew out when 
I descended for a moment to the ground. She, like the male, flew to the tip of 
a nearby oak. There were four eggs in the nest, two of them fairly well incubated, 
the other two claw-marked and seemingly infertile. 

The male taken near Littleshot Cabin in the Santa Ritas on June 1, 1931, 
was noticed, quite by accident, perched at the tip of a small dead sycamore and 
in the full glare of the mid-day sun. Just before he was shot he was attacked 
furiously by a female Rivoli Hummingbird. 

The only other Pigmy Owls that I have been able to examine from extreme 
southern Arizona are a male from the Huachuca Mountains (32378 Museum 
of Vertebrate Zoology) and a male from the Santa Ritas (406 San Diego Soc. 
Nat. Hist.). 

For over ten years I have been keeping notes and measurements of Pigmy 
Owls whenever opportunity arose. The total examined has been in excess of 200 
specimens and over 100 have been assembled at one time. The conclusions 
reached differ in several respects with some previously published opinions, not 
only as to the systematic status of birds from certain regions but as to the ranges 
currently accorded several subspecies. 

In the first place, it is very evident that the matter of color phases has not 
received anywhere near the consideration it deserves. Secondly, size and propor- 
tions are of much more value than has been realized. Among recent writers Dr. 
L. B. Bishop (1931) has been practically alone in his contention that the race 
pinicola of Nelson is only the extreme gray phase of calijornicum. With this 
belief I am in agreement. However, I cannot subscribe to placing the Pigmy Owls 
of the northern Rocky Mountains with calijornicum. Compared with northwest 
coast examples of grinnelli, the interior birds unquestionably average paler or 
grayer, but certain interior birds are just as red or as richly colored as coastal 
grinnelli. The appended table of measurements shows the size agreement between 
coastal and interior Pigmy Owls and there remains only an incomplete phase 


San Diego Society of Natural History 

segregation by which to distinguish series from the two areas. I suggest to any- 
one interested a study of the geographic phase behavior of the closely related 
Glaucidium brasilianum ridgwayi as illuminating in the present connection. 

To return to the Pigmy Owls of southern Arizona, I must confess at the 
outset to having misled Dr. Bishop on the subspecific status of the Atasco Moun- 
tains pair, for at that time I had not made the critical comparisons which subse- 
quently proved to be necessary. Briefly, the southern Arizona specimens differ 
from "pinicola" \ = calif ornicum] in their decidedly smaller size, more conspicu- 
ously spotted upper parts and slightly darker (phase for phase) coloration. The 
four males are all in the gray phase, the female in the brown or "mongrel" phase. 
I cannot distinguish them in any way from gnoma of the Mexican highlands and 
the range of gnoma should be corrected to read "north to the Huachuca, Santa 
Rita, and Atasco Mountains in extreme southern Arizona." 

Glaucidium minutissimum 1 Males 



4 gnoma 





4 gnoma 





6 calif ornicum 





4 californicum 





1 grinnelli 



6 grinnelli 





8 grinnelli 





10 grinnelli 





17 grinnelli 





3 swarthi 

84-89 (86.3) 61-63 (61.7) 


California Sierras 
New Mexico 
Wrangle, Alaska 
Puget Sound Region 
Western Oregon 
N. W. California 
Interior Brit. Columbia 
and northern Idaho 
Vancouver Island 

Micropallas whitneyi whitneyi (Cooper) 

Perhaps the most noticeable of night noises in the cottonwood groves along 
the Santa Cruz River in the vicinity of Tumacacori Mission, in early May, 1931, 
were the familiar call notes of the Elf Owl. This point is some 14 miles from the 
nearest giant cactus (a small grove near Continental), and nearly 40 miles from 
the extensive groves to the west of Tucson. On the night of May 14, I had no 
trouble in locating, with the aid of a flashlight, two pairs within a hundred yards 
of camp, nor in collecting three specimens in almost as few minutes. I saw at 

1 Mr. Ludlow Griscom (1931) has recently reviewed the Central and South American 
races. He is unquestionably correct in his contention that gnoma and minutissimum are con- 
specific and that the two supposed species blend through griseiceps and cobanense. A criti- 
cal examination of the material in the British Museum, including the types of both races, 
shows that the Guatemalan lowland race griseiceps (13 specimens) is distinct from cobanense 
(5 specimens) by reason of the definitely shorter tail and the lesser number of tail bands. 
Incidentally griseiceps does have a red phase which is (in the single instance known to me) 
indistinguishable in color from cobanense. In such rare cases the measurements are, of course, 
diagnostic. Measurements of these two interesting races are appended. 

Males (Some griseiceps not Marked on Tags as to Sex) 

Wing Tail Tail bands (excluding tip) 

3 cobanense 67-88 59-62 6 to 7 

13 griseiceps 84-90 48-58 3 to 6 

van Rossem— Birds of South-central Arizona 133 

least a dozen birds within half a mile and could have taken all of them had it 
been desirable to do so. They were always in pairs, even though the breeding 
season was at its height, and for the most part they were located first by call notes 
and then observed with a flashlight. By far the greater number were in the medium 
height cottonwoods, perched or flying about in the branches below the crown 
foliage. Two were seen in a pile of drift-wood in the dry stream bed, and one in 
a mesquite thicket. The oviduct of the single female collected contained an egg 
ready to be laid. One pair was seen repeatedly to enter a small natural cavity, 
some thirty feet above the ground, in the trunk of a half-dead Cottonwood. In 
June, 1932, our party found Elf Owls to be not uncommon at Pefia Blanca 
Spring, Pajarito Mountains, in the live oak association of the Upper Sonoran 
Zone at about 4000 feet. One pair, accompanied by grown young, was found with- 
in a hundred yards of the spring, and enough additional birds were heard to 
convince us that several pairs were in the vicinity. This locality was one in which 
spotted and Mexican screech owls, poor-wills, acorn woodpeckers and other char- 
acteristic Upper Sonoran species were present — an unexpected place, surely, in 
which to find the supposedly cactus-restricted Elf Owl! Campbell (1934) has 
previously recorded Elf Owls from this same locality. 

A single Elf Owl was found half hidden in a mistletoe clump in a mes- 
quite thicket at Bates Well on June 22, 1932. There was no giant cactus in the 
immediate vicinity of this spot, but scattered plants were noted a half mile or so 
away. This bird is now in the collection of the University of Arizona. 

Strix occidentalis lucida (Nelson) 

The barking calls of Spotted Owls were frequently heard at night in the 
Santa Ritas, invariably between altitudes of 6000 and 8000 feet. At Littleshot 
Cabin, shortly before dusk on June 8, 1931, a Spotted Owl, subsequently found 
to be a female, suddenly appeared on the roots of a fallen tree, ten feet from the 
spot where I sat watching a whip-poor-will catch low-flying insects along the 
course of a small stream. This owl was not in the least disturbed by my presence, 
but obviously was intensely curious; indeed I believe it came down with the sole 
purpose of a close range inspection. At about 9 P. M. another Spotted Owl was 
heard barking from higher up the canon. When I answered, it came immediately 
and perched in a pine tree not twenty feet distant. Like the first bird it was per- 
fectly fearless and I had excellent opportunity to test it for eye shine. Though 
it was looking straight at me for several minutes I could get no reflection of any 
sort, the eyes appearing perfectly black at all times. This bird, a male, was very 
probably the mate of the female taken earlier in the evening. 

In 1932 another pair of Spotted Owls were found to have come into the 
territory opened up by the taking of the pair in 1931. Both birds were seen several 
times, and still more frequently were heard at night as they barked to each other 
from opposite sides of the canon. These 1932 birds were not nearly so tame as 
the first pair, but usually could be decoyed to reasonably close distances by an imi- 
tation of their calls. This pair was not molested. 

A Spotted Owl was seen, at dusk, to fly out of a grove of oaks near a spring 
at about 7000 feet in Stone Cabin Canon on June 25, 1932. Although this bird 
exchanged barks with me for more than an hour, it refused to come closer than 

134 San Diego Society of Natural History 

a hundred yards. It was evidently disturbed by the flashlight and would promptly 
shift location at every attempt to find it with this means. 

Bailey (1923) and various other observers have also found Spotted Owls 
in Stone Cabin and Madera Canons. The species is evidently fairly common in 
the Santa Ritas. 

Asio otus wilsonianus (Lesson) 

According to Swarth (1929) the Long-eared Owl has been detected but 
once as a breeding bird in Arizona, the record being that of a young bird taken 
at 5000 feet altitude in Stone Cabin Canon on the west side of the Santa Ritas. 
It was of interest, therefore, to find a family of these owls in the mesquite (Lower 
Sonoran Zone) along the dry stream bed at Bates Well. The young were nearly 
full grown, but partly in juvenal plumage. One specimen, a juvenal male, was 
collected on June 23, 1932. 

Caprimulgus vociferus arizonae (Brewster) 

The vertical range of Stephens's Whip-poor-will is not limited to the higher 
mountains. Above 6000 feet in the Santa Ritas many birds were heard on every 
occasion that we stayed out after dark, but I also heard a whip-poor-will several 
times at 5000 feet in the Atascos and Dr. Miller collected a specimen at Peha 
Blanca Spring, Pajarito Mountains, in June, 1931. These last two localities are 
well down in the Upper Sonoran Zone. 

I took, all told, six specimens of this common, though seldom collected, 
whip-poor-will and saw or heard several times that number. In the Santa Ritas 
they showed a decided preference for groves of oaks and sycamores in the canon 
bottoms, and nearly all of those which were found at night were feeding in the 
immediate vicinity of running water. A pair to the mile seemed to be normal in 
most canons which contained water and it was obvious that each pair had its own 

Though males, and sometimes, before eggs were laid, mated pairs, were 
invariably found in the canon beds, the two nests discovered were on hillsides at 
least a quarter of a mile from water. On the night of June 6, 1931, I caught the 
red eye-shine of a whip-poor-will some distance away (estimated by daylight at 
150 yards) and across a steep-banked canon. With no expectation of collecting 
anything I followed the trail to a point where I estimated the shine to have been, 
but could locate nothing and supposed that the bird had gone. On my return 
to the original spot the eye was seen in the same location as before, and this time, 
after a little search, I found a female sitting under the protection of a 
fallen spray of leafless twigs which lay on a steep bank beside the trail. Three 
of us had passed within five or six feet of this sitting bird on two occasions the 
day before. The sitting bird made no effort to escape and was picked up by hand. 
There was one nearly fresh egg in the shallow depression in the gravelly soil 
which served as a nest, and stuck to the ventral plumage of the incubating female 
were several small pieces of shell, showing that another egg had been laid and 
somehow broken. This single egg was by no means immaculate white, but was 
clouded and mottled with brown and lilac, mostly in the nature of semi-concealed 
shell markings. It was similar to but very much less highly colored than eggs of 
the eastern vociferus, however. 

van Rossem — Birds of South-central Arizona 135 

A second nest found on the night of June 27, 1932, within two feet of the 
same trail was, like the first, under a slight canopy of dead oak twigs. This nest 
contained one pure white egg and a newly hatched chick clothed with a very 
respectable covering of down — in color between "cinnamon" and "orange-cinna- 
mon" of Ridgway. 

A, to me, surprising circumstance, was the marked erectability of the 
feathers above the eyes. Both of the sitting females carried these tufts constantly 
erect the entire time they were under observation. A male seen from directly in 
front alternately raised and flattened them. On one previous occasion, when night 
hunting in El Salvador, I had observed the eastern subspecies (vociferus) to 
have markedly erectile tufts; in fact until I picked the bird up I was certain that 
I had shot some small "eared" owl. About 45% from the horizontal was the 
maximum elevation, though when viewed from directly in front the tufts appear 
nearly vertical. 

Phalaenoptilus nuttallii nuttallii (Audubon) 

Poor-wills were heard at night chiefly in the oak association of the Upper 
Sonoran Zone but we also found them to be common on the Lower Sonoran 
desert. Specimens were taken in the Atasco Mountains, Pajarito Mountains and 
at Bates Well. This last locality is interesting as indicating the approximate area 
of intergradation between nuttallii and hueyi of the lower Colorado River Val- 
ley. The series of 4 birds from Bates Well is certainly referable only to nuttallii, 
but an approach to hueyi is seen in the slightly browner and paler coloration. 
These specimens were taken on June 22 and 23, 1932, a date which precludes 
the possibility of their being migrants. The two males were in breeding condition; 
the two females were evidently incubating. All were collected in an arrowweed- 
mesquite association along the borders of the dry stream bed. Conditions, both 
as to habitat and temperatures, closely approximated those found along the Lower 
Colorado River Valley, beyond the confines of which hueyi has not been detected. 

Previous to this time I had anticipated that nuttallii would be found to be 
confined, during the breeding season, to the Upper Sonoran Zone and that hueyi 
would be found to be the race of the Lower Sonoran deserts. That the distribu- 
tion of the two races is not dependent on zonal considerations is obvious by rea- 
son of the presence of nuttallii at Bates Well. 

Selasphorus rufus (Gmelin) 

One specimen was taken at 4500 feet Stone Cabin Canon in the Santa Ritas 
on April 27, 1931. I believe the Rufous Hummingbird has not been previously 
detected in these mountains during the spring migration. 

Trogon elegans canescens van Rossem 

Regardless of its status in former years, this trogon may now be counted a 
fairly common summer visitant in the Santa Ritas. Possibly it has always been 
more numerous than was supposed, for one of the rangers, who has been stationed 
for many years in the Santa Ritas, knew the bird well and told me of having seen 
as many as five or six feeding together at a single patch of manzanita. At any 
rate there were several pairs in Madera Canon in the summers of 1931 and 1932. 

136 San Diego Society of Natural History 

The loud, hoarse, "koa-koa-koa" call of a male trogon was first heard in 
the pines at the head of Madera Canon on May 1, 1931, but he was moving con- 
tinually and we never were able to catch sight of him. On May 30 of the same 
year we occasionally saw, and more frequently heard, at least two pairs in the 
right hand (south) fork of the canon about two miles above the resort, and at 
altitudes of between 6000 and 6500 feet. The association in which they were 
noted was the oak-sycamore growth near the juncture of Upper Sonoran and 
Transition. Two more males (both of which presumably had mates) were heard 
in the left (north) fork — one at 7000, the other at 8000 feet altitude. These alti- 
tudes are in the pine-oak association in the Transition Zone. 

On June 8, 1931, while tramping through a bed of fallen sycamore leaves 
at 6000 feet, I found a female trogon flopping about but unable to rise from the 
ground. Dissection showed that she had been shot in the body with what appeared 
to be an air gun pellet. The spot where she was found was a frequently occupied 
camp site, and the assumption is that she had been wounded by one of the many 
campers present the previous week-end. In 1932, trogons were still more plenti- 
ful. On June 27, Mr. Gorsuch and I saw or heard eight birds between the forks 
of the canon, at 6000 feet, and Littleshot Cabin at 7000. On that date a fully 
adult male was collected by Mr. Gorsuch for the museum at the University of 
Arizona. On June 28, a very young trogon, about two-thirds grown and evidently 
just out of the nest, was shot, quite unintentionally, in a patch of oaks at 6000 
feet. It was perched about two feet from the ground in fairly thick undergrowth 
and was mistaken for a juvenile robin. This specimen has been donated to the 
University of Arizona, for trogons are, properly, on the list of species the col- 
lecting of which is prohibited. 

Dryobates villosus icastus Oberholser 

Hairy Woodpeckers were not uncommon in the higher parts of the Transi- 
tion Zone in the Santa Ritas, where one specimen was collected at 8500 feet alti- 
tude on the east slope of Mt. Wrightson on June 5, 1931. The record is appar- 
ently the first for the Santa Ritas, though the occurrence of the species was to 
be expected. 

Dryobates arizonae arizonae (Hargitt) 

Like that of many other Upper Sonoran species, the range of the Arizona 
Woodpecker extends west (Bruner, 1926) to the Baboquivaris. We found it 
to be common in those mountains (specimen taken in Thomas Canon at 5500 
feet altitude) and also in the oak association of the Atascos and Pajaritos. 

Myiarchus tyrannulus magister Ridgway 

But one specimen of the Arizona Crested Flycatcher was taken and this is 
recorded only because of the nesting of the species at a point some ten or eleven 
miles from the nearest giant cactus. 

Though Swarth (1929) noted the first arrivals on May 15, the season of 
1931 must have been somewhat advanced, for two pairs were seen in the cotton- 
woods along the dry bed of the Santa Cruz River two miles south of Tumaca- 
cori Mission on May 12. Both of these pairs were engaged in carrying nesting 

van Rossem — Birds of South-central Arizona 137 

material into natural cavities in tall cottonwoods, and it must be supposed that 
they had arrived in the locality some days at least prior to that date. The male 
of one of these pairs was collected on the 13th. 

Myiarchus tuberculifer olivascens Ridgway 

In common with many other birds whose habitat is mainly in the Upper 
Sonoran Zone, the Olivaceous Flycatcher extends west through the oak covered 
hills along the international boundary to the Baboquivari Mountains. We found 
the species to be common in the oaks of the Atascos, Pajaritos and Baboquivaris, 
and also found it nesting, not uncommonly, in the cottonwoods at Tumacacori. 
This last locality has an altitude of 3300 feet, and is well within the Lower So- 
noran life zone. At no other point in Arizona, so far as I am aware, have the 
Olivaceous Flycatcher, Ash-throated Flycatcher, and the Arizona Crested Fly- 
catcher been found nesting in the same locality and environment, although Camp- 
bell ( 1934) records all three species at Pena Blanca Spring in the Pajaritos "be- 
fore July 15," a date which indicates that they were breeding birds. 

In southern Sonora and southward the Olivaceous Flycatcher is widely 
distributed zonally, and I believe that its usual close adherence to the Upper 
Sonoran in northern Sonora and Arizona is because of preference for a woodland 
habitat and not because of any intolerance for Lower Sonoran temperatures. 

Specimens were collected in the Santa Ritas, Atascos, Baboquivaris (Thomas 
Canon, May 27, breeding), and Tumacacori (nesting pair taken May 12). 

Empidonax difficilis difficilis Baird 

Four breeding specimens of the Western Flycatcher were collected in Ma- 
dera Canon, Santa Rita Mountains, at altitudes varying from 6000 to 7000 feet. 
These are all of the large, dull-colored (as yet unnamed) form which breeds in 
the southern Rocky Mountains north to Wyoming and Montana and west to 
the Warner Mts., Oregon. In measurements the males from the interior average 
about 70 mm. in wing length and 60 in tail length, as opposed to about 64 and 
55 mm. for far western birds from Alaska south, coastwise, to southern Califor- 
nia. The coloration of the interior birds is grayer and duller when specimens in 
equal stages of wear are compared. 

A migratory specimen of typical difficilis was collected in the Atasco Moun- 
tains on May 19, 1931. 

Camptostoma imberbe ridgwayi (Brewster) 

The Beardless Flycatcher is not new to the territory of this paper, indeed 
the Santa Cruz River Valley is evidently the center of its range in Arizona. But 
since it has been supposed to be rare, there is justification for placing our obser- 
vations on record. 

I believe this species to be common in southern Arizona, and that the chief 
reason why it has not been detected more often is its close resemblance in color, 
size, and call notes to the Verdin. 

I first heard the "verdin" notes of this species at Continental on April 24, 
1931, and caught occasional flashes of a bird in a dense patch of mesquite and 
second growth cottonwoods along the nearly dry stream bed. On the 26th, Dr. 

138 San Diego Society of Natural History 

Miller and I went back to the same spot and succeeded in taking both members 
of a pair which was nearly ready to breed. We next met the Beardless Flycatcher 
in a grove of oaks, alders, and sycamores near the mouth of Madera Canon in 
the Santa Ritas, at an altitude of about 4000 feet and just at the juncture of Lower 
and Upper Sonoran Zones. A pair of these birds was obviously much interested 
in one section of a sycamore grove but, though we watched them for several hours, 
we could find no nest. Both birds kept well up in the foliage at from twenty to 
forty feet from the ground, though one made frequent flights to a small clump 
of young mesquites on the hillside nearby. This pair was certainly breeding, for 
the condition of the female showed her to be incubating and she had but recently 
finished laying. At Tumacacori we found Beardless Flycatchers to be common in 
the groves of cottonwoods and willows along the dry river bed. I estimated the 
population to average a pair to every quarter-mile for at least two miles either 
way from our camp. However, pairs were by no means regularly spaced 

One might easily have missed seeing any or all of these birds, for they were 
decidedly partial to perches just under the crown foliage of the cottonwoods at 
heights of from twenty to fifty feet. Were it not for the tell-tale, verdin-like pip- 
ing, a search in such an environment would not be likely to produce results, for 
the diminutive size of the birds, their inconspicuous coloring, and rather seden- 
tary habits combine to make them exceedingly difficult to detect, even after their 
general location has been ascertained. I found that sitting quietly and waiting 
for a flycatcher to become concerned about my continued presence was a much 
better method of locating pairs than to search at random. Every grove of good- 
sized cottonwoods contained at least one pair and sooner or later, if one kept 
quiet, the male would begin to sound his sharp alarm notes and to flit nervously 
about the grove. As often as not his mate would join him for a few minutes and 
then disappear. 

A nest was found on May 13. It was about twenty-five feet above the ground 
and was tucked into the pendent stems of a clump of mistletoe which grew at 
the tip of a Cottonwood branch. Certain that a nest was near at hand, I spent two 
hours in trying to follow the more elusive member of the pair and finally saw 
her disappear into one of several clumps of mistletoe. No nest was visible from 
the ground, but by climbing an adjacent tree I could see it plainly enough. In 
the absence of a sufficiently long rope the site was inaccessible, and I finally shot 
off the branch in order to examine the nest at close range. It was a thick-walled, 
rather loosely packed, four-inch globe of grasses and fine weeds with the interior 
well padded with plant-down, feathers and a small amount of rabbit fur. The 
entrance was in the side, slightly above center. The single egg, badly broken in 
the fall, was white, thickly spotted with tiny reddish-brown dots about the larger 
end and more sparingly elsewhere. 

Specimens of the Beardless Flycatcher were taken at Continental on April 
26; at the mouth of Madera Canon on April 29, and at Tumacacori on May 12, 
13, 15 and 16, 1931. Though a total of eleven birds was collected, it would have 
been possible to have taken twice that number at Tumacacori alone, along the 
four or five miles of river bed which was explored. At Fresnal, on the west side 
of the Baboquivari Mountains, on June 20 and 21, 1932, I repeatedly saw a pair 
of Beardless Flycatchers in a grove of small mesquites. I collected one of the- 

van Rossem — Birds of South-central Arizona 139 

birds and, as I remember, gave it to Moore, but I do not know its location at the 
present time. Fresnal is considerably to the west of any previously recorded sta- 
tion for the species. 

I have elsewhere (1930, 1934) given my reasons for believing that the 
northwestern race, ridgway'i, should be recognized. 

Otocoris alpestris adusta Dwight 

The Scorched Horned Lark does not reach its western limit at the Santa 
Ritas, but continues west along the mesa lands to, and all along, the east base 
of the Baboquivaris. Bruner (1926) has previously provisionally referred birds 
from this locality to the race adusta. Nineteen specimens were collected at the 
following points, though the distribution is really practically continuous, even 
in the grasslands of the oak association: Nogales, 1 (specimen lost en route); 
Atasco Mountains, 11; 5 miles west of Ruby, 1; Arivaca, 1; Altar Valley 15 miles 
north of the boundary, 1; east base of the Baboquivari Mountains at 5, 10, and 
15 miles north of the Boundary, 5. These are all typical of the race adusta except 
that four of the nine males from the Atasco Mountains are variously paler. I 
incline to the belief that this paleness is a local tendency rather than evidence of 
intergradation with leucansiptila. There is as yet no evidence that any horned lark 
occurs in the breeding season between the Baboquivaris and the Colorado River 

Corvus cryptoleucus Couch 

White-necked Ravens were found to be fairly common at Continental in 
late April, common at Tumacacori in mid-May and still more numerous all along 
the east side of the Baboquivaris the latter part of May. Bruner (1926) records 
them from the west side of the mountains at Fresnal, and I suspect from various 
other data that they extend even further west. We saw no nests nor any indica- 
tions of breeding other than that the birds were usually in pairs and a female 
taken at Tumacacori on May 13, 1931, was nearly ready to lay. Dr. Vorhies 
(1934) has recently called attention to the absence of recent records of the 
White-necked Raven in the valley of the Santa Cruz and suggests that the species 
no longer occurs there. The above records are therefore of interest as showing 
that it is still a common bird in the upper Santa Cruz valley (at certain times of 
the year at least) , and also that it ranges west at least to the Baboquivaris. 

Arizona winter records of this raven are not numerous. I found them to 
be very common all over the Sulphur Spring Valley in the winter of 1914-1915. 
The Dickey collection contains specimens taken by myself 4 miles west of Light 
on February 22, 1915, and 12 miles southeast of Willcox, February 24, 1915. 

Auriparus flaviceps ornatus (Lawrence) 

Verdin material from the border is unsatisfactory both as to quantity and 
quality and the present tentative determinations of individuals must be verified 
by series of fresh-plumaged specimens. Two adult males from Continental (April 
24 and 26, 193 1 ) and one from Tumacacori (May 12, 193 1 ) are in badly abraded 
plumage and do not provide much in the way of color values. However, their 
measurements most closely approximate those of ornatus. A single juvenile (May 
24, 1931) from the east slope of the Baboquivaris appears to be exactly like juve- 

140 San Diego Society of Natural History 

niles of ornatus from Saric, in northern Sonora. Compared with juveniles of 
acaciarum it is darker, grayer, and lacks the slightly buffy suffusion of acaciarum. 
Midwinter specimens from Tucson and Fort Lowell are intermediates between 
ornatus and acaciarum, though closer in both size and color to ornatus. Presum- 
ably, verdins from the southern localities given above ( i. e. west to the east slope 
of the Baboquivaris) are in this same category. 

A single adult female from Fresnal (June 20, 1932) and another from 
Bates Well (June 22, 1932) are so worn that no comment on them is possible. 
The latter is presumably of the subspecies acaciarum. 

Progne subis hesperia Brewster 

Purple Martins were seen about Arivaca, where they were apparently nest- 
ing in certain buildings. It was, of course, not possible to collect specimens in the 
town, but an adult male was taken on June 19, 1932, over a shallow pond, bor- 
dered by willows, about a mile south of that place. The measurements of this 
specimen (wing, 140; tail, 69 mm.) place it as hesperia. I have previously (1931) 
called attention to the fact that hesperia is the subspecies present at Tucson in 
May and June, and the assumption is that this name will be found to apply to 
all Purple Martins breeding in the Lower Sonoran Zone of southern Arizona. 

Aphelocoma sordida arizonae (Ridgway) 

Arizona Jays were found to be common throughout the oak belt of the Upper 
Sonoran Zone west to, and including, the Baboquivari Mountains. The last 
named range is the most westerly outpost of territory suitable for this species in 
southern Arizona. 

Two breeding specimens were collected in the Atasco Mountains on May 
18, 1931, and one at 5500 feet altitude in Thomas Canon in the Baboquivaris 
on May 27, 1931. 

Sitta carolinensis nelsoni Mearns 

The Rocky Mountain Nuthatch ranges west through the oak regions of 
the Atascos and Pajaritos to the Baboquivaris. A breeding male was taken at 
5500 feet in Thomas Canon in the Baboquivaris on May 27, 1931. Two speci- 
mens (an apparently mated pair) were taken in the Lower Sonoran cottonwoods 
at Tumacacori on May 13, 1931, but whether these birds were transients or were 
preparing to breed in the locality is unknown. 

Toxostoma curvirostre palmeri (Coues) 

The series of thrashers collected leaves no doubt that the dividing line be- 
tween palmeri and curvirostre in extreme southern Arizona is the ridge formed 
by the Santa Ritas and their southern continuation, the San Cayetanos. In the 
valley of the Santa Cruz, palmeri occurs in typical form at Continental (5 speci- 
mens). Two of the three specimens from Tumacacori, some 20 miles up stream 
(south) from Continental, show slight but definite tendencies toward curvirostre, 
as does a single specimen from just southwest of Nogales in Sonora. Two birds 
from the east slope of the Baboquivaris are in this same category. It is likely that 
most individuals from the higher ground between Nogales and the Baboquivaris 

van Rossem — Birds of South-central Arizona 


Map 3. Distribution of Toxostoma curvirostre curvirostre (squares), and Toxostoma curvirostre palmen 
(dots) in southern Arizona and northern Sonora. Bars indicate intergrades. 

will be found to show certain curvirostre tendencies, but all those from this region 
which I have examined personally are certainly much better referable to palmeri. 
I have included on the map certain Sonora stations from which I have ex- 
amined specimens as an aid to a better understanding of the behavior of the 
species on the Arizona side of the line. 

Turdus migratorius propinquus Ridgway 

Robins have been reported from the Santa Ritas in winter, but not previously 
as breeders. They were not to be classed as common, but several nesting pairs were 
seen during June, 1931, in Madera Canon, and at least four pairs were found 
in the quaking asp thickets near the ranger station on Mt. Wrightson. Speci- 
mens were collected on June 1,3, and 5, at altitudes of 7000, 7000 and 8500 feet, 

Hylocichla guttata polionota Grinnell 

Two specimens of the Great Basin Hermit Thrush were taken in the Santa 
Ritas. A female, probably a migrant, was collected at 4500 feet in the oak asso- 
ciation in Stone Cabin Canon on April 25, 1931. A single male, whose condition 
and actions made it almost certain that it was breeding, was taken at 6500 feet 
in Madera Canon on June 1, 1931. Other hermit thrushes were heard singing 
(in June) in Madera Canon and its tributaries at altitudes up to 8000 feet, but 

142 San Diego Society of Natural History 

they were extremely shy and I was able to collect only the specimens recorded 
above. The (presumably) breeding male is typical of the subspecies polionotd 
in size and color. The wing and tail measurements are 99 and 71 mm. respec- 

Geothlypis trichas chryseola van Rossem 

Five specimens of the Golden Yellowthroat were collected at Continental 
between April 30 and May 4, three at Tumacacori on May 15 and one in the 
Altar Valley, 15 miles north of the International Boundary on May 26, 1931. 
All of these birds were breeding, or about to do so, and may be taken as repre- 
sentative of the yellowthroat populations of the localities in which they were 
collected. For the most part they are easily referable to chryseola and have wide, 
yellow-tinged, post-frontal bands, yellowish green dorsal coloration, and golden 
yellow underparts. The series as a whole, however, does not show the extreme char- 
acters of the race, and is slightly intermediate toward scirpicola. 

It is still not possible to outline the ranges of the yellowthroats of southern 
Arizona with any degree of finality, save that scirpicola ranges east up the valley 
of the Gila and its tributary the Santa Cruz as far as Tucson, and that chryseola 
crosses the boundary in the valleys of the San Pedro and Santa Cruz at least to 
Fairbank and Continental and also occurs in the upper Altar Valley. Material 
to show whether or not chryseola occurs at the isolated reservoirs and water holes 
in the extreme southeastern corner of the state is lacking. The distribution of 
chryseola in Sonora and Chihuahua, however, would indicate that such is the 


Sturnella magna lilianae Oberholser 

The unmistakable song of this species was occasionally heard (though 
neglecta was certainly more common) at Continental, where Dr. Miller took a 
specimen on April 29, 1931. So far as I could determine, magna was the only 
species of meadowlark in the grasslands on the west slopes of the San Cayetano 
Mountains (specimen taken May 14, 1931), in the grasslands near Tumacacori 
(specimen taken May 15, 1931), and on the mesas along the Atasco and Babo- 
quivari Mountains. 

The subspecies lilianae appears to be an excellent race and easily distinguish- 
able from hoopesi of the Atlantic (Rio Grande) drainage. In color, it is an almost 
exact duplicate of Sturnella neglecta, and some specimens would be virtually 
impossible to place with certainty were it not for the sharp division between the 
yellow throat and buffy white jugal area of lilianae. 

Xanthocephalus xanthocephalus (Bonaparte) 

Yellow-headed Blackbirds were observed as not uncommon transients in 
the cultivated ground at Continental. An adult male was collected on April 29, 


Agelaius phoeniceus sonoriensis Ridgway 

The breeding red-winged blackbirds of the Santa Cruz Valley are similar 
to those from Tucson. They are not appreciably different in color from speci- 
mens from the Lower Colorado River Valley (the metropolis of the subspecies) 
but have, on an average, slightly thicker and shorter bills; in fact certain extreme 

van Rossem — Birds of South-central Arizona 143 

individuals have bills almost as stubby as those of fortis. All in all, they are essen- 
tially like the type of sonoriensis which, as has repeatedly been noted, is a winter 
bird from Camp Grant, a locality well within the breeding territory of A. p. 
nevadensis. It is not improbable that the type of sonoriensis was simply a winter 
vagrant from the Santa Cruz Valley. 

I agree with Swarth that the breeding redwings from east of the Santa Ritas 
should be called nevadensis. 

Specimens of sonoriensis were collected at Continental (17), Tumacacori 
(2) and Arivaca (3). 

Molothrus ater artemisiae Grinnell 

One specimen of the Nevada Cowbird was collected at Continental on May 
4, 1931. The bird, a female, was not in breeding condition and was obviously a 
transient in the locality. 

Tangavius aeneus milleri van Rossem 

Bronzed Cowbirds were occasionally seen about the ranch buildings at Los 
Encinos at the east base of the Baboquivaris, though invariably in the cattle cor- 
rals where it was not practicable to collect them. Several males were noted at a 
water tank at Fresnal at the western base of the Baboquivaris, but these birds 
were extremely shy and I never succeeded in getting within gunshot. Bruner 
(1926) has also noted the species at Fresnal. Both of the above localities are 
considerably to the west of the currently supposed range of this species. 

Richmondena cardinalis superba (Ridgway) 

Save for the old record of "Colorado River, Arizona," collected by Bischoff, 
the Arizona Cardinal has apparently not been detected (save by Bruner, 1926) 
west of the Santa Cruz River. We found it to be fairly common at Fresnal (spec- 
men collected June 20, 1931) on the west side of the Baboquivaris, and also saw 
probably a dozen birds in the mesquites at Bates Well. An adult male was taken 
in this last named locality on June 23, 1932. Without investigation of the terri- 
tory between Bates Well and the Colorado River it is not possible to suggest how 
much farther west the cardinal ranges, nor why it is (apparently) absent from 
the Lower Colorado River Valley. 

On April 23, 1931, during a stop at a service station about 10 miles east 
of Gila Bend on the Gila Bend-Tucson highway, I saw a male cardinal in the 
thick growth of desert vegetation (chiefly sahuaro and mesquite) near the road, 
and learned from the station operator that cardinals were often seen there. There 
is an abrupt break in the character of the vegetation just east of Gila Bend and 
one would assume that the range of the cardinal does not extend west of that 

Hesperiphona vespertina californica Grinnell 

A solitary female, whose condition indicated that she was incubating, was 
shot at 8500 feet altitude near the summit of Mt. Wrightson, in the Santa Ritas, 
on June 5, 1931. I cannot distinguish this single bird from females from the 
mountains of California and southern Nevada. It is, definitely, not montana. I 

144 San Diego Society of Natural History 

shall elsewhere (MS. in press) advance reasons for the recognition of the subspe- 
cies calif ornica as distinct from brooksi. 

Pipilo fuscus mesoleucus Baird 

The Canon Towhee in typical form ranges west to the east slope of the 
Baboquivari Mountains (specimen taken May 24, 1931). West of these moun- 
tains we found towhees to be common at Fresnal (2 specimens taken June 21 
and 22, 1932) and not rare at Bates Well (3 specimens taken June 23, 1932). 
These birds from west of the last Upper Sonoran outpost in the Baboquivaris are 
slightly darker and grayer than typical mesoleucus in comparable plumage, but 
larger series in fresh plumage are necessary in order to verify the comparative 
characters shown in worn, breeding specimens. 

Why this species ranges across a hundred miles of Lower Sonoran Desert 
between the Baboquivaris and Bates Well and does not continue on to the Colo- 
rado River is a question which cannot be answered until more is known about the 
character of the country for the remaining distance. 

Aimophila ruficeps scottii (Sennett) 

In the San Cayetano, Atasco, Pajarito and Baboquivari Mountains, this 
sparrow is a common inhabitant of all grass or cactus-grown slopes and speci- 
mens were collected in all localities except the Baboquivaris. However, the species 
is, as above stated, common there. 

Spizella carpalis carpalis (Coues) 

Moore (1932) has already recorded the collecting of four specimens of the 
Rufous-winged Sparrow at Fresnal, on June 20 and 21, 1932. My familiarity 
with this species in Sonora indicated that Fresnal provided an association, (a 
mesquite-cactus-grass growth on broken, gravelly ground) eminently suitable for 
its presence and I described the bird to Moore in hopes that one of us might de- 
tect one. Moore took three individuals on the afternoon of the 20th and one the 
following morning. I, personally, saw only a single individual and collected none. 

The purpose of mentioning this previously published record of the sup- 
posedly extinct (in Arizona) species is to call attention to the bird's typical habi- 
tat. It is not an inhabitant of grassland plains, other than as a possible casual, 
but occupies, normally, very much the same ecologic niche as Amphispiza bill- 
neat a. 

I do not think that the Rufous-winged Sparrow can be considered rare in 
south-central Arizona. It is, however, obviously very local and this, combined 
with its close resemblance to Spizella passerina, has doubtless led to an erroneous 
assumption of rarity. 

This species is a typical Spizella in almost every respect and why it has been 
considered an "Aimophila" is incomprehensible. 

van Rossem— Birds of South-central Arizona 145 

1907 Mearns, E. A. 

Mammals of the Mexican Boundary of the United States. Bull. U. S. 
Nat. Mus., 56. 

1914 Swarth, H. S. 

A Distributional List of the Birds of Arizona. Pacific Coast Avifauna, 
No. 10. 

1923 Bailey, Florence M. 

Birds Recorded from the Santa Rita Mountains in Southern Arizona. 
Pacific Coast Avifauna, No. 15. 

1926 Bruner, Stephen C. 

Notes on the birds of the Baboquivari Mountains, Arizona. Condor, 28, 
pp. 231-238. 

1928 GrinnellJ. 

A Distributional Summation of the Ornithology of Lower California. 
Univ. Calif. Publ. Zool., 32, No. 1. 

1929 Swarth, H. S. 

Faunal Areas of Southern Arizona; a Study in Animal Distribution. 
Proc. Calif. Acad. Sci., 4th Ser., 18, No. 12. 

1930 van Rossem, A. J. 

The Sonora Races of Camptostoma and Platypsaris. Proc. Biol. Soc. 
Wash., 43, No. 18, pp. 129-132. 

193 1 Bishop, L. B. 

Sexual Dichromatism in the Pygmy Owl. Proc. Biol. Soc. Wash., 44, 
No. 24, pp. 97-98. 

1931 Griscom, Ludlow 

Notes on Rare and Little Known Neotropical Pygmy Owls. Proc. New 
England Zool. Club, 12, pp. 37-43. 

1931 Peters, J. L. 

Check-List of Birds of the World. Vol. 1. Harvard Univ. Press. 

1931 van Rossem, A. J. 

Report on a Collection of Land Birds from Sonora, Mexico. Trans. San 
Diego Soc. Nat. Hist., 6, No. 19. 

1932 Moore, Robert T. 

A New Race of Aimophila carpalis from Mexico. Proc. Biol. Soc. Wash., 
45, No. 63, pp. 231-234. 
1934 Campbell, Berry 

Bird Notes from Southern Arizona. Condor, 36, pp. 201-203. 

1934 van Rossem, A. J. 

Critical Notes on Middle American Birds. Bull. Mus. Comp. Zool., 77, 

No. 7. 
1934 Vorhies, Charles T. 

The White-necked Raven, a Change of Status? Condor, 36, pp. 118-119. 

146 San Diego Society of Natural History 


Fig. 1. Upper Sonoran oak and grass association in the Atasco Mountains. 
View looking northward down Piskiorski Canon. Although this region 
is inhabited chiefly by Upper Sonoran species, certain Lower Sonoran 
birds, such as Otocoris alpestris adusta and Sturnella magna lilianae, 
were found to be fairly common along the bare-topped ridges. 

Fig. 2. Grass plains along the eastern base of the Baboquivari Mountains. Here 
terminate, westwardly, the ranges of several Lower Sonoran species 
and subspecies of birds, among them Callipepla squamata pallida, 
Otocoris alpestris adusta, Corvus cryptoleucus, and Sturnella magna 
lilianae. The Upper Sonoran (oak) Zone of these mountains forms 
the western outpost of that association along the boundary. 

van Rossem — Birds of South-central Arizona 

Plate 18 



Fi 2 . 1 

Fig. 2 



Volume VIII, No. 19, pp. 149-184, plates 19-20, figs. 1-3, map 



Laurence M. Klauber 

Curator of Reptiles and A mphibians, San Diego Society of Natural History 


Printed for the Society 

May 29, 1936 




Laurence M. Klauber 

Curator of Reptiles and Amphibians, San Diego Society of Natural History 


Crotalus mitchellii, a rattlesnake of the southwestern United States 
and Lower California, is characterized by great variability of head scala- 
tion, pattern, and color. This morphological plasticity has rendered the 
determination of its status and relationships somewhat difficult. It is 
the purpose of this paper to present the results of a reinvestigation of the 
form, largely based on specimens lately acquired; especially worthy of 
mention is a series from the Cape region of Lower California, the first 
available from that area in sufficient numbers to permit a statistical 


Crotalus mitchellii was first described by Cope in 1861 ' as Caudisona 
Mitchellii, based on a specimen (USNM 5291^) collected by John 
Xantus at Cape St. (San) Lucas, Lower California. The snake was 
named in honor of Dr. S. Weir Mitchell, the famous Philadelphia neu- 
rologist, scientist, and author, who was at that time engaged in researches 
on rattlesnake venoms. In 1866 2 Cope described as Caudisona pyrrha a 
specimen (USNM 6606) collected by Dr. Elliott Coues at Canyon Prie- 
to, near Fort Whipple, Yavapai County, Arizona. 3 Cope recognized the 
affinity of this new form with mitchellii. 

Writing in 1893 (published in 1895) Stejneger 4 considered pyrrhus 
a subspecies of mitchellii. In 1894, with new material available from 
Lower California, Van Denburgh 5 showed that the differences which 

1 Proc. Acad. Nat. So. Phila., 1861, p. 293. 

2 Proc. Acad. Nat. Sci. Phila., 1866, p. 308. 

3 A graphic contrast between the difficulties faced by the collectors of the past, as com- 
pared with the facilities of the present, is afforded by Coues' reference to this specimen: "It is 
not in the best order, as it was procured under the untoward circumstances of a hasty retreat 
from hostile Indians." 

^ Rept. U. S. Nat. Mus. for 1893, 1895, p. 456. 
5 Proc. Cal. Acad. Sci., Ser. 2, Vol. 4, p. 450. 

152 San Diego Society of Natural History 

Cope had used to segregate pyrrhus from mitchellii failed to hold in a 
larger series; he therefore placed pyrrhus in the synonymy of mitchellii, 
where it has since remained. 

A closely allied species, Cro talus tigris, must now be surveyed; this 
was first described by Kennicott from southern Arizona in 1859. In 
reporting the results of the Death Valley Expedition, Stejneger 7 classified, 
as belonging to this species, certain rattlesnakes collected in east-central 
California and southern Nevada. In 1929 Amaral, 8 noting the similarity 
of this California-Nevada material with mitchellii, considered the latter a 
subspecies of tigris. 

The present writer 9 reviewed the situation in 1930 and reached the 
conclusion that these California-Nevada snakes were different from the 
true Arizona tigris. Noting border-line specimens from Mineral and 
Esmeralda Counties, Nevada, which showed certain affinities to Crotalus 
confluentus lutosus, I considered these as coming within the confluentus 
group, and applied to the snakes from California and Nevada, formerly 
considered tigris, the name Crotalus confluentus stephensi. As the rela- 
tionship between this form and mitchellii was evident, the latter became 
Crotalus confluentus mitchellii. 

This classification led to a curious anomaly, namely, the presence 
of two subspecies in the same territory without intergradation, for, in 
southern California, Crotalus confluentus oreganus and C. c. mitchellii 
occupy extensive areas together. Of the ring oreganus-lutosus-stephensi- 
mitchellii connecting these forms, clearly the lutosus-stephensi link was 
the weakest. With the considerable additional material that has come 
to hand I am now inclined to the belief that the Mineral-Esmeralda 
(Nevada) specimens are pure stephensi, rather than lutosus-stephensi in- 

Gradually new material, especially in the Museum of Vertebrate 
Zoology, has brought the lutosus and stephensi ranges together in other 
areas and this without any approach toward intergradation; that is, the 
border specimens of one species do not show a tendency toward the other 
form, each retaining its individuality. While an actual overlap in ranges 
has not yet been shown, thus finally settling the lutosus-stephensi non- 
intergradation beyond question, still such an overlap is indicated as 

6 Rept. U. S. Mex. Boundary Survey, Vol. 2, 1859, p. 14. 

7 North American Fauna, No. 7, 1893, p. 214. 

8 Bull. Antivenin Inst. Am., Vol. 2, 1929, p. 82. 

9 Trans. S. D. Soc. Nat. Hist., Vol. 6, No. 3, 1930, p. 95. 

Klauber — The Speckled Rattlesnake 153 

probable, particularly in the Belted Range in Nevada. Thus it is my 
present conclusion that mitcbellii (and with it stephensi) should be di- 
vorced from the conflnentus group. 

At the same time the augmented collections of the past four years 
from other areas, particularly the large series from the Cape region of 
Lower California, together with a study of additional characters, have 
convinced me that these Cape specimens are different in several par- 
ticulars, especially head size and rattles, from those found in Arizona 
and California and therefore pyrrhus should be revived as the subspecific 
name for the latter form. 

Concerning the relationship of the mitchellii group with tigris I have 
not changed my previously expressed opinions. 10 While mitchellii (espe- 
cially as exemplified by the Cape subspecies) is closely allied to tigris, 
there is an actual overlap of the ranges of the two species, without inter- 
gradation, in central Arizona. Tigris is a species which, as far as now 
known, is restricted to central and southern Arizona, and Sonora; its 
differences from stephensi are quite definite and consistent. 

Thus at present I view mitchellii as comprising three subspecies as 
follows : 

Crotalus mitchellii mitchellii: Cape and central areas of Baja 

Crotalus mitchellii pyrrhus: Central and southwestern Arizona; the 
Californias from central San Bernardino County south to the 
Sierra San Pedro Martir of Baja California. 

Crotalus mitchellii stephensi: California, east of the Sierras from 
Inyo County south to central San Bernardino County; south- 
western Nevada. 

In the outline which follows there are presented first, descriptions 
of these forms, their ranges and morphology, followed by a discussion of 
the characters in which they differ from, or resemble, each other and 
closely allied species. In these data advantage is taken of the enlarged 
collections now available and of studies of variation which have been 
made preparatory to a general summary of rattlesnake characters. 

Mitchellii is a peculiar snake in that a tendency to subdivision of 
its head scales renders their classification difficult; also it is, amongst all 
the rattlesnakes, the most variable in color and pattern. The separation 

10 Trans. S. D. Soc. Nat. Hist., Vol. 6, No. 3, 1930, p. 106; ibid. Vol. 6, No. 24, 
1931, p. 353. 

154 San Diego Society of Natural History 

of the rostral from the prenasals by a row of granules or small scales, long 
thought to be a simple and universal criterion of mitchellii, is not always 
present in this form, even if we omit the subspecies stephensi from con- 
sideration; also this separation sometimes occurs in certain other species, 
notably C. c. oreganus from Arizona. Thus a simple and invariable key 
character for this group is not available. 

Mitchellii has often been called the White, Bleached, or Faded Rat- 
tlesnake, especially based on specimens from southwestern Arizona, which 
happen to have been among the first to be brought east alive. But from 
many sections it is brightly colored and strongly, although indefinitely, 
patterned. I deem it best to refer to the two more typical subspecies as 
Speckled Rattlesnakes, for the punctations which constitute the pattern 
are its most outstanding characteristic. 

Crotalus mitchellii mitchellii (Cope) 

San Lucan Speckled Rattlesnake 

Plate 19, fig. 1 

1861 Caudisona Mitchellii Cope, Proc. Acad. Nat. Sci. Phila., 1861, p. 293. 

(Type locality: Cape St. (= San) Lucas, Lower California, Mexico. 

Type specimen: USNM 5291V2 1 ! collected by John Xantus.) 
1875 Crotalus mitchellii Cope, in Yarrow Surv. W. 100th Merid., Vol. 5, p. 535. 

1882 Crotalus mitchelli Yarrow, Bull. U. S. Nat. Mus., No. 24, pp. 12, 73. 

1883 Crotalus oreganus var. mitchelli Garman, Mem. Mus. Comp. Zobl., 

Cambr., Vol. 8, No. 3, p. 173. 
1887 Crotalus mitcheli Belding, West American Scientist, Vol. 3, No. 24, p. 

1887 Crotalus mitchillii Cope, Bull. U. S. Nat. Mus., No. 32, p. 90. 

1894 Crotalus mitchellii (part) Van Denburgh, Proc. Cal. Acad. Sci., Ser. 2, 

Vol. 4, p. 450. 

1895 Crotalus Mitchellii Mitchellii Stejneger, Rept. U. S. Nat. Mus. for 1893. 

p. 454. 

1929 Crotalus tigris mitchellii (part) Amaral, Bull. Antivenin Inst. Am., Vol. 

2, No. 4, p. 82. 

1930 Crotalus confluentus mitchellii (part) Klauber, Trans. S. D. Soc. Nat. 

Hist., Vol. 6, No. 3, p. 108. 
Material. — Of this subspecies 81 specimens from the Cape region of Baja 
California have been available for study, of which fifty were seen alive. In addi- 

1 ' This specimen has apparently disappeared; in fact, Dr. Stejneger says it had been 
removed from the USNM collection before his coming to the museum fifty years ago. It has 
not been located amongst the Gape material in the collection of the Academy of Natural Sci- 
ences of Philadelphia. 

Klauber — The Speckled Rattlesnake 155 

tion there are 5 specimens from three different Gulf of California islands; the 
latter however are omitted from the descriptive summary as it is not desired to 
complicate the statistics with possible incipient island subspecies. No specimens 
are available to me from the central area of the peninsula (the vicinity of Mulege, 
Santa Rosalia, and San Ignacio). There are five specimens in the Paris Museum 
from this area; of these Mr. F. Angel has kindly supplied me the measurements. 
These indicate that the speckled rattlesnakes of that vicinity, while intermediate 
between C. mitchellii mitchellii and C. m. pyrrhus, are somewhat closer to the 
former, so they have been included therewith. The affiliation is not unexpected, 
for in many kinds of reptiles the Cape species extend to the central area with a 
break between these and the San Diegan forms further to the north. Thus in the 
present instance we consider C. m. mitchellii to extend as far north as San Igna- 
cio. The most southerly available specimen of pyrrhus is from the Sierra San 
Pedro Martir; 12 we assume that, in this gap of about 250 miles, intergradation, 
which is begun in the San Ignacio area will be found complete, for this interven- 
ing territory is of a character suitable to C. mitchellii. 

Range. — C. m. mitchellii has been collected at the following localities in the 
southern and central areas of Lower California, and adjacent islands on the Gulf 
of California and Pacific coasts. (See map) . For a few localities approximate lati- 
tudes are given where the identity, through duplication of names or obscurity, 
may be confusing. 

Cape San Lucas (Type locality) San Evaristo (24°50') 

San Jose del Cabo Santo Domingo (25°30') 

Sierra El Taste Punta Escondido (25° 50') 

Sierra San Lazaro Mulege 

Miraflores Santa Rosalia 

Todos Santos (23° 30') San Ignacio 

La Rivera (Riberia) Ceralvo Island 

Agua Caliente (23°30') Espiritu Santo Island 

Ensenada de los Muertos (24°) San Jose Island 

La Paz Santa Margarita Island 

Lepidosis and Form. — Size medium among rattlesnakes. Scale rows at mid- 
body usually 25 (90 per cent) , rarely 23 (6 per cent) or 27 (4 per cent) . The 
scales are strongly keeled, except the first row on each side. Scale bosses are rather 
conspicuous. Ventrals: males, (52 specimens) max. 181, min. 165, av. 176.69± 
0.32 interquartile range 174.4 — 179.0, coefficient of variation 2.0 per cent; 
females, (27 specimens) max. 186, min. 172, av. 179.44±0.45, interquartile 
range 177.1 — 181.8, coefficient of variation 1.9 per cent. Anal entire. Caudals: 
males 28 to 22, average of 44 specimens 25.1 ±0.12; females 23 to 16, average of 
27 specimens 20.7±0.21. The caudals, while generally entire, may have a few 
at either end of the series divided. 

The supralabials usually number 15 (24 per cent), 16 (49 per cent), or 17 

12 An intermediate specimen from Las Huevitas (sometimes given as Huavitas or Cue- 
vitas) near San Fernando Mission, contained in the California Academy of Sciences collec- 
tion, was destroyed in the fire of 1906. 

156 San Diego Society of Natural History 

(16 per cent) ; occasionally 14 (4 per cent) or 18 (6 per cent) ; rarely 13 or 19 
(less than 1 per cent of each) . The infralabials generally number 15 (39 per cent) 
or 16 (37 per cent) ; occasionally 14 (10 per cent) or 17 (13 per cent) ; rarely 13 
(less than 1 per cent) . The posterior supralabials are often divided horizontally, 
thus being lower than the scales next above. 

The rostral is usually wider than high, as is characteristic of this species; how- 
ever in 10 per cent the width and height are equal and in 7 per cent it is higher 
than wide. As is usually the case with this species the prenasals are separated from 
the rostral by rows of scales or granules; only 3 per cent make contact. 

The prenasals are also usually separated from the supralabials by the exten- 
sion to the rostral of the small scales anterior to the pit; this separation is evident 
in 83 per cent of the specimens examined. The internasals cannot be counted with 
accuracy as they form part of the row of scales which, continued down along the 
sides of the rostral, separate that scale from the prenasal. In fact this tendency, 
which is so characteristic of mitcbellii, of splitting such scales as the prenasals, 
loreals, canthals, and preoculars, renders a statistical analysis of the head scales 
difficult and to some extent useless. Therefore, only the more important items are 

The scales on the crown anterior to the supraoculars vary from 25 to 46, in- 
terquartile range 31.5 to 37.9, average 34.7. The minimum scale rows between 
supraoculars vary from 1 to 8, interquartile range 4.7 — 6.3, with an average of 

Supraocular sutures or indentations are present in only 3 per cent of the 
specimens. The nasals are 2 — 2. About 60 per cent of the specimens have two 
loreals, the rest from 1 to 4. The scales along the canthus rostralis from rostral to 
supraocular usually number from 5 to 7. 

The upper preocular is frequently split horizontally, vertically, or both; there 
results confusion with both loreals and canthals so that the classification of these 
scales and the determination of their contacts, useful in the classification of some 
species, is here usually impossible. There is often a small, circular scale between 
the two preoculars; this is quite characteristic of this subspecies, but is not inva- 
riably present. 

The scale rows from labials to orbit usually number 3 or 4; the scales in the 
orbital ring average about 9. 

The first infralabials are undivided; normally, 2 or 3 are in contact with the 
genials on each side. 

The mental is triangular. The genials are in a single pair, relatively short 
and obtuse. Intergenials are present in only one per cent of the specimens and 
submentals in 2 per cent. 

In shape the head is subtriangular and depressed, as compared with most 
rattlesnake species. The average ratio of body length to head length in adults (over 
700 mm. in length) is 24.6. The ratio of the distance across the supraoculars to 
the space between averages 2.5. 

The average ratio of the length of tail to total length exclusive of rattle is 
0.080 in adult males and 0.063 in females. 

The largest specimen examined measured 958 mm. (39 in.) when alive. 

Klauber — The Speckled Rattlesnake 157 

Color and Pattern.— This snake is light-gray or tan in ground color with a 
dorsal series of large and highly irregular blotches which almost obscure the 

The blotches are neither clearly outlined nor of regular form; they consist 
partly of darker scales, but more conspicuously of large aggregations of black or 
dark-brown punctations. The blotch centers are usually somewhat lighter than 
the edges. Toward the tail the blotches become rings. Secondary and, occasion- 
ally, tertiary series of blotches are in evidence on the sides. The inter-blotch light 
areas are freest of punctations along the mid-dorsal line, but even here the ground 
color is seldom spotless. As a result, the live snake usually gives the impression 
of a quiet, gray or brown neutrality. The extreme color variations found in pyrrhus 
in California and Arizona do not seem to be present in this southern race. Below, 
the color is buff, with aggregations of dark spots. 

The head is spotted but is neither conspicuously nor regularly marked. 
There is a dark line from the eye to a point above the angle of the mouth. The 
outer edge of each supraocular usually has a conspicuous, light tip. 

The tail rings are alternating ash-gray and black, in some contrast to the rest 
of the body. The light are usually wider than the dark rings, thus following 
scutulatus rather than atrox. 

In number the body blotches vary from 26 to 39, the average being 31.7 
±0.23 in the males and 32.0±0.36 in the females. The coefficient of variation 
is about 8 per cent in this characteristic. 

The tail rings number 3 to 5 in the males, with an average of 3. 8 ±0.06, and 
3 or 4 in the females (average 3. 2 ±0.05) . 

Crotalus mitchellii pyrrhus (Cope) 

Southwestern Speckled Rattlesnake 

Plate 19, fig. 2 and Plate 20, fig. 1 

1866 Caudisona pyrrha Cope, Proc. Acad. Nat. Sci. Phila., 1866, p. 308. (Type 

locality: Canyon Prieto, Yavapai County, Ariz. Type specimen: 

USNM 6606 collected by Dr. E. Coues) . 
1875 Crotalus pyrrhus Cope, in Yarrow, Surv. W. of 100th Merid., Vol. 5, 

p. 535. 
1883 Crotalus conjluentus var. pyrrhus Garman, Mem. Mus. Comp. Zobl., 

Cambr., Vol. 8, No. 3, p. 173. 

1894 Crotalus mitchellii (part) Van Denburgh, Proc. Cal. Acad. Sci., Ser. 2, 

Vol. 4, p. 450. 

1895 Crotalus Mitchellii pyrrhus Stejneger, Rept. U. S. Nat. Mus. for 1893, 

p. 456. 

1896 Crotalus mitchelli (part) Boulenger, Cat. Snakes, Brit. Mus., Vol. 3, p. 

1922 Crotalus goldmani Schmidt, Bull. Am. Mus. Nat. Hist., Vol. 46, Art. 
11, p. 701. (Type locality: El Pinon, 5300 ft., San Pedro Martir Mts., 
Lower California. Type specimen: USNM 37573 collected by Nelson 
and Goldman) . 

158 San Diego Society of Natural History 

1929 Crotalus tigris mitchellii (part) Amaral, Bull. Antivenin Inst. Am., Vol. 

2, No. 4, p. 82. 

1930 Crotalus confluentus mitchellii (part) Klauber, Trans. S. D. Soc. Nat. 

Hist., Vol. 6, No. 3, p. 108. 
Material. — I have had a considerable field and reptile-house experience with 
this form, particularly in southern California, and have seen in excess of four hun- 
dred live specimens. The following preserved specimens have been available for 


Yavapai County 14 

Maricopa County 7 

Yuma County 3 

Mohave County 3 27 


San Bernardino County 
Riverside County 
Imperial County 
San Diego County 

Northern Lower California: 
Angel de la Guarda Island: 

Total 185 

Range. — C. m. pyrrhus is extensively distributed in rocky situations in cen- 
tral and western Arizona, southern California, and northern Lower California. 
(See map) . It is certainly present but has not yet been recorded in northwestern 
Sonora. The known localities of collection are as follows: 

Yavapai County: Yuma County: 

Canyon Prieto, near Ft. Whipple Mohawk Mts. 

( Type local ity ) Tule Mts . 

Drake Tinajas Altas Mts. 

Hillside Gila Mts. 

Date Creek Gonzales Well 

Congress Junction 20 mi. N. of Picacho 

Maricopa County: 5 mi. N. of Mohawk, near 

Wickenburg Gila River 

8 mi. SE. of Wickenburg Mohave County: 

Hot Springs Junction 2 mi. SW. of Valentine 

Cave Creek Chemehuevis Mts. 

Black Canyon, 25 mi. N. of Foot of The Needles 

Phoenix Topock 

Estrella Mts. 

Mts. S. of Phoenix 

Near Phoenix 









Klauber — The Speckled Rattlesnake 


San Bernardino County: 
3 mi. N. of Topock Bridge 

Mountain Spring 

(N. end Piute Mts.) 
14 mi. NE. of Blythe Junction 
Gilroy Canyon, Providence Mts. 
Turtle Mts. 
Twentynine Palms 
Key's Ranch 
Windmill Tank 
Old Woman Spring 

9 mi. S. of Old Woman Spring 
Cushenbury Spring 
Cushenbury Grade 
Bet. Cushenbury Spring and 

Lake Baldwin 
Johnston Ranch below 

Lake Baldwin 
Forest Home 
Lucerne Valley 
Deadman's Point 
12 mi. S. of Barstow 
Oro Grande 

Riverside County: 
Palen Mts. 

10 mi. N. of Desert Center 
6 mi. SW. of Shaver's Well 
5 mi. E. of Mecca 

1 mi. S. of Indian Wells 

Palm Springs 

Murray Canyon near Palm Canyon 

Mouth San Andreas Canyon 


Santa Rosa Mts. 

Coahuila Mt. 

Nightingale Ranch 


Asbestos Spring 

Poppet Flat 

Vanderventer Flat 

Pinyon Flat 



Riverside County (continued) : 
Schain's Ranch Road, 5 mi. W. 

of Banning 
Between Idyllwild and Keen Camp 
S. Fork, San Jacinto River 

Temescal Canyon 
San Juan Canyon, Elsinore Mts. 
5 mi. E. of San Juan Hot Springs 

Orange County: 

3 mi. E. of Hot Springs in 
Elsinore Mts. 
Imperial County: 

Near Picacho 

Near Seeley 

Coyote Wells 

Myers' Creek Bridge 

Mountain Spring 

Boulder Park 

San Diego County: 
Point Loma 
Mission Valley 
Fall brook 

Mission Gorge 
El Monte 

Lakeview (Johnstown) 
EI Cajon 

Warners Ranch 
Valley Center 
Lake Wohlford 
Santa Ysabel 
San Vicente 
Shady Dell 
Padre Barona 
EI Capitan Dam 
Boulder Creek 


San Diego Society of Natural History 

San Diego County (continued) 
Twin Brooks 
Pine Valley 
Glen Lonely 

Lawson Valley 
Lyons Valley 
Barrett Dam 
Deerhorn Flat 
Clover Flat 
Live Oak Springs 

San Diego County (continued) 
Palomar Mt. 
Cuyamaca Mt. 
Laguna Mt. 
Laguna Junction 
Coyote Canyon 
Collins Valley 
Culp Valley 
Borego Palm Canyon 
Tubbs' Spring 
San Felipe Valley 
Sentenac Canyon 
La Puerta (Mason Valley) 
Grapevine Spring 
Yaqui Well 
The Narrows 
Carrizo Gorge 
Near Mountain Spring 
Dos Cabezas 


There are, in the literature, two Los Angeles County records, namely, Fair- 
mont and Lovejoy Springs. It is not improbable that these are the results of inac- 
curate identifications, particularly the former, and therefore they are not regu- 
larly listed above. 

Baja California 

East Base, Cocopah Mts. 

Volcano Lake 

Garcia (S. D. & A. Ry.) 

Redondo (S. D. & A. Ry.) 


8 mi. E. of Valentin 

4 mi. N. of San Pedro (32°5') 

Laguna Hanson 

Descanso (32° 15') 

San Antonio (near Socorro) 
San Matias (31° 15') 
San Jose (31°) 
Parral (30°40') 

El Pinon, San Pedro Martir Mts. 
(Type locality of goldmani) . 
Las Huevitas (Lat. 30°) 
Angel de la Guarda Island 

Lepidosis and Form. — Size large among rattlesnakes. Scale rows at midbody 
usually 25 (71 per cent), occasionally 23 (19 per cent), or 27 (10 per cent). 
The scales are strongly keeled, except the first row on each side. Ventrals: males 
(110 specimens), max. 185, min. 168, av. 177.56±0.21, interquartile range 175.3 
— 179.8, coefficient of variation 1.9 per cent; females (53 specimens), max. 187, 
min. 163 (170 if one aberrant specimen be omitted), av. 179.25 ±0.39, inter- 
quartile range 176.4 — 182.1, coefficient of variation 2.3 per cent. Anal entire. 
Caudals: males 28 to 20, average of 110 specimens 23.7±0.10; females 23 to 
16, average of 51 specimens 19.1zh0.17. The caudals, while generally entire, may 
have a few at either end of the series divided. 

Klauber — The Speckled Rattlesnake 161 

The supralabials vary from 13 to 19; they usually number 15 (25 per cent), 
16 (36 per cent) , or 17 (27 per cent) ; occasionally 14 (5 per cent) or 18 (6 per 
cent) ; rarely 13 or 19 (less than 1 per cent of each) . The infralabials generally 
number 15 (20 per cent), 16 (39 per cent), or 17 (26 per cent); occasionally 
14 (7 per cent) or 18 (6 per cent) ; rarely 13 or 19 (less than 1 per cent of each) . 

The rostral is usually wider than high; however, this is by no means the im- 
portant characteristic it has been often assumed, since 18 per cent of the speci- 
mens examined were higher than wide and 30 per cent were equal in the two di- 
mensions. The prenasals are normally not in contact with the rostral, this lack of 
contact being highly characteristic of the present species. However, in 6 per cent of 
the specimens examined, contact is made on one side and in 12 per cent on both; 
thus 82 per cent run true to form, but in the others this key character partly or 
entirely fails. The prenasals are in contact with the supralabials in 24 per cent of 
the cases; the contact is entirely prevented in 32 per cent by the extension to the 
rostral of the small scales anterior to the pit, and the contact is partly prevented 
by these same scales in the remaining 44 per cent. The internasals are indetermin- 
ate since the nasals do not ordinarily contact the rostral. 

The scales on the crown anterior to the supraoculars number at least 21 and 
average about 35. The minimum scale rows between supraoculars vary from 4 to 
8, interquartile range 5.2 — 6.5, average 5.9. 

Sutures or indentations are present in 9.3 per cent of the supraoculars. The 
nasals are 2 — 2. The Ioreals are often identified with difficulty; they vary from 
to 4, with an average of about 1.5. 

The upper preocular is frequently split, horizontally, vertically, or both; in 
fact, in only 28 per cent of the cases is it entirely intact, and in no less than 34 
per cent is split in both directions. 

The scale rows from labials to orbit usually number from 2 to 4. There are 
from 7 to 12 scales in the orbital ring, most specimens having 8 or 9. 

The first infralabials are often divided (22 per cent divided, 78 per cent un- 
divided) . 

The mental is triangular. The genials are in a single pair, relatively short 
and obtuse. Intergenials are present in 7.8 per cent of the specimens; submentals 
in only 1.4 per cent. 

In shape the head is subtriangular, and depressed as compared with most 
species of the genus. The average ratio of body length to head length in adults 
(over 700 mm. in length) is 21.5. 

The ratio of the length of tail to total length exclusive of rattle is approxi- 
mately 0.072 in adult males, and 0.057 in females. 

The largest specimen examined measured 1295 mm. (51 in.); the smallest 
303 mm. (12 in.). 

Color and Pattern. — This is the most variable of all the rattlesnakes in color 
and pattern; its bewildering variety renders any considerable accuracy or consis- 
tency of description quite impossible. 

The ground color may be white, cream, tan, buff, drab, gray, brown, pink, 
orange, or salmon. On this there is superimposed a series of blotches which may 
approach hexagons, hour-glasses (with transverse axes), diamonds, rectangles, or 
cross-rings. The blotches consist partly of dark-colored scales and partly of dark 

162 San Diego Society of Natural History 

punctations; these two forms of color application may blend or be in strong con- 
trast. The blotches may be pink, salmon, red, brown, gray, black, or mixtures of 
these. They are usually highly irregular and indefinite in outline. There is also 
present a secondary series of blotches on each side; these may be of the same or 
twice the frequency of the main series. Caudad the two series are confluent to 
form transverse rings. A tertiary series is sometimes present. 

The ground color is usually freest from punctations along the mid-dorsal 
line; along the sides gray punctations and even a gray suffusion are likely to be 
present, this being particularly the case with Mohave Desert specimens toward 
the stephensi range. Black scale tips are often present as in stephensi. Sometimes 
the blotches have light centers. The lower surfaces are cream, buff, or pink, usually 
blotched or punctated. The head is irregularly blotched or spotted. A postocular 
dark line is sometimes present. Supraocular light cross-dashes or light outer edges 
are often in evidence. 

The tail rings are frequently in considerable color contrast with the body 
blotches, the terminal rings being usually black, even though the body be pink. 
An ash-gray ground color on the tail, of the atrox and scutulatus type, may be 

In some areas this snake maintains a rather consistent coloration. Thus, in 
Yavapai County, Arizona, from Wickenburg north it is almost invariably a beau- 
tiful pink or salmon-red, a color which, by-the-way, fades badly in preservation, 
so that these snakes must be seen alive to observe the full effect. Pink, however, 
is not a universal Arizona coloration, since the mountains to the south of Phoenix 
produce light grayish-green specimens, while those from the Yuma sector are tan. 
In the Elsinore Mountains of Orange and Riverside counties, California, there 
is found a color variety in a beautiful shade of orange, with blotches of burnt- 
orange. In some sections of the Mohave Desert the snakes are russet-brown. 

The lightest specimens I have seen came from the Tinajas Altas Mts., Yuma 
County, Arizona. In these the ground color is creamy-white; individuals from this 
area originally caused mitchellii to be known as the "White Rattlesnake." (Plate 
20, fig. 1 ) . It is said that these snakes blend well with a white granite found in that 

However, that these color varieties are not always territorially consistent is 
shown by the snakes of San Diego County, where, although grays and browns pre- 
dominate, pink, salmon-red, and buff individuals are also present. Occasionally 
specimens are found with black scales or blotches on a light-gray ground; this form 
has locally been well termed the "granite rattler." 

The body blotches number from 23 to 42, the average for the males being 
33.7±0.18 and for the females 32.9±0.28. The tail rings average 5.6±0.06 in 
the'males (range 4 to 9) , and 4.4dz0.08 in the females (range 3 to 6) . 

Crotalus mitchellii stephensi Klauber 

Panamint Rattlesnake 
Plate 20, fig. 2. 
1893 Crotalus tigris Stejneger, North American Fauna, No. 7, p. 214. 
1896 Crotalus tigris (part) Boulenger, Cat. Snakes Brit. Mus., Vol. 3, p. 580. 

Klauber — The Speckled Rattlesnake 163 

1929 Crotalus tigris tigris (part) Amaral, Bull. Antivenin Inst. Am., Vol. 2, 

No. 4, p. 82. 

1930 Crotalus conflucntns stepbensi Klauber, Trans. S. D. Soc. Nat. Hist., 

Vol. 6, No. 3, p. 108. (Type locality: 2 mi. W. of Jackass Springs, 

6200 ft., Panamint Mts., Inyo County, Calif. Type specimen: MVZ 

6699 collected by Dr. Jos. Grinnell) . 
1933 Crotalus mitcbellii (part) Stejneger and Barbour, Check List N. A. 

Amph. and Repts., 3d Ed., p. 136. 
Material. — Of this subspecies there have been available for study, the fol- 

Nevada : 

Mineral County 1 

Esmeralda County 6 

Nye County 6 

Clark County 3 16 

California : 

Mono County 1 

Inyo County 43 

Kern County 1 

San Bernardino County 5 50 

Total 66 

About thirty individuals of this form have been seen alive. 
Range. — C. m. stepbensi is an inhabitant of the desert-mountain region lying 
east of the crest of the Sierra Nevada from Round Valley, Mono Co., California 
and southern Mineral County, Nevada, south to central San Bernardino County, 
California. (See map) . The southern boundary may be roughly indicated by a 
line drawn from Barstow, California to Searchlight, Nevada along which line it 
intergrades with pyrrhus. Eastward it ranges at least to the Belted Mts., Nye 
County, Nevada, and Boulder Dam, Nevada; its presence beyond the Colorado 
River is not yet established. The following are the known localities of collection: 

Mineral Cou nty : Nye Cou nty : 

Endowment Mine, Excelsior Mts. Grapevine Mts., above Salt Wells 

Esmeralda County: Bullfrog 

7 Mi. N. of Arlemont Near Oak Spring, Belted Range 

McAfee Ranch (near Mono O/2 Mi - S., 1 M. S., 4 Mi. 

Co. border) SE., l/ 2 Mi. NW.) 

1.7 Mi. S. of Goldfleld Clark County: 

Lida Indian Spring Valley 

7 Mi. S. of Tonopah Las Vegas Valley 

Las Vegas Wash 
Boulder Dam Site 
Harris Spring, Charleston Mts. 

164 San Diego Society of Natural History 


Mono County: Inyo County (continued): 

N. end Round Valley Junction Ranch 

Inyo County: Hanaupah Canyon, Panamint Mts. 

Rocky Creek above Round Valley (Hananpole or Hannopee) 

Birch Creek, W. of Bishop Johnson Canyon, Panamint Mts. 

Bishop Creek Goler Canyon, Panamint Mts. 

(6, 6V2, 7, 71/2, 9, 10, 1 1 and Near Ballarat (6 mi. S.; 7 and 

12 mi. W. of Bishop) 10 m i. SW.) 

2 Mi. S. of Aberdeen Shepherd Canyon, Argus Mts. 

2 Mi. W. of Independence Little Lake 

Independence Creek San Bern ardino County : 

Mesquite Spring (N. end Slate Range, NW. of Borax Flat 

Death Valley) Willow Creek, Panamint Mts. 

Beveridge Canyon 5 Mi. S. of Cave Spring 

Lone Pine 4 Mi. NE. of Randsburg 

Carroll Creek Odessa Canyon, Calico Mts. 

2 Mi. W. of Jackass Spring Near Baker 

(Type locality) Coolgardie 

Emigrant Canyon, Panamint Mts. Yermo 

Coso Valley Kern County : 

Dante's View, Black Mts. Last Chance Canyon 

Wild Rose Spring Near Mohave 

Maturango Spring 6 Mi. E. of Brown 

Lepidosis and Form. — Size medium among rattlesnakes. Scale rows at mid- 
body usually 23 (73 per cent), occasionally 25 (25 per cent), rarely 21 (2 per 
cent) . The scales are keeled, except the first row on each side. Ventrals : males 
(32 specimens), max. 181, min. 162, av. 174.44±0.50, interquartile range 171.6 
— 172.2, coefficient of variation 2.4 per cent; females (22 specimens), max. 182, 
min. 173, av. 178.72±0.40, interquartile range 176.8 — 180.6, coefficient of vari- 
ation 1.6 per cent. Anal entire. Caudals: males 28 to 23, average of 33 specimens 
25.0±0.16; females 22 to 17, average of 22 specimens 19.4zb0.21. The caudals, 
while generally entire, may have a few at either end of the series divided. 

The supralabials usually number 13 (20 per cent), 14 (33 per cent), or 15 
(39 per cent); rarely 12 or 16 (4 per cent of each). The infralabials generally 
number 14 (38 per cent), 15 (42 per cent), or 16 (11 per cent); occasionally 
13 (4 per cent) , 17 (3 per cent) , or 18 (2 per cent) . 

The rostral is usually wider than high (equal in 13 per cent) and is in con- 
tact with the prenasals, although in some cases the prenasals are sutured, thus 
starting the granules which are so characteristic of mitchellii and pyrrhus. The 
prenasals are normally in contact with the supralabials but such contact is pre- 
vented, in 17 per cent of the specimens examined, by the extension to the rostral 
of the small scales anterior to the pit; there is partial interference in an additional 4 
per cent. The internasals (scales in contact with the rostral between the nasals, 
regardless of size or relative position) usually number two; owing to the splitting 

Klauber — The Speckled Rattlesnake 165 

off of upper corners of the prenasals there is one instance of three and another of 
four; this out of a total of 60 specimens. 

The scales on the crown anterior to the supraoculars vary from 13 to 38 and 
average 25.3. The minimum scale rows between supraoculars vary from 3 to 8, 
interquartile range 4.9 to 6.1; the average is 5.5 

Supraocular sutures are highly characteristic of this subspecies; they are 
present in 96 per cent of the specimens. These sutures are of considerable variety; 
often they are whorls or longitudinal cuts, particularly at the outer edge. Some- 
times these edges are rough as if pieces of scale had been broken away (text fig. 
2). The nasals are divided. The loreals vary from 1 to 5, averaging 1.98; there 
are two in 41 per cent of the specimens, one in 37 per cent, and greater numbers 
in 22 per cent. The scales along the canthus rostralis, from internasal to supra- 
ocular, usually number 3, sometimes 2 or 4. 

The upper preocular is divided horizontally in 4 per cent, and vertically in 
18 per cent of the specimens. This is a division which is still more evident in the 
other members of the mitchellii group. 

The upper preocular, which is the larger, is not in contact with the postnasal. 
In 46 per cent such contact is prevented by the contact of the postcanthal with 
the loreal, in 54 per cent by the presence of a small upper loreal. 

The scale rows from labials to orbit usually number 2 or 3. Generally the 
third and fourth, or fourth and fifth supralabials are in contact with the pit bor- 
ders; there is no conspicuous difference in size amongst the supralabials. 

The first infralabials are undivided. The mental is triangular. The genials 
are in a single pair, relatively short and obtuse. Intergenials are present in 4 per 
cent of the specimens; submentals are not in evidence. 

In shape the head is subtriangular, flat-topped and low. The average ratio 
of body length to head length in adults is 22.8. 

The ratio of the length of tail to total length, exclusive of rattle, averages 
0.079 in adult males and 0.059 in females. 

The largest specimen examined measured 885 mm. (35 in.), the smallest 
257 mm. (10 in.) . A specimen 860 mm. long weighed 376 grams. The smallest 
female with eggs measured 674 mm. 

Color and Pattern. — This subspecies is highly variable in color and pattern, 
as if the several desert mountain ranges which it inhabits had produced individual 
races. The ground color may be straw, tan, buff, yellow-brown, red-brown, gray, 
or blue-gray. Upon this there is superimposed a series of darker dorsal blotches 
which, while usually subhexagonal in shape, may approach circles, diamonds, 
squares, or rectangles. They are buff, gray, brown, or deep red-brown, and may, 
or may not, be sharply contrasting with the ground color. Some of the contrasts 
and harmonies, particularly in the browns, are very striking, rendering this one of 
the handsomest of rattlesnakes, exceeded only by enyo and molossus. 

There is a secondary series of blotches on the sides; at about mid-body the 
main dorsal series contacts these so as to form rings, which become narrower and 
less sharply in contrast with the ground color toward the tail. At the neck several 
dorsal blotches may coalesce to form a longitudinal band. The inter-blotch areas 
are lighter dorsally than laterally. Anteriorly the sides are often suffused with a 

166 San Diego Society of Natural History 

punctated application of gray (even on the brown specimens), a character found 
in pyrrhus as well. The dorsal blotches are sometimes even-edged, but are more 
often serrated by a border of unicolor scales in the manner of scutalatus. Some- 
times the light scales bordering the blotches are conspicuously lighter than the 
rest of the ground color. Often the light scales, immediately anterior to the 
blotches, are posteriorly tipped with black; this is, in fact, quite characteristic of 
the present subspecies. 

Northern specimens are darker and more conspicuously and definitely 
marked than those from the south; the latter more nearly approach the amorphous 
punctations of pyrrhus. Red-brown or dark-brown specimens are the rule from the 
northwestern corner of the range in the vicinity of Bishop Creek, California. In 
the northeastern corner, in Nevada, blues and grays predominate. The south- 
eastern specimens are generally tan, blotched with brown. 

The ventral surface is usually buff or tan, with aggregations of darker punc- 

The head is spotted, but less conspicuously and regularly than the body. 
Supraocular light cross-marks are usually absent, but the outer edge of each supra- 
ocular may be light. There is often a dark dash from the eye to a point above the 
corner of the mouth, but in many specimens this is obscured by a characteris- 
tic suffusion of gray, which, in the tan or brown specimens, is in rather sharp 
contrast with the dorsal color. 

The tail rings are usually distinct, but the last two or three, which are 
generally black and in strong color contrast with the rest of the body, are not 
evenly outlined, and may be partly confluent. 

The body blotches vary in number from 30 to 43, the average being 37.0. 
The tail rings number 6 to 9 (average 6.8) in the males, and 3 to 6 (average 4.6) 
in the females. 


Having presented separately, for each of the three subspecies of mitchellii. 
the essential details of lepidosis, form, and appearance, I deem it desirable to 
combine the enumerations of other characteristics, in order to avoid repetition 
under each subspecies, since the differences between the three forms are not con- 

Habitat. — All three subspecies of mitchellii are essentially rock-dwelling 
snakes, and to a large extent, but not entirely, are restricted to the wastes of the 
southwestern deserts. Here, however, they are not universally distributed; from 
the level, sandy plains and the great alluvial fans they are generally absent, 
their habitat being in the rocky mountains and buttes which rise above the plains. 
Thus, while the range of mitchellii is roughly coincident with those of C. cerastes, 
C. scutulatus, and C. atrox in parts of Arizona and California, they are not par- 
ticularly active competitors, for locally there is often a rather sharp distinction in 
the habitat which each species prefers. 

For example, in the Panamint and Death Valley regions we find stephensi 
in the lava flows, rocky buttes, and the mountains themselves (to altitudes of at 
least 7000 ft.) rather than in the plains between, the latter being inhabited by 
cerastes. In the Mohave Desert we find cerastes and scutulatus on the flats> 

Klauber — The Speckled Rattlesnake 167 

and pyrrbus in the mountains, and the same is true in southwestern Arizona 
where atrox is included with the plains species. From the Imperial and Coa- 
chella Valleys pyrrhus seems absent, although present in all the surrounding 

In the Peninsula ranges of southern and Lower California, including the 
San Bernardinos, San Jacintos, Cuyamacas, Sierra de Juarez, and Sierra San 
Pedro Martir, we find pyrrhus ranging into Upper Sonoran habitats and even 
touching Transition. Yet, in the chaparral and oaks, it continues to show its 
preference for rocky outcrops, although not restricted to them. Here it reaches 
altitudes of at least 5300 ft. in areas shared with Crotalus ruber and C. c. 
oreganus. But while the latter are equally common westward to the ocean, 
pyrrhus rarely is found below the 1200 ft. contour on the coastal slope. 

It is in these mountains also that mitchellii, in the subspecies pyrrhus, reaches 
its maximum mainland size, specimens slightly exceeding 1220 mm. (48 in.) hav- 
ing been accurately measured. Such an individual would weight about 2 1 /2 lb. 
(1130 g.). This size is not approached by either stephensi or m. mitchellii. 13 
These, of course, are exceptional specimens; I would consider 1100 mm. (43 in.) 
as representing a "large" adult male pyrrhus in the foothills of San Diego 
County. In Arizona and, in fact, throughout the truly desert areas, this 
size is not reached, few adults exceeding 920 mm. (36 in.). The Angel de la 
Guarda Island specimens are very large, reaching at least 1240 mm. (49 in.). 
Stephensi is a smaller snake than pyrrhus, specimens exceeding 850 mm. (33 J /2 
in.) being exceptional. M. mitchellii, while smaller than pyrrhus, sometimes ex- 
ceeds 900 mm. (35 in.) and one specimen 939 mm. (39 in.) has been noted. 
However, most of the adult males run about 850 mm. (331/> in.) in length. 

From a few areas we have some statistics as to the relative frequency of oc- 
currence of mitchellii compared with the other species with which it shares the 
range. Thus, in four lots of rattlers brought up from the vicinity of Cape San 
Lucas there were 69 m. mitchellii or 17.6 per cent, out of a total of 391 rattlers; 
6.4 per cent were enyo, the rest being lucasensis. 

In San Diego County, out of 2006 rattlesnakes recorded, only 260 or 12.6 
per cent were pyrrhus, for this form was greatly exceeded in number by ruber and 

Along the line of the Santa Fe Ry. between Hillside and Wickenburg, Yava- 
pai County, Arizona, a collection of 454 rattlers yielded 19 pyrrhus or 4.2 per 
cent. The majority were atrox and scutulatus (55.6 and 31.9 per cent) with a few 
molossus and oreganus (4.6 and 3.7 per cent). Since these specimens were col- 
lected by track maintenance crews it is to be expected that the rock inhabiting 
forms would not be as well represented as those preferring the flat-lands. 

Thus, it is seen that where mitchellii is in competition with other forms it 
usually constitutes only a small part of the rattlesnake population. Several species 
of rattlers occur in most of the territory inhabited by pyrrhus and mitchellii 

13 A specimen (USNM 64588) 1295 mm. (51 in.) long is contained in the National 
Museum. The locality of collection is uncertain, but it is recorded tentatively as Cedros Island. 
This I rather doubt, since subsequent collectors have failed to find it there, although other 
rattlers (C. exsul) are not uncommon. 

168 San Diego Society of Natural History 

although they may have the rocky prominences to themselves. Stephensi, on the 
other hand, is the sole possessor of most of its range, or, at least, shares it only 
with cerastes, with which it is hardly in competition, so clearly does the one prefer 
valleys and sand, and the other rocky slopes and mountains. 

There is evidence that where mitchelln shares its territory with others it does 
not den with them. Thus, a reliable observer reported having seen some twenty 
pyrrhus in a shallow cave under a large granite boulder in San Diego County; 
there seemed to be no other rattlers present, although both ruber and oreganus 
are more common than pyrrhus in that particular place. In another instance, in 
December, nine rattlers were found under a single stone; all were young adult 

Breeding. — Specimens of stephensi have been noted with 6 and 8 eggs; 
pyrrhus with 3, 4 (2 specimens) , and 5. Experience with other species leads us to 
believe that these pyrrhus broods were smaller than the average for the species. 

Habits. — In the field pyrrhus is a distinctly more nervous species than ruber; 
rather it resembles oreganus in its alert readiness to defend itself or escape. It will 
usually rattle if alarmed. Mrs. G. O. Wiley, however, reports it as easily tamed 
as other species, if not more so. 

As is the case with most of the western rattlers, particularly the desert forms, 
mitchellii is largely nocturnal, especially in summer. However, it will be found 
abroad in the daytime in spring, and to a less extent in autumn. Thus, a specimen 
of stephensi was seen to issue from a hole at 6:30 p. m., in August. In late March 
specimens were observed crossing the road at Bullfrog, Nevada, at 3:00 p. m., 
and near Ballarat, at 6:25 p. m. In the spring, in San Diego County, specimens 
of pyrrhus have been observed sunning themselves before rock clefts in which 
they took refuge so promptly as to indicate advance consideration of these retreats. 

Food. — The snakes of the mitchellii group, as is usual with the larger rattle- 
snakes, subsist principally upon rats, mice, and other small mammals. Amongst 
these, Dipodomys has been recognized. Young specimens are probably more ac- 
customed to lizards, and even the adults do not scorn this prey. Amongst the 
species noted in the stomach contents were Uta stansburiana, Cnemidophorus 
tessellatus, and Eumeces skiltonianus . One large specimen contained both mam- 
mal hair and a Uta. One large mitchellii was observed to eat a ground squirrel 
which had been shot some three hours before. 

Birds are eaten occasionally. Thus Mr. Dean E. Batchelder informed me he 
had found 8 birds, presumably goldfinches, in the stomach of a pyrrhus. All had 
been swallowed head first. This was near an aqueduct construction camp where a 
lawn, garden, and bird bath had been installed on the desert 10 miles north of 
Desert Center, Riverside Co., Calif. Birds had been attracted from great distances 
and evidently the snake had lain in wait for them. 

Hemipenes. — The hemipenes of the mitchellii group may be described thus: 
Completely bifurcate with divided sulcus. Base covered with short, stiff spines, 
particularly at the outer shoulders; the branches covered with reticulate fringes. 
There is a sharp cleavage between spines and fringes (a characteristic of Crotalus 
as compared with Sistrurus, with the partial exception of C. lepidus) . The apices 
are calyculate. In shape the two lobes are of medium weight (ratio of length to 
diameter about 2.6) similar to the conjluentus group; this is in contrast to the 

Klauber— The Speckled Rattlesnake 169 

attenuated organs which characterize the atrox group, or the opposite extreme, 
illustrated by the short heavy lobes of molossus. The spines cannot be counted 
with accuracy, since they vary by imperceptible degrees from mere pustules to full 
points. The fringes vary from 27 to 34, averaging about 31. 

One of the important variations in the hemipenes of Crotalus is the presence 
or absence of spines in the crotch between the lobes. These spines are present in 
all subspecies or mitchellii. They are especially conspicuous in mitchellii mitchelln 
and stephensi, somewhat less so in the intermediate specimens (pyrrhus) from 
California, and are least perceptible in pyrrhus from Arizona. 

Fangs. — The fangs of mitchellii, in shape and size, follow closely the con- 
jluentus group, that is, they are shorter proportionately than those of terrificus, 
adamant eus, and the atrox group. Average adult ratios of body and head length 
to fang length (measured from the lower edge of the upper lumen to the tip) 
are as follows: 

Fang Length Ratios 

L/F H/F 

Mitchellii 151 5.74 

Pyrrhus 107 5.01 

Stephensi 128 5.59 

Here again we have a substantial difference between mitchellii and pyrrhus, 

although one which is obviously correlated with proportionate head size. 

Venom. — The physical characters of the venoms of these three subspecies 
are indicated in the following table: 

Venom Characteristics 

Specimens milked 

Average yield dry venom per adult snake, mg. 

Maximum yield, mg. 

Specific gravity 

Average MLD 

The MLD is taken from data kindly furnished by Dr. Thos. S. Githens of 
the Mulford Biological Laboratories of Sharp and Dohme, and represents the 
fatal dose for a pigeon of 350 g. weight. From results in other species we would 
expert mitchellii, with its small head and short fangs, to have a relatively power- 
ful venom, as compared with pyrrhus. 14 

Rattles. — Studies, as yet unpublished, indicate that the characteristics, and 



















H Githens (Journal of Immunology, Vol. 29, p. 171, Aug. 1935) reports two qualities 
of mitchellii venom : a "strong" with an MLD of 0.04 and a "weak" with an MLD of 0.50. 
The venoms which yielded these results were sent to the biological laboratory from San Diego. 
Through correspondence with Dr. Gthens I have ascertained the lot numbers involved in the 
several assays and checking the original data I find that three assays yielding an average 
MLD of 0.50 contained only pyrrhus venom. One assay resulting in an MLD of 0.04 was 
based exclusively on mitchellii venom. Only one assay tends in any way to upset the theory 
that the venoms of pyrrhus and mitchellii differ considerably, the latter being the more toxic. 
This assay, on a mixture of mitchellii and pyrrhus venoms sent in before the subspecies were 
distinguished, also yielded an MLD of 0.04, whereas some figure between the two might 
have been expected. I am unable to explain this unless the more powerful venom tends to 
mask the weaker. 











































170 San Diego Society of Natural History 

particularly certain measurements, of the rattles, are of considerable interest in 
classification. These dimensions are found to be quite consistent, and from them 
it is possible to verify relationships and differences. As an instance, the width of 
each rattle (where the string is complete and the rattle-number in the sequence 
is therefore known) is found of value in classification. In the present study we 
have the following data: 

Average Width of Rattle in Tenths of Millimeters 

Rattle No. 
Pyrrhus (Cal.) 
Pyrrhus (Ariz.) 

The figures are not particularly trustworthy beyond the fifth rattle for two 
reasons; first, a sufficient number of specimens is not available to afford accurate 
averages; and, secondly, sexual dimorphism begins to affect the result beyond the 
sixth rattle, and therefore for accuracy of diagnosis it is necessary to treat the sexes 
separately. However up to the sixth rattle, except in the case of stephensi, of which 
comparatively few specimens, even of the first rattles, are available, the figures 
are quite reliable. These studies, to be presented elsewhere, show that, within geo- 
graphically homogeneous groups, the coefficient of variation of the rattle widths 
of the first five rattles usually runs from 5 to 7 per cent and rarely exceeds 9 per 
cent. Under such circumstances differences such as those indicated in these tables, 
particularly between mitchellii and pyrrhus, can be shown mathematically to be 
highly significant. 15 

While it is not believed advisable to attempt any species differentiation exclu- 
sively based on rattle divergences, these certainly should not be neglected as con- 
firmatory evidence. 

As I have stated before, mitchellii cannot always be distinguished from the 
other rattlesnakes by its best known character, namely, the separation of the ros- 
tral from the prenasals by small scales or granules; for this character is not uni- 
versally positive in either of the southern races, m. mitchellii and pyrrhus, and it 
is always negative in stephensi. Furthermore, this prevention of contact is of 
sporadic occurrence in other forms, particularly in oreganus from central Arizona. 
Nor is the depression of the head, while somewhat evident, the clear-cut and obvi- 
ous character which it has been occasionally considered in the past. The same is 
true of the shape of the rostral, which is not always wider than high. 

In general, it can be said that these snakes are conspicuous amongst the rat- 
tlesnakes for the tendency to subdivision of certain head scales (prenasals, can- 
thals, loreals, and preoculars in mitchellii and pyrrhus; and supraoculars in 
stephensi) , and for the punctated application of color and pattern. Yet there is 
sufficient variation and divergence in all of these characters so that simple and 
invariable keys are impossible; therefore, considerable judgment and some experi- 

15 Employing the usual formula for significance by evaluating the ratio which the dif- 
ference between the means bears to the standard error of the difference. 

Klauber — The Speckled Rattlesnake 171 

ence are necessary in their application if errors are not to be made. However I do 
not think that this is an adverse criterion of the validity of these forms, since the 
summation of the differences is impressive; and some differences which are not 
particularly useful as key characters, such as the form of the hemipenes, never- 
theless are important in the final determination of validity. 

The nearest existing relatives to mitchellii are tigris, cerastes, and confluen- 
tus (the latter through the subspecies oreganus and lutosus), probably in the or- 
der named. These affinities are shown in form, scalation, pattern, and hemipenes, 
as will be pointed out. 

The characters of lepidosis which are usually employed statistically in dis- 
tinguishing species of snakes, are not of particular value in separating the mem- 
bers of the jjiitchellii group, either from each other or from some of their near 
relations, for a number of rattlers, including the several subspecies of confluentus 
and mitchellii, as well as scutulatus and tigris do not differ greatly in these charac- 
ters. However, a few tendencies are to be noted. 

In scale rows, 25 is the mode in m. mitchellii and pyrrhus, but this is reduced 
to 23 in stephensi, which, in this particular, resembles tigris. About half of the 
Arizona specimens of pyrrhus have 23 scale rows, a much higher percentage than 
is encountered in the California specimens. 

Mitchellii is a rough-scaled species, the dorsal scales being sharply ridged 
and with raised bosses posterior to the middle of each scale. The latter are more 
conspicuous in mitchellii mitchellii, and in southern California pyrrhus, than in 
pyrrhus from Arizona or stephensi. However, in none of these snakes are the 
bosses so extreme as in cerastes, which is outstanding in this character amongst 
our southwestern rattlers; and even this form is exceeded in the prominence of 
these dorsal scale protuberances by dunssus. The extreme spinal ridge of durissus 
is also absent in mitchellii, enyo most nearly approaching the tropical rattler in 
this character. 

In ventrals mitchellii and Arizona pyrrhus average slightly lower than Cali- 
fornia pyrrhus, but the difference is not conspicuous; stephensi is also low. In this 
character tigris is conspicuously different from stephensi, as set forth in the fol- 
lowing table: 

Average Ventral Scale Counts 

Males Females 
Mitchellii 176.7 179.5 

Pyrrhus (Southern California) 178.3 180.0 

Pyrrhus (Arizona) 174.4 175.4 

Stephensi 174.4 178.7 

Tigris 165.2 168.5 

The extent of the difference between stephensi and tigris, as indicated by the 
ratio of the difference between the means to the standard error of the difference, is 
9.2 for the males and 9.3 for the females. In calculations of this kind any ratio 
above 3 is usually considered significant; 9 may be taken as conclusive on this 
point, that is to say the difference is real and cannot be attributed to chance differ- 
ences in sampling. The differences in caudal scale counts are not important; ste- 
phensi and mitchellii are slightly higher than pyrrhus. 

It is interesting to note that there is less sexual dimorphism in ventral scales 


San Diego Society of Natural History 

in mitchellii than is apparent in most rattlesnake species. In the mitchellii sub- 
species the females average about 3 more ventrals than the males; the confluentus 
group and many other rattlers run from 4 to 7. 

In labial counts we find pyrrhus somewhat higher than mitchellii, which, in 
turn, exceeds stephensi. None of the differences is sufficient to be of interest as a 
key character. 

Divided first infralabials are quite characteristic of Arizona pyrrhus, being 
present in 70 per cent of the specimens; in California pyrrhus they are frequent 
in San Bernardino County but are rarely met with elsewhere in the range of 
pyrrhus, or in mitchellii or stephensi. Intergenials and submentals are occasion- 
ally noted, but are not characteristic of any of these forms. 

Of the other head scales, such as the internasals, canthals, inter-supraoculars 
(frontals), nasals, loreals, and oculars, whose numbers, contacts, and arrange- 
ments are so frequently of value in rattlesnake diagnosis, we can in the mitchellii 
group make little use because of their tendency to be split to such a degree that 
their classification becomes impossible. For instance, where there is so frequently 
a row of scales between prenasal and rostral (as in mitchellii and pyrrhus) the 
internasals are indeterminate; canthals and loreals cannot be separated; preocu- 
lars are broken vertically, horizontally, or both (text fig. 1), and thus are con- 
fused with loreals. But a few differences of importance may be noted in these 
scales which may be summarized as follows: 

1. Supraocular sutures (text fig. 2) are highly characteristic of stephensi 
and will serve to differentiate this form from tigris, and, to a lesser extent from 
lutosus. Stated statistically we have the following: 

Fig. 1 

Fig. 1. Crotalus mitchellii mitchellii. Lateral view of head showing subdi- 
vision of preoculars, loreals, and supralabials. 

Fig. 2. Crotalus mitchellii stephensi. Dorsal view of head showing supra- 
ocular sutures. 

Fig. 3. Crotalus mitchellii pyrrhus. End view of head showing absence of 
contact between rostral and prenasals. 

Klauber — The Speckled Rattlesnake 173 

Supraocular Sutures 
(two counts per specimen) 




Sutures Doubtful 


2 2 

82 2 


168 4 


Lutosus (Cal. and Nev. only) 
Supraocular sutures are not infrequent in mitchellii and pyrrhus; they are 
quite prevalent in specimens from northern Baja California. 

2. Internasals are determinate in stephensi and will distinguish this form 
from lutosus. Thus in 60 specimens of stephensi, one had 3 and one had 4 inter- 
nasals; all others had 2. On the other hand, in 103 specimens of lutosus from 
California and Nevada there were only 4 specimens with less than 3 internasals, 
the complete score being as follows (the first number indicating the number of 
internasals, the second the number of individuals) : 1/1, 2/3, 3/28, 4/62, 
5/6, 6/3. Only California and Nevada specimens of lutosus were used in this 
comparison, instead of snakes from all areas, upon the theory that the former, 
occupying a territory close to that of stephensi, are the ones most necessary to 
differentiate from stephensi. It may be of interest to note that the two aberrant 
specimens of stephensi (having more than two internasals) are from near pyrrhus 
territory and have an increased number of internasals by reason of the splitting of 
the prenasals (a characteristic of pyrrhus) , rather than of the internasals them- 
selves, as in lutosus and all the confluentus group. 

3. The rostral-prenasal interruption (text fig. 3), while not a universal key 
character to mitchellii and pyrrhus as was once supposed, is, nevertheless, usually 
indicative. Thus, it is positive in 97 per cent of mitchellii, and about 88 per cent 
of pyrrhus. Most of the aberrant specimens of pyrrhus occur in southern Califor- 
nia, approaching the territory of stephensi; this is to be expected in view of the 
intergradation of the two forms. In stephensi there are no cases of the failure of 
the rostral prenasal contact, and the same is true of tigris. The interruption of 
this contact, thus shown to be characteristic (but not universally so) of mitchellii 
and pyrrhus, does occur in some other species, particularly the subspecies of con- 
fluentus; it is most prevalent in oreganus from central Arizona, where about 12 
per cent of the specimens have this contact interrupted. Lack of contact has also 
been observed in abyssus and in California oreganus. 

4. It is characteristic of all the mitchellii subspecies and tigris that the ros- 
tral is wider than high; the contrary is usually true in all the confluentus sub- 
species. However, the two dimensions are not infrequently equal, especially in 
specimens of pyrrhus from southern California, so that at best this is but a con- 
firmatory character. Occasionally specimens of pyrrhus have the rostral higher 
than wide. 

The great variability in pattern and color of all subspecies of mitchellii ren- 
ders these characteristics of little use in classification. Outstanding are the punc- 
tulate application of color in mitchellii and pyrrhus and the indefiniteness of the 
pattern; the gray suffusions on the lateral areas of stephensi and northern pyrrhus; 
the black posterior scale tips in many stephensi and some pyrrhus; the scutulatus 

174 San Diego Society of Natural History 

type of tail rings in many specimens of pyrrhus and mitchellii. But none of these 
characters is the sole possession of mitchellii subspecies; even the punctations are 
seen to be highly developed in tigris, atrox (although in a different manner), 
confluentus from New Mexico (Cope's pulverulentus) , triseriatus, and others. 

In number of body blotches and tail rings both pyrrhus and mitchellii fall 
below stephensi, and this, in turn, below tigris. However, the dispersion of these 
characters is so great that they are not useful as keys. 

Coming now to form, we observe that pyrrhus is a somewhat heavier bodied 
snake than the other two subspecies. Proportionately it has a slightly shorter tail 
than either mitchellii or stephensi. But none of these is conspicuously different 
from the atrox or confluentus groups. 

It is in head size that really important differences amongst the mitchellii sub- 
species are evident; here we have probably the most essential divergence between 
mitchellii and pyrrhus, and this in a character which must be presumed to be rela- 
tively stable. 

The determination of the statistical significance of differences in such 
a character as the ratio of the head to body length is a somewhat involved 
problem. Such an investigation has been made, but its exposition would be out of 
place at this point, and is reserved for future publication. The conclusions con- 
cerning rattlesnakes may be summarized as follows: 

1. The relationship of the size of the head to the body conforms closely to 
a linear equation of the form H = a L+b, the constants a and b being different 
for each of the several species or subspecies. 

2. Owing to the presence of the constant term b in the regression equation, 
the ratio of body length to head length is not constant throughout life; young 
rattlesnakes have proportionately larger heads than adults. 

3. There is a close correlation between head and body size in any one form, 
the coefficient of correlation probably being about +0.85 to 0.90. The dispersion 
about the regression line approximates the normal curve of error. The dispersion 
remains nearly proportional throughout life, although increasing slightly with 
age. The coefficient of variation about the regression line is from 2.5 to 3.5 per 
cent in homogeneous groups. 

To simplify the problem in the present instance we can restrict our investiga- 
tion to adult groups of approximately the same body size; if this be done the con- 
stant term b may be neglected, since the ratio L/H will be practically constant 
for each subspecies in any short length-range. The results of such a study of re- 
stricted adult groups are as follows, all specimens having been measured under 
the same conditions of preservation: 

Adult Head-Length Ratios 



Head Length 

Number of 


A verage 

A verage 














Pyrrhus (Calif.) 






Pyrrhus (Ariz.) 


















Klauber — The Speckled Rattlesnake 175 

Thus it is seen that there is an essential difference in this character (which 
our investigations lead us to consider stable and consistent) between pyrrhus and 
mitchellii, and between stephensi and tigris. It will be noted that the average size 
of the specimens of mitchellii is slightly less than the average for California 
pyrrhus, and the tigris average is less than stephensi. Since L/H increases with 
growth, if the average length were the same in the two cases the differences in 
the L/H ratios would be slightly greater than is shown in the table. 

As a further proof of the difference between mitchellii and pyrrhus the re- 
gression lines for the L — H relationships were determined; all specimens were re- 
duced to the same standard body size and the hypothetical head length for each 
specimen was calculated on the assumption that the deviation of any individual 
from the regression line for that subspecies remains constant (in percentage) 
throughout life. By this method it was determined that the ratio of the difference 
in the means of the head sizes to the standard error of the difference was over 17, 
which indicates that the difference is a real one and not attributable to the acciden- 
tal composition of the group available for study (i. e., sampling errors) . On the 
other hand between Arizona and California pyrrhus there was found to be no 
significant difference, the corresponding ratio being only 0.4. 

Tigris has proportionately the smallest head amongst all the rattlesnakes; 
it is of interest to note that it is closely approached by mitchellii; on the other hand 
pyrrhus and stephensi approximate the rattlesnake mode in the body-head ratio. 

That these differences in the head dimensions between mitchellii and pyrrhus, 
and between stephensi and tigris are not merely an exaggeration of a minor sta- 
tistical deviation will be at once apparent to any one having the opportunity to 
compare adult specimens of these forms in life. It is to be remembered that there 
are proportionately similar differences in head width and in depth as well as in 
length; there is therefore a difference in bulk approximately equal to the cube of 
any linear dimension. The result is so striking that there is no difficulty in segre- 
gating these forms at a glance when adults of approximately the same body length 
are available. 

As might be expected, fang length is closely correlated with head length 16 
and thus the fangs repeat the differences observed in the ratio of head size to 
length of body overall. But here we find that the differences are even more im- 
pressive, for tigris and mitchellii not only have shorter fangs than stephensi and 
pyrrhus because of their smaller heads, but have, in fact, fangs which are dispropor- 
tionately shorter even where snakes of the same head size are taken. With relation 
to the body lengths overall, the discrepancy is still greater. This is apparent from 
the following table showing data for the subspecies under consideration and their 

Average Fang-Length Ratios 
L/F H/F L/F H/F 

Mitchellii 151 5.74 Tigris 165 6.11 

Pyrrhus 107 5.01 Lutosus 131 5.49 

Stephensi 128 5.59 Cerastes 98 4.83 

(L = body-length overall; H = length of head; F= length of fang, upper lumen 
to point) . 

16 The regression equation is of the form F=a H - b. 

176 San Diego Society of Natural History 

The resemblance of mitchellii to tigris rather than to pyrrhus, and of 
stephensi to lutosus rather than to tigris is at once evident. 

Venom differences, at present imperfectly known, have already been set forth. 
I would anticipate that mitchellii would again show an affinity to tigris rather 
than pyrrhus, but its MLD has only been tentatively determined. Githens reports 
tigris venom to be the most toxic of any of the 22 subspecies of rattlers thus far 

The study of the rattles of the rattlesnake, from the standpoint of species 
differences is so involved that publication of these data must be reserved for an- 
other place. Suffice it to say that mitchellii is well differentiated from pyrrhus. 
Tigris more nearly resembles pyrrhus than stephensi. Mitchellii has the largest 
rattles of any known rattlesnake, up to and including the fifth rattle, a most sur- 
prising fact in view of its relatively small size. It considerably exceeds its relative 
pyrrhus and is in fact approached most closely by adamanteus and lucasensis. The 
first of these being the largest of rattlesnakes, its possession of large rattles is not 

The hemipenial characteristics of the mitchellii group afford the clearest 
differentiation from the confluentus group, for although both of these have organs 
of somewhat the same ratio of lobe length to diameter (thus sharply different 
from the attenuated organs of the atrox group on the one hand and the globular 
habitus of molossus on the other) the confluentus group is without spines in the 
central crotch while these are always present in the mitchellii group. Here pyrrhus 
shows a closer affinity to lutosus and oreganus than do mitchellii or stephensi, for 
the latter have patches of spines in the cleft while in pyrrhus (especially those 
from Arizona) the spines in this area are usually few and not particularly large. 
In this character tigris resembles stephensi; cerastes also has a cleft-patch. 

The following Gulf of California island specimens are available for study: 
Angel de la Guarda 5 

San Jose 1 

Espiritu Santo 3 

Ceralvo 1 

It is interesting to survey these for indications of incipient differentiation 
from the mainland forms, although there are not sufficient specimens upon which 
to premise descriptions of new species unless the differences be very wide. 

First, we find, based on the most important difference (head proportion), 
between mitchellii and pyrrhus, that the Angel de la Guarda specimens are 
pyrrhus, while those from San Jose and Ceralvo are mitchellii, which is quite what 
would be expected geographically. The Espiritu Santo specimens are between the 
two mainland forms, although somewhat nearer mitchellii than pyrrhus; in ratde 
dimensions they are somewhat closer to the latter, but for the present we will con- 
sider them as mitchellii. 

Klauber — The Speckled Rattlesnake 177 

In ventrals the Angel de la Guarda specimens are distinctly higher than most 
of the specimens of pyrrhns, while those from the other islands have fewer scales 
than the average of mitcbellii. There are indications that both differences would 
be significant, were enough specimens available to determine the dispersions of 
the island specimens more accurately. The caudals show no deviations of interest. 
The labials are rather low in the Angel de la Guarda specimens, and supraocular 
sutures are prevalent, showing in this last character an affinity for the specimens 
from the mountains of Lower California. 

The Angel de la Guarda specimens are very large; they are pink or straw 
and usually have black scale tips; in color they resemble Arizona specimens rather 
than those from California. It is quite possible, I think, that USNM 64588, the 
largest known pyrrhus, may have come from Angel de la Guarda instead of 
Cedros Island, as has been presumed. 17 

The other island specimens are also pink, with the exception of that from 
Ceralvo, which is gray. We conclude that these island forms have been so long 
separated from the mainland that differentiation has begun, but it is not sufficient 
to permit taxonomic separation, at least not until much larger series are available. 

It is unfortunate that the type specimen of mitcbellii (USNM 5291 1 /2) 
has been lost and that Cope did not give the head measurement. He records this 
snake as being 44 in. (1118 mm.) long, which is so much larger than any other 
specimens from the Cape that we are disposed to doubt the accuracy of the local- 
ity. As was so often the case in those days the specimen was probably recorded 
from the locality from which it was sent, rather than the point of collection. The 
color and pattern descriptions also do not fit mitcbellii. 

We know from the tail length that the specimen was a male. The ventral 
scale count, given as 198, is so much higher than any other Cape male specimen 
(max. 181) that the specimen was either a freak or there has been a miscount. 
Calculations indicate an extremely small chance of such a high count, assuming 
that the dispersion follows the normal curve of error (which other studies indi- 
cate to be the case) , for the deviation of this specimen is over six times the stand- 
ard deviation. We conclude that the derivation of this specimen from Cape San 
Lucas is exceedingly doubtful, and if all the facts were known a shifting of names 
might be necessary. 


Crotalus mitcbellii is a valid species of rattlesnake containing three 
well differentiated subspecies. C. m. mitcbellii, C. m. pyrrhus, and C. m. 
stephensi. Of other existing species their closest relatives are C. tigris, 
C. cerastes, C. c. oreganus, and C. c. lutosus. C. m. mitcbellii shows an 
affinity to tigris; while pyrrhus and stephensi have important resemblances 
to oreganus and lutosus. 

17 Schmidt, 1922, p. 700. 

178 San Diego Society of Natural History 


I am indebted to the following individuals and institutions for the 
loan of important material, or other favors in connection with this inves- 
tigation: Mr. J. R. Slevin of the California Academy of Sciences; Drs. 
Leonhard Stejneger and Doris M. Cochran of the United States Na- 
tional Museum; Drs. A. W. Herre and W. H. Rich of Stanford Uni- 
versity; Drs. Jos. Grinnell and J. Linsdale of the Museum of Vertebrate 
Zoology, University of California, Berkeley; Mr. K. P. Schmidt, Field 
Museum of Natural History; Mr. H. R. Hill, Los Angeles Museum; 
Dr. R. B. Cowles and Mr. Chas. M. Bogert, University of California at 
Los Angeles; Pomona College; M. F. Angel, Museum National D'His- 
toire Naturelle, Paris; Dr. A. H. Wright, Cornell University; Mr. R. 
Conant, Philadelphia Zoological Society; Dr. T. Barbour and Mr. A. 
Loveridge of the Museum of Comparative Zoology, Harvard University; 
Dr. G. K. Noble, American Museum of Natural History; Dr. E. H. 
Taylor, Kansas University; Mrs. H. T. Gaige and Mr. H. K. Gloyd, 
University of Michigan; Mr. H. W. Fowler, Academy of Natural Sci- 
ences, Philadelphia; Mrs. Belle Benchley and Mr. C. B. Perkins, Zoologi- 
cal Society of San Diego. 

The following have presented me with live or preserved specimens 
which have been of the greatest service in this investigation: C. C. Lamb, 
J. R. Pemberton, Miss Ada Meling, the late A. H. Schlanze, Maj. Chap- 
man Grant, Mrs. G. O. Wiley, A. P. Artran, C. M. Perkins, Chas. M. 
Bogert, L. H. Cook, T. C. Biggs, G. W. Kuns, M. E. Spivey, Thos. 
Fitzmorris, C. L. Evans, Dr. C. E. Burt, F. E. Walker, C. L. Davis, E. L. 
Bulpitt, H. K. Gloyd, P. M. Klauber, O. N. Arrington, A. N. Handley, 
R. R. Humphrey, Dr. E. H. Taylor, C. B. Perkins, Dr. C. T. Vorhies. 

Of the utmost importance have been the specimens received from 
the Santa Fe Railway Company in central Arizona, and the live collec- 
tion brought to the San Diego Zoological Society from the Cape region 
of Lower California by Fred Lewis of the Yacht Stranger. Without the 
latter large and uniform series the differences between mitchellii and 
pyrrhns would not have been so apparent. 

I have been greatly assisted by Messrs. L. H. Cook, Robert Hoard, 
and P. M. Klauber in the making of scale counts, and by Mrs. Elizabeth 
Leslie and Miss Eileen Carmody in tabulations and computations. The 
sketches are by Mr. Norman Bilderback, and the map and photographs, 
by Mr. L. C. Kobler. 

Klauber — The Speckled Rattlesnake 179 


There are listed here certain papers important in the history of the classifi- 
cation of Crotalus mitchellii, and our knowledge of the character, range, and 
habits of the species. 

Amaral, A. do 

1927 Crotalus goldmani Schmidt, 1922, A Synonym of C. mitchelli Cope, 
1861. Bull. Antivenin Inst, of Amer., Vol. 1, No. 2, pp. 47-48. 

1929 Studies of Nearctic Ophidia. Ill — Notes on Crotalus tigris Kenni- 
cott, 1859. Bull Antivenin Inst, of Amer., Vol. 2, No. 4, pp. 82-85. 

Belding, L. 

1887 Reptiles of the Cape Region of Lower California. West Am. Scien- 
tist, Vol. 3, No. 24, pp. 97-99. 
Blanchard, F. N. 

1925 A Key to the Snakes of the United States, Canada and Lower Cali- 
fornia. Papers of the Michigan Academy of Science, Arts and Let- 
ters, Vol. 4, Part 2, pp. 1-65. 


1896 Catalogue of the Snakes in the British Museum, Vol. 3. 
Brown, A. E. 

1901 A Review of the Genera and Species of American Snakes, North of 
Mexico. Proc. Acad. Nat. Sci. Phila, Vol. 53, Part 1, pp. 10-110. 

Cope, E. D. 

1861 Contributions to the Ophiology of Lower California, Mexico and 
Central America. Proc. Acad. Nat. Sci. Phila., 1861, pp. 292-306. 

1866 On the Reptilia and Batrachia of the Sonoran Province of the Nearc- 
tic Region. Proc. Acad. Nat. Sci. Phila., 1866, pp. 300-314. 

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

Coues, E. 

1875 Synopsis of the Reptiles and Batrachians of Arizona; with Critical 
and Field Notes, and an Extensive Synonymy. Report upon Explo- 
rations and Surveys West of the 100th Meridian. Vol. 5, Chap. 5, 
pp. 585-633. 

Garman, S. 

1883 The Reptiles and Batrachians of North America. Mem. Mus. Comp. 
Zool., Vol. 8, No. 3, pp. xxxi+185. 
Kennicott, R. (in Baird, S. F.) 

1859 United States and Mexican Boundary Survey. Reptiles of the Boun- 
dary, pp. 1-35. 

Klauber, L. M. 

1927 Some Observations on the Rattlesnakes of the Extreme Southwest. 
Bull. Antivenin Inst, of Amer., Vol. 1, No. 1, pp. 7-21. 

1930 New and Renamed Subspecies of Crotalus confluentus Say, with 
Remarks on Related Species. Trans. San Diego Soc. of Nat. Hist., 
Vol. 6, No. 3, pp. 95-144. 

180 San Diego Society of Natural History 

Klauber, L. M. (continued) 

1931a A Statistical Study of the Snakes of the Southern Border of Califor- 
nia. Bull. Zool. Soc. of San Diego, No. 8, pp. 1-93. 

1931b Crotalus tigris and Crotalus enyo, Two Little Known Rattlesnakes 
of the Southwest. Trans. San Diego Soc. of Nat. Hist., Vol. 6, No. 
24, pp. 353-370. 

Mearns, E. A. 

1907 Mammals of the Mexican Boundary of the United States. Bull. U- 
S. Nat. Mus., No. 56, pp. xv+530. 

Meek, S. E. 

1905 An Annotated List of a Collection of Reptiles from Southern Cali- 
fornia and Northern Lower California. Field Mus. Zool. Ser., Vol. 
7, No. 1, pp. 1-19. 


1899 Contribution a la Faune Herpetologique de la Basse-Calif ornie. 
Nouv. Arch. Mus. Nat. Hist., Ser. 4, Vol. 1, pp. 297-344 

Schmidt, K. P. 

1922 The Amphibians and Reptiles of Lower California. Bull. Am. Mus. 
Nat. Hist., Vol. 46, Art. 11, pp. 607-707. 

Stejneger, L. 

1891 Crotalus pyrrhus in California. West Am. Scientist, Vol. 7, No. 59, 
pp. 165-167. 

1893 Annotated List of the Reptiles and Batrachians Collected by the 
Death Valley Expedition in 1891, with Descriptions of New Species. 
North American Fauna, No. 7, pp. 159-228. 

1895 The Poisonous Snakes of North America. Rept. U. S. Nat. Mus. 
for 1893, pp. 337-487. 

Stejneger, L. and Barbour, T. 

1933 A Check List of North American Amphibians and Reptiles. 3d Ed. 
Harvard University Press, pp. xiv+185. 
Streets, T. H. 

1877 Contributions to the Natural History of the Hawaiian and Fanning 
Islands and Lower California. Bull. U. S. Nat. Mus., No. 7, pp. 
VanDenburgh, J. 

1894 Notes on Crotalus Mitchellii and "Crotalus Pyrrhus." Proc. Cal. 
Acad. Sci., Ser. 2, Vol. 4, pp. 450-455. 

1922 The Reptiles of Western North America. Occas. Papers Calif. 

Acad, of Sci., No. 10, Vol. 1, Lizards; Vol. 2, Snakes and Turtles, 

pp. 1-1028. 
Yarrow, H. C. 

1875 Report upon the Collections of Batrachians and Reptiles. Report 

upon Explorations and Surveys West of the 100th Meridian, Vol. 

5, Chap. 4, pp. 509-584. 

Klauber — The Speckled Rattlesnake 

Plate 19 


■ m 


Fig. 1. Crotalus mitchellii mitchellii. San Lucan Speckled Rattlesnake. 
Adult male, collected at La Rivera, Baja California, by C. C. Lamb. 

;, • ■; 


^■•:. •■ 

Fig. 2. Crotalus mitchellii pyrrhus. Southwestern Speckled Rattlesnake. 
Adult male, collected at Yaqui Well, San Diego County, California, by E. E. 

Klauber — The Speckled Rattlesnake 

Plate 20 

Fig. 1. Cro talus mitchellii pyrrhus. Southwestern Speckled Rattlesnake. 
Adult male, collected in Tinajas Altas Mts., Yuma Co., Arizona, by Dr. C. T. 

Fig. 2. Crotalus mitchellii stephensi. Panamint Rattlesnake. Adult male, 
collected 2 mi. S. of Aberdeen, Inyo Co., California, by F. E. Walker. 

&*8*6 ^ 



Volume VIII, No. 20, pp. 185-276, figs. 1-112 



Laurence M. Klauber 

Curator of Reptiles and Amphibians, San Diego Society of Natural History 


Printed for the Society 

December 7, 1936 



Volume VIII, No. 20, pp. 185-276, figs. 1-112 



Laurence M. Klauber 

Curator of Reptiles and Amphibians, San Diego Society of Natural History 


Printed for the Society 

December 7, 1936 



Introduction 187 

The Status of the Rattlesnakes 187 

List of Species and Subspecies 190 

Validity 193 

Nomenclature 193 

Condensed Alphabetical Synonymy 195 

Summary of Characters 197 

Use of Key 198 

Rattlesnake Ranges 200 

Summary of Rattlesnake Life History 201 

Habitats 201 

Economic Status 202 

Habits 202 

Food 204 

Size 205 

Reproduction 205 

Control 205 

Rattlesnake Enemies 206 

Commercial Value 206 

Senses 207 

Rattles 208 

Fangs 208 

Venom 209 

Snake Bite and Treatment 211 

The Biting Mechanism of the Rattlesnake (figures) 216 

Glossary 220 

Method of Employing the Key 226 

Acknowledgments 226 

Key 227 

Maps 250 

Photographs 257 

Index 276 



Laurence M. Klauber 

Curator of Reptiles and Amphibians, San Diego Society of Natural History 


The presentation of an identification key to the rattlesnakes appears 
desirable at this time since none is now available which incorporates the 
changes and additions resulting from recent taxonomic researches. The 
papers describing new subspecies are scattered in the literature and it would 
seem well to co-ordinate the results in a single publication in order that 
they will be more generally recognized by the nonspecialist in this field. 
Hence the publication of the key at this time, even though further changes 
resulting from investigations in taxonomy and nomenclature are to be 
expected in the future. 

Recent researches in rattlesnake venoms have disclosed some rather 
surprising species differences both in toxicity and physiological effects. 
Differences in venom strength of an order of 60 to 1 have been indicated ; 
and there appear to be differences in the relative proportions of hematoxins 
and neurotoxins. Under such circumstances, if our remedial technique 
is to be improved, it is important that species be differentiated in case 
reports. It is hoped that this key will serve a useful purpose in permitting 
physicians and others interested in the snake-bite problem to identify ac- 
curately the snake involved. 


The following summary concerning the status of the rattlesnakes is 
presented to permit the non-herpetologist to orient himself with respect 
to the position of the group herein discussed. 

Rattlesnakes comprise a number of species of venomous snakes be- 
longing to the family Crotalidae. They are found only in the Western 
Hemisphere and reach their greatest profusion in the southwestern United 
States and northern Mexico. All rattlesnakes have at least one rattle* 
even when born, and the user of this key is presumed to be certain that he 
has a rattlesnake at hand, since the key is not intended to be of service in 
identifying other snakes, whether venomous or harmless. 

Rattlesnakes and other members of the family Crotalidae are pit 
vipers, so called because of their possession of a sensory organ, probably 
auditory in purpose, in the form of a pit or deep depression, plainly 
visible on either side of the head below and back of the nostril (fig. 6) . 

* Of course occasionally a rattlesnake may lose the end of his tail through accident and with it his 
identifying rattles, which thereafter will not be regenerated. But the tail will remain only as a stump; 
no sharp tailed snake is a rattlesnake in disguise, as a surprising number of people seem to think possible. 

San Diego Society of Natural History 

In this the members of this family (Crotalidae) differ from the Old World 
vipers Of the family Viperidae, sometimes called the true vipers, which do 
not possess pits. The true vipers and pit vipers have this in common: 
the venom fangs are seated in a rotatable bone (the maxillary) whereby 
when not in use they lie folded back against the roof of the mouth, from 
which position they may, at will, be rotated forward and downward into 
the biting position perpendicular to the upper jaw. There are many forms 
of dangerously venomous snakes which are not vipers, including the 
cobras, mambas, coral snakes, etc.; these have shorter, permanently erect 
fangs, at the front of the upper jaw. 

By no means all of the members of the family Crotalidae are rattle- 
snakes. The pit vipers of this family are divided into the following 
genera, of which only the first two are rattlers: 





Lac bests 

Common Name 

Ground Rattlesnakes 

Characteristics Habitat 

Possess rattles; North & South 

scales on crown. America 

Possess rattles; North America 
plates on crown. 

Neotropical Pit Vipers Without rattles ; North & South 

scales on crown; America 
large posterior 


Trimeresurus Asiatic Pit Vipers 
Agkistrodon Moccasins 

Without rattles ; Central & South 
scales on crown; America 
small posterior 

Without rattles; Asia 

scales on crown. 

Without rattles; North America; 
plates on crown. SE. Europe; 

Some herpetologists, feeling that the genera Bothrops and Trime- 
resurus are insufficiently distinct to warrant separation, combine them 
under the latter name. 

The rattlesnakes comprise the most important group of venomous 
snakes to be found in the United States. The only dangerously venomous 
snakes occurring in this territory except rattlesnakes are the following: 

Mtcrurus fulvius fulvius (Linne), 1766. 
Southeastern Coral Snake. 

North Carolina to southern Florida; westward and southward along 
the Gulf lowlands into Mexico; northward in the Mississippi Valley 
to Ohio and Indiana. 

Klauber — Key to the Rattlesnakes 189 

Micrurus julvius barbouri Schmidt, 1928. 
South Florida Coral Snake. 

Extreme southern Florida. 
Micruroides euryxanthus (Kennicott), I860. 
Sonoran Coral Snake. 

Southern border of Arizona and New Mexico, and south into Mexico. 
Agkistrodon mokasen mokasen Beauvois, 1799. 
Eastern Copperhead. 

Massachusetts west to Illinois and from these south to northern 

Florida and eastern Texas, including the intervening states. 
Agkistrodon mokasen laticinctus Gloyd and Conant, 1934. 
Broad-banded Copperhead. 

Western and central Oklahoma southward to western and central 

Agkistrodon piscivorus (Lacepede), 1789. 
Water Moccasin 

Virginia south throughout Florida, westward along the Gulf to the 

Rio Grande and north in the Mississippi Valley to Illinois. 
Thus it will be seen that there are found in the United States four 
species (six subspecies) of venomous snakes other than rattlesnakes. It 
is true that there are other, moderately venomous snakes in the southwest, 
including certain species of the genera Leptodeira and Trimorphodon. 
However it is doubtful whether the bite of one of these would be dan- 
gerous to a human being. The fangs are in the back of the mouth and 
are grooved rather than hollow ; they are therefore not mechanically well- 
designed to inject venom into a wound in a large animal, although no 
doubt effective in subduing such small prey as lizards. 

The copperheads and moccasins, being pit vipers, can be readily 
recognized by the presence of the pit and by their possession of rotatable 
fangs in the front of the upper jaw. These criteria will serve to distin- 
guish them from various harmless snakes with which they are often con- 
fused, particularly certain water snakes of the genus Natrix. The coral 
snakes, small inoffensive-appearing creatures, may in their turn be recog- 
nized by the arrangement of the rings of their pattern. The coral snakes 
are ringed with black, red, and yellow; and it is characteristic of them 
that there is always a yellow ring between successive red and black rings. 
This is different from the color sequence in various harmless forms which 
are often taken for coral snakes, such as certain king snakes of the genus 
Lampropeltis, and other snakes of the genera Rhinocheih/s, Sonora, Chilo- 
meniscus, and Cemophora. 

The above remarks on the recognition of dangerous snakes apply 
only to the United States. In Mexico, and more especially in Central and 
South America, there are many more species of venomous snakes other 
than rattlesnakes ; for the latter do not occupy the dominant position there 
which they do in our country. As we go southward we encounter an 
increasingly complex variety of coral snakes (Micrurus) and a bewildering 
array (possibly 50 forms) of pit vipers of the genus Bothrops* to say 

* One of the most important of these tropical pit vipers is the fer-de-lance, Bothrops atrox, 

190 San Diego Society of Natural History 

nothing of the bushmaster, Lachesis muta, and another moccasin, Agkistro- 
don bilineatus. Also in these areas various back-fanged snakes become so 
large that they may well be considered dangerous to man, although I have 
heard of no serious accidents from their bites. Thus it is not as simple, 
south of our borders, to determine which snakes are venomous and which 
harmless, although the pit vipers can still be recognized by their pits, and 
most of the coral snakes by the color arrangements of their patterns. 

Having given this brief nontechnical survey of venomous snakes 
in the Americas other than rattlesnakes, we now return to the consideration 
of this group alone. 


In this key the following species and subspecies of rattlesnakes of 
the genera Crotalus and Sistrurus are recognized as valid: 

1. Crotalus durissus durissus Linne, 1758. 

Central American Rattlesnake. 
Southern Mexico to Costa Rica. 

2. Crotalus durissus terrijicus (Laurenti), 1768. 

South American Rattlesnake. 
Costa Rica to Argentina. 

3. Crotalus unicolor van Lidth de Jeude, 1887. 

Aruba Island Rattlesnake. 

Aruba Island, Dutch West Indies. 

4. Crotalus basiliscus (Cope), 1864. 

Mexican West-Coast Rattlesnake. 

West Coast of Mexico from Sinaloa to Oaxaca. 

5. Crotalus enyo (Cope), 1861. 

Lower California Rattlesnake. 
Central and southern Baja California. 

6. Crotalus molossus molossus Baird and Girard, 1853. 

Northern Black-tailed Rattlesnake. 

West Texas to central Arizona and south to northern Durango. 

7. Crotalus molossus nigrescens Gloyd, 1936. 

Southern Black-tailed Rattlesnake. 
Central Mexico from Durango to Puebla. 

8. Crotalus adamanteus Beauvois, 1799. 

Eastern Diamond Rattlesnake. 

Coastal plains of the southeastern states. 

9. Crotalus cinereous Le Conte in Hallowell, 1852. * 

Western Diamond Rattlesnake. 

Arkansas to southeastern California and south to San Luis 
10. Crotalus tortugensis Van Denburgh and Slevin, 1921. 
Tortuga Island Diamond Rattlesnake. 
Tortuga Island, Gulf of California. 

* Previously known as Crotalus alrvx Baird and Girard, 1853. 

Klauber — Key to the Rattlesnakes 191 

11. Cro talus lucasensis Van Denburgh, 1920. 

San Lucan Diamond Rattlesnake. 
Southern Baja California. 

12. Crotalus ruber Cope, 1892. 

Red Diamond Rattlesnake. 

Coastal southern California, and northern and central Baja 

13. Crotalus exsul Garman, 1883. 

Cedros Island Diamond Rattlesnake. 

Cedros Island off Baja California, Pacific side. 

14. Crotalus scutulatus (Kennicott), 1861. 

Mohave Rattlesnake. 

Southeastern California to west Texas, and south to the central 
Mexican plateau. 

15. Crotalus viridis viridis (Rafinesque) , 1818.* 

Prairie Rattlesnake. 

Western Great Plains from Alberta and Saskatchewan to extreme 
northern Mexico. 

16. Crotalus viridis nuntius Klauber, 1935. 

Arizona Prairie Rattlesnake. 
Northeastern Arizona. 

17. Crotalus viridis abyssus Klauber, 1930. 

Grand Canyon Rattlesnake. 
Grand Canyon of Arizona. 

18. Crotalus viridis lutosus Klauber, 1930. 

Great Basin Rattlesnake. 

The Great Basin between the Rockies and the Sierra Nevada. 

19. Crotalus viridis concolor Woodbury, 1929. 

Midget Faded Rattlesnake. 

Eastern Utah and western Colorado. 

20. Crotalus viridis oreganus Holbrook, 1840. 

Pacific Rattlesnake. 

Pacific Coast from British Columbia to central Baja California. 
Also Arizona. 

21. Crotalus mitchellii mitchellii (Cope), 1861. 

San Lucan Speckled Rattlesnake. 
Southern half of Baja California. 

22. Crotalus mitchellii pyrrhus (Cope), 1866. 

Southwestern Speckled Rattlesnake. 

Southern California, western Arizona, and northern Baja Cali- 

23. Crotalus mitchellii stephensi Klauber, 1930. 

Panamint Rattlesnake. 

Southern Nevada and east-central California, 

* Previously known as Crotalus confluentus cunfluentus Say, 182 3, 

192 San Diego Society of Natural History 

24. Crotalus tigris Kennicott, 1859. 

Tiger Rattlesnake. 

Southern Arizona, and northern and central Sonora. 

25. Crotalus cerastes Hallowell, 1854. 

Horned Rattlesnake; Sidewinder. 

Deserts of the southwestern United States and northwestern 

26. Crotalus polystictus (Cope), 1865. 

Mexican Lance-headed Rattlesnake. 
Tableland of central Mexico. 

27. Crotalus horridus horridus Linne, 1758. 

Timber Rattlesnake. 

Eastern United States, Maine to Oklahoma. 

28. Crotalus horridus atricaudatus Latreille, 1802. 

Canebrake Rattlesnake. 

Coastal plain of South Atlantic and Gulf states; lower Missis- 
sippi Valley. 

29. Crotalus lepidus lepidus (Kennicott), 1861. 

Eastern Rock Rattlesnake. 

From west Texas south to northern San Luis Potosi. 

30. Crotalus lepidus klauberi Gloyd, 1936. 

Green Rock Rattlesnake. 

Mountains of southern Arizona, southern New Mexico, and 
extreme west Texas south to Jalisco. 

31. Crotalus triseriatus triseriatus Wagler, 1830. 

Mexican Spotted Rattlesnake. 
Central Mexican plateau. 

32. Crotalus triseriatus pricei Van Denburgh, 1895. 

Arizona Spotted Rattlesnake. 
Southeastern Arizona to Durango. 

33. Crotalus stejnegeri Dunn, 1919. 

Long-tailed Rattlesnake. 

Mountains of eastern Sinaloa and western Durango. 

34. Crotalus willardi Meek, 1905. 

Ridge-nosed Rattlesnake. 
Southern Arizona to Zacatecas. 

35. Sistrurus ravus (Cope), 1865. 

Mexican Ground Rattlesnake. 
Central Mexican plateau. 

36. Sistrurus miliarius miliarius (Linne), 1766. 

Carolina Ground Rattlesnake. 

From North Carolina to central Alabama. 

37. Sistrurus miliarius barbouri Gloyd, 1935. 

Southeastern Ground Rattlesnake. 

The Gulf lowlands from Georgia to Mississippi ; Florida. 

Klauber — Key to the Rattlesnakes 193 

38. Sistrurus miliarias streckeri Gloyd, 1935. 

Western Ground Rattlesnake. 

Southern Missouri to Louisiana and west to central Texas. 

39. Sistrurus catenatus catenatus (Rafinesque), 1818. 

Eastern Massasauga. 

Central New York west to eastern Oklahoma. 

40. Sistrurus catenatus tergeminus (Say), 1823. 

Western Massasauga. 

Southwestern plains from central Kansas to northern Tamaulipas 
and southeastern Arizona. 

While the ranges of the several forms are broadly indicated in the 
above table, more specific and detailed range limits will be found under 
each species or subspecies in the key itself. For page references to key 
characters and descriptions, and cross references to the appropriate maps 
and photographs of each species see the index. 


Decisions as to the validity of the several species and subspecies 
of rattlesnakes recognized are based on studies carried on by the writer 
during the past eight years. Scale counts and other data have been avail- 
able on about 8000 specimens. All except four of the forms have been 
seen alive. 


Although during the past two years I have given considerable time 
to a study of rattlesnake nomenclature, in the preparation of this key it 
was first decided to make no fundamental departures from current prac- 
tice in the technical names employed, even though the validity of several 
might be questioned. It was feared that shifting some of the names long 
established in the literature might be a handicap to the adoption of the 
key; which I hoped would prove of practical value. Therefore the key 
first went to press with only minor changes in current usages. 

But meanwhile I have had a further opportunity to discuss this 
phase of the situation with some of my herpetological friends. They 
have pointed out that as the key is presumed to clarify certain species 
differences, and as a long time must elapse before a second edition can 
be issued, such changes as are certain to be required eventually should 
be made at this time. To make the changes later, in a subsequent paper, 
would render the key obsolete only a short time after its issuance, and 
add to the confusion. This seems to be a logical view. 

I know that to some these changes will appear a useless and over- 
technical imposition. But this is not hair-splitting nor the evidence of a 
contentious desire to disturb the peace. The changes will inevitably be 
made some day as studies of the species and literature continue, and "it is 
wiser for the present generation to bear with the temporary inconvenience 
of a few changes than to transmit to future generations our nomen- 

194 San Diego Society of Natural History 

clatorial problems, augmented a hundred fold by the addition of the 
ever-increasing number of systematic units made possible by the like 
increase in the amount of literature."* 

The important changes from current usage which I find necessary are 
as follows: (1) substitute Crotalus cinereous Le Conte in Hallowell, 
1852 for Crotalus atrox Baird and Girard, 1853; (2) substitute Crotalus 
vtridis (Rafinesque), 1818 for Crotalus conjluentus Say, 1823, affecting 
all the conjluentus subspecies; (3) employ Crotalus durissus Linne, 1758 
as the species name of the neotropical rattlesnake, relegating terrificus 
(Laurenti), 1768 to subspecific status as the name of the South American 
form; and finally (4) use tergeminus (Say), 1823 as the subspecific 
name of the western massasauga rather than edwardsii Baird and Girard, 


The reasons which have dictated these decisions are summarized here- 
under. I deem it undesirable at this time to give more than a brief out- 
line of the factors involved, reserving for future publication a more com- 
plete exposition of these and other questions affecting rattlesnake nomen- 
clature. It is not impossible that other changes will be found necessary 
as the work proceeds; and in any case it will be advisable to place on 
record the bases of decision where some currently used, but disputed 
names, have been retained. 

(1) Cinereous antedates atrox. Although the description (Proc. 
Acad. Nat. Sci. Phila., Vol. 6, No. 5, pp. 177-182, 1852) is included 
under Hallowell's description of Crotalus lecontei (an invalid synonym 
of Crotalus riridis), nonetheless the description of Crotalus cinereous, as 
LeConte sent it to Hallowell, is printed in full in this publication. The 
snake which LeConte describes is not the same as Hallowell's lecontei, 
which the latter thought to be the case; on the contrary it is an excellent 
description of the western diamond rattlesnake and it was collected in an 
area (along the lower Colorado Riverj) where no other species could be 
confused with it. Opinion No. 4 of the Commission seems to be exactly 
in point: "Manuscript names acquire standing in nomenclature when 
printed in connection with the provisions of Art. 25, and the question as 
to their validity is not influenced by the fact whether such names are 
accepted or rejected by the author responsible for their publication." 

(2) Rafinesque's description of his Crotalinus viridis (Am. Mon. 
Mag. and Crit. Rev., Vol. 4, No. 1, p. 41, Nov. 1818) leaves no question 
as to his meaning. The description, though brief, is clearly recognizable, 
especially when reinforced by the type locality. The upper Missouri 

* International Commission on Zoological Nomenclature, Opinion No. 12, Smithsonian Special 
Publication No. 1938, p. 20, 1910. . 

t The description concludes with the words "Colorado, March. 1851 from which one might 
suppose that this specimen was taken in what is now the state of Colorado. But we know from LeConte s 
paper on Coleoptera in Ann. Lye. Nat. Hist. N. Y., Vol. 5, pp. 125-216 at p. 125, that he was 
collecting along the Colorado River in Dec. 1850 and Mar. 1851, and in the valley of the Gila in Jan. 
and Feb. 1851. In assigning type localities to his insects he uses such terms as "Deserta fluminis Colo- 
rado; ad flumina Colorado et Gila, Martio; ad (lumen Colorado circa millia XXX a mare;" or simply 
"Colorado." Thus it is clear that the type locality of cinereous is the Colorado Desert in the Yuma 
area. From the high dorsal and ventral scale counts we may even venture the guess that the specimen 
came from the California side of the river. 

Klauber — Key to the Rattlesnakes 195 

Valley is the center of population of the prairie rattlesnake; is was ex- 
tremely plentiful in those days and, in fact, still is in many areas. No other 
rattler is found on the upper river. Two other rattlesnakes occur on the 
lower Missouri, the timber rattler and the massasauga, but we know that 
Rafinesque, in describing liridis, had neither of these in mind, for he de- 
scribed them as C. cyanurus and C. catenat/a, respectively, in the same 
paper in which he described viridis. 

(3) I have no desire at this time to revive the horridus-durissus- 
terrificus—adamanteus nomenclatorial discussion, always a fruitful source 
of argument. It is evident, however, that durissus Linne, 1758 must 
either take precedence over terrificus Laurenti, 1768 or it must fall entirely 
as unrecognizable; it is not a nomen nudum since a description is given 
and there was a type specimen, although it has been lost. Therefore it is 
either a species name or it should not be used at all; it cannot be revived 
with a date subsequent to 1758 to become a subspecies of terrificus, as 
has been done by some authors. For the present I retain durissus as the 
species name of the neotropical rattler, for while the caudal scale count 
(24) of the type seems low for this form, the black rhombs with light 
centers are characteristic of it over large areas in Mexico. This leaves 
terrificus as the South American subspecies. 

(4) I prefer tergeminus to edwardsii as the name for the western 
massasauga, since I think it describes this rather than the eastern subspecies. 
1 have discussed this with Mr. H. K. Gloyd, who has in preparation an 
extensive work on Sistrurus and we are in agreement on this point, which 
he will cover fully in one of his papers. 


In a work of this character it is impossible to include a synonymy 
under each species. The following condensed alphabetical list of synonyms 
is given in order that users of the key may ascertain the disposition 
which the present writer has made of the specific names hitherto proposed 
which he does not recognize as valid. The reasons dictating the decisions 
will be offered more fully in a subsequent publication. 

In the case of each specific name only the intention of the original 
describer is considered ; the shifts and reallocations made by subsequent 
authors, often giving the name a scope foreign to the purpose of the 
original describer, are omitted from presentation. Several species are 
listed which are not rattlesnakes but, being described under the genus 
Crotalus. are sometimes listed in synonymies of this genus. 

Americana Catesby, 1743. Syn. atricaudatus. Pre-Linnean name with- 
out validity. 
Atrox Baird and Girard, 1853. Syn. cinereous. 
Boicjuira Lacepede, 1789. Syn. durissus. 
Cascavella Wagler in Spix, 1824. Syn. terrificus. 
Catesbaei Fitzinger, 1826. Nomen nudum. Syn. atricaudatus. 
(Ex Hemprich, 1820?) 

196 San Diego Society of Natural History 

Cerberus Coues, 1875. Syn. oreganus. May later be recognized as a 
subspecies covering the southern half of the range of this form. 

Collirhombeatus Amaral, 1926. Syn. terrificus. 

Collilineatus Amaral, 1926. Syn. terrificus. 

Concolor Jan, 1859. Nomen nudum. Not to be confused with concolor 
Woodbury, 1929, which is valid. 

Confluentus Say, 1823. Syn. viridis. 

Consors Baird and Girard, 1853. Syn. tergeminus. 

Cumanensis Humboldt, 1833. Syn. terrificus. 

Cyanurus Rafinesque, 1818. Syn. horridus. 

Decolor Klauber, 1930. Syn. concolor Woodbury. 

Dryinas Linne, 1758. Probably syn. durissus. 

Edwardsii Baird and Girard, 1853. Syn. tergeminus. 

Elegans Schmidt, 1922. Syn. ruber. 

Exalbidus Boddaert, 1783. Syn. durissus. 

Fasciatus Higgins, 1873. Composite syn. horridus and others. 

Goldmani Schmidt, 1922. Syn. pyrrhus. 

Gronovii Laurenti, 1768. Description too brief for recognition. 

Hallow elli Cooper in Cronise, 1868. Nomen nudum. 

Helleri Meek, 1905. Syn. oreganus. : 

Immaculatus Latreille, 1802. Probably syn. durissus. 

Intermedins Troschel in Miiller, 1865. Syn. triseriatus. 

Intermedins Fischer, 1882. Syn. triseriatus besides being preoccupied by 
intermedins Troschel above. 

]imenezii Duges, 1877. Syn. polystictus. 

Kelly i Amaral, 1929. Syn. scutulatus. 

Kirtlandi Holbrook, 1842. Syn. catenatus. 

Lecontei Hallowell, 1852. Syn. viridis. 

Loeflingii Humboldt, 1833. Syn. terrificus. 

Lucifer Baird and Girard, 1852. Syn. oreganus. 

Lugubris Jan, 1859- Composite syn. triseriatus and polystictus. 

Melanurus Jan, 1859. Nomen nudum. Syn. atricaudatus by locality. 

Messasaugus Kirtland, 1838. Syn. catenatus. 

Mexicana Jan, 1863. Nomen nudum. 

Minor Catesby, 1743. Syn. miliarias. Pre-Linnean name without 

Mnltimaculata Jan, 1863. Nomen nudum. Syn polystictus by figure 
published in 1874 but this was subsequent to description of 
polystictus Cope. 

Mutus Linne, 1766. Not a rattlesnake. 

Omiltemanus Giinther, 1895. Syn. triseriatus; may have subspecific 
validity for extreme southern area where ventral scale counts 
are high. 

Orientalis Laurenti, 1768. Description too brief for recognition; prob- 
ably not a rattlesnake. 

Omatus Hallowell, 1854. Syn. molossus. 

Pallidas Giinther, 1895. Syn. triseriatus. 

Table 1. Summary of Rattlesnake Characters 

' 8*d« .. 5.1-p™. 



™w.® " ' 

Venlr.1 Sola 

S U hc. U d.l S„l« 





T... Eh... 

C. duri&sus durissus 
C. dunK-us terrificus 



(25) 27—29—31 
(25) 27— M (31) 

170— 175— 184 1 

171 — 181 — 190f 





4— 6—11 











1— 5— 9 

1—2.3 — 1 



3 — 1.5— 6 




















18— 21—29 



2— 5— in 




1- 6.7—9 

C. enyo 













4- 1.6—7 

C. molossus molossus 




(23) 25—27 







4— 7—16 




16— 2(1—2 1 



4— 6—10 

2— 2.1— 5 


C. adamanteus 




27—29 (31) 
(28) 25— 27 (29) 




22—2 1—26 







25—35 — 15 

5—6.9 1" 





25—2? 1 180—184—19(1 








4— 1.4— 6 


4— 5.3— 7 


C. lucasensis 

C. ruber 
C. exsul 



(25) 27—29 (31) 179— 189— 199 183—193—2113 

22— 25— 21i 





(25) 27—29—31 185—193—203 ! 188—197—206 







_>;.— 36— 11 

3 — 50— 7 




27—29 (31) 188—191—195 J 192— 194— 196 








3 1.1—5 

3—3 7—1 



(21) 23 25 27 (29) 166—178—190 167—181—192 

21— 25— 29 






27—36 — 14 



C viridis viridia 



,23) 25—27—29 1164—178-189 170-185-196 




11 — 16—19 



3:1 — 13—55 





(21) 2:'— 25 (27) 163—171—181 i 169—177—182 







C viridis abyasua 



(23) 25—27 1173—178—185 179—184—191 





23—34 — 18 

4- 6 6—8 

38—42 — 18 


6— 6.9— 8 

C. vindis lutosus 



23—25—27 (29) 171—179—189 J 174—184—196 








5—7 1 10 


C viridis concotor 



23—25 (27) 166—178—180 173—179—182 








7 — 3 .5-11 

6—7 1-1" 

C. viridis oreganus 



23— 25— 27 (29) 158—174—191) > 165—178—191 






2— 5.0— S 

23 34 46 



C mitchellii mitchellii 



(23) 25-27 171—177—183 172—179—186 





20—3; — 16 



3— 3.8— 5 


C. mitchellii pvrrhus 



(21) 23—25—27 168—178—185 168—179—185 







23—33 — 12 

1 5 6- 9 

3—4 5—6 

C. mitchellii stephenai 



(21) 23—25 166—174—181 







3— 5.. 5 — 8 




C. tigris 



21-23-25 (27) 






U— lj— 16 

11 — 17 — 37 





C. cerastes 



(19) 2;— 23— 25 






Hi— 13— 16 




3— 4.7— 1 


18 1 


2:.— 21 







6— 8—10 

1— 2.9— 5 

35 — 12—1? 



C horridui horridua 

123 J 










3—6.4 in 

15—23— 28 

1 . hon ! - atricmidatiu 



(21) 23—25 

lfil — 171 — 17*> 







5—6.6— 9 


C. lepidui lepidua 




l 18—160 -166 




11 — 12 — 13 


6— 9—16 

1— 2.9— t 


3 — 1.0—5 





(21) 23 (25) 






9—11 — 13 

:-,— 8—13 





r. triaeriatus triseriatus 









4— 8—12 


22—38 -57 


2 — 1.7—9 

C tnseriatus pricei 



2/— 23 



43 — 14 — 16 

18—21 -23 

8— 9—12 

8— Kt— 13 

:,_ 7—11 




1—2.3— 3 




C itejnegeri 



25—27 (29) 






in— 11— 12 


40 — 12 — 13 




C. willardi 





25— 28— 32 










10 — 12 — 14 


25 -28—31 
25—32 — 11 

22—30 — 10 


4— 5.2— 6 
•.'■ L0.6 1) 
8- 10.2- 13 

3— 3.5 — 1 


S miliarius miliariua 



21-23 (25) 











(19) 21—23 

(21) 23—25—27 






»The figure in il 



2£ S3MA 

-T'fiTur,.",'- th'-'mm. 






Klauber — Key to the Rattlesnakes 197 

Palmeri Garman, 1887. Syn. lepidus. 

Piscivorus Lacepede, 1789. Not a rattlesnake (A. piscivorus). 

Pulverulentus Cope, 1883. Syn viridis. 

Pulvis Ditmars, 1905. Probably based on an albino C.d. durissus, but 
might be C. unicolor. More material is required for a decision. 

Rhombifer Latreille, 1802. Syn. adamanteus. 

Salvini Giinther, 1895. Syn. scutulatus. May be a valid southern sub- 

Simus Latreille, 1802. Syn. durissus. 

Sonoraensis Kennicott, 1861. Syn. cinereous. 

Strepitans Daudin, 1803. Description inadequate; probably syn. durissus. 

Tesselatus Hermann, 1804. Cannot be recognized; resembles t riser iat us 
in scale counts, and durissus or adamanteus in size and pattern. 


To permit an additional check on the determinations which may be 
made by the use of this key, Table 1 is presented, giving some of the 
more important scale counts of the various species and subspecies. If it 
be found that the specimen under consideration differs extensively in some 
of these characters from the numerical range given in the table the result 
of the determination may be viewed with suspicion and a recheck should 
be made. 

The setting forth of scale-count data in abbreviated form presents 
certain difficulties. The usual method of giving minimum, maximum, and 
average shows nothing as to dispersion, that is, how closely most of the 
specimens cluster about the mean; and if either the maximum or mini- 
mum represents a freak or defective individual (as is not infrequently the 
case) or an error in counting or sexing, the mental picture of the disper- 
sion is distorted, overemphasis being placed on a single specimen out of 
the many which may have been examined. This is particularly true where 
broods of young have been included, since they often seem to contain 
freaks (especially if bred in captivity) which probably would not survive 
in nature. As an example of such a freak we have a defective juvenile 
female lucasensis with 170 ventrals; the lowest normal individual has 183. 
Again as a sample adult freak we note an oreganus with 33 scale rows, 
while no other specimen out of 1343 has more than 29 and only 6 have 
that many. 

Even the average leaves much to be desired when we deal with a 
form which is territorially variable, since the resulting figure is dependent 
on the origin of the individuals comprising the group averaged. Thus 
male cinereous average 185.0 ventrals in California and 177.7 in southern 
Texas ; obviously the relative numbers from each area which may be con- 
tained in the composite cinereous group will affect the average. 

Only a graphic presentation, or a tabulation of the variation of each 
item in percentages, will give a true picture of the dispersion, but in a 
condensed table this is impossible. So also is the presentation of the in- 

198 San Diego Society of Natural History 

terquartile range and the probable error of the mean, which are of interest 
in a complete statistical statement. 

For these reasons, while I have set forth in Table 1 the minimum, 
average, and maximum of each item, the minima and maxima are not 
always the extremes recorded; rather they are what might be termed the 
normal extremes, eliminating the solitary individuals here and there which 
seem to be freaks. 

The numbers of scale counts available are also given as an indication 
of the validity of the figures. Obviously, where only a few specimens 
have been at hand, neither numerical ranges nor averages can be considered 
of much value. 


The key is prepared in the usual dichotomous form, in which the 
selection is consecutively limited to one of successive pairs of alternatives. 
In the present instance, to facilitate reference, the alternatives are desig- 
nated by the letters "a" and "b." Where one of the two leads directly 
to a species (and thus to a conclusion) , the arrangement is such that the 
"b" alternative is selected to take this course. 

No identification key can be made infallible when the forms are 
closely related or intergrade; and, in the case of the rattlesnakes, because 
of the great variability in their lepidosis and patterns, it has been par- 
ticularly difficult to select key characters which lead invariably to the cor- 
rect conclusion. This is not an argument against the validity of the species 
and subspecies which have been recognized; many characters which are 
important in taxonomic studies are unsuitable for use in keys because they 
presuppose the availability of other specimens for comparative purposes. 
Other characters require special preparation and are therefore seldom at 
hand, as, for instance, venom and extruded hemipenes. Even the scale 
counts, which are of primary importance in classification when handled 
statistically, are often of little use as key characters because of overlapping. 
Thus, while it can be shown that the difference between the number of 
ventral scales in scutulatus and cinereous is highly significant mathematical- 
ly, there is a sufficient overlap so that the character becomes virtually useless 
in a key. Besides, a key must usually shunt out a single species from a 
group (those remaining undetermined at each point) and there is seldom 
a case in which some member of the group fails to overlap considerably, 
in these statistical characters, the single species it is desired to key out. 

In accuracy of determination the present key leaves much to be de- 
sired. This is because of the frequency of aberrant specimens which 
deviate from the mode. For example, in pyrrhi/s we have occasional 
specimens which have prenasals in contact with the rostral; and some 
specimens of ruber have undivided first infralabials, while conversely, 
some cinereous specimens have these scales divided. The internasal criterion 
(see 23b) in selecting viridis and its subspecies, although the best 
character available, fails in an appreciable number of cases, as shown 
in the following table: 

























Klauber — Key to the Rattlesnakes 199 

Viridis viridis 
Viridis nuntius 
Vhidis abyssus 
Viridis lutosus 
Viridis concolor 
Viridis oreganus 

Total 3548 239 6.3 

Oreganus, it will be noted, is conspicuously the worst offender. 

Where possible, more than one key character has been set forth so 
that the selected path may be verified. As a result the key is no doubt 
subject to criticism on the score of prolixity; yet I know of no other way 
to secure even approximate accuracy with a group such as the rattlers, in 
which a single universally consistent key character is so seldom available 
for any species. Condensed keys often lead to unsatisfactory determina- 
tions in large genera, especially where a single specimen (rather than a 
large series from one area) is to be identified, as anyone will testify who 
has tried to use the existing keys for such genera as Pituophis, Thamno- 
phis, Sceloporus, and Ufa. In such keys long experience in the genus 
must be had before the key can be applied with accuracy. 

In order to minimize further the effect of the key failures, a brief 
color description is incorporated with each alternative which leads finally 
to a species or subspecies; but the antithetical description is not given 
unless the pattern, or color, constitutes a part of the key. Occasionally 
footnotes direct attention to special deviations or to certain precautions 
which will reduce inaccurate findings. 

Where color and pattern are a part of the key one should not only 
allow for the ordinary fluctuations within a species but must be on the 
lookout for melanistic or albinistic individuals (fig. 111). Partial albinos, 
with one or more color-principles lacking, are also met with; these are 
particularly confusing unless normal specimens from the same area are 
available for comparison. Preserved specimens, which are faded or from 
which the epidermis has been rubbed away through continued handling or 
drying, thus changing both color and pattern, must be guarded against in 
making comparisons. Preservation tends to dull the brighter colors into 
neutral grays and browns; this is especially true of red and yellow. Speci- 
mens which have been long in captivity sometimes rub their snouts on 
the barriers so continuously as to deform the rostral and internasals in 
shape and arrangement; such changes should be noted where the key uses 
these scales. 

Under each species the range has been set forth, as far as known, 
and there should be no hesitancy in using this as a check on the determina- 
tion, when the locality of collection of the specimen is available. Of 

200 San Diego Society of Natural History 

course if the specimen keys to a species known to occur only a short 
distance from the place of collection, the identification may be presumed 
to be accurate, for range extensions are to be expected in the future as 
larger and more thorough collections become available. On the other 
hand, if, for example, a Pacific Coast specimen is found to be horridus or 
adamanteus, the error must be in the key or its use, since neither of these 
species occurs within a thousand miles of that territory. The key is a 
working tool; imperfections are to be expected and therefore all informa- 
tion available on the specimen, including the locality, should be used to 
verify the determination. 


The ranges are described as closely as the available specimens and 
records permit. To facilitate these descriptions tabulations and large-scale 
maps were prepared for each species or subspecies; altogether .several 
thousand locality records were available, although comparatively few were 
at hand in the case of some of the rarer forms. In some areas — this is 
particularly true of Mexico — the authoritative records are rather scanty 
and the results have been somewhat generalized. Usually territories listed 
as within the range of a species are limited to those from which specimens 
or authentic records have been available; but in some instances where it is 
evident that a species occurs in intervening territory between two records, 
its presence there has been assumed. 

Due allowance has been made for the character of certain early 
records. In the days of the Indian wars in the West, material was often 
gathered from a considerable area, yet was labeled by the recipient museum 
with the locality of the fort or army post from which it was sent. As a 
further complication these posts were sometimes moved for considerable 
distances without change of name. Therefore these old records have been 
neglected if they appear questionable, as indicated by currently available 
specimens and the present knowledge of habitat preferences. 

Published locality records of species whose definitions have been con- 
fused in the literature have been discarded unless verified by specimens. 
Examples: trtseriatus confused with polystictus; cinereous with scutulatus; 
durissus with basiliscus or molossus, and the latter with each other. 

It should be understood that when an area is specified as the range 
of a species this does not mean that it is to be found universally dis- 
tributed throughout that area. In some sections the encroachments of 
civilization, such as industrial and agricultural developments, irrigation, the 
cutting of forests, or drainage of marshes, have caused the disappearance 
of species from whole districts which they once inhabited. And even 
where natural conditions remain, a species is often not spread evenly 
throughout a territory. This is particularly true in the southwest, where 
ecological conditions vary greatly within short distances owing to changes 
in altitude or topography. Thus while three rattlesnakes may be said to 
occupy the same geographical area (a county, for example), one may 

Klauber — Key to the Rattlesnakes 201 

occur only on the plains, another amongst rocky foothills, and the third 
upon the higher peaks. None is found uniformly distributed over the 
entire area, yet the county would be included in the designated range of 
all three. Space limitations have made necessary the same type of gen- 
eralization in the range maps which accompany this key. 


While it is impossible, within the scope of this key, to present a 
complete account of the life history of the rattlesnakes, it is desirable not 
to leave this feature untouched. The following paragraphs outline some 
of the more important or interesting facts concerning rattlesnakes and their 
ways. The necessity for abridgment has required rather broad generali- 
zation on some phases, and therefore this brief discussion should not be 
judged too severely by those already familiar with the subject. 

Habitats. — While most species of rattlers thrive in arid or semi-arid 
areas, particularly in rocky or brushy country, there are others which are 
not averse to swamps or timbered lands. Some forms, such as C. tri- 
seriatus. seek mountain areas; Cm. Stephens'! prefers rock-strewn canyons; 
C. cerastes is at home in sandy desert wastes. In California, rattlers (C.i . 
oreganus) are found at an altitude of 11,000 ft., and in parts of Mexico 
C.t. triseriatus occurs up to 14,500 ft. They have been found on the sands 
of the seashore only a few feet from the waves and occasionally have been 
seen swimming in the sea as well as in lakes. While no rattlesnake is as 
aquatic as the water or garter snakes, they will take to water at times, and 
the little known C. polystictus may be semiaquatic. Rattlers being heavy 
bodied, are less arboreal than many other species of snakes ; however they 
occasionally climb trees, no doubt in search of squirrels or birds. They 
are sometimes found crawling through chaparral well above the ground. 
Clumps of cacti are favorite refuges of some of the western forms, which 
are not impeded by the spines. 

An adherence to a certain ecological niche cannot always be specified 
for a given rattlesnake. It is not only that some forms are more tolerant 
than others, which is the case, but that a species may be restricted in 
ecological range in one area and not in another, owing possibly to com- 
petition or other more obscure conditions. Thus scutulatus is a snake of 
the flatlands in the Mohave Desert; in Arizona it is not only a desert, 
but a foothill and even a mountain form. In Arizona cinereous is an Upper 
as well as Lower Sonoran form, but in southern California it is exclusively 
a desert inhabitant ; here it has never secured a foothold even on the lower 
mountain slopes, probably owing to the competition of the closely related 
ruber. Hence in these brief notes it has often been impossible to cite the 
habitat preferences of a form because of this variability within a species. 

It is probable that in pre-Columbian times at least one species of 
rattlesnake was found in every part of what is now the United States 
excepting eastern Maine, upper Michigan, northern Wisconsin, central 
and northern Minnesota, eastern North Dakota, and Washington and 

202 San Diego Society of Natural History 

northern Oregon west of the Cascade Mountains. The higher mountains 
were likewise untenanted, the altitude not invaded depending on local 
conditions. While in southern California rattlers have been observed at an 
altitude of 11,000 feet, further north and in the Rockies the more rigorous 
climate holds them to lower levels. 

Where a species is approaching its toleration limit (in habitat con- 
ditions) its range may be highly irregular and intermittent. Thus only the 
lowlands in mountainous territory may be inhabited (as is the case, for 
example, in central Idaho and western Montana) and the range may be as 
irregular and broken as are the contours of the river- valleys, with further 
modification resulting from the nature of the exposure, for snakes range 
higher on slopes with a southern exposure. 

As far as numbers of forms are concerned Arizona is the headquarters 
of the rattlesnakes, no less than 15 subspecies being found within the 
limits of that state. However, because of their restriction to certain eco- 
logical niches, as previously mentioned, it is doubtful whether more 
than 5 or 6 species actually meet in any one locality. 

While rattlers have been exterminated in many industrial and agri- 
cultural areas, it is probable that in a few places they are now even more 
numerous than in primitive times. This may be the case where agri- 
cultural development, or the destruction of competitive predators, have 
served to increase the supply of rats, mice, gophers, ground squirrels, 
and the other rodents upon which most of the species feed. 

Economic Status. — Rattlers are of considerable economic importance 
in many areas, since they serve as a check on destructive rodents. How- 
ever, one would hardly recommend their protection, particularly adjacent 
to cities, because of the danger to humans inherent in their presence. It 
would be best if they could be replaced by other snakes such as the bull 
snakes, gopher snakes, and rat snakes, which have an equal economic 
value (in the destruction of harmful rodents) but, being harmless, do not 
have the one outstanding objectionable quality of the rattlesnakes, namely, 
their venomous character. Incidentally it may here be stated that when 
a person, through mental laziness, refuses to discriminate between harm- 
less and venomous snakes, and kills all snakes at sight, he is defeating the 
very purpose of his act. Every time he kills a harmless gopher or king 
snake he is making room in the economic scheme of things for one more 
rattler, for the number of these snakes is limited by the available food 

Habits. — While rattlesnakes are diurnal in spring and autumn, they 
are largely nocturnal in summer, this being especially true of the species 
found in the Upper and Lower Sonoran Zones. In the spring when they 
first issue from hibernation, they are abroad in search of food and mates, 
and are rather careless of concealment. At this season they are occasionally 
found in pairs, but the widespread belief that if one rattler be killed its 
mate will shortly appear at the scene of the tragedy is quite erroneous. 
Later they become secretive, their activities being restricted largely to the 

Klauber — Key to the Rattlesnakes 203 

evening hours or night, when the heat is not excessive, and when the 
rodents which constitute their principal food are abroad. In the daytime 
they seek refuge in ground holes, in the shade of dense thickets, or under 
rocks. In the arid Southwest no rattlesnake can stand the direct heat of 
the summer sun. Under such conditions even the desert sidewinder will 
succumb within ten or fifteen minutes. 

In the colder climates rattlers hibernate together in large numbers, 
going into hibernation about mid-October and emerging in mid-April, the 
dates varying somewhat with latitude and altitude. In some areas rocky 
retreats are preferred; in others, prairie dog towns or other ground holes. 
In milder climates the snakes do not gather in large groups for hibena- 
tion but seek separate refuges. Here they may come out briefly at any 
time during the winter if there is a warm spell. 

Rattlers are secretive and timid. When approached they will usually 
remain quiet in order to avoid detection, and when discovered will en- 
deavor to escape if given an opportunity. It is only when they are 
frightened and cornered that they will stand their ground with a strident 
warning to the intruder. Stories of rattlers chasing a person are probably 
the outgrowth of instances wherein the man stood between the rattler 
and his natural refuge, usually the nearest bush or rock cleft. Rattlers 
will not strike unless they are disturbed or frightened; they are not in- 
nately vicious, but seek only to defend themselves or escape. They do not 
always rattle before striking, as this depends on the disposition of the 
individual snake and the nature of the disturbance which has alarmed 
him. They can bite without coiling, but cannot strike without first 
throwing themselves into the loose S-shaped coil with raised forebody 
which constitutes the striking posture (fig. 108). The strike is merely a 
forward lunge of the head and rarely exceeds half the length of the snake ; 
if the snake is violently excited, it might reach three quarters. At the end 
of the strike the mouth is widely opened and the forward pointing fangs 
(fig. lc) are driven into the victim. The strike is made with such rapidity 
that the forward drive of the head cannot be followed with the eye; one 
can see only the white blur of the open mouth where the direction of 
motion is reversed. The head is retracted more slowly. The mouth is 
not opened until near the end of the strike; rattlers do not threaten their 
enemies with open mouth as sometimes pictured. 

The resting coil (fig. 109) in which rattlers are usually found is 
quite different from the striking coil. A rattler cannot strike when in the 
resting coil, but if alarmed rears up and quickly throws himself into his 
characteristic defensive posture. The striking coil not only facilitates a 
possible forward lunge but also bodily maneuver, for in this posture the 
snake can move backward or to either side where a safe retreat may offer; 
but meanwhile he faces his foe, ready to strike if the enemy comes within 
range. A big rattler, thoroughly alarmed, is something both to see and 
hear. Not only is the rattle sounded continuously but the cornered snake 
inhales and exhales with a violent hiss; the posterior body is flattened; 

204 San Diego Society of Natural History 

and the protruding tongue is held alternately pendent, and vertically erect, 
with the tips widespread. 

Rattlers, in the striking coil, rear up to about 6 to 12 inches with the 
head a trifle lower than the lateral curve of the neck; they strike slightly 
downward, usually well below 12 inches, so that heavy leather boots or 
puttees afford good protection for the legs. A large rattler can puncture 
thin flexible leather. Crotdus durissus has a striking posture resulting in 
a higher strike than that of our nearctic rattlers. Since rattlesnakes are not 
naturally vicious and do not attack unless disturbed, the principal danger 
to hiker or hunter results from walking along a trail without watching his 
step so that a rattler which has not been seen may be trod upon. Under 
such circumstances a rattler would bite without coiling or a warning rattle. 
Occasionally accidents happen to persons climbing about amongst rocks 
and placing their hands in a fissure in which a rattler lies concealed, or in 
walking abroad at night without a light. Farmers trimming shrubbery 
sometimes suffer from having disturbed a rattler lurking there. To city 
folk the best advice is, "Watch where you place your hands and feet; don t 
put them into places you can't see." 

Rattlers, being thick of body, crawl rather slowly. Usually they adopt 
a sinuous motion (fig. 110) ; sometimes progression is caterpillar-like and 
they leave an almost straight trail. The sidewinder has a peculiar rolling 
motion, developed for efficient transit over loose sand. In this, with head 
anchored, the body is thrown to the side in a loop, after which the head 
is moved. The resulting track is not continuous, but is a series of short 
lines advancing en echelon. 

Food. — Rodents, such as rabbits, ground squirrels, prairie dogs, 
gophers, rats, and mice, comprise the natural food of most species. Birds 
are occasionally eaten. Some of the smaller species, such as lepidits, tn- 
seriatus, and cerastes, are largely lizard feeders, although they do not scorn 
small rodents when obtainable. Rarely other snakes are eaten. 

Rattlesnakes always strike and poison their prey; that is what the 
venom is for — to kill the prey; as a means of defense it is secondary and 
incidental. The prey is struck but not held by the rattler. The small 
animals quickly succumb to the venom and are then eaten, usually head 
first. The stomach juices of the rattler are very powerful and every part 
of the prey is digested except hair or feathers. It is probable that in their 
natural state they feed at approximately weekly intervals, if a full meal is 

Rattlers do not feed well in captivity and many die of self-imposed 
starvation if disease does not supervene. They do not charm their prey 
but secure it by stealth, lying in wait for it to come within range of the 
deadly stroke. Most animals placed in a cage with rattlers show no fear 
of them. A hungry rat with no other food available will sometimes kill 
a rattler in whose cage it has been placed as food. Zoo visitors are heard 
to express surprise that two or more rattlers can be placed in the same 

Klauber — Key to the Rattlesnakes 205 

cage without an immediate fight. As a matter of fact being peaceful, they 
get along well with each other and with other snakes. 

Size. — The largest of rattlesnakes is the Eastern Diamond, C. ad am mi - 
tens, which reaches a length somewhat in excess of 2400 mm. (8 ft.) and 
a weight of more than 15 pounds. This is the largest, that is the heaviest, 
of all venomous snakes; the longest is the king cobra. Other very large 
species are C. cinereous and C. durissus. The smallest rattler is probably 
C. willardi or C. stejnegeri. which scarcely reach 590 mm. (2 ft.) ; the 
latter is also the rarest (in collections), only 3 specimens being available 
to date. For a rough classification of rattlesnakes as to length, see the 
appropriate column in Table 1. 

Adult male rattlesnakes average larger than females by about 8 per 
cent; they are also somewhat more plentiful. Males can be distinguished 
from females by their relatively thicker and longer tails ; in the females 
there is a distinct reduction of diameter where the tail begins, while in the 
males the taper at this point is more gradual. Rattlesnake growth is quite 
rapid during the first two years of life, after which it is much slower. 
The young of rattlesnake species having an average ultimate length of 
1220 mm. (4 ft.) will be about 265 mm. (IOV2 in.) at birth. Rattlesnake 
lengths determined from skins are inaccurate owing to stretching. 

Reproduction. — Our nearctic rattlesnakes mate in the spring, soon 
after leaving hibernation. The young are born between mid-August and 
early October, depending on the species and the geographical area in- 
habited. Although many kinds of snakes lay eggs, others, including 
rattlesnakes, are born alive. Broods vary in size from 2 or 3 to 30 or 
more, but average about 10. Female rattlesnakes give birth to their first 
broods when three years of age. Young mothers have smaller broods; 
also the smaller species have fewer young. Young rattlesnakes shift for 
themselves immediately after birth; occasionally a mother is found with 
her young, which probably have been born but a few hours before, or 
they are using a common retreat. The long-existent theory that parent 
rattlesnakes swallow their young for protection is not true. 

Control. — Various methods of rattlesnake control have been tried, 
such as the use of snake-proof fences, blasting them out of dens, or poison- 
ing them in such dens with liquids or gases. These means are not often 
effective; results are much more likely to be achieved by curtailing their 
food supply, that is, by eliminating the rodents upon which they feed. 
However, the great concentration of rattlesnakes at hiberating time does 
offer an opportunity to destroy large numbers within a short period. This 
is especially true in the colder climates. A. M. Jackley of South Dakota 
has devised means for capturing rattlers leaving hibernation, and has been 
successful in securing great numbers of C.i. viridis. Whether such de- 
struction is to be recommended on economic grounds is an open question. 
Bounties have been proposed but cannot be recommended, since they often 
lead to the importation of snakes from other areas. Hair ropes, of course, 
are entirely ineffective in keeping rattlesnakes out of a camp; a snake 

206 San Diego Society of Natural History 

which does not hesitate to crawl over cactus could hardly be expected to 
notice a hair rope. 

Forest and brush fires cause great destruction of rattlesnakes, as they 
do of all animals occurring in the devastated area. Autos kill many 
snakes upon the highways. 

Rattlesnake Enemies. — Aside from man, the principal enemies of the 
rattlesnakes are birds and other snakes. Hawks and owls are sometimes 
observed carrying rattlers or other snakes in their talons. Ravens have 
been seen to attack young rattlers. A number of kinds of harmless 
snakes, including especially king snakes and racers, feed frequently or 
occasionally on other species of snakes, and rattlers are amongst those 
which fall prey to them. The attitude of the king snake toward rattle- 
snakes is often misunderstood. King snakes do not range about spoiling 
for a fight with a rattlesnake; however, when in search of food a young 
rattlesnake is as tempting a morsel as any other. Some mammals, in- 
cluding a South American skunk, are known to kill snakes. The mortality 
amongst young rattlesnakes is high, for they are especially vulnerable to 
birds and snakes. When they have reached maturity they are too much for 
some of their erstwhile enemies to handle. 

Various of the hoofed animals, especially pigs, deer, and goats, are 
known to kill rattlesnakes, as well as other species. Snakes are said to be 
scarce where goats habitually graze. 

Commercial Value. — Rattlesnake skins are used in the manufacture of 
such ornaments as belts, slippers, hat bands, purses, and the like. The 
flesh is edible and in Florida has been canned on a commercial basis. The 
venom when carefully segregated by species, purified by centrifuging and 
dried, is used in the treatment of horses in preparation of antivenomous 
serum; and has certain other medicinal uses upon which research is now 
being carried forward. Rattlesnake oil is sometimes in demand in certain 
oriental trade circles. 

The snakes themselves are in sporadic demand for snake shows, car- 
nivals, zoological gardens etc. Prices, when there is a market, range from 
$.50 to $10.00 each, depending on the size and species. The commoner 
kinds are often sold by weight, bringing from twenty to fifty cents per 
pound. With the exception of a few experienced and well organized 
firms which supply much of the American market for these products, few 
persons are successful in making a livelihood from any phase of the rattle- 
snake business, since the demand for snakes or snake products is at best 
sporadic and uncertain. 

Finding snakes and catching them are matters of experience. Various 
kinds of noose-sticks are used for picking them up.* A practiced eye 
can discover snakes which the novice will overlook, as has often been 
demonstrated in the field by pointing to a patch of rocks amongst which 

* For methods of catching and shipping rattlesnakes, see Klauber: Notes on Herpetological Field 
Collecting. S. D. Soc. Nat. Hist., Collecting Leaflet No. 1, pp. 1-10, 1935. 

Klauber — Key to the Rattlesnakes 207 

a snake lies in plain sight. A person who does not know what to look 
for will have great difficulty in locating the reptile. 

The largest catches can be made about dens in the autumn or spring. 
In desert areas one catches snakes at night by driving on black paved 
roads, against which background the light-colored snake shows strongly 
under the glare of the headlights. 

Rattlesnake farms, except for their possible exhibition value, are not 
often successful. In these farms the population is not replenished by 
breeding, but by acquiring fresh material continuously in the wild, it 
being impractical to breed snakes in captivity. Caged rattlesnakes are 
afflicted with diseases and a variety of internal and external parasites which 
shorten their lives. Rattlers seldom take food naturally in captivity, but 
often live for a year or more without it. However they must have water. 

Rattlers in captivity quickly become lethargic; after a short time they 
are accustomed to human beings and do not greatly resent handling. Un- 
due familiarity with rattlesnakes, however, is to be decidedly discouraged, 
as a sudden fright may result in a serious accident. A considerable pro- 
portion of snake-bite cases in this country results from handling captive 
snakes. Pulling the fangs out is an easy, but not a lasting, safety measure, 
since they will be shortly replaced ; and if the operation is carried deeper 
and the reserve fangs or venom glands are removed, the snake usually dies 
within a short time.* 

Senses. — Zoo vistors, unfamiliar with snakes, frequently are heard 
to confuse the snake's fangs (which cannot be seen unless the snake 
yawns) with its tongue. The tongue is a harmless and delicate organ, 
probably auxiliary to the sense of smell; or it may be partly auditory in 
function. It is frequently advanced and retracted, especially if the snake 
is in an unusual situation; with it the snake seems to be sensing his sur- 
roundings. An angered or defensive rattlesnake not only protrudes the 
tongue to the fullest extent but also alternately points it vertically up- 
ward and downward. These motions are not hurried, but are made with 
deliberation. The rattler's tongue is bifurcate and, in nearly all species, 
is black. When on the alert the tips are spread wide apart. 

Rattlers, like other snakes, have ears, but they are without external 
aural openings. The hearing seems to be dull. The sight also is not of 
the best, judging by the frequency which captive rattlers are seen to miss 
their prey, even when it is within easy range. Possibly they see better at 
night. A short time prior to changing its skin a snake's eye coverings 
appear bluish and almost opaque. There is an exudation of a liquid 
between the old and new skins designed to facilitate shedding. At such 
times the snake is nearly blind. A day or so before exuviation the eye 
clears up. 

The unique pit (fig. 6) which characterizes the snakes of the family 
Crotalidae (the pit vipers) is another sense organ, the purpose of which is 
uncertain, but is thought to be auditory. 

* Re effect of captivity on rattlesnakes, see Klauber: A Herpetological Review of the Hopi Snake 
Dance. Bull. Zool. Soc. S. D., No. 9, p. 32, 1932. 

208 San Diego Society of Natural History 

Rattles. — The rattle is used for the purpose of warning away enemies 
which are large enough to cause possible injury to the snake. It is not 
employed (as has been sometimes stated) to warn prey; this would be 
contrary to reason. It is not a mating call as has been suggested; in ex- 
cursions through snake-infested country during the mating season I have 
never heard a rattle sounded except by a snake that has been disturbed by 
my companions or myself. 

In vibrating the rattle the tail is shaken at the rate of from 45 to 
60 cycles per second, the speed depending both on the individual snake 
and the temperature. The sound, which is caused by the impingement 
against each other of the interlocking segments of the rattle string, cannot 
be distinguished as discrete impacts, for it is much too rapid; it is a hiss 
not unlike escaping steam. A large rattler can be clearly heard at a dis- 
tance of a hundred feet or more; to have one of these suddenly let go 
immediately under one's feet is startling indeed. On the other hand, some 
species — S. miliarius for example — have such small rattles that they can 
scarcely be heard at a distance of a few feet. 

The first rattle, or prebutton, is always lost with the first shedding 
of the skin, which occurs within a day to a week after birth, leaving the 
permanent button exposed ; subsequently the snake acquires an additional 
rattle with each shedding. Snakes at an age of one year have from 3 to 
6 rattles, the number depending on the species of snake and the duration 
of its seasonal activity. Thus the age of a rattler can only be approxi- 
mated from the number of rattles, and then only if the button be present, 
showing that the string is complete. Rattles are lost through wear, so that 
strings exceeding 15 segments are extremely rare. I have not seen one 
with more than 13 segments which still retained the original button. 
Adult snakes usually have from about 5 to 10 rattles; the loss of the ad- 
ditional segments is not a detriment to the snake since long strings are 
inefficient vibrators. Rattlers living in places where they are seldom dis- 
turbed — for example on Tortuga Island — tend to have long strings. The 
same is true of species — such as the sidewinder — which inhabit sandy 
areas, for here the rattles do not often catch in the clefts of rocks or 
shrubs, nor are they abraded by rough objects. Long sets of rattles can 
be easily faked, which accounts for most of the phenomenal strings which 
have been reported. 

The rattle consists of a horn-like material which is exuded and 
solidified on a corrugated matrix of tissue prior to the shedding of an old 
skin. The method whereby each new rattle is advanced one corrugation 
ahead of its predecessor and yet remains interlocked with it, is mechani- 
cally quite complex. It is effected by a wave action of tissue. 

I wish to repeat that rattlers do not invariably sound their rattles be- 
fore striking. Some are so peaceful — C. ruber for example — that they will 
neither rattle nor strike except under great provocation. 

Fangs. — The fangs are two greatly elongated curved teeth at the front 
of the upper jaw. Normally they are folded back against the roof of the 

Klauber — Key to the Rattlesnakes 209 

mouth (fig. lb), whence they may be rotated forward and downward into 
the striking or biting position (figs. Id and le) . Of this fang rotation the 
snake has voluntary control; the fangs are not automatically tilted as the 
snake opens his mouth. The movements of the two fangs on the op- 
posite sides of the head are independent of each other. When in the 
resting position they are covered by a white protective sheath of tissue, 
which is partially pushed back from the points as the fangs are advanced. 
The fangs are hollow and have an upper and lower opening, the latter 
just above the point (fig. lg) . The venom is conducted from the venom 
gland, which lies back of the eye, through a duct to the upper opening 
(fig. ll) and thence through the tubular fang to the orifice above the 
point, thus constituting a perfect natural hypodermic needle. Rattlesnake 
fangs are replaced at regular intervals even though they are unbroken. On 
each side of the head there is a pair of maxillary sockets which the active 
fangs occupy alternately (figs, lg and lh). While replacement is under 
way the old fang may remain in place while the new fang is being 
ankylosed in the adjacent socket; thus for a short time the rattler may 
have two fangs on a side. The reserve fangs, from which replacements 
are made, lie in an orderly series behind each functional fang. There are 
approximately eight on each side in successive stages of development from 
rudiments to almost complete fangs. As the reserve is drawn on for re- 
placements additional buds appear, so there is no decrease in the number 
in reserve. In each individual fang, development takes place from the 
point upward. The fang develops as a tube, notwithstanding the central 
longitudinal suture (fig. lg) which indicates that a remote ancestral form 
had fangs with open grooves, as is the case with some groups of venomous 
snakes today. The fangs of the several species of rattlesnakes differ some- 
what in curvature and in length proportionate to the size of the head. The 
curvature aids in imbedding the fangs as the mouth closes. 

Venom. — Rattlesnake venom is a yellow liquid having a specific 
gravity of about 1.08. It may be dried, without serious modification in 
toxic properties, by heating to 100° F; when dry it will retain its potency 
indefinitely. In drying, the venom loses about three-quarters of its 
weight. Dried venom has a yellow crystalline appearance, although the 
flakes are not true crystals. 

The venom is primarily a means of securing food; venomous snakes 
practically always secure their prey by striking and poisoning it. In ef- 
fect the venom not only kills the prey but is said to aid in its digestion. 
There are considerable differences, both in toxicity and physiological ef- 
fects, in the venoms of the several rattlesnake species, some being 60 
times as powerful, drop for drop, as others. Some are primarily hema- 
toxic; others neurotoxic. 

The yield of venom per snake varies greatly amongst the different 
species of rattlesnakes, even though the snakes may be of similar size. 
The following are a few adult averages, the figures representing the yield 
in milligrams of dried venom: C. cinereous. 270; C. v. viridis, 80; C. r. 

210 San Diego Society of Natural History 

oreganus, 140; C. m. mitchellii, 33; C. m. pyrrhus, 215; C. cerastes, 32; 
C. tigrif, 11. A general, but not universal rule, is that rattlers which give 
low quantitative yields proportionate to their sizes have the most power- 
ful venoms. So far as now known C. tigris has the most powerful and 
C. ruber the weakest venom. C. durissus seems to have a combination of 
high yield with powerful venom and therefore may be considered the most 
dangerous of the rattlesnakes. The maximum venom recovery from a 
single snake at a single milking was from a large C. cinereous; this pro- 
duced 3.9 cc. of liquid or 1145 mg. of dried, purified venom. This is 
the highest record among over 4000 rattlesnakes that I have milked, and 
1 know of no greater quantity reported from any kind of venomous snake. 

A satisfactory procedure in "milking" large rattlers is as follows: An 
assistant catches the snake immediately behind the head by means of a 
noose-stick, and holds it with the head resting on the edge of the table. 
When the snake is so caught and held it has no opportunity to reach 
any object with its fangs and thus waste venom. The operator by means 
of a metal hook catches the snake's upper jaw under the rostral plate 
and tips the head back. Then the rim of a porcelain cup is introduced 
below the fang points, and the fangs are drawn downward and forward 
into the erected position. Since the head is tipped back and steadied 
by the hook while the cup approaches, the snake can neither see the cup 
nor slash at it until it is in position to catch any venom expelled. 

As the fangs are drawn forward, the edge of the cup is pressed 
steadily against them; this tends to hold the head firmly and gives the 
snake a feeling of something yielding on which to bite. The hook is 
now withdrawn, and the operator, further forcing the head against the cup 
with his index finger, presses the venom glands with the thumb and third 
finger. The snake will usually eject some venom in an attempt to bite 
when it feels the steady pressure of the cup against the fangs, but in all 
cases the flow is increased by the mechanical manipulation of the glands. 

Holding the snake with a noose-stick is not a suitable method for the 
smaller rattlesnakes, whether juveniles of the larger species or adults of 
the smaller, since their heads will not protrude far enough beyond the 
holding strap to permit manipulation. With these small specimens (say 
under 800 mm.) an operator can work most efficiently alone. In this process 
a centrifuge tube or test tube is firmly attached to a stand or vise. The 
snake is caught by pressing the head against the table with a short straight 
stick and is then grasped behind the head with the left hand. Using the 
metal hook to tip the head back, the fangs are hooked over the edge of the 
stationary tube and the glands are manipulated with the fingers of the 
right hand. The operator can keep the tail of the snake from thrashing 
about by placing it between himself and the work table, and then leaning 
against it. 

Many operators recommend the use of a thin rubber or parchment 
cover for the venom cup or tube, to be bitten through by the snake. This 
may be justified for short-fanged snakes, but I have not found it efficient 
with rattlesnakes, since it impedes rapidity of operation, and does not 

Klauber — Key to the Rattlesnakes 211 

increase the yield. When a diaphragm is used, venom is frequently spilled 
on it before the fangs penetrate; and as the fang points cannot be seen 
after they have gone through the diaphragm, it is impossible to observe 
the effectiveness of the manipulation of the glands, or when the flow 
has ceased.* 

In the wild, snakes are presumed to use only a small quantity of their 
venom in securing their prey, for no more is needed to cause the death 
of these small creatures within a minute or so. Rattlers have complete 
muscular control of the quantity of venom discharged, and naturally will 
not waste it. In biting an enemy in anger they probably eject from one 
half to two thirds of the quantity which may be secured by manipulation 
in the milking process. 

It is presumed (but not definitely known) that a snake in the wild 
would replenish empty venom glands within two weeks or less. In cap- 
tivity the secretion is much slower; snakes milked at successive intervals 
of two weeks show a sharply declining supply. For this reason a con- 
tinual accession of fresh specimens is necessary for an adequate supply of 

Young snakes have venom at birth, but because of their small size it 
is quite limited in quantity; the bite of such an infant would probably be 
painful but not dangerous. Very old snakes show evidence of a declining 
venom secretion. 

Snake Bite and Treatment. — Although rattlesnakes are moderately 
plentiful in many areas in the United States which are frequented by large 
populations, especially on week-end excursions, hunting or fishing trips, 
or by hikers or campers, rattlesnake bite constitutes a relatively small ac- 
cident risk; not to be compared, for example, to the chance of a highway 
accident. The naturally inoffensive and secretive character of the snakes, 
and the fact that people going abroad are usually well protected about the 
legs, reduce accidents. Only in a few areas of our country is the snake 
bite problem sufficiently important to warrant much attention. 

The gravity of a rattlesnake bite is something which cannot be closely 
defined or predicted, any more than one might predict the seriousness of a 
fall, without knowing the exact circumstances surrounding the accident, 
such as the height of the fall, the character of the surface struck, etc. 
And in a snake-bite case the conditions are even more obscure, since there 
are important factors which cannot be ascertained, even after the accident 
has occurred. So no one can give an off-hand opinion as to the gravity of 
such a case; and correspondingly, while there should be no desire to ex- 
aggerate the gravity, it will be best, in the interest of safety, to over-treat 
rather than under-treat the case, provided a proper treatment is used. In 
any event the victim should remain under close observation for at least 
48 hours. 

Some of the more important variable factors involved in snake-bite 
cases are the following: 

* For the methods used in venom recovery and purification see Klauber: The Collection of Rattle- 
snake Venom. Bull. Antivenin Inst, of America, Vol. 2, No. 11, 1928. 

212 San Diego Society of Natural History 

( 1 ) The size, vigor, and health of the victim, these being important 
in determining absorptive power and resistance to venom. 

(2) The allergy complex of the victim; his susceptibility to protein 
poisoning; sensitization (anaphylaxis), or partial immunity imposed by 
previous bites and treatment. Some individuals are so susceptible to 
venom that the mere handling of it causes typical asthmatic symptoms 
lasting for 24 hours or more ; most persons under similar circumstances are 
entirely unaffected. 

(3) The psychological condition and nature of the victim; extreme 
fear and apprehension will affect heart action and therefore rapidity of ab- 
sorption; and it is not impossible that there may be more direct reactions. 

(4) The site of the bite, which will be less dangerous in the ex- 
tremities, or in tissues where absorption will be less rapid (fat, for ex- 
ample) , as compared to a bite near the vital organs or penetrating a vein. 

(5) The nature of the bite, whether a direct stroke with both fangs 
fully imbedded, or a glancing blow or scratch. The movement of the vic- 
tim (jumping backward, for instance) may cause a partially ineffective 
bite; or a bone may be struck, thus causing imperfect penetration. The 
snake may misjudge his distance and have the fangs only partially erected 
at contact, thus resulting in only slight penetration; or he may, for the 
same reason, eject venom before the fangs are imbedded. 

(6) The protection afforded by clothing, which, by interposing 
thickness, will permit less depth of fang penetration, and will cause the 
external and harmless absorption of part of the venom. Only the point 
of the fang may penetrate the skin, in which case there will be no venom 
injection, for the orifice is well above the tip (fig. li). 

(7) The number of bites; occasionally an accident involves two or 
more distinct bites. 

(8) The length of time the snake holds on; it may withdraw or be 
torn loose before injection takes place. This is likely to be more im- 
portant with the elapine snakes, with their less specialized fangs, than 
with such long- and hook-fanged snakes as the rattlers. 

(9) The extent of the anger or fear upon the part of the snake. The 
muscles which wring the venom glands and thus inject venom are sep- 
arately controlled from the biting mechanism. The snake's natural ten- 
dency is to withhold venom, since this is his means of securing prey; but 
if hurt or violently angered he is likely to inject a large part of the venom 
contained in the glands. 

(10) The species and size of the snake, affecting venom toxicity and 
physiological effects, venom quantity, and (by reason of length and 
strength of fangs) depth of injection. The age of the snake is likewise 
important; not only are young snakes less dangerous because of their 
smaller size (and therefore reduced quantity of venom) but also the 
venom is less toxic, judging from the reduced proportional recovery of 
solids upon evaporation. Snakes which have passed their prime also 
probably secrete less venom and of a reduced quality. 

Klauber — Key to the Rattlesnakes 213 

(11) The condition of the venom glands, whether full, or partially 
depleted or evacuated by reason of recent feeding, defense, ill health, or 
captivity. The season of the year (proximity to aestivation or hibernation) 
may also cause a variation, but this is not definitely known. 

(12) The condition of the fangs, whether entire or broken, lately 
renewed or ready for shedding. 

(13) The presence, in the mouth of the snake, of various micro- 
organisms, some of which, gaining access to the wound, may, abetted by 
the anti-bactericidal effect of the venom, entail serious sequelae. 

(14) The nature of the instinctive first aid treatment, if any, such 
as suction, or circulation stoppage by pressure. 

To conclude, with variable factors of such importance, it is to be ex- 
pected that some cases will prove extremely grave, whereas others may 
cause little or no discomfort. It is the latter class (which really require 
no treatment) that have given an entirely fictitious value and reputation 
to some of the remedies which have been proposed, for the patient re- 
covers in spite of the remedy, rather than through its use. 

In general, it can be said that even with the crudest treatment, or 
with no treatment of any kind, rattlesnake bite would probably not be 
fatal in more than 10 per cent of the cases, although greater with some 
especially dangerous species. Snake bite is likely to be more serious in 
the case of children, since the ability of a body to absorb venom without 
fatal results, varies with the weight. With proper treatment the mortal- 
ity from rattlesnake bite should be less than 3 or 4 per cent. 

In the case of an accident of this kind, be sure that the snake which 
has inflicted the wound is a venomous snake. Many harmless snakes will 
bite fiercely when trod upon or captured, but their bites are without any 
untoward effects; they are no more serious than a scratch and should be 
given a like antiseptic treatment. Nevertheless, there are authentic in- 
stances in which grave results and even death have been caused by fear 
following the bite of a harmless snake. 

The actual injection of rattlesnake venom into a wound is followed 
immediately by severe local pain in almost every case, and this should be 
used as a criterion in determining whether the bite is that of a rattler, and 
if venom has actually been injected. With most species a marked swell- 
ing is also evident within a very short time. 

Assuming that a person has actually been bitten by a rattlesnake, the 
following procedure should be adopted by the victim and his companions, 
if any be present: 

( 1 ) The victim should not become unduly alarmed or excited, and 
should not run, for to do so will speed up the circulation and the rapidity 
with which the venom is absorbed. Remember that few cases of rattle- 
snake bite are fatal. 

(2) Apply a tourniquet between the bite and the heart. This may 
be a shoe string, necktie, or a rubber band. Rubber tubing makes the 

214 San Diego Society of Natural History 

best tourniquet. Do not tie it too tightly. Complete stoppage of the 
circulation is unnecessary and undesirable, but the venous flow should 
be impeded. Loosen the tourniquet briefly at 15 minute intervals. 

(3) With a sharp instrument, such as a razor blade or a knife, make 
a cross-incision over each fang mark, or connect the two with a single in- 
cision. The depth should be about equal to that of the fang, say a quarter 
of an inch if the snake is of moderate size. Before using, sterilize the 
cutting instrument if possible, using iodine, alcohol, or the flame of a 

(4) Apply suction to the wound and the incisions thus made, either 
with the mouth or using one of the cupping or suction devices* which 
have been placed in first-aid kits for this purpose. Apply this continu- 
ously for at least half an hour. In a healthy person with good teeth 
there need be no fear of getting venom into the mouth or stomach with 
untoward results. 

(5) If antivenin is available, use it in accordance with the instruc- 
tions accompanying the syringe. However, do not depend upon it as a 
cure-all. Remember that antivenin and suction are not mutually exclu- 
sive; use antivenin if available, but the suction procedure should be car- 
ried through in any case. 

(6) If swelling or discoloration progresses up the limb, additional 
cross incisions should be made above this point and suction should be ap- 
plied there, the tourniquet having been moved above the swelling. It is 
best to put on a second tourniquet before removing the first. 

(7) If the patient is faint, give a cup of strong coffee or a teaspoon- 
ful of aromatic spirits of ammonia in a glass of water. 

(8) Get the patient to a doctor or hospital as soon as possible, secur- 
ing a physician experienced in previous snake-bite cases if one be avail- 

(9) Do not do any of the following things: Do not use potassium 
permanganate. Do not give whiskey. Do not burn or cauterize the 
wound, since this will interfere with the all-important suction and drain- 
age. Don't use "folk-lore'' remedies; they are a waste of time when 
time is valuable. 

(10) If the physician in charge of the case has not had previous 
experience he can secure advice from the United States Public Health 
Service by wire. The case should be closely watched for the first 24 
and preferably the first 48 hours. Some cases have been lost because the 
decline in the prominent hemorrhagic symptoms (evidenced by local swell- 
ing and discoloration) seemed to indicate that the danger was past, to be 
followed by a sudden and unexpected onset of neurotoxic symptoms. It 

* The rubber-bulb type is probably to be preferred since it will continue its action without an 

Klauber— Key to the Rattlesnakes 215 

is suggested that physicians called upon to treat rattlesnake bite, study the 
publications of the United States Public Health Service, or those of Dr. 
Dudley Jackson of San Antonio, who has had a wide experience in this 
field ; also the literature accompanying some of the suction devices now 
on the market in safety-first kits,* and the publications accompanying 
antivenin ampuls contain much useful information. It should be remem- 
bered, however, that these directions may be slightly biased as there has 
been some factional disagreement concerning the relative merits of anti- 
venin and suction. I repeat that antivenin and suction are not mutually 
exclusive remedies ; both should be used extensively in serious cases. The 
victim should always be typed so that a blood transfusion, if necessary, 
may be made without delay. Neurotoxic symptoms, frequently involving 
paralysis of the respiratory center, call for additional antivenin treatment 
The physician will use intravenous injections of glucose and normal salt 
solution as necessitated. 

The carrying of kits containing suction devices (there are several 
good ones on the market) is to be recommended to campers, hunters, or 
others going into rattler infested country. This is said without any de- 
sire to frighten people or to exaggerate the chance of snake bite, which is 
indeed remote. It is, however, a reasonable insurance precaution. 

The above brief remarks on the treatment of rattlesnake bite do no 
more than skim the surface. It must be remembered that most of the 
experience in this country has been in the treatment of cases of C. cine- 
reous bite and that of closely allied species. Rattlesnake venom is an 
exceedingly complex protein poison, having a variety of effects, neurotoxic, 
hemolytic, cytolytic, anti-bactericidal, etc. These effects probably differ 
considerably in the several species. It is well known that the venom of 
C. durissus differs extensively in its effects from that of C. cinereous and 
some of our more common nearctic species. We may well expect that 
future research will show that others of our North American rattlers have 
quite different effects than has C. cinereous. This in turn may influence 
the development of antivenins and otherwise profoundly change the pres- 
ent recognized methods of treatment. Polyvalent antivenins cannot be 
made as effective as those to counteract the bite of specific snakes. Prob- 
ably this is one of the reasons why Brazilian anticrotalus serum has been 
so successful; as there is but one species of rattlesnake in that country the 
antivenin is specific. In our country the situation is quite different. I 
anticipate that the future will see the venoms of our rattlers grouped in 
classes, with an antivenin for each class, although this could not be a suc- 
cessful commercial venture. In extensive areas of the country, where only 
one or two species of rattlers occur, only a single class would be required. 

* The directions accompanying the Dudley First Aid Kit are particularly complete with respect 
to the procedure of the suction treatment, both in the field and hospital. 

216 San Diego Society of Natural History 


Explanations of figs. 1 to In inclusive. 

Fig. 1. Dorsal View of Skull of Crotalus (C. ruber). 

The bones of the rattlesnake skull are thin and delicate. One of the 
distinguishing characters of snakes is the lack of a bony connection 
between the anterior ends of the mandibles, there being only an elastic 
ligament between these outer ends. This arrangement greatly facilitates 
the distension of the jaws, and by the independent action of the two 
sides, permits swallowing objects which are large relative to the 
size of the head. 

Fig. la. Ventral View of Skull. 

Note that the fangs are shown in the two outer maxillary sockets. 
This location is one of pure chance; with equal justification both 
might have been shown in the inner sockets, (see figs, lg and lh), 
or one in an inner and the other in an outer socket. As explained 
below, the fangs occupy the sockets alternately, and there is no 
synchronism in occupancy between the two sides of the head. 

Fig. lb. Lateral View of Skull. 

The fang is folded against the roof of the mouth in its resting posi- 
tion. The lower jaw is dropped slightly, and the reserve fangs are 
omitted for clarity. This is not a cross section of the skull; it is a 
view of the outside from the left. 

Fig. lc. Lateral View of Skull. 

The mouth is wide open and with the fangs directed forward in the 
position assumed at the end of the strike. 

(Figs. 1 to le, inclusive) 

1 Prexamilla (premaxillary) 12. Mandible (mandibular) 

,,,,,, 12a Dentary 

2. Prefrontal (lachrymal of some au- 12b Am Cu i ar 

thors ) 13. Pro-otic 

3. Frontal !4 Exoccipital (lateral occipital) 

4. Parietal 15. Poison fang 

5. Basisphenoid 16. Mandibular teeth 

6. Squamosal (supratemporal of some H. Pterygoid teeth 
authors) 18- Palatine teeth 

7. Maxilla (maxillary or supermax- 19. Supraoccipital 

illary) 20. Stapes (or columella auris) 

8. Palatine (palatal) 21. Postfrontal 

9. Pterygoid (internal pterygoid) 22. Basioccipital 

10. Ectopterygoid (external pterygoid or 23 - Nasal 
transpalatine) 24. Turbinal 

11. Quadrate 25. Vomer 

Klauber — Key to the Rattlesnakes 


23 2 3 21 10 4 13 6 20 II 14 

Fig. la. 

Fie. lc. 

218 San Diego Society of Natural History 

Figs. Id and le. The Fang Tilting Mechanism. 

The bones of the skull are indicated as a linkage. Fig. id shows the 
fang at rest against the roof of the mouth; in fig. le the change in 
the angle between the frontal-parietal-supratemporal and the quadrate 
tilts the fang forward into the biting position by pushing forward on 
the maxillary. The fang is not rotated forward automatically by the 
opening of the mouth; the tilting is controllable, otherwise the fangs 
would interfere with swallowing food. Each fang may be tilted 
independently, as the bones on one side of the head are independent 
of those on the other. 

Fig. If. Lateral View of Fang Seated in Maxillary. 

Figs, lg and lh. Front View of Fang. 

Fig. lg shows a fang in the inner maxillary socket; in fig. lh the 
next succeeding fang has been seated in the outer maxillary socket, 
while the fang shown in fig. lg has dropped out. In this way suc- 
ceeding active fangs occupy the two sockets alternately. (It should 
be noted that, because of the curvature of the fang, the lower part, in 
these figures, is viewed at an angle. This makes the orifice appear 
closer to the point than is really the case. See fig. li). 

Fig. li. Point of Fang (Perpendicular view). 

Figs, lj and Ik. Cross Sections of Fang. 

Fig. lj shows a cross section of the fang at the upper end just below 
the lower edge of the maxillary in which it is anchored. Fig. lk is a 
cross section just above the lower orifice. 

Fig. ll. Phantom Lateral View of Rattlesnake Head. 

This illustrates the location of the venom gland and duct in relation 
to the fang. 

Fig. lm. The Position of the Reserve Fangs. 

The reserve fangs do not tilt with the active fang but remain in place 
against the roof of the mouth. 

Fig. In. Diagram Showing Fang Succession. 

The two large circles represent the maxillary sockets, the left-hand 
being occupied by a functional fang. When this is ready to drop 
out, reserve fang No. 1 advances and becomes anchored in the vacant 
socket on the right. Later, when No. 1 is about to be superseded, No. 
2 advances into the left-hand socket. Thus, the replacements are 
made periodically. New buds also appear periodically, so that, in all, 
about 8 reserves, in consecutive stages of development, are always 
present. The reserves which will ultimately be seated in the left- 
hand socket are separated from those intended for the right by a mem- 
branous wall. It should be understood that this discussion has had 
reference exclusively to the condition on one side of the upper jaw. 
There is a duplicate set of sockets and reserve fangs on the other side. 
In this diagram the reserve fangs have been spread out for reasons of 
clarity. Actually they lie closely bunched in a membranous sac above 
the active or functional fang, in the position indicated in fig. lm. 

Klauber — Key to the Rattlesnakes 


Fig. If- 



Fig. 1 

Fig. lh. 



Fig. lj. Fig. Ik. 


Fig- Li. 




Fie. U. 

Fig. lm. 


San Diego Society of Natural History 


While many users of this key will be familiar with the technical 
terms employed, such as the names of the scales used in herpetological 
classification, there may be others, especially physicians interested in snake- 
bite cases, who may not have had occasion to employ them. In addition 
there are certain peculiarities of the rattlesnakes which render it desirable 
to explain a few of the terms to those who have not worked in this group. 
For these reasons an illustrated glossary has been included. 

Anal plate: The large plate covering the vent (fig. 3). It marks the division 
between body and tail. 

Apical scale-pit s: A pair of depressions faintly evident on the posterior end of 
each scale ; usually most evident dorsally near the tail. 

Body blotches: These are counted from the posterior edge of the head to the anus; 
the tail rings are not included. On the sides there are usually additional 
series of smaller blotches known as the lateral or secondary blotches, often 
in several rows, one below the other. In many species of rattlesnakes, 
especially on the posterior half or third of the body, the main dorsal 
blotches merge with the laterals to form crossbars or rings. 

Button: See rattles. 

Canthals: The border scales of the crown between the internasals and the supra- 
oculars (fig. 5). For intercanthals see prefrontals. 

Canthus rostralis: The outer edge of the flat area of the crown where it turns 
downward on the side, extending from the rostral to the supraocular 
(fig- 5). 

Caudals: See subcaudals. 

Frontal: The large plate between the supraoculars in Sistrurus (fig. 4). In 
Crotalus, this space is filled with scales more or less irregularly disposed 
(fig. 5) and is referred to as the frontal area, and the scales as the inter- 
supraoculars.' When the "minimum scales between the supraoculars" 
are specified, the path traversing the fewest scales is meant; this is usually 
at the anterior part of the frontal area. 

Genials: The genials or chin-shields are a pair of enlarged scales back of the first 
infralabials (figs. 7 and 8). Occasionally the posterior tips of the first 
infralabials are cut off to form an extra pair of triangular scales which 
are called intergenials (fig. 46). Snakes other than rattlesnakes often 
have two pairs of genials, an anterior and posterior. 

Gulars: The small scales covering the underside of the head between the two rows 
of infralabials and not otherwise specifically named (fig. 8). 

Head marks: Although there is a considerable variation in the head marks of the 
rattlesnakes there are some which occur in many species ; these are indi- 
cated as to general position and direction in figs. 9 and 10. The light 
supraocular crossbars in some species are present only on the supraoculars 
and not in the intervening frontal area. 

Infralabials: See labials. 

Klauber — Key to the Rattlesnakes 




Fig. 2. Methods of measurement. 

Fig. 2a. Method of counting dorsal scale rows. 

l ,ANAL 


Fig. 3. Ventral view of tail with nomenclature. 



Fig. 4. Nomenclature of head scales of Sistrurus. 

222 San Diego Society of Natural History 

Intercanthals: See prefrontals. 

lnternasals: The scales in contact with the rostral from nasal to nasal regardless 
of size (figs. 4 and 5). In most rattlesnakes there are two; in viridis 
generally three or more (fig. 49). 

Inter supraoculars: See frontal. 

Labials: Bordering the mouth above are the supralabials (or upper labials) which 
extend from the rostral to the rictus of the mouth or commissure (figs. 6 
and 7). The rostral is not counted as one of the supralabials. Similarly 
the infralabials (lower labials) extend along the lower lip from the 
mental to the angle of the mouth (figs. 6, 7, and 8). The first pair 
of infralabials are sometimes divided transversely (fig. 42). 

Lengths: The length over-all (or body length) is measured from the tip of the 
snout (rostral) to the forward edge of the proximal rattle (fig. 2). 
Head length is measured from the rostral to a line joining the posterior 
tips of the mandibular bones. Tail length is from the anus to the for- 
ward edge of the proximal rattle. 

Loreals: The scales (one or more) on the side of the head between the postnasal 
and the preocular (fig. 6). No species of rattlesnake is regularly without 
at least one loreal on each side, although rarely an individual will have 

Mental: The triangular scale at the anterior tip of the lower jaw (figs. 7 and 8). 
Occasionally the posterior tip of the mental may be cut off to form a 
submental (fig. 47). 

Nasals: A pair of scales on either side of the nostril, called respectively, the 
prenasal and postnasal (figs. 6 and 7). 

Oculars: The scales surrounding the eye. The supraoculars are large and jut over 
the eyes (fig. 6). In front of the eyes there are usually two preoculars, 
the upper larger, the lower narrow and crescent shaped. Back of, and 
below the eye, are the postoculars and suboculars. It is usually difficult 
to determine which scale should be considered the lowest postocular, and 
which the first subocular. 

Parietals: A pair of large plates posterior to the supraoculars and frontal in 
Sistrurus (fig. 4). This area is occupied by irregular scales in Crotalus 
(fig. 5). 

Pit: A deep depression on the side of the head below and back of the nostril 
(fig. 6) ; this is the external opening of a sensory organ, probably 
auditory in function. Where the pit-border scales are mentioned, those 
constituting the internal rim or lip are meant, rather than those which 
are completely external to the pit. 

Postnasals: See nasals. 

Postoculars: See oculars. 

Prefrontals: In Sistrurus the two large plates posterior to the internasals (fig. 4). 
In Crotalus, with a few exceptions of which durissus is an example, 
this space (often referred to as the prefrontal area) is filled with irregu- 
larly disposed scales called the intercanthals (fig. 5). 

Prenasal s: See nasals. 

Klauber — Key to the Rattlesnakes 







Fig. 5. Nomenclature of head scales of Cro talus, dorsal view. 









Fig. 6. Nomenclature of head scales of Crotalus, lateral view. 


PIT — 




Fig. 7. Nomenclature of head scales of Crotalus, front view. 

224 San Diego Society of Natural History 

Preoculars: See oculars. 

Rattles: The rattle terminology is illustrated in figs. 11 and 12. The proximal 
rattle is that next the tail and is the one most recently added to the string. 
The button (or rattle-button) is the first permanent rattle acquired by a 
young snake, the rattle present at birth (the prebutton) being invariably 
lost with the first exuviation. The button remains as the posterior 
terminus of the rattle-string until lost by breakage; it is usually present 
in juveniles or young adults, but rarely in older specimens. In some 
snakes, particularly if only one or two rattles have been lost, it is a 
trifle difficult to tell whether the button is still present; however this can 
be ascertained quite definitely with a little practice in the study of 
juvenile termini. The rattle width of any segment is measured as indi- 
cated in fig. 12. It is to be noted that the measurement (to secure the 
greatest width) is not exactly vertical, this being the result of the asym- 
metry in the rattle shape designed to prevent the rattle from dragging on 
the ground as the snake crawls. In this key the widths of the proximal 
rattle and the button are occasionally used. 

Rostral: The large scale on the front of the nose (figs. 4, 5, 6 and 7). 

Scale-boss: A knobby prominence or swelling on the posterior part of each scale, 
particularly evident on the middorsal rows of some species. It is to a 
certain extent independent of the keeling (or central ridge on each 
scale) which is present on all but the one or two lowest dorsolateral rows 
in all rattlesnake species. 

Scale roics: Where the number of dorsal scale rows is given, the number at mid- 
body is meant, beginning with the row next to the ventrals on one side 
and ending with the corresponding row on the other (fig. 2a). Oc- 
casionally the scale rows at the center of the tail are referred to; these 
should be counted midway between the anal plate and the anterior edge 
of the proximal rattle. 

Subcaudals: The subcaudals (caudals or urosteges) are counted beginning with 
the first scale on the mid-ventral line posterior to the anal plate and end- 
ing with the last scale anterior to the proximal rattle (fig. 3). Divided 
scales, that is, those having mid-ventral sutures, frequently found toward 
the beginning or end of the series, are treated as if undivided. The 
fringe of small and irregular scales which occasionally covers the an- 
terior edge of the rattle is not counted as a caudal. 

Supralabials: See labials. 

Supraoculars: The large plates above each eye (figs. 4, 5, 6 and 7). Also see 

Suture: A division or crease between two scales or plates, or the parts of a plate. 

Tail rings: The dorsal rings between the anus and the proximal rattle. They are 
often obscure and can only be counted approximately. 

Ventrals: The large plates on the belly. In counting the ventral plates (sometimes 
called the gastrosteges) the count is begun with the first scale on the 
underside of the head which is distinctly wider than long (fig. 8), and 
ends with the scale anterior to the anal plate, but does not include the 
the latter (fig. 3). 

Klauber — Key to the Rattlesnakes 









Fig. 8. Nomenclature of head scales of Crotalus, ventral view. 


Fig. 9. Head pattern of Crotalus, Fig. 10. Head pattern of Cro- 
dorsal view. talus, lateral view. 



Fig. 11. Complete rattle with 


Fig. 12. Incomplete rattle. 

226 San Diego Society of Natural History 


Start at 1 and decide whether the specimen to be identified is cor- 
rectly described by paragraph la or by lb. If the former be the case, 
the snake belongs to the genus Sistrurus; if the latter, to the genus Cro- 
talus. Assume, in this instance, that it is a -Crotalus. Note that the bold- 
faced figure 7 appears at the end of the final line in paragraph lb ; this is 
an instruction to proceed to the paragraphs headed 7a and 7b. Now 
decide which of these alternative descriptions fits the specimen in hand. 
If it fits description 7a, note that the bold-faced figure 8 appears at the end 
of .the last line of the description; you are therefore next to choose 
between paragraphs 8a and 8b. If, on the contrary, course 7b is found to 
be the proper one, then the specimen is Crotalus stejnegeri and the identi- 
fication is concluded. Thus by successive selections of one of pairs of 
alternatives a final decision is reached. 

When an identification has been made it is recommended that the 
color description, the range, and the photograph (if one is given) be also 
checked against the specimen and its data, so that an inaccurate con- 
clusion may be avoided, even if some peculiarity of the specimen, or 
ambiguity in the key, has caused a wrong turning at one of the branch 
points. The table of scale counts may also serve as a check. 

It should be observed, with reference to the line drawings in the 
text (figs. 1-64), that the stippling is usually not for the purpose of indi- 
cating punctations in the pattern of the snake but more often serves to 
suggest the even application of some color other than black. 
****** * * 

The writer will welcome correspondence with users of this key 
calling his attention to errors or discrepancies. New locality records, es- 
pecially if verified by live or preserved specimens (heads or skins are 
usually sufficient for identification), will be much appreciated in order that 
the range maps may be more accurate in future publications. 


. To the many persons and institutions which have assisted me by the 
gift and loan of specimens during this investigation of the rattlesnakes I 
shall make acknowledgments in subsequent papers. At this time however 
I wish to give credit for the line drawings in this key to Mr. Norman 
Bilderback, and for the photographs, maps, and also the lettering on the 
drawings to Mr. Leslie C. Kobler, both of San Diego. 

Klauber — Key to the Rattlesnakes 



1 a. Top of head with large plates anteriorly (usually 9 in number) 
including a single frontal and a pair of large, symmetrical parietals 
in contact (fig. 13).* Genus Sistrurus 2 

1 b. Top of head with scales of varying size; more than one scale in 

the frontal area; parietals, if enlarged, not in contact, nor sym- 
metrical (fig. 14). Genus Crotalus 7 

2 a. Rostral not curved over the snout (fig. 15) ; canthus rostralis 

sharply angled ; dorsal series of body blotches about equal in width 
and length, or shorter (along the snake) than wide. 3 

2 b. Rostral curved over the snout (fig. 16) ; canthus rostralis rounded, 

not sharply angled ; dorsal series of body blotches distinctly longer 
than wide (fig. 17). A pattern of dark-brown blotches on a light- 
brown ground-color, with a lateral series on each side shorter 
(longitudinally) and usually darker than the dorsal series. 

Sistrurus ravus 
A small area of the Mexican plateau including: eastern Mexico (state), 
southern Hidalgo, Tlaxcala, northeastern Puebla, west-central Veracruz, 
and northern Oaxaca (fig. 73). 

3 a. Upper preocular usually in contact with postnasal ; usually 3 supra- 

labials in contact with the pit-border scales (fig. 15) ; 11 or more 
dorsal scale rows at center of tail ; rattles larger, width of proximal 
rattle being contained in body length (over-all) less than 90 

* Deviations of snakes of the genus Sistrurus from the standardized arrangement of the nine large 
plates on the crown (as illustrated in fig. 13) are not particularly rare, but such deviations usually consist 
of small and insignificant extra scales detached from the posterior end of the frontal or, less often, from 
the rostral. It is quite rare to find the nine large plates themselves seriously distorted in arrangement, 
although specimens have been noted with a central prefrontal and a canthal on either side. None of 
these aberrant specimens will be confused with any Crotalus individual if the criteria given in the key- 
be followed, for although there are Crotalus species (durissus for example) which have large paired inter- 
nasals and prefrontals (fig. 56), they lack the single frontal, and the paired and conterminous parietals 
of Sistrurus. 

Fig. 13. Dorsal head plates of 
Sistrurus (S. c. catenatiis). 

14. Dorsal head scales of 
Crotalus (C. cinereous). 


San Diego Society of Natural History 

times, or in head length less than 6 times ; no red or orange in the 
interblotch spaces on the middorsal line. Sistrurus catenatus 

(For subspecies continue on to 4) 

3 b. Upper preocular not in contact with postnasal; usually 2 supra- 

labials in contact with pit-border scales (fig. 18) ; 10 or less dorsal 
scale rows at center of tail ; rattles smaller, width of proximal rattle 
being contained in body length more than 90 times, and in head 
length more than 6 times ; usually with red or orange between the 
blotches on the middorsal line. Sistrurus miliarius 

(For subspecies continue on to 5) 

4 a. Undersurface dark, heavily clouded with black blotches, often 

almost solid black (fig. 19) ; scale rows usually 25 ; body blotches 
usually less than 37. A pattern of square, red-brown, or black 
blotches on a gray-brown ground; sometimes unicolor black. 

Sistrurus catenatus catenatus 
(Fig. 106). 
From central New York westward to eastern Oklahoma including: 
New York from Madison County west; southern Ontario, south and 
west of the line Spanish — North Bay — Port Hope; extreme western 
Pennsylvania; northern and central Ohio; lower Michigan (including 
Bois Blanc Island); central and northern Indiana; Illinois; southern 
and western Wisconsin; eastern and southern Iowa; extreme south- 
eastern Nebraska ; Missouri ; eastern Kansas and eastern Oklahoma, 
intergrading here with S.c. tergeminus (fig. 73). 

Fig. 15. S. c. catenatus, showing 
rostral not curved over snout; 
also preocular contacting post- 

Fig. 16. S. ravus, showing rostral 
curved over snout. 

Fig. 17. S. ravus; body pattern. 

Fig. 18. S. miliarius, showing 
preocular separated from post- 

Klauber — Key to the Rattlesnakes 


4 b. Undersurface mottled or spotted, the dark areas being less exten- 

sive than the light (fig. 20) ; scale rows often 23; body blotches 
usually more than 36. A pattern of dark red-brown blotches on a 
gray-brown ground. Sistrurus catenatus tergeminus 

(Fig. 107). 
The southwestern plains including: central and southwestern Kansas; 
extreme southeastern Colorado; central and western Oklahoma; the 
plains areas of central and southern New Mexico; extreme south- 
eastern Arizona; Texas, west of the Brazos River; and extreme 
northern Tamaulipas, Mexico. Intergrades with S.c. catenatus in east- 
ern Kansas and Oklahoma (fig. 73). 

5 a. Dorsal coloration brown or light-gray ; ventral surface cream, mod- 

erately flecked with brown or gray ; head markings distinct ; lateral 
spots in 1 or 2 series. 6 

5 b. Dorsal coloration dark-gray to black; ventral surface white, heavily 

blotched with dark-brown or black; head markings obscure; lateral 
spots in 3 series Sistrurus miliarius barbouri 

(Fig. 104). 
From extreme southern South Carolina (where it intergrades with 
S.m. miliarius) and southern Georgia, south throughout Florida and 
westward across southern Alabama to southeastern Mississippi, inter- 
grading with S.m. streckeri in the Pearl River Valley (fig. 73). 

6 a. Dorsal scale rows usually 21; dorsal spots wider than long and 

with irregular edges ; lateral spots usually higher than wide ; ven- 
tral spots confined to individual plates. 

Sistrurus miliarius streckeri 
(Fig. 105). 
From the Pearl River Valley of southeastern Louisiana and western 
Mississippi (where it intergrades with S.m. barbouri) westward 
through Louisiana and eastern Texas (north of Lat. 28°) to Long. 
98° ; also north through Arkansas to southern Missouri and west to 
central Oklahoma; also southwestern Tennessee (fig. 73). 

6 b. Dorsal scale rows usually 23; dorsal spots oval or subcircular, 
edges even; lateral spots usually round; ventral spots usually oc- 
cupying two adjacent plates. Sistrurus miliarius miliarius 

(Fig. 103). 

Fig. 19. S. c. catenatus; ventral 

Fig. 20. S. c. tergeminus ; ventral 


San Diego Society of Natural History 

7 a. 

7 b. 

8 a. 

8 b. 

From extreme southern South Carolina (where it intergrades with 
5./;?. barbouri) north throughout South Carolina and eastern North 
Carolina to the Pamlico River. Also central Georgia and central 
Alabama (fig. 73). 

Males with less than 40 subcaudals; females with less than 35 
subcaudals; adults with proximal rattles wider than 3^mm. 8 

Males with more than 40 subcaudals; females with more than 35 
subcaudals; rattles very small, width less than 3V2 mm. in adults. 
A pattern of about 40 black-edged, olive-gray diamonds on a gray- 
brown ground. Crotalus stejnegeri 

The mountains of southeastern Sinaloa and western Durango in 

Mexico (fig. 71). 

Outer edges of supraoculars not extended into raised and flexible 
hornlike processes (fig. 21). 9 

Outer edges of supraoculars extended into raised and flexible 
hornlike processes distinctly pointed at the tip (figs. 22 and 23). 
Dorsal scales strongly keeled and with posterior bosses. Ground 
color cream, straw, pink, or light-gray, with a central series of 
square, brownish blotches, often with yellow or orange on the 
middorsal line. Crotalus cerastes 

(Fig. 95). 

The deserts of the southwest including: California east of the Sierra 
Nevada and the southern California coastal ranges, from Lat. 37° 30' 
southward; Nevada south of Lat. 37° 30'; west-central and south- 
western Utah; Arizona south and west of the line Kingman — Miami — 
Nogales; extreme northwestern Sonora; the transmontane desert area 
of northeastern Baja California from the U. S. border south to Lat. 
29°40' (fig. 71). A species preferring sandy deserts but sometimes 
found on rock-strewn flats. (Note: In the Southwest the range of 
cerastes, is popularly supposed to be considerably more extensive than 

Fig. 21. Lateral head scales of 
Crotalus (C. cinereous). 

Fig. 22. C. cerastes, showing 

hornlike supraocular (lateral 

view) . 

Fig. 23. C. cerastes (front view). 

Klauber — Key to the Rattlesnakes 


is here depicted, since in many areas all young rattlesnakes are termed 
"sidewinders", leading to confusion with the real cerastes. While 
further collecting may be expected to develop some range extensions, 
this most highly specialized of our desert snakes cannot possibly exist 
in some areas where it has been said to occur, but from which no 
authentic specimens have ever been forthcoming). 

9 a. Tip of snout and canthus rostralis not raised into a sharp ridge 
(fig. 24) ; no central light line on rostral and mental. 10 

9 b. Tip of snout and canthus rostralis raised into a sharp ridge, by 
bending up of the outer edges of internasals and canthals (fig. 
25) ; rostral and mental usually marked vertically by a narrow, 
light line on a red-brown ground (fig. 26). Body pattern of 
large, brown or red-brown blotches separated by narrow light 
areas, the blotches being often without definite outlines on the 
sides; tail pattern terminating in logitudinal bands rather than 
crossbars. Crotalus willardi 

(Fig. 102). 
Highland areas from extreme southern Arizona to Zacatecas, Mexico 
including: the Santa Rita and Huachuca mountains in Arizona; and 
the Sierra Tarahumare and Sierra Madre in eastern Sonora, western 
Chihuahua, Durango, and western Zacatecas (fig. 72). 

Fig. 24. Cross-section of Crotalus 

head (C. m. mitchellii), showing 

absence of internasal ridge. 

Fig. 26. Head of C. willardi, 
showing light line usually pres- 
ent on rostral and mental. 

Fig. 25. Cross-section of C. wil- 
lardi head, showing internasal 

Fig. 27 C. poly st ictus, showing 

indentation on upper edge of 



San Diego Society of Natural History 

10 a. 

10 b. 

11 a. 

11 b. 

Head width usually greater than 60 per cent of the length; pre- 
nasal not deeply indented by a lateral bulge in the first canthal 
(fig. 21) ; paired intercanthals, if present, not distinctly longer 
than wide. 11 

Head long and narrow, maximum width less than 60 per cent of 
length; prenasal deeply indented by a lateral bulge in the first 
canthal (fig. 27) ; paired intercanthals longer than wide (fig. 28) ; 
usually 8 scales in two transverse rows on top of the head anterior 
to the supraoculars and intersupraoculars (maximum variation 6 to 
10) ; dorsal pattern usually of parallel rows of dark-brown ellipti- 
cal blotches, the major axes of which are longitudinal to the snake 
and with narrow gray-brown interspaces (fig. 29) ; ventral surface 
heavily mottled with dark-brown or black. Crotalus polystictus 
The tableland of central Mexico from Zacatecas to Oaxaca including: 
southern Zacatecas, eastern Jalisco, Guanajuato, Michoacan, Distrito 
Federal, west-central Veracruz, Oaxaca, and probably the intervening 
states of Aguascalientes, Queretaro, Hidalgo, Mexico, Morelos, Tlax- 
cala, and Puebla (fig. 69). Unverified reports from southern Jalisco 
and eastern Colima. 
No paired dark vertebral stripes on the neck; or if present, not 
extending as much as 1% head-lengths before meeting the first 
dorsal blotches; vertebral process less sharp and prominent; dorsal 
scales without such prominent posterior bosses.* 13 

A pair of dark vertebral stripes 1 to 3 scales wide on the neck, 
bounding a lighter middorsal stripe about 3 scales wide, and ex- 
tending posteriorly 1% head-lengths or more before meeting the 
first dorsal blotches ; spinous process sharp and ridge-like ; posterior 
head scales and dorsal scales strongly keeled and with conspicuous 
posterior bosses ; usually with only 4 and rarely with more than 6 
large flat scales on the crown anterior to the supraoculars and 
intersupraocubrs. Crotalus durissus 

(For subspecies continue on to 12) 

* There are rare instances of paired vertebral stripes in viridu; these can readily he differentiated 
from durissus by the presence of more than 6 scales in the internasal — prefontal area. Similarly occa- 
sional specimens of durissus occur in which the characteristic anterior paravertebral stripes are replaced 
by blotches. As far as we now know any rattlesnake from Central or South America ;hould be classified 
as durissus, except that umcolor may occur in Nicaragua. 

Fig. 28. C. polystictus. showing 
paired and elongated inter- 

Fig. 29. Dorsal pattern of C. 

Klauber — Key to the Rattlesnakes 


12 a. Light scale rows (one scale wide), laterally bounding the dark 
paravertebral stripes (or dorsal blotches) in strong color-contrast 
with the next scales below on the sides (fig. 30) . A pattern of 
brown dorsal diamonds on a brown ground color, without strong 
contrast between the dorsal and lateral areas. 

Crotalus durissus terrificus 
(Fig. 75). 
South America and eastern Central America including: Argentina, 
Uruguay, Paraguay, Brazil, Bolivia, eastern Peru, eastern Ecuador, 
the Guianas, Venezuela, Colombia, and central and eastern Costa 
Rica where there is intergradation with C. durissus durissus (figs. 65 
and 66). Presence in Panama and Canal Zone somewhat doubtful. 

12 b. Light scale rows (one scale wide), laterally bounding the dark 

paravertebral stripes (or dorsal blotches) but little or no lighter 
than the lateral areas next below (fig. 31). A pattern of dark- 
brown or black dorsal diamonds (usually with light centers) in 
strong contrast to the cream, yellow, gray, or light-brown ground 
color of the sides. Crotalus durissus durissus 

(Fig. 74). 

Southern Mexico and Central America including: the Mexican states 
of Michoacan, Guerrero, Distrito Federal, central Puebla, central and 
southern Veracruz, Oaxaca, Tabasco, Chiapas, Campeche, Yucatan, and 
probably also Morelos, Mexico, Tlaxcala, and Quintana Roo ; Guate- 
mala, British Honduras, Honduras, El Salvador, and Nicaragua (fig. 
66). The area of intergradation with C. durissus terrificus seems to 
be approximately central Costa Rica, although the change is so gradual 
that specimens between Honduras and Panama may be difficult to allo- 
cate, the pattern being somewhat dependent on altitude. 

13 a. Upper preocular not split vertically; or, if split,* the anterior sec- 

tion not conspicuously higher than the posterior and not curved 
over the canthus rostralis in front of the supraocular ; prenasal not 

As is frequent in mitchellti and occasional in triseriatus. 


Fig. 30. Dorsal neck pattern of 
C. d. terrificus. 

Fig. 31. Dorsal neck pattern of 
C. d. durissus. 


San Diego Society of Natural History 

curved under the postnasal (fig. 21) ; the pattern not of widely 
separated crossbars or rings. 15 

13 b. Upper preocular split vertically, the anterior section being higher 

than the posterior and curved over the canthus rostralis in front 
of the supraocular; prenasal curved under the postnasal (fig. 32) ; 
usually a pattern of widely separated crossbars (fig. 33). 

Crotalus lepidus 
(For subspecies continue on to 14). 

14 a. A dark stripe passing backward from the eye to the angle of the 

mouth (fig. 32) ; dorsal pattern of crossbars often not strongly 
differentiated from the ground color; ventral surface mottled. A 
pattern of 13 to 22 brown or black blotches or crossbars on a 
punctate background of gray, brown, or pink flecked with gray, 
the blotches being separated by a much greater distance (neglect- 
ing rudimentary* or obsolescent blotches which are often present) 
than the longitudinal extent of the blotches; tail usually pink or 
reddish, crossed by 4 or less widely separated brown rings. 

Crotalus lepidus lepidus 
(Fig. 98). 

West Texas and northeastern Mexico including: the trans-Pecos region 
of Texas (especially the Davis and Chisos mountains, but excluding 
the El Paso area, where it is replaced by C.l. kla-uberi), and Valverde, 
Real, and Maverick counties; Coahuila, northern San Luis Potosi, and 
probably northern Zacatecas (fig. 69). 

14 b. No dark postocular stripe; dorsal pattern of crossbars sharply con- 
trasting with the ground color; ventral surface punctated. A 
pattern of 14 to 21 dark reddish-brown, or black blotches, or 
crossbars on a background of green, blue-green, or blue-gray, the 
blotches being separated by a much greater distance than their 
longitudinal extent (fig. 33) ; tail usually cream or pink, crossed 
by 4 or less widely separated brown rings. A subspecies pre- 
ferring mountains. Crotalus lepidus klauberi 

(Fig- 99). 
Southeastern Arizona, southern New Mexico, the El Paso area in 
Texas, and north-central Mexico including: the Santa Rita, Huachuca, 

* Occasionally these are almost as evident as the fundamental bars thus producing a pattern of 
closely adjacent bands. 

Fig. 32. C. I. lepidus, showing 
split upper preocular and pre- 
nasal curved under postnasal. 

Fig. 33. C.l. klauberi; dorsal 

Klauber — Key to the Rattlesnakes 


Dragoon, and Chiricahua mountains of southeastern Arizona ; the 
Franklin, Magdalena, Pinos Altos, Mimbres, Animas, Big Hatchet, 
and Dog mountains of southern New Mexico; El Paso County, Texas; 
and mountain areas in the Mexican states of Chihuahua, Durango, 
southern Zacatecas, Aguascalientes, eastern Nayarit, and northern and 
eastern Jalisco (fig. 69). 

15 a. Prenasals in contact with rostral (fig. 34) ; upper preocular not 
divided, or if divided (as in a few triseriatus), the loreal con- 
spicuously longer than high. 17 

15 b. Prenasals usually separated from the rostral by small scales or 

granules (fig. 35) ; upper preoculars often divided, horizontally, 
vertically, or both (fig. 36); rostral usually wider than high; a 
pattern of dorsal blotches essentially comprising aggregations of 
punctations. 16 

16 a. Head smaller; length of head contained in adult body length more 

than 24 times; original rattle-button (fig. 11), if present, more 
than 7.5 mm. wide. A pattern of dark-gray or brown, punctated 
blotches on a gray or tan background. 

Crotalus mitchellii mitchellii 
(Fig. 91). 

Distrito del Sur of Baja California, Mexico, intergradation with C. 
mitchellii pyrrhus occurring approximately along the border of the 
two districts; also the islands of Ceralvo, Espiritu Santo, San Jose 
(Gulf Coast), and Santa Margarita (Pacific Coast). Fig. 69. 

16 b. Head larger; length of head contained in adult body length less 
than 24 times; original rattle-button, if present, less than 7.5 mm. 
wide. A pattern of red, gray, brown, or black, punctated blotches 
on a cream, tan, buff, gray, pink, salmon, fawn, or brown back- 

Fig. 34. Head scales of Crotalus; 
front view (C. cinereous). 

Fig. 35. C. m. pyrrhus; front 
view, showing separation of ros- 
tral from prenasals; also rostral 
wider than high. 

Fig. 36. Cm. mitchellii; lateral 
view showing split upper pre- 


San Diego Society of Natural History 

ground; often with posterior black tips on some dorsal scales 
between blotches Crotalus mitchellii pyrrhus 

(Fig. 92). 
Southern California, western Arizona, and northern Baja California 
including the following: California south of the line Barstow— 
Ivanpah (approximate line of intergradation with Cm. stepbensi) 
and east of Long. 118° (but absent from the San Gabriel Mountains 
and the coastal plain) ; the southern tip of Nevada; Baja California 
south to the northern boundary of Distrito del Sur (where inter- 
gradation with mitchellii mitchellii is to be expected) ; Isla Angel de la 
Guarda; extreme northwestern Sonora; west-central and southwestern 
Arizona, inside the line Peach Springs — Williams — Casa Grande — Ajo 
(fig. 69). A species preferring rocky habitats. 

17 a. Tail of alternating black, and light ash-gray rings, both colors 
being in sharp contrast with the posterior body color (fig. 37), 
which may be gray, dark-gray, cream, pink, red, red-brown, or 
olive-brown.* 18 

17 b. Tail not of alternating black, and light ash-gray rings in strong 

color-contrast to the body color immediately anterior to the tail. 


18 a. Dark and light tail rings of approximately equal width (fig. 37) ; 

postocular light stripe, if present, intersects the supralabials from 
1 to 3 scales anterior to the angle of the mouth (fig. 38) ; minimum 
scales between supraoculars 3 or more (fig. 14) ; no definite line 
of demarcation between the scales in the frontal and prefrontal 
areas ; proximal rattle black. Cinereous group 19 

* Many other species of rattlers have barred tails but it is characteristic of the cinereous group that 
there is a sharp transition in color at the beginning of the tail, whereas in the others there is no sharp 
contrast between the posterior body rings and the anterior tail rings. The cinereous-type tail has the ap- 
pearance of having been attached to the wrong snake. 

Fig. 37. Tail pattern of cinereous 

Fig. 38. C. cinereous; lateral 
head marks. 

Fig. 39. C. scutulatus; lateral 
head marks. 

Klauber — Key to the Rattlesnakes 


18 b. Dark tail rings narrower than light; postocular light stripe, if 
present, passes backward above the angle of the mouth (fig. 39) ; 
minimum scales between the supraoculars rarely more than 2 ; a 
definite division line or suture between the scales in the frontal 
and prefrontal areas (fig. 40) ; lower half of proximal rattle light 
in color. A pattern of brown hexagons or diamonds on a green, 
olive-green, or brown background ; light scales bordering the dark 
diamonds are unicolor, it being characteristic of this species that 
the blotch edges follow the scales and do not cut them. 

Crotalus scutulatus 
(Fig. 84). 

From the Mojave Desert in California southeastward to south-central 
Mexico including: Kern, Los Angeles (Antelope Valley), and San 
Bernardino counties in California but only eastward of the Tehachapi, 
San Gabriel, and San Bernardino mountains ; the desert valleys of 
Lincoln and Clark counties, Nevada ; extreme southwestern Utah ; 
Arizona south and west of the line Williams — SafFord; extreme south- 
western New Mexico ; trans-Pecos Texas ; northeastern Sonora, Chi- 
huahua, eastern Durango, southern Coahuila, southwestern Tamaulipas, 
Zacatecas, San Luis Potosi, and southeastward at least to Tlaxcala, no 
doubt including the intervening Mexican plateau states of Guanajuato, 
Queretaro, and Hidalgo (fig. 66). 

Fig. 40. Dorsal head scales of C. 

scutulatus showing paired inter- 


Fig. 41. Chin shields of Cro- 
talus; first infralabials undivided. 

Fig. 42. Chin shields of Cro- 
talus; first infralabials divided. 


San Diego Society of Natural History 

19 a. First infralabials usually not divided transversely (fig. 41) ; general 
color cream, buff, gray, or gray-brown (sometimes pink or red in 
central Arizona or New Mexico) ; dark punctations conspicuous 
in markings. 20 

19 b. First infralabials usually divided transversely (fig. 42) ;* general 

color pink, red, brick-red, red-brown, or olive-brown; dark punc- 
tations weakly in evidence or absent from markings. 21 

20 a. Upper preocular usually not in contact with the postnasal and no 

upper loreal present (fig. 43) ; head smaller in proportion to 
body. Pattern of dark-brown, punctated diamonds with lighter 
centers on a gray background, and with light borders of the 
diamonds often absent laterally. Crotalus tortugensis 

(Fig- 81). 
Tortuga Island in the Gulf of California (fig. 67). 

20 b. Upper preocular usually in contact with the postnasal (fig. 44), 
or such contact prevented by an upper loreal (fig. 45) ; head pro- 
portionately larger. Pattern of brown dorsal diamonds consisting 

* A considerable proportion of California cinereous have divided first infralabials and one must em- 
ploy the pattern criteria to judge California specimens or he will be improperly led to course 21 instead of 
20. The blotches of cinereous are always strongly punctate, those of ruber almost unicolor. 

Fig. 43. Snout of Crotalus; no 

contact between postnasal and 


Fig. 44. Snout of Crotalus; con- 
tact between postnasal and pre- 
ocular; also between prenasal 
and first supralabial. 

Fig. 45. Snout of Crotalus; up- 
per loreal present; no contact 
between prenasal and first supra- 

Klauber — Key to the Rattlesnakes 


of aggregations of punctations on a cream, buff, gray, gray-brown, 
or (rarely) reddish background. Crotalus cinereous 

(Figs. 80 and 108). 
From Arkansas and Texas south to central Mexico and west to Cali- 
fornia including the following: extreme southeastern Missouri; north- 
eastern, central, and west-central Arkansas; western and southern 
Oklahoma; Texas (except the Panhandle and east of Long. 95°) ; New 
Mexico south of Lat. 36° but not including the mountains of the 
central-west; Arizona south and west of the line Ash Fork — Clifton; 
the southern tip of Nevada ; the desert areas of Riverside and Im- 
perial counties, California, touching extreme eastern San Diego 
County; extreme northeastern Baja California; and the Mexican states 
of Sonora, Chihuahua, Coahuila, Nuevo Leon, Tamaulipas, San Luis 
Potosi, and the northern tip of Veracruz; also Tiburon Island in the 
Gulf of California; probably present in northeastern Durango and 
northern Zacatecas (fig. 67). Has been introduced and seems to 
have gained a foothold in Vernon County, Wis. 

21a. Intergenials usually absent (fig. 41) ; prenasals generally in contact 
with first supralabials (fig. 43) ; tail rings complete, or broken 
only on the middorsal line; adults exceed 900 mm. 22 

21b. A pair of intergenials usually present (fig. 46) ; generally no con- 
tact between the prenasal and first supralabial (fig. 45) ; dark tail 
rings often broken laterally; size smaller, adults rarely exceeding 
900 mm. A pattern of red, circular, and ill-defined blotches on a 
pinkish ground color. Crotalus exsul 

Cedros (Cerros) Island off the Pacific Coast of Baja California (fig. 


22 a. General color pink, red, brick-red, or red-brown; usually no light 
areas present within the diamonds; light preocular stripe 1 or 2 
scales wide,* dull and often obscure; supraocular light crossbars 
usually absent. A pattern of reddish, almost unicolor, diamonds 
on a pinkish ground color. Crotalus ruber 

(Fig. 83). 

* Preferably to be determined at the second row of scales above the supralabials if the scales are regular. 

Fig. 46. Chin shields of Crotalus ; 
intergenials present. 

Fig. 47. Chin shields of Crotalus; 
submental present. 


San Diego Society of Natural History 

The Californias from Lat. 34° to Lat. 26° including southeastern Los 
Angeles County, Orange County (excluding the coastal plain) River- 
side County (west of the desert), San Diego County, and extreme 
southwestern Imperial County, California; Baja California, Mexico, 
from the northern border to the vicinity of Comondu, but excluding 
the deserts of the north lying east of the Sierra Juarez and Sierra San 
Felipe; also the adjacent Gulf of California islands of Angel de la 
Guarda, Pond, South San Lorenzo, San Marcos, and Monserrate 
(fig. 67). 

22 b. General color brown, olive-brown, or yellow-brown; light areas 

usually present within the diamonds; light preocular stripe 3 or 
more scales wide, bright and conspicuous ; supraocular light cross- 
bars usually in evidence. Dorsal pattern of brown or olive-brown 
blotches on a buff background. Crotalus lucasensis 

(Fig. 82). 
The southern part of the peninsula of Baja California, Mexico, from 
Lat. 25° 30' to the Cape; the adjacent islands of Santa Margarita and 
San Jose (fig. 67). 

23 a. Two internasals (fig. 48). 29 

23 b. More than two internasals i. e. scales between nasals, and in 

contact with rostral, regardless of size or position* (fig. 49). 

Crotalus viridis 

(For subspecies continue on to 24). 

24 a. Light postocular stripe 1 or \ x k scales wide and clearly outlined 

(fig. 50) ; body blotches commonly subrectangular, with even 
edges and usually with a narrow light border (fig. 52). 


* I have already stated (p. 198) that the internasal criterion will occasionally fail properly to key out 
viridis and its subspecies. It may be noted at this point that if the snake is from California west of 
the Sierra Nevada, from Oregon, Washington, British Columbia, northern Nevada, Idaho, Montana, 
Alberta, Saskatchewan, Wyoming, Utah (except the extreme southwest), Colorado, the Dakotas, western 
Nebraska, or western Kansas one may safely take course 24 even tnough the snake have only 2 internasals. 

Fig. 48. Crown of Crotalus 
showing two internasals. 

Fig. 49. Crown of Crotalus 
showing four internasals. 

Klauber — Key to the Rattlesnakes 


24 b. Light postocular stripe 2 or more scales wide, often indefinite or 

absent (figs. 10 and 51) ; body blotches, if in evidence, commonly 
diamonds, ellipses, or if rectangles, with edges rough or serrated, 
and often without narrow light borders. 26 

25 a. Color usually green or olive-green; less often olive-brown or 

brown; scale-rows 27 or 25 ; dorsal scale rows at the center of the 
tail 13 or more; adult size exceeding 850 mm. A pattern of even- 
edged dark-brown rectangular or subhexagonal blotches usually 
surrounded by a thin light line. Crotalus viridis viridis 

(Fig- 83). 
The Great Plains from Long. 97° to the Rocky Mountains and from 
southern Canada to extreme northern Mexico including the following: 
southwestern Saskatchewan (south of the South Saskatchewan River 
and west of Long. 107°30') ; southeastern Alberta (south of the Red 
Deer River and east of Long. 113°); Montana, except the higher 
mountains in the west; the Lemhi Valley in Idaho; Wyoming east of 
the Rockies; Colorado (except in the higher mountains, and in the 
basins of the Colorado and Green rivers west of the Continental 
Divide) ; extreme southeastern Utah and northeastern Arizona (San 
Juan River basin) ; New Mexico, except the mountains of the west- 
center; extreme northeastern Sonora and northern Chihuahua near the 
U. S. boundary; southwestern North Dakota (west of the Missouri 
River but including the first tier of counties on the eastern bank) ; 
western South Dakota (limiting counties Hand and Jerauld) ; western 
Nebraska; central and western Kansas (limiting counties Riley and 
Geary); Oklahoma west of Woods and Custer counties; Texas west 
of Long. 97° and north of Lat. 30° (fig. 68). The rattler of the 
western Mississippi basin plains. 

Fig. 50. Lateral head pattern of 
C. v. viridis. 

Fig. 51. Lateral head pattern of 
C. v. oreganus. 

Fig. 52. Dorsal pattern of C. v. 

242 San Diego Society of Natural History 

25 b. Color, pink, red, or red-brown;* scale rows 25 or 23; dorsal scales 

at the center of the tail 12 or less; adult size rarely exceeding 
650 mm Crotalus viridis nuntius 

(Fig. 86). 

Northeastern and north-central Arizona from the New Mexican line to 
Cateract Creek including the following: the basin of the Little Colo- 
rado River; the southern part of the Apache Indian Reservation; the 
Hopi Indian Reservation; and the Coconino Plateau, from the south 
brink of the Grand Canyon south to U. S. Highway 66 (rig. 68). 

26 a. Color darker, not straw, cream, or yellow; adult size larger, over 

650 mm. 27 

26 b. Color straw, cream, or yellow; blotches often only faintly in evi- 

dence or obsolete in adults; adult size smaller, usually under 
650 mm Crotalus viridis concolor 

(Fig. 89). 
The basins of the Colorado and Green rivers including: a small area 
in southwestern Wyoming; Utah east of Long. 111° and north of the 
San Juan River (intergradation with viridis viridis is indicated in this 
river valley) ; western Mesa, southwestern Delta, and northern Mont- 
rose counties in western Colorado (fig. 68). 

27 a. Adult color other than vermilion or salmon; body blotches in 

evidence, or body black. 28 

27 b. Adult color vermilion or salmon; body blotches tending toward 

obsolescence in adults. Crotalus viridis abyssus 

(Fig. 87). 
The Grand Canyon of the Colorado River, Grand Canyon National 
Park, Arizona from the north to the south rim (fig. 68). 

28 a. Ground color lighter, usually buff or drab; body blotches occupy 

less or but little more longitudinal space than interspaces; sec- 
ondary series of lateral blotches little in evidence; a pattern of 
dark-brown dorsal blotches (often with light centers) on a buff 
or drab ground color. Crotalus viridis lutosus 

(Fig. 88). 
The Great Basin between the Rocky Mountains and the Sierra Nevada 
including: Idaho south of Lat. 44° ; Utah west of Long. 111° ; Arizona 
north and west of the Colorado River and the north rim of the Grand 
Canyon; all of Nevada except Esmeralda, southern Nye, and Clark 
counties; California, east of the Sierra Nevada, from Lower Klamath 
Lake south to below Mono Lake; Oregon south and east of the line 
Upper Klamath Lake — Fort Rock— Burns— Council (Idaho), this 
being the approximate line of intergradation with C.v. oreganus 
(% 68). 
28 b. Ground color darker, usually dark-gray, olive, brown, or black; 
dark-brown or black dorsal blotches (usually diamonds or hexa- 
gons) occupying considerably more longitudinal space than the 

* Specimens from the plateau south of the Grand Canyon, Arizona, are usually greenish or grayish, 
but should be referred to this subspecies. 

Klauber — Key to the Rattlesnakes 243 

interspaces; a secondary series of lateral blotches conspicuously in 
evidence. Some mountain specimens nearly uniform black, only 
patches of yellow scales representing the interspaces on the mid- 
dorsal line. Crotalus viridis creganus 

(Figs. 90, 109, and 110). 

The Pacific slope from British Columbia to central Lower California 
including the following: the basins of the Fraser and Okanogan Rivers 
in south-central British Columbia, south of Lat. 51° and between 
Long. 119° and 122°, Washington east of the Cascade Mountains; 
the western edge of Idaho from Coeur d'Alene south to Lat. 44° , 
Oregon west of the line Upper Klamath Lake — Fort Rock — Burns — 
Council (this being the approximate line of intergradation with 
lutosus) but absent from northwestern Oregon west of the Cascades 
and from southwestern Oregon immediately contiguous to the 
coast;* all of California west of the Sierra Nevada (including 
these mountains), but excluding a narrow coastal fringe in the ex- 
treme northwest and the entire desert (transmontane) area of the 
southeast ; the west coast and mountains of Baja California from the 
U. S. border south to Punta Maria (Lat. 29°); the mountain areas 
of central and southern Arizona south of the line Peach Springs — 
San Francisco Peak — Springerville but not including the desert areas 
of the southwest; the mountains of extreme west-central New Mexico 
(vicinity of Steeple Rock) and of extreme northern Sonora;t the 
Pacific Coast islands — Mono Rock, Santa Catalina, and Los Coronados 
(fig. 68). 

29 a. No vertical light line on the posterior edge of the prenasals and 
first supralabials. 30 

29 b. A vertical light line on the posterior edge of the prenasals and 
first supralabials (fig. 53). A pattern of black, or dark-brown 
diamonds, with lighter centers, surrounded by single rows of 
yellow scales on a dark-brown background. Crotalus adamanteus 

(Fig. 79). 
The coastal plains of the following southeastern and gulf states: North 
Carolina south of Albemarle Sound; South Carolina; Georgia; all 

* An occasional specimen is found at the river mouths, evidently carried down by floodwater. 

t The Arizona — Sonora range of oreganus does not conjoin the coastal range; there is an unoccupied 
desert gap between. Nevertheless I do not find consistent differences warranting the recognition of the 
Arizona form as a separate subspecies, C. v. cerberus. 

Fig. 53. C. adamanteus, showing 

vertical light marks on prenasal 

and first supralabial. 


San Diego Society of Natural History 

of Florida, with many of the adjacent keys; Alabama; Mississippi; 
and extreme southeastern Louisiana (fig. 67). A lowland species. 

30 a. Supraoculars not pitted, sutured, nor with broken outer edges 
(fig. 14). 31 

30 b. Supraoculars pitted, sutured, or with outer edges broken (fig. 54) . 
A pattern of buff, gray, brown, or deep red-brown blotches on a 
background of straw, tan, buff, brown, or gray; often with gray 
suffusions on the sides of head and body, and with black-tipped 
scales scattered on the dorsum particularly at blotch edges. 

Crotalus mitchellii stephensi 
(Fig. 93). 

The mountain and rocky desert areas of east-central California and 
southwestern Nevada including: the eastern slopes of the Sierra 
Nevada from southern Mono County to southern Kern County and 
eastward in California through the desert mountain ranges to the 
Nevada line; Nevada west and south of the line Hawthorne — Tonopah 
— St. Thomas. Intergradation with Cm. pyrrbus occurs approximately 
along the line Barstow — Ivanpah, which thus constitutes the southern 
limit of stephensi (fig. 69). A rock inhabiting form. 

31a. No distinct and evenly-outlined light supraocular crossbars curving 
forward inwardly; scales on the crown and frontal area smooth 
and flat. 32 

Fig. 54. C. m. stephensi, showing 
sutured supraoculars. 

Fig. 55. Dorsal head pattern, C. 
enyo, showing light marks on 
supraoculars curving forward in- 

Klauber — Key to the Rattlesnakes 


31 b. Distinct and evenly-outlined light supraocular crossbars curving 

forward inwardly (fig. 55) ; scales on the crown and frontal area 
rough, ridged, or knobby; outer edges of the supraoculars raised 
above the crown (particularly evident in life) forming a depres- 
sion in the frontal area; dorsal scales sharply keeled and with 
prominent posterior bosses ; ridged spinous process sharply evident. 
A pattern of dark-brown blotches on a fawn background, usually 
with black in the lateral corners of the blotches at midbody. 

Crotalus enyo 
(Fig. 77). 

The peninsula of Baja California, Mexico, from the Cape north to 
Lat. 30°, together with the adjacent islands of San Francisco, Carmen, 
and Partida, in the Gulf of California, and Santa Margarita on the 
Pacific side (fig. 66). 

32 a. General color dark and with conspicuous blotches 33 

32 b. General color white, cream, or yellow, with grayish tail; a mid- 

dorsal light line faintly in evidence, especially at the neck ; paired 
apical scale-pits rendered conspicuous (particularly near the tail) 
by gray dots. Crotalus unicolor* 

Aruba Island, Dutch West Indies, off the coast of Venezuela (fig. 

33 a. Head larger; head length contained less than 25 times in adult 

body length ; proximal rattle width contained in head length more 
than 2 x /2 times. 34 

33 b. Head notably small for a rattlesnake; head length in adults con- 
tained in body length (over-all) 25 times, or more; proximal 
rattle width contained in head length less than 2Y2 times; pattern 

* Not to be confused with albino specimens of other species (fig. 111). 

Fig. 56. C. molossus, showing 
enlarged scales in anterior part of 
frontal area. 

Fig. 57. C. horridus, showing 

lack of enlarged scales in 

frontal area. 

246 San Diego Society of Natural History 

a series of cross-rings or blotches comprising brown punctations 
on a pink, buff, or gray ground. Crotalus tigris 

(Fig. 94). 
The rocky desert foothills of south-central Arizona, and northeastern 
and central Sonora, from the vicinity of Phoenix, Arizona, via Tucson, 
to Guaymas, Sonora, Mexico, including the following Arizona moun- 
tain ranges: Phoenix, Salt River, Estrella, Santa Catalina, Tucson, 
Coyote, Baboquivari (Verde), Sierrita, and Santa Rita (fig. 69). 
A rock inhabiting form. 

34 a. Usually a definite division between the scales in the frontal and 
the prefrontal areas; scales in the anterior part of frontal area 
larger than those behind (fig. 56) ; anterior body pattern not in 
chevron-shaped bands or not all black. 36 

34 b. No definite division or continuous suture between the scales in 

the frontal and the prefrontal areas ; scales in the anterior part of 
frontal area not conspicuously larger than those behind (fig. 57) ; 
normal pattern a series of chevron-shaped crossbands sometimes 
broken (fig. 58), or with the body all black. Crotalus horridus 

(For subspecies continue on to 33 ). 

35 a. Dorsal scale rows usually 23; postocular dark stripe indistinct; no 

middorsal reddish-brown stripe evident anteriorly; sometimes en- 
tirely black Crotalus horridus horridus 

(Fig. 96). 

The northeastern and north-central United States including: all of 
the Atlantic states from southwestern Maine to southern Virginia, 
(possibly exterminated in Delaware); West Virginia; Ohio; central 
and eastern Kentucky; the mountains of western North Carolina, 
northwestern South Carolina, northern Georgia, central and eastern 
Tennessee, and extreme northern Alabama; Indiana; Illinois (except 
the southern tip); southwestern Wisconsin; extreme southeastern 
Minnesota; eastern and southern Iowa; Missouri (except the southeast 
corner) ; extreme southeastern Nebraska; Kansas (east of Long. 97°) ; 

Fig. 58. Dorsal pattern of C. h 

Klauber — Key to the Rattlesnakes 247 

the mountainous area of northwestern Arkansas; the eastern half of 
Oklahoma; north-central Texas (fig. 68). Originally present in 
southwestern Ontario but now said to be extinct.* 

35 b. Dorsal scale rows usually 25 ; postocular dark stripe distinct and in 

contrast with the ground color; a middorsal reddish-brown or 
brown stripe evident anteriorly. Crotalus horridus atricaudatus 

(Fig. 97). 
The lowlands of the South Atlantic and Gulf states and the lower 
Mississippi Valley including: eastern North Carolina ^South Carolina; 
central and southern Georgia; Florida north of Lat. 29° 30'; central and 
southern Alabama; Mississippi; extreme western Kentucky and Ten- 
nessee; Louisiana; southern and eastern Arkansas; extreme south- 
eastern Missouri and southern Illinois; and Texas east of Long. 99° 
and north of Lat. 28° (fig. 68). On the Atlantic Coast this sub- 
species may range as far north as southern New Jersey.* 

36 a. Dorsal scale rows usually less than 24; scale rows at the center 

of the tail 11 or less; ventrals rarely exceed 168; 1 or 2 scales 
between bottom-center of orbit and supralabials ; usually a single 
loreal, longer than high (fig. 59) ; size small, adults rarely exceed 
600 mm. Crotalus triseriatus 

(For subspecies continue on to 37). 

36 b. Dorsal scale rows usually exceed 24; scale rows at the center of 

the tail 12 or more; ventrals rarely less than 169; 3 or more scales 
between bottom-center of orbit and supralabials ; usually more than 
one loreal, but if single, then higher than long (fig. 60) ; size 
large, adults over 600 mm. 38 

37 a. Dorsal pattern a single series of dark-brown blotches, often edged 

with black and not sharply contrasting with the ground-color of 

* This rattlesnake, once indigenous to what are now the most populous areas of the United States, 
has largely been forced out of industrial and agricultural districts, but still remains in adjacent mountains 
and forests, where rocky or wooded retreats are available. Thus, its present range is intermittent and it is 
extinct in many areas over which it once ranged. 

The boundaries between the subspecies C.h. horridus and C.h. atricaudatus are as yet not definitely 
determined. The division of the species C. horridus into these two subspecies considerably complicates 
the statement of the range so that it is difficult to visualize. For the species C. horridus as a whole the 
range can be more succinctly given thus: All of the states east of the Mississippi River except Michigan; 
also with a limited range in five other states, being confined to southwestern Maine, southern New Hamp- 
shire and Vermont, southern and western Wisconsin, and Florida north of Lat. 29°30'; it is possibly 
extinct in Delaware. West of the Mississippi the range includes: extreme southeastern Minnesota; eastern 
and southern Iowa; probably extreme southeastern Nebraska; Missouri; Kansas, east of Long. 97°; the 
eastern half of Oklahoma; Arkansas; Louisiana; and Texas east of Long. 99° and north of Lat. 28° 
(fig. 68). Of course to all of these areas the previous remarks concerning possible extermination in 
industrial and agricultural territories apply. 

Fig. 59. Lateral head scales of Fig. 60. Lateral head scales of C. 
C. t. triseriatus. basiliscus. 


San Diego Society of Natural History 

brown or dark-gray (fig. 61) ; usually 3 labials in contact with 
pit-border. Crotalus triseriatus triseriatus 

(Fig. 100). 
The central Mexican plateau including: the higher areas in extreme 
southern Durango, Nayarit, southern Zacatecas, San Luis Potosi, south- 
western Tamaulipas, Jalisco, Aguascalientes, Guanajuato, Queretaro,. 
Hidalgo, Michoacan, Mexico (state), Distrito Federal, Morelos, 
Guerrero, Tlaxcala, northern Puebla, and west-central Veracruz (fig. 
71). Intergrades with C.t. pricei in southern Durango. 

37 b. Dorsal pattern of two parallel rows of small brown blotches on a 

steel-gray ground color (fig. 62) ; usually 2 labials in contact with 
the pit-border. Crotalus triseriatus pricei 

(Fig. 101). 
The mountains of southeastern Arizona and northwestern Mexico in- 
cluding: the Pinalino (Graham), Santa Catalina, Santa Rita, Huachuca, 
and Chiricahua mountains in Arizona; and the Sierra Tarahumare and 
Sierra Madre in eastern Sonora, western Chihuahua, and western Dur- 
ango, intergrading with C.t. triseriatus in the southern part of the 
last named state (fig. 71). 

38 a. Usually a single loreal; tail rings sharply contrasting in color. 

Crotalus scutulatus 
(See under 18b).* 

38 b. Usually two or more loreals; tail unicolor or with rings rather 

faint — not sharply contrasting. 39 

39 a. Tail often black, or with rings faintly in evidence against a dark 

background ; vertebral process not conspicuous ; tail shorter, ap- 
proximately 7.1 per cent of body length (over-all) in the males 
and 5.8 per cent in females; subcaudals rarely more than 27 in 
males, or 23 in females; initial rattle-button (if present), over 
5 mm. wide; body color primarily olive-green, or yellow-green 
with dark-brown blotches, often with a light interior blotch on 
each side of the center; blotch-bordering scales unicolor. 

Crotalus molossus 
(For subspecies continue on to 40). 

* This is the only species which is double-keyed, for scutulatus may or may net have a 
like tail, and hence may take either course 17a or 17b. 

Fig. 61. Dorsal pattern of C. t. Fig. 62. Dorsal pattern of C. t. 
triseriatus. pricei. 

Klauber — Key to the Rattlesnakes 


39 b. Tail not black but with gray rings on a darker gray background; 

vertebral process prominent; tail longer, approximately 9.2 per 
cent of body length in the males and 7.4 per cent in the females; 
subcaudals in males rarely less than 28, or 23 in females; initial 
rattle-button (if present) less than 5 mm. wide; body pattern a 
series of red or red-brown diamonds (outlined with buff) on a 
pinkish background. Crotalus basiliscus 

(Fig. 76). 
The west coast of Mexico between Lat. 17° and 25° including southern 
Sinaloa, Jalisco, Colima, and central Oaxaca certainly, and probably 
Nayarit, Michoacan, and Guerrero; restricted to the coast and ad- 
jacent higher areas (fig. 66). 

40 a. Dark dorsal blotches (on the anterior half of the body) open on 

the sides and extending to the ventrals (fig. 63). A pattern of 
dark-brown blotches (often with a light interior blotch on each 
side of the center) on an olive-green or yellow-green ground 
color; blotch bordering scales unicolor. 

Crotalus molossus molossus 
(Fig. 78). 
From west Texas to Arizona and south in Mexico to northern Durango, 
including: the limestone area north and west of San Antonio, and 
trans-Pecos Texas; New Mexico southwest of the line Gallup — Otto — 
Carlsbad ; Arizona, from the Grand Canyon and Little Colorado River 
south, but not including Mohave and Yuma counties; central and 
eastern Sonora, and western Chihuahua, intergrading with Cm. 
nigrescens in northern Durango ; also San Esteban Island in the Gulf 
of California (fig. 70). 

40 b. Dark dorsal blotches (on the anterior half of the body) closed 
laterally by light borders (fig. 64) . Body color primarily olive- 
brown, or brownish-black, with dark-brown diamonds bounded 
by grayish or yellowish unicolored rows of light scales. 

Crotalus molossus nigrescens 
The tableland of Mexico from northern Durango (where it inter- 
grades with Cm. molossus) south and east through Durango, southern 
Coahuila, Zacatecas, western San Luis Potosi, Aguascalientes, eastern 
Jalisco, Guanajuato, northern Michoacan, Mexico (state), and Dis- 
trito Federal, to central Veracruz and southeastern Puebla (fig. 70). 

Fig. 63. Lateral pattern of C. 


Fig. 64. Lateral pattern of C. m. 


San Difgo Society of Natural History 

Fig. 65. Ranges of Crotalus durissus terrificus (in part) and Crotalus unicolof. 

(The range of the South American rattlesnake is not known with accuracy, and this 

map is to be considered only a generalized indication of the area covered. Its presence 

in the central basin of the Amazon is doubtful). 

Note. The grouping of the forms in the several maps is not necessarily an indication of rela- 
tionship, the arrangement being primarily selected to reduce the required number of maps. How- 
ever, subspecies of a single species always appear on the same map so that the approximate lines of 
intergradation can be seen, such lines being indicated by stars. 

Klauber — Key to the Rattlesnakes 


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San Diego Society of Natural History 


Fig. 69. Ranges of Crotalns mitcbellii mitchellii. C. m. pyrrhus. C. m. stephensi. 
Crotalns tigris. Crotalns lepidns lepidns. C. I. klauberi, and Crotalns polystictus. 

(Stars indicate approximate lines of intergradation between subspecies). 

Klaurer — Key to the Rattlesnakes 



San Diego Society of Natural History 

^ M- 

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Ki.awber — Key to the Rattlesnakes 


Fig. 74. Crotalus durissus durissus. 

Central American Rattlesnake (12b).* 

(Specimen from Vera Cruz, Mexico. West-coast specimens have less color in the lateral 

areas between dorsal blotches). 

Fig. 75. Crotalus durissus terrijicus. 

South American Rattlesnake (12a). 

(Specimen from Central Brazil. Photo by courtesy of the New York Zoological Society). 

* This number refers to the item (not page I number in the text of the key. under which the key 
characters, color description, and range will be found. 


San Dif.go Society of Natural History 

Fig. 76. Crotalus basiliscus. 

Mexican West-Coast Rattlesnake (39b) 

(Specimen from Retes, Sinaloa, Mexico). 

Fig. 77. Crotalus enyo. 

Lower California Rattlesnake (31b). 

(Specimen from La Rivera, Baja California, Mexico). 

Klauber — Key to the Rattlesnakes 


Fig. 78. Crotalus molossus molossus. 

Northern Black-tailed Rattlesnake (40a). 

(Specimen from Entro, Yavapai County, Arizona). 

Fig. 79. Crotalus adamant eus. 
Eastern Diamond Rattlesnake (29b). 
(Specimen from Eureka, Marion County, Florida. The blur at the tail is the vibrating rattle). 


San Diego Society of Natural History 

Fig. 80. Crotahts cinereous. 

Western Diamond Rattlesnake (20b). 

(Specimen from Date, Yavapai County, Arizona). 

Fig. 81. Crotalus tortugensis. 
Tortuga Island Diamond Rattlesnake (20a). 
(Specimen from Tortuga Island, Gulf of California). 

Klauber — Key to the Rattlesnakes 


Fig. 82. Crotalus lucasensts. 

San Lucan Diamond Rattlesnake (22b). 

(Specimen from La Rivera, Baja California, Mexico). 

Fig. 83. Crotalus ruber. 

Red Diamond Rattlesnake (22a). 

(Specimen from Santa Margarita, San Diego County, California). 


San Diego Society of Natural History 

. ■< -Or* - 

Fig. 84. Crotalus scutulatus. 

Mohave Rattlesnake (18b). 

(Specimen from Date, Yavapai County, Arizona). 

Fig. 85. Crotalus viridis viridis. 

Prairie Rattlesnake (25a). 

(Specimen from near Jetmore, Hodgeman County, Kansas). 

Klauber — Key to the Rattlesnakes 


■* I MM 






Fig. 86. Cvotalus viridis nuntius. 

Arizona Prairie Rattlesnake (25b). 

(Specimen from Canyon Padre, Coconino County, Arizona). 


Fig. 87. Crotalns viridts abyssus. 

Grand Canyon Rattlesnake (27b). 

(Specimen from Tanner Trail in Grand Canyon, Coconino County, Arizona). 


San Diego Society of Natural History 

Fig. 88. Crotalns v'tridis lutosus. 

Great Basin Rattlesnake (28a). 

(Specimen from near Delta, Millard County, Utah). 

,-- . . . 


Fig. 89. Cro talus viridis concolor. 

Midget Faded Rattlesnake (26b). 

(Specimen from Jensen, Uintah County, Utah). 

Klauber — Key to the Rattlesnakes 


Fig. 90. Crotalus liridis oregamis. 

Pacific Rattlesnake (28b). 

(Specimen from near Wenatchee, Chelan County, Washington). 

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Fig. 91. Crotalus mitckellii mitchellii. 

San Lucan Speckled Rattlesnake ( 1 6a) . 

(Specimen from La Rivera, Baja California, Mexico). 


San Diego Society of Natural History 

Fig. 92. C total us mite hell ii pyrrhus. 

Southwestern Speckled Rattlesnake (16b). 

(Specimen from Yaqui Well, San Diego County, California). 

Fig. 93. C to talus mitchellii Stephens}. 
Panamint Rattlesnake (30b). 
(Specimen from near Aberdeen, Inyo County, California). 

Klauber — Key to the Rattlesnakes 


Fig. 94. Cvotalus tigris. 

Tiger Rattlesnake (33b). 

(Specimen from Squaw Peak, Maricopa County, Arizona). 

Fig. 95. Cvotalus cerastes. 

Horned Rattlesnake or Sidewinder (8b). 

(Specimen from Borego Valley, San Diego County, California). 


San Dif.go Society of Natural History 

Fig. 96. Crotalus horridus horridus. 

Timber Rattlesnake (35a). 

(Specimen from near Baraboo, Sauk County, Wisconsin). 

Fig. 97. Crotalus horridus atricaudatus. 

Canebrake Rattlesnake (35b). 

(Specimen from Imboden, Lawrence County, Arkansas). 

Klauber — Key to the Rattlesnakes 


Fig. 98. Crotalus lepidus lepidus. 

Eastern Rock Rattlesnake (14a). 

(Specimen from Mt. Locke, Jeff Davis County, Texas), 

Fig. 99. Crotalus lepidus klauberi. 

Green Rock Rattlesnake (14b). 

(Specimen from Pinos Altos Mts., Grant County, New Mexico). 


San Diego Society of Natural History 

Fig. 100. Crotalus triseriatus triseriaius. 

Mexican Spotted Rattlesnake (37a). 

(Specimen from Jacala, Hidalgo, Mexico). 

Fig. 101. Crotalus triseriatus pricei. 

Arizona Spotted Rattlesnake (37b). 

(Specimen from Ramsey Canyon, Huachuca Mts., Cochise County. Arizona). 

Klauber — Kfv to thi: Rattli-snakhs 


Fig. 102. Crotalus willardi. 

Ridge-nosed Rattlesnake (9b). 

(Specimen from Ramsey Canyon, Huachuca Mrs., Cochise County, Arizona). 

Fig. 103. Sistrurus miliarias miliarius. 

Carolina Ground Rattlesnake (6b). 

(Specimen from Leesville, South Carolina. Photo by courtesy of Howard K. Gloyd). 


San Dif.go Society of Natural History 

Fig. 104. Sistrurus miliar ins barbouri. 

Southeastern Ground Rattlesnake (5b) 

(Specimen from Marion County, Florida). 

Fig. 105. Sistrurus miliarias streckeri. 

Western Ground Rattlesnake (6a). 

(Specimen from Gentilly, Orleans Parish, Louisiana). 

Klaubi-r — Key to the Rattlksnaki-s 


Fig. 106. Sist runts catenates catenatus. 

Eastern Massasauga (4a) . 

(Specimen from Proud Lake, Oakland County, Michigan. The flattened posture is one 

often assumed by an angered rattler). 

Fig. 107. Sistrurus catenatus tergeminus. 

Western Massasauga (4b) . 

(Specimen from China Spring, McLennan County, Texas). 


San Diego Society of Natural History 




Fig. 108. A rattlesnake in a striking coil or typical defensive posture. 
(Crotalus cinereous, Western Diamond Rattlesnake, Riverside County, California). 

Fig. 109. A rattlesnake in its resting coil. 
{Crotalus viridis oreganus, Pacific Rattlesnake, San Diego County, California). 

Fig. 110. A rattlesnake crawling; note how the rattle is elevated to prevent dragging. 
(Crotalus viridis oreganus, Pacific Rattlesnake, San Diego County, California). 

Ki auber — Key to the Rattlesnakes 


Fig. 111. A juvenile albino rattlesnake, compared with one of normal coloration. 
{Crotalus viridh viridis, Prairie Rattlesnake, Weld County, Colorado). 

Fig. 112. The biting mechanism of the rattlesnake. 
These are poses of a freshly killed specimen of Crotalus cinereous, with mouth 
opened to show the fangs. In the left-hand figure the fangs are folded against the 
roof of the mouth and are covered by their sheaths. In the central figure the 
fangs are slightly advanced and the sheaths have been cut away. The right fang 
has been outlined in white to bring it out. In the right-hand figure the mouth has 
been widely opened and the fangs advanced, as at the end of the forward drive of 

a strike. 


San Diego Society of Natural History 



and Range 




Key No. Page 






27b 242 
29b 243 






Atricaudatus 1 





































31b 245 
21b 239 






35a 246 
14b 234 







14a 234 
22b 240 

























40b 249 
25b 242 






28b 242 
10b 232 
















2b 227 
22a 239 

























4b 229 

12a 233 
































The Scientific Publications of the 

San Diego Society of Natural History 

comprise the following series: 

Transactions — descriptive and taxonomic contributions. 

Memoirs — lengthy monographs. 

Occasional Papers — reports primarily of interest to specialists. 

A complete list of publications will be furnished on application to 


Natural History Museum 

Balboa Park, 

San Diego, California 

& ri*$ 



Volume VIII, No. 21, pp. 277-304, plate 21 



Steve A. Glassell 

Research Associate in Crustacea, San Diego Society of Natural History 


Printed for the Society 

December 7, 1936 



U. S. Grant, IV, Chairman 

Fred Baker Clinton G. Abbott, Editor 



Steve A. Glassell 

Research Associate in Crustacea, San Diego Society of Natural History 


The littoral fauna of the west coast of Mexico, during the last 
few years, has produced many interesting additions to the known list 
of marine invertebrates. With the descriptions of these new Pacific 
species the foundations are being laid for a more intelligent study of the 
analogies which exist between faunas of the tropical Pacific and Carib- 
bean seas. This study, revealing as it does relationships, still close, be- 
tween many of our new species and those already known from Carib- 
bean and Atlantic waters, clearly demonstrates the comparatively recent 
intrusion of a land barrier separating the Atlantic and Pacific oceans. 

This paper, then, is an attempt to bring our knowledge of a small 
portion of this western fauna up to date by describing a number of 
newly found forms of those crab-like Anomurans, the Porcellanids. 
More than fifty years have passed since Lockington described a num- 
ber of this group from the Gulf of California. Since that time very 
little attention has been paid to the Porcellanids in this area. It is 
hoped that, at this time, the addition of thirteen new species from those 
waters may make amends for this neglect. 

In addition to these Porcellanids, three new species of Pinno- 
therids are also described: one from Panama, the other two from the 
Gulf of California. 

I wish to express my thanks to Dr. Mary J. Rathbun and Dr. 
Waldo L. Schmitt of the U. S. National Museum, and to Dr. Isabella 
Gordon of the British Museum (Natural History), for their encour- 
agement, valuable suggestions and assistance. I am indebted to Mr. 
Anker Petersen of Beverly Hills, California, for the drawings of the 
text figures. 

280 San Diego Society of Natural History 

Petrolisthes schmitti, new species 

Type. — Female, holotype, Cat. No. 71534, and male, paratype, U. S. Na- 
tional Museum: from San Felipe, Baja California, Mexico, low tide; June 8, 
1933; collected by Steve A. Glassell. Two paratypes, female and male, Cat. Nos. 
765 and 766, S. D. S. N. H.; from same series. 

Diagnosis. — Carapace quadrilateral. Chelipeds short; carpus short, unarmed. 
First antennal peduncle lobed. A small species. 

Description. — Carapace as long as wide, sides subparallel, margined with a 
distinct ridge; epibranchial regions roughened with rugae, remaining regions 
smooth, but having a few scattered tufts of tomentum. Front depressed, a single 
undivided process, not trilobate; line of front sinuous over antennules, slightly 
tomentose. Eyes large. Upper ocular margin smooth. First antennal peduncle 
with a horizontally depressed, distal lobe; second peduncle lightly granulate. 

Chelipeds short, stout; merus short, a small lobe at distal inner angle, not 
protruding as far as outer edge of carpus; carpus 2/3 as long as wide, unarmed, 
with two shallow, longitudinal sulci dividing the surface into three granulated 
ridges; hands short, thick, subequal, similar, a longitudinal median swelling from 
proximal end to gape, bordered by indistinct sulci; outer part of hand and fingers 
granulate, with a light film of tomentum near margins; palms of hands smooth, 
with tufts of sponge-like hair in the gape of the fingers; pollices blunt at tips, up- 
turned; dactyli long, sinuous, granulate, with prehensile edges closely meshing 

Ambulatory legs longer than body length, decorated with irregular tufts of 
tomentum, sparingly setose. 

Abdomen and epimera fringed with tomentum. 

Color in alcohol. — A red-brown mottled with white. 

Measurements. — Female holotype: length of carapace 4.7 mm., width 4.7 
mm. Male paratype: length 4.4 mm., width 4.1 mm. 

Range. — Upper part of the Gulf of California. 

Material examined. — San Felipe, Baja California, June 8, 1933; 4 males and 
7 females; collected by the author. The types for this species were selected from 
this series. 

Habitat. — Taken at low water from under rocks. 

Remarks. — This proposed species is allied to P. gracilis, Stimpson, 1859, in 
which the shape of the front is similar, but it differs from that species in that 
P. schmitti has short heavy chelipeds, with longitudinal sulci and granulated 
ridges on the carpus, while in P. gracilis the chelipeds are long, and the carpus is 
long, smooth and narrow. P. schmitti has tufts and fringes of tomentum, while 
P. gracilis is nude. 

Glassell — New Porcellanids and Pinnotherids 281 

This species, being of such small size, might be considered to be in the 
adolescent stage, if it were not for the fact that most of the females were gravid, 
and that intensive collecting failed to produce larger specimens. 

This species is dedicated to Dr. Waldo L. Schmitt, of the U. S. National 
Museum, with feelings of warm personal regard and admiration. 

Petrolisthes sanfelipensis, new species 

Type. — Male, holotype, Cat. No. 71535, and female, paratype, U. S. 
National Museum: from San Felipe, Baja California, Mexico, low tide; June 
8, 1933; collected by Steve A. Glassell. Two paratypes, male and female, Cat. 
Nos. 767 and 768, S. D. S. N. H.; from same series. 

Diagnosis. — Carapace striated, bearing spines on the front, ocular margin, 
and on the hepatic, branchial and protogastric regions. Nearly the entire dorsal 
and ventral surfaces covered with either striations or squamae. 

Description. — Carapace nearly as long as wide, regions well marked, 
transversely plicated with micro-piliferous striae, interrupted by the cervical 
groove and cardiac margins; a row of spines completely across the carapace be- 
hind the eyes, interrupted by the median sinus and cervical groove; a group of 
spines on the epibranchial region and a spine on the margin posterior to the 
hepatic region. The front is triangular, depressed, and margined with thick-set 
blunt teeth, separated from the ocular margin by a sinus. The supraocular 
margin is armed with sharp, forward-pointing teeth, increasing in length anteri- 
orly; a single sharp, postorbital tooth. Other teeth may be present on the forward 
part of the carapace in some specimens, but, for the most part, those described 
here are not subject to change in location. The eyes are large, the distal end of 
the stalk being fringed with red hair. The distal end of the first antennal 
peduncle bears a horizontally compressed plate, armed with teeth on its outer 
margin; the second peduncle is roughened on its outer surface with blunt tuber- 
cles; the third is plain. 

The chelipeds are long and narrow; merus armed at its distal inner angle 
with a long rugose lobe surmounted with a sharp spine, its distal margins armed 
with spines, its surface transversely crossed with interrupted striae; the carpus is 
nearly twice as long as wide, armed on its inner margin with four or five major 
multiformed teeth; these teeth are covered on their long proximal edges with 
smaller denticles, the distal margins being short and tuberculate; the carpus 
may be divided into three longitudinal parts; the outer part is armed with a series 
of 7 or 8 forward- and upward-pointing spines, which are inside the margin and 
are connected with transverse striae which come up from the under side of the 
carpus; these spines border an interrupted sulcus; the median portion of the 
carpus is raised and formed of transverse laminae broken up, and this is sepa- 
rated from the inner side by another interrupted sulcus; the remaining portion 
is covered with piliferous, sharp-pointed, granulated rugae; the distal end has 
several teeth; the manus is long and slender, and oblique plications from the 
inner palm come over the crest to an inner-marginal sinus; the median ridge is 

282 San Diego Society of Natural History 

formed of longitudinally oblique, granulate-edged plications which connect with 
the marginal sinus; from this ridge to the outer margin the surface is covered with 
lamellar rugae, the interspaces filled with a short tomentum; the outer border 
is tomentose proximally and armed with a series of sharp, short spines; the pollex 
is long, thin and upturned at the tip; the dactyli fit closely to the pollices, and 
they are armed on their upper inner crest with a row of upward- and forward- 
pointing, sharp spines; their tips are compressed and multiunguiculate; on the 
inner side the fingers are covered with a sponge-like tomentum, short cropped; 
the inner side of the hand and fingers is lined with oblique, granulated stria- 
tions. The under side of the carpus, merus and propodus is transversely striate. 

The ambulatory legs are rather long, the merus in all three being trans- 
versely striate, and the upper crests armed with three or four sharp, outward- 
pointed teeth; the distal posterior end of the merus of the first and second legs 
has spines, as has also the upper distal end of the carpus on these legs; the 
propodus is nearly IV2 times as long as the dactylus; the dactyls are curved, with 
a corneous tip, armed on their under sides with four corneous spines. 

The abdomen is transversely striated on all segments, bordered with setae. 

The first three segments of the outer maxillipeds are striated. 

Color in life. — A faint pinkish tint, but when freshly preserved in alcohol 
is beautifully colored with rich reds and purple. 

Measurements. — Male holotype: length of carapace 8.7 mm., width 9 mm. 
Female paratype: length 7.6 mm., width 7.9 mm. 

Range. — Upper end of the Gulf of California. 

Material examined. — A series of 10 males and 10 females, collected at San 
Felipe, Baja California, June 8, 1933, by the author; the types were selected 
from this series. A series of 10 males and 10 females, collected at Punta Penasco 
(Rocky Point) , Sonora, May 2, 1935, by the author. 

Habitat. — Taken at mean low tide level and below, from among sponges 
and gorgonians. 

Remarks. — This proposed species is allied to P. hirtispinosus Lockington, 
1878, but differs from that species in the greater number of spines on the cara- 
pace and front, the more distinctive markings of the carapace, the less equal 
distribution of spines on the carpus of the chelipeds, and in the hand not being 
covered with a short pile of tomentum. Also, the ventral part of P. hirtispinosus 
is not striated. 

Petrolisthes nigrunguiculatus, new species 

Type. — Male, holotype, Cat. No. 71536, and female, paratype, U. S. 
National Museum: from Santa Catalina Island, Gulf of California, Mexico, 
low tide; December 14, 1931; collected by Steve A. Glassell. Two paratypes, 
male and female, Cat. Nos. 769 and 770, S. D. S. N. H.; from same series. 

Glassell — New Porcellanids and Pinnotherids 283 

Diagnosis. — Carapace flattened, smooth in central regions, squamoso-granu- 
late on front and anterior margins, front deflexed. Chelipeds of male dissimilar, 

Description. — Carapace about equally wide as long, depressed, central 
regions smooth, anterior margins and front squamoso-granulate and microscopi- 
cally covered with tomentum; a transverse hepatic lobe; a well-defined protogas- 
tric ridge; front deflexed, triangular, with a wide ridge-bordered median sulcus; 
the upper ocular margin paralleled by a groove which terminates in a notch, this 
notch separates the front from the preorbital lobes. First antennal peduncle with 
a prominent lobe distally; second peduncle serrated with granules on outer 
surface; third smooth; flagellum smooth. 

Chelipeds of males, dissimilar; of females, similar in some, in others as in 
the males; merus with inferior distal end extending nearly 1/3 the distance to the 
hooked tip of the carpus; the inner distal end has a long subvertical, blunt, granu- 
lated lobe; carpus short, about twice as long as wide, covered with squamoso- 
granules along the median ridge, with irregular tuberculate granules on the re- 
maining surfaces, the inner margin armed with three or more clustered, blunt- 
ended teeth, the distal posterior end a long, forward-curving tooth; hands covered 
with various sized squamous granules and tubercles, the inner margin rounded, 
the outer sharp and fringed with an even length of fine pinnate hair, growing 
out of flat, lamellar serrations to a point near the blunt tip of the pollices; also 
extending over the fringe is a series of short, subequally spaced, transverse ridges, 
ending as teeth; the major hand of the male has a median swollen ridge to the 
gape; the pollex is thick, stubby; the dactylus stubby, distorted, extending past the 
pollex and bluntly curved at the tip; these fingers gape widely and cannot be 
brought together, due to a large, distorted lobe on the proximal cutting edge 
of the dactylus; the inner surface of the dactylus has a heavy growth of pinnate 
hair, which is absent or present in less quantity on the minor chela; in the females 
with similar hands this growth may be entirely absent. The minor chela of the 
males is similar to both chelas in the majority of females, in which the tip of the 
pollex is truncate, thin and wide, the dactyl long, curving, and engaging the 
pollex for its entire length; the surface is as that of the major chela. 

The ambulatory legs are short; merus compressed, wide, crested with minute 
granules distally; carpus and propodus longitudinally ridged with rows of small 
teeth, lightly fringed with setae; dactyli with curved corneous tips; a distinctive 
feature of these dactyli is their color — a deep, dark brown, almost black. 

Color in life. — Thickly mottled with dark brown and reds. Tips of chela 
a light red. Tomentum and fringes a dirty white. 

Measurements. — Male holotype: length af carapace 7.2 mm., width 7 mm. 
Female paratype: length 7.5 mm., width 7.3 mm. 

Range. — Gulf of California. 

Material examined. — Approximately 20 males and 20 females, Santa Cata- 
lina Island, Gulf of California, December 14, 1931, collected by the author. The 
type material was selected from this series. 

284 San Diego Society of Natural History 

Habitat. — Under side of rocks at low and half tide levels. 

Remarks. — This proposed species is closely allied to P. hirtipes Lockington, 
1878, which is found in the same region, but the latter may be distinguished from 
P. nigrunguicidatus by some of the following characters: P. nigrunguiculatus is 
smooth in the central regions of the carapace, instead of rugose; the carpus of the 
chelipeds is armed with several lobular spines, instead of having but one lobular 
group at the proximal end; the front is unbordered with setae, instead of being 
fringed with fine thick hair; only the outer margins of the hands are fringed, 
instead of the entire margins of the chelipeds and ambulatory legs. P. hirtipes is 
also the larger species of the two, a male collected at Magdalena Bay, Baja 
California, December 4, 1931, measured: length of carapace 10 mm., width 
10 mm. 

Petrolisthes tiburonensis, new species 

Type. — Female, holotype, Cat. No. 71537, and male, paratype, U. S. 
National Museum: from south end of Tiburon Island, Gulf of California, 
Mexico, low tide; December 31, 1931; collected by Steve A. Glassell. Two para- 
types, female and male, Cat. Nos. 771 and 772, S. D. S. N. H.; from same 

Diagnosis. — Carapace convex anteriorly, very rugose; front trinoduled. 
Antennal peduncles nodulous. Merus of ambulatory legs carinate. Dimorphic. 

Description of female. — Carapace a little wider than long, convex anteriorly, 
with regions well denned, anteriorly covered with lobate granules, posteriorly 
with well defined, disconnected striae; borders rounded over, high, except for 
posterior margin which is bimarginal and concave. The front is trinodulous, 
the nodules blunt and upturned, the median the largest and more extended. 
Eyes large. First antennal peduncle with a large single or bilobed process extending 
past the joint of the second peduncle; the latter also has two or three prominent, 
blunt lobes posteriorly, one distal, also many small granules; the third peduncle 
has a single distal lobe. 

The chelipeds are similar; merus with a single upward-pointed lobe at the 
distal anterior margin, the surface is rugose; the carpus has subparallel margins, 
is rugose, with a median ridge, armed with a forward-pointed, hook-like tooth 
at the posterior distal end; the anterior border is armed with a series of unequal, 
small teeth, more numerous at the proximal end; hands similar, rounded on the 
inner margin, compressed on the outer, which is armed with a series of sharp, 
forward-pointing teeth, their anterior base setose; viewed from underneath the 
teeth are unequally spaced and of irregular length; on the outer surface the hands 
are granulate, with a wide rounded median ridge to the gape of the fingers, 
thence continues around the cutting edge of the pollex, and terminates at the 
base of the upturned tip; the prehensile fingers are distorted, with a striated upper 
carina, a median striated or granulated ridge, channeled on each side; the palms 
of the hands are smooth, the inner gape with a bunch of fine, long, pinnate hair. 

Glassell — New Porcellanids and Pinnotherids 285 

The ambulatory legs are short; merus stout, armed on the upper surface 
with irregular, blunt, conical lobes, a median ridge with sulci on each side; the 
carpus of the first leg is bilobed on its upper surface, a lobe at each end, a large 
double-ended lobe on the distal inner side; carpi of second and third legs bilobed, 
eroded; propodi with longitudinal ridges; dactyli with curved corneous tips, 
armed underneath with short, supplemental, corneous spines; legs sparsely mar- 
gined with small tufts of setae, and granulate. 

Description of male. — Differs from the female in being much larger, and 
having the carapace, front, antennal peduncles and chelipeds much smoother. 
The margins of the carpus of the chelipeds are not parallel, being widest at the 
distal end, unarmed on the inner margin, or at best vaguely indicated. The 
hands are unarmed on the outer margin with either teeth or setae, being smooth 
and rounded. The ambulatory legs differ from those of the female in that the 
merus of the first leg has a smooth crest; the other legs differ only in that they 
are less heavily margined with lobes. 

Color in life. — A chocolate brown. 

Measurements. — Female holotype: length of carapace 9.2 mm., width 10 
mm.; length of carpus 7.3 mm., width 2.6 mm.; length of hand 12 mm.; width 
at base of finger 5.8 mm. Male paratype: length of carapace 10.5 mm., width 
10.8 mm.; length of carpus 12 mm., width 3.8 mm.; length of hand 21 mm.; 
width at base of finger 8.1 mm. 

Range. — -It seems peculiar that this species has so far been taken in the 
Gulf of California only between latitudes 28° 30' N., and 30° N., although 
extensive collecting has been done on each side of these boundaries. 

Material examined. — South end of Tiburon Island, Gulf of California, 
December 31, 1931: approximately 20 males and 20 females, collected by the 
author. The types have been selected from this series. Also from Angeles Bay, 
Baja California, January 4, 1932: 1 male and 3 females. Puerto Refugio, Angel 
de la Guardia Island, Gulf of California, January 6, 1932 : 2 males. 

Habitat. — Collected under rocks at low tide. Plentiful. 

Remarks. — This proposed species is allied to P. crenulatus Lockington, 
1878, which it somewhat resembles in the shape of the carapace and ratio of 
measurements (both species being wider than long) . They differ in nearly every 
other respect, P. tiburonensis being brown instead of red and white, slightly setose 
instead of quite tomentose, etc. The chelipeds of the adult males somewhat 
resemble those of P. gracilis Stimpson, 1859, but here the resemblance ceases. 

Stimpson in his description of the genus Petrolisthes 1 states: "Carapax 
depressed subovate, not broader than long." Strictly followed, neither P. 
crenulatus nor P. tiburonensis would be placed in the genus Petrolisthes, although 
they both agree in other particulars, for both of these species are subquadrate, 

1 Rept. on Crust. Coll. by N. Pac. Expl. Exped. 1853-56. Smithsonian Misc. Coll. 
Part vol. XLIX, 1907, p. 181. 

286 San Diego Society of Natural History 

instead of subovate, and are broader than long. A normal, large specimen of 
P. crenulatus, which I collected at the type locality, has the following measure- 
ments: length of carapace 13.5 mm.; width 15 mm. 

Without attempting to amend the genus or create a subgenus for the recep- 
tion of this and other species, I have placed it in the genus Petrolisthes, awaiting a 
much-needed, thorough revision of the Porcellanids. 

I selected the female for the holotype of this species, as most of the juvenile 
males bear a closer resemblance to the female form than they do to the mature 

Pisosoma smithi, new species 

Type. — Female, holotype, Cat. No. 71538, and female, paratype, U. S. 
National Museum: from Miramar Beach, near Guaymas, Sonora, Mexico, low 
tide; December 23, 1931; collected by Steve A. Glassell. Female paratype, Cat. 
No. 773, S. D. S. N. H.; from same series. 

Diagnosis. — Carapace suboval, lightly marked with transverse plications. 
Carpus of chelipeds nearly flat on upper surface, unarmed and rugose; hands with 
three ridges between the margins. 

Description. — Carapace suboval, convex toward the front, nearly as long as 
wide, regions denned, cervical groove fairly deep near margins, the posterior 2/3 
crossed with light transverse plications and punctae, more prominent near the 
sides and posteriorly. The front sinuous and entire in dorsal view, more promi- 
nent in front of median depression. Eyes small. 

Chelipeds short, stout; merus short on dorsal side, extending further ven- 
trally, a granulated raised carina on both upper margins, the distal inner end 
not protruding as far as the inner margin of the carpus; carpus rather flat on top, 
with both inner and outer margins projecting outward from beneath; margins 
unarmed, but serrated with granules, a granulated median ridge, separated from 
the outer margin by a granulated sinus, distally crossed by granulated, trans- 
verse striations, distal end granulate, ventral side striate; hands subequal, granu- 
late, with three longitudinal, granulated ridges, the upper short, the lower longer, 
but not extending onto the pollex, which is slightly upturned at the tip; dactyli 
long, sinuous, granulate, with a median ridge, tip hooked; the fingers gape 
evenly from base to tip; under side of hand punctate, granulate in gape. 

Ambulatory legs granulate on upper crest; dactyli slightly curved, tip corne- 
ous, supplemental spines on lower margin. 

Color in alcohol. — Carapace cream color. Ambulatory legs a light pink. 
Abdomen and sternum iridescent. 

Measurements. — Female holotype: length of carapace 6.4 mm., width 6.8 
mm. Male paratype: width 4.7 mm., length 4.7 mm. 

Range. — Length of the Gulf of California. 

Glassell — New Porcellanids and Pinnotherids 287 

Material examined. — Miramar Beach, near Guaymas, Sonora, December 
23, 1931: 3 females, 1 male, collected by the author. The types were selected 
from this small series. Also one specimen was collected at each of the following 
localities: Espiritu Santo Island, Gulf of California, December 6, 1931; Carmen 
Island, Gulf of California, January 21, 1932; Seargent's Point, Sonora, January 
2, 1932; Punta Pefiasco (Rocky Point) , Sonora, May 3, 1935. 

Habitat. — This appears to be a shore form occupying the lower intertidal 
zone with P. sinuimanus Lockington, but is not plentiful at any locality. 

Remarks. — This proposed species is allied to P. sinumianus Lockington, 
1878, but differs from that species by the shape and form of the carpus of the 
chelipeds, and by the carpus not being armed on its inner margin with a proximal 
lobe. It differs also in that the upper surface of the carpus is flattened instead 
of rounded as in P. sinuimanus. 

That Lockington was sensible to a difference between this species and 
P. sinuimanus is evident, but he attributed this difference to a variation in his 
species. He was no doubt influenced in his judgment by a lack of material. To 
quote from his notes (the italics are mine) : "This species varies considerably: 
some jew specimens are without a trace of the lobe upon the meros or of the tooth 
upon the anterior margin of the carpus; in others they are small, in others large 
and prominent. One specimen combines with the want of these teeth a carapax 
the surface of which is plicate upon the margins. The rolling ridges of the manus 
and carpus, and the deeply punctate surface of both, are constant characters." 

This species is named in honor of Sidney I. Smith, whose work in carcinology 
on the west American coast added so much to our knowledge of this fauna. 

Pisosoma lewisi, new species 

Type. — Female, holotype, Cat. No. 774, San Diego Society of Natural 
History; male, paratype, Cat. No. 775, S. D. S. N. H: from Tenacatita Bay, 
Jalisco, Mexico, low tide; December, 1932; collected by Captain Fred O. Lewis, 
of the yacht "Stranger." 

Diagnosis. — Carapace subquadrate, as wide as long, punctate. Chelipeds 
heavy; carpus short, wide, deeply bisculate; hands heavy, trisulcate. 

Description. — Carapace depressed, subquadrate, convex anteriorly, punctate; 
lateral margins subparallel, plicate, with plications continuing but a short dis- 
tance on dorsal surface; front trilobed, not strongly advanced, median lobe de- 
pressed, triangular, with a deep median sulcus continued from the gastric regions; 
protogastric lobes distinct; anterior to these are another pair of lobes, trans- 
versely striate; these lobes are separated from the raised, transversely striate 
ocular margins by deep sulci which join the anterior hepatic sulcus posterior to 
the eye; a transverse plicated hepatic lobe, laterally margined by an anteriorly 
distinct cervical groove; on each side of the cardiac region is a crescentic whorl, 
opening posteriorly. The eye stalks are large, the cornea is small. Antennal 
peduncles heavy, stout, unarmed. 

288 San Diego Society of Natural History 

Chelipeds heavy, stout, subequal, granulate, tuberculate, rigid and deeply 
furrowed; merus stout, with carpal articulation transversely straight, armed at 
anterior distal end with a high granulated lobe; carpus including teeth, nearly as 
wide as long, short, wide (length 3.9 mm., width 3.3 mm.), armed on anterior 
margin with three or four stout, granulated, blunt, conical teeth, the distal the 
smallest; the carpus is deeply furrowed by two wide grooves, dividing the upper 
surface into three wide crests; the anterior and median crests are composed of 
obliquely placed, close set, rounded plications, or, in other specimens, these 
ridges are formed of a pavement of close set granulations; these two ridges are 
connected at the proximal end, open distally; the third ridge, on the outer margin, 
is not connected with the other two and is composed of obliquely placed tubercles 
and granules, which continue for a short distance on the ventral side; the hands 
are heavy, short, distorted, thick, with three longitudinal, deep, wide, granulated 
furrows, the median not separating its marginal ridges proximally; these two 
ridges thus form a long V, they being high, rather flat on top and composed of 
thickly crowded granules; the outer margin on the upper surface is a tumid, 
obliquely plicated ridge; these close set, beaded plications are scarcely continued 
on the inner side of the palm; the fingers of the major hand are short, thick, 
blunt, not crossing at the tips; the dactyl is armed with a large proximal lobe; 
the fingers of the minor hand are contorted, furrowed, granulate, with tips 

Ambulatory legs short, heavy; merus wide, compressed; carpus, propodus 
and dactylus clothed with short, sparse setae; under side of propodi and dactyli 
armed with a row of short, sharp spines. 

Sexual variation. — In the female the abdomen covers the sternum, in the 
male it does not. The sex may be determined at a glance by noting the relative 
difference in the size of the plates of the telson, those of the female being much 
the larger. 

Color in alcohol. — A uniform light red. 

Measurements. — Female holotype: length of carapace 5.5 mm., width 5.5 
mm. Male paratype: length 4.9 mm., width 4.8 mm. 

Range. — West Mexican coast, from Acapulco to Tenacatita Bay. 

Material examined. — Three males, one female, from Tequepa Bay, N. of 
Acapulco, Guerrero, December 18, 1932, collected by Captain Fred O. Lewis, of 
the yacht "Stranger." One male and two females, from Tenacatita Bay, Jalisco, 
December, 1932, collected by Captain Fred O. Lewis. The type specimens were 
selected from this latter series. 

Habitat. — Littoral. 

Remarks. — This proposed species is allied to P. curacaoensis Schmitt, in 
the general shape of the carapace and chelipeds. It differs from that species in 
the carpus being bisculate, instead of trisculate, by the hands being deeply 
trisculcate, instead of fairly smooth, and by the hands being nude, instead of 
with a peculiar bunch of tomentum on the upper surface. The carapace of P. lewisi 

Glassell — New Porcellanids and Pinnotherids 289 

greatly resembles that of the figure of Petrolistbes quadratus Benedict (Bull. 
U. S. Fish Comm, vol. 20, pt. 2, 1900 [ 1901 ] , pi. 3, fig. 4) . 

This species is dedicated to Captain Fred O. Lewis of Newport Beach, 
California, from whose yacht "Stranger" the specimens were collected. 

Pisosoma erosa, new species 

Type. — Female, holotype, Cat. No. 71539, and male, paratype, U. S. 
National Museum: from Magdalena Bay, Baja California, Mexico, 12 fathoms; 
December 2, 1931; collected by Steve A. Glassell. Female and male paratypes, 
Cat. Nos. 776 and 777, S. D. S. N. H.; from same series. 

Diagnosis. — Carapace suboval, convex, heavily eroded on margins. Carpus 
of chelipeds and ambulatories heavily eroded. 

Description. — Carapace subcircular, about as wide as long, convex fore and 
aft, regions outlined with sulci; branchial regions heavily eroded, lined and 
pitted; posterior third with prominent, outstanding, transverse carina, converging 
and ceasing at borders of intestinal region; antero-lateral margins forming a 
decided subcrenulate, granulated ridge; gastric regions lobed. Front in dorsal 
view subtruncate, entire, with a median sinus, in a front view, sharply upturned 
for the antennules. Eyes small. 

Chelipeds short, heavy; ischium pitted and toothed; merus reticulated, pitted 
and eroded, a gnarled granulated lobe at distal inner end; carpus a third longer 
than wide, armed with a single, proximal, serrated tooth on inner margin, granu- 
lated serrations on outer surface, reticulated, eroded, pitted and scored, pits 
bordered with granules, under surface transversely striate, pitted; hands with 
two granulated ridges from proximal end to base of fingers; on both sides of these, 
toward the margins, are reticulations and granulate bordered pits and erosions; 
pollex with a granulate tumid ridge supporting its cutting edge; dactyli with a 
longitudinal median sinus, granulate; fingers close fitted; under side of hands 

Ambulatory legs with merus sharp crested, granulate; carpus sharply ridged 
and eroded, as is the propodus; dactyli rather long, slightly curved at tip and 
with a row of supplementary spines on under surface. 

Abdomen fringed with hair. 

Color in alcohol. — A light pink. 

Measurements. — Female holotype: length of carapace 5 mm., width 5.2 
mm. Male paratype (a small specimen) : length 3.4 mm., width 3.4 mm. 

Range. — Magdalena Bay, Baja California, and Gulf of California. 

Material examined. — Two males and two females, taken at Magdalena Bay, 
December 2, 1931, 12 fathoms, by the author. The types for this species were 
selected from this material. Two small specimens from the north end of Tiburon 
Island, Gulf of California, January 1, 1932, 20 fathoms, collected by the author. 

290 San Diego Society of Natural History 

Habitat. — Evidently not a shore form, as it has so far been taken only in 
depths ranging from 5 to 20 fathoms. 

Remarks. — This proposed species is allied to P. sinuimanus Lockington, 
1878, but differs from that form in the extreme roughness of its surfaces and by 
the hands being less thick. 

The difficulty of obtaining undamaged specimens is due not only to its 
small size, but also to the hard materials which are brought up in the dredge 
with it. 

Pachycheles marcortezensis, new species 

Type. — Female, holotype, Cat. No. 71540, and male, paratype, U. S. 
National Museum: from off SE end of Angel de la Guardia Island, Gulf of 
California, Mexico, 20 fathoms; January 8, 1932; collected by Steve A. Glassell. 
Female paratype, Cat. No. 778, S. D. S. N. H.; from same series. One male 
paratype, in the collection of Steve A. Glassell, Beverly Hills, California. 

Diagnosis. — Carapace convex fore and aft, having scattered, short bristles 
on anterior 2/3 to orbital area, orbital area covered with short, sparse tomentum. 
Carpus with three long and two shorter, falcate spines. Telson of abdomen with 
five plates. 

Description. — Carapace wider than long, convex fore and aft, regions lightly 
outlined, scattered bristles, single and small groups on anterior 2/3; front cov- 
ered with sparse, short tomentum, subtruncate in dorsal view, sinuous in front 
view; a sharp postocular spine separates the eye from the antennae; anterior 
margin with a distinct carina which comes onto the carapace a third of the dis- 
tance from the posterior border. The first antennal peduncle is armed distally 
with a sharp, forward-pointing spine, the second is armed with two spines, one 
median, one distal. 

The chelipeds are subsimilar though unequal; merus short, with a distinct 
distal, anterior, spinose lobe, carpal articulation tuberculate, lined with bristles, 
a sharp spine at distal ventral terminus; carpus short, with four or more 
rows of longitudinally placed tubercles, those on the posterior half having radi- 
ating setae; the anterior half is covered with flat, squamoso-granulated, bristle- 
bearing groups; the anterior margin armed with three large scythe-like, forward- 
and outward-pointing teeth and two smaller distal teeth; hands triangular, short, 
thick, covered on the outer surface with lobes and bristle-bearing tubercles, and 
armed on outer margin with a close-setting row of short, dull-pointed teeth; inner 
surface punctate, with more roughness near outer margin; fingers with a slight 
gape, tips curved, overlapping. 

Ambulatory legs short, slightly setose; merus wide; carpus and propodus 
stout, with longitudinal rows of squamae, bearing bristles; dactyli short, heavy, 
curved, tips corneous, under side with several corneous spines; merus of second 
ambulatory leg has a few transverse striae on inner surface. 

Glassell — New Porcellanids and Pinnotherids 291 

Abdomen with five plates on the terminal segment. 

Color in alcohol. — Red mottled with white; bristles a straw color. 

Measurements. — Female holotype: length of carapace 4.8 mm., width 5.2 
mm. Male paratype: length 3.7 mm., width 3.7 mm. 

Range. — Gulf of California, at one time called the Sea of Cortez. 

Material examined. — Two females and two males, dredged off the SE end 
of Angel de la Guardia Island, Gulf of California, January 8, 1932, in 20 fathoms, 
by the author. The type specimens are from this series. 

Habitat. — Evidently not a shore form. 

Remarks. — This proposed species is allied to P. rugimanus A. M. Edwards, 
1880, but differs in that it has five teeth on the carpus instead of three or four, 
and these are longer in the first three and more hooked at their tips; also the 
entire surface of the chelipeds is covered with setae instead of being simply granu- 
lated and furrowed. 

Pachycheles sonorensis, new species 

Type. — Male, holotype, Cat. No. 71541, and female, paratype, U. S. 
National Museum: from Miramar Bay, near Guaymas, Sonora, Mexico, low 
tide; December 23, 1931; collected by Steve A. Glassell. Two paratypes, male 
and female, Cat. Nos. 779 and 780, S. D. S. N. H.; from same series. 

Diagnosis. — Chelipeds unequal, covered with short and long bristles. Cara- 
pace smooth in central area, punctate, with small bristles on remaining portions. 
Telson composed of seven plates. 

Description. — Carapace nearly as long as wide, subquadrate; punctate and 
sparsely bristled to central area, which is smooth; a pair of transverse, setose 
ridges behind the frontal region; front depressed, margined with short bristles, 
convex, not tomentose. First antennal peduncle rough at distal end, the others 

Chelipeds unequal; merus with a large anterior, distal lobe, upper surface 
rugose, sparsely bristled; carpus short, wide, tuberculate on proximal third and 
posterior border, the entire surface covered with squamae, bearing a group of 
short bristles; anterior margin tridentate in an arc; hands covered as in carpus, 
longest bristles on outer margin; inside of palm with tubercles and squamae 
except for central portion, tuberculate and with long bristles on inner gape of 
major hand; fingers of major hand unarmed, gaping; of minor hand, dentate, 
close-fitting, hooked at tips. 

Ambulatory legs short, heavy, sparsely covered with bristles; dactyli long, 
curved at corneous tip, armed on under side with a row of supplementary spines. 

Telson with seven plates, as in P. pubescens Holmes, 1900. 

292 San Diego Society of Natural History 

Color in alcohol. — Red mottled with white; bristles a straw color. 

Measurements. — Male holotype: length of carapace 7 mm., width 7.5 mm. 
Female paratype, length 7.6 mm., width 8 mm. 

Range. — Known from type locality only. Gulf of California. » 

Material examined. — A series of nearly a hundred specimens of both sexes, 
from Miramar Bay, Sonora, December 23, 1931, collected by the author. 

Habitat. — Found at low tide, under moss and sponge incrusted stones. 

Remarks.— This proposed species is allied to P. setimanus (Lockington) , 
1878, both as to size and general appearance, and in also having 7 plates in the 
telson; but it differs in a marked degree by not being tomentose, but setigerous 
instead, by having the inner side of the hand roughened and setose in the gape, 
instead of smooth and in having the first antennal peduncle rough at its distal end, 
instead of smooth. 

It seems remarkable that this species was not obtained at other collecting 
stations in the Gulf of California, where similar collecting conditions were found. 

Porcellana cancrisocialis, new species 

Type.— Male, holotype, Cat. No. 71542, and female, paratype, U. S. 
National Museum: from Punta Penasco (Rocky Point), Sonora, Mexico, low 
tide; May 2, 1935; collected by Steve A. Glassell. Two paratypes, male 
and female, Cat. Nos. 781 and 782, S. D. S. N. H.; from same series. 

Diagnosis. — Carapace slightly convex, sides slightly rounded. Front sharply 
tridentate. Shoulder posterior to terminus of cervical groove armed with a row 
of fine teeth. Lower orbital margin armed with a long, sharp, forward-pointing 
tooth. Hands tomentose. 

Description. — Carapace slightly convex in both directions, highest in central 
regions, with a polished appearance, but with front tomentose; transverse ridge 
behind front lined with tomentum; several short, transverse striations on branchial 
regions, tomentose. Front sharply tridentate, horizontal, median tooth equilater- 
ally triangular, largest, and separated from the laterals by a deep V-shaped 
notch; the outer margin of the lateral teeth forms the upper ocular margin in 
a long sweeping curve; the lateral teeth are sharp-pointed and nearly as long as 
the median tooth. The eyes may be partly retracted under the shelter of the 
upper ocular margin, which ends in a sharp-pointed postorbital tooth. The lower 
ocular margin extends forward, forming a flat, sharp, horizontal tooth at its 
inner angle, which extends as far forward as the base of the median rostral tooth. 
A shoulder at the posterior terminus of the cervical groove; this shoulder ex- 
tends from the margin a short distance onto the carapace and is armed on its 
anterior margin with a series of sharp denticles. The posterior margin of the 
carapace is nearly straight. 

Chelipeds short, stout, subequal, the left hand usually the larger; merus 
short, dorsal ly triangular, with a vertically compressed plate-like process armed 

Glassell — New Porcellanids and Pinnotherids 293 

on its forward surface with sharp thin teeth; on its distal inner end, this process 
terminates below the plane of the upper part of the carpus; the carpus is short 
and broad, about as wide as long, armed with a single, flattened, sharp-pointed 
tooth which occupies the proximal third of the inner margin, lightly transversely 
striate, with tufts of tomentum posteriorly; hands curving outward on both 
margins, short, thick, convex on upper surface, fingers heavier than pollices, 
covered on outer half with long pinnate tomentum; outer margin granulate- 

Ambulatory legs stout; merus wide, crested with a few setae as are the car- 
pus, propodus and dactylus; dactyli curved at the corneous tip and with a row of 
supplementary spines on the lower margin. 

Color in life. — Ground color an ivory yellow, overcast with lavender and 
blood red spots. Protogastric regions lighter (white in alcohol). Chelipeds same 
as carapace. Ambulatory legs banded on propodus with white. 

Measurements. — Male holotype: length of carapace 5.7 mm., width 4.8 
mm. Female paratype: length 5.4 mm., width 4.8 mm. 

Range. — Gulf of California to Magdalena Bay, Baja California. 

Material examined. — Thirteen specimens of both sexes were collected by 
the author at Punta Penasco (Rocky Point), Sonora, May 2, 1935. From this 
series the type specimens were selected. Three specimens were collected by the 
author at San Felipe, Baja California, June 20, 1936. 

Habitat. — This species, like P. paguriconviva, is commensal with the large 
Pagurid, Petrochirus calif orniensis Bouvier, 1895, and enjoys the same associa- 
tion. Usually a single pair occupies a shell, but one or three may be present. 

Remarks. — This proposed species is allied to P. sayana (Leach), 1820, but 
differs from that species in that the base of the antennae is armed with spines 
instead of being smooth, by the hands being more covered with hair and tomen- 
tum, and by the tip of the rostrum not being decurved. Specimens so far are 

Porcellana paguriconviva, new species 

Type. — Male, holotype, Cat. No. 71543, and female, paratype, U. S. 
National Museum: from Punta Penasco (Rocky Point), Sonora, Mexico, low 
tide; May 1, 1935; collected by Steve A. Glassell. Two paratypes, male and 
female, Cat. Nos. 783 and 784, S. D. S. N. H.; from same series. 

Diagnosis. — Carapace smooth, or with tomentum on front. Front tridentate, 
the median tooth largest, triangular, equilateral. Chelipeds with carpus unarmed; 
hands dorsally flat, wide, fringed on outer margin or nude. 

Description. — Carapace longer than wide, depressed, sides subparallel, 
regions ill-defined. Front horizontal, tridentate, the median tooth the largest 
and most prominent, triangular, equilateral (almost as in Petrolisthes, except not 

294 San Diego Society of Natural History 

depressed, and without a median sulcus) , the lateral teeth truncate in front, 
forming a right angle with the upper ocular margin; the ocular margin partly 
covers the eyes. There is a postocular tooth; a shoulder behind the terminus of 
the cervical groove, unarmed; a pair of lunate pits between the gastro-cardiac 
regions. There may or may not be tomentum on the frontal region, usually not 
more than microscopic. 

The chelipeds are short and heavy, depressed on upper surface; the merus is 
short, the distal inner end with a blunt compressed upward-pointed lobe; the 
upper surface of the merus is triangular, widest posteriorly, its apex the hinge 
of the carpus; the carpus is short, wide, flattened, lightly rugose, widest at proxi- 
mal end, at which point it is as wide as long, 2 mm.; the hands are short, stout, 
microscopically rugose, flattened on upper surface, subequal, with tips of pollices 
slightly upturned; dactyli with truncate, flattened tips, crossing tips of pollices 
on inner margin, one hand slightly the larger, outer margin fringed with tomen- 
tose hair, or nude. 

Ambulatory legs short, stocky, margined with tomentum and light setae; 
dactyli stout, curved at corneous tip, armed on under surface with a row of 
supplemental corneous spines. 

Color in life. — Ground color, in longitudinal stripes, a bright lavender, a 
uniform design of bright orange overlaid on this. Chelipeds same as carapace, 
but not patterned; legs with a white spot on propodus. Ventral side iridescent, 
pinkish white, with longitudinal pattern of carapace continued on first three 
segments of abdomen. 

Measurements. — Male holotype: length of carapace 5.8 mm., width 5.1 
mm. Female paratype: length 5.6 mm., width 5.1 mm. 

Range. — Gulf of California. 

Material examined. — A series of about 25 specimens, collected by the 
author, at Punta Penasco (Rocky Point) , Sonora, May 1, 1935. 

Habitat. — These little crabs are commensal with the large hermit crab, 
Petrochirus californiensis Bouvier, 1895. The usual association is: the Pagurid 
host, occupying the shell of Phyllonotus nigritus (Philippi), accompanied by a 
large Pollonoid worm and a pair of these little Porcellanids. At times the inner 
face of the shell may have a Crepidula nivea Gould, attached, and this in turn 
may be commensalized with the Pinnotherid, Fabia granti Glassell. 

Remarks. — This proposed species is allied to P. sayana (Leach), 1820, but 
differs from that species in the shape of the front, which does not have its lateral 
teeth separated from the angles of the orbits by deep incisions. Also in that the 
carpus of the chelipeds in diis species has its inner margin in an unbroken oblique 
line, while P. sayana has a proximal lobe on the inner margin. In this species 
there may be individual variations in the size and shape of the lateral teeth of the 
front, some protruding a little more than in the holotype and having a rounded 
sinus on the margin at the base of the median tooth. 

Glassell — New Porcellanids and Pinnotherids 295 

Porcellana magdalenensis, new species 

Type. — Female, holotype, Cat. No. 71544, U. S. National Museum: from 
Magdalena Bay, Baja California, Mexico, 12 fathoms; December 2, 1931; col- 
lected by Steve A. Glassell. One paratype female, Cat. No. 785, S. D. S. N. H.; 
from same series of 5 specimens. 

Diagnosis. — Carapace nearly as long as wide, suboval, convex, regions 
denned; lateral, hepatic and upper and lower ocular margins dentate. Front 
trilobate, margined with denticles. Chelipeds with median longitudinal ridges, 
dentate. Legs long. 

Description. — Carapace suboval, regions defined, convex, crossed with trans- 
verse rugae; protogastric lobes distinct, with anterior border prominent, tomen- 
tose. Lateral margins bordered with sharp, forward-pointing spines that are con- 
tinued onto the margin of the shoulder on the carapace, which is behind the ter- 
minus of the cervical groove; hepatic regions bordered with forward-pointing 
spines, the proximal the largest; the front has three dentate lobes; the median 
triangular, with a median sulcus and slightly depressed at the tip; the lateral 
lobes are rounded at their apices, separated by a wide V-shaped groove from the 
median lobe, the outer margin extending backward, forming the upper ocular 
margin. The frontal lobes and both the upper and lower ocular margins are lined 
with small sharp teeth. 

The chelipeds are long and narrow, subequal; merus fringed with small teeth 
on upper carpal articulation, and having a wide, compressed, inner distal lobe, 
fringed with teeth, and one or more teeth at ventral distal angle; carpus longer 
than wide, armed on inner margin with a row of small, sharp teeth, the proximal 
ones the largest; a median spinate ridge, the surface between this ridge and the 
inner margin lightly tomentose and concave, while toward the outer margin the 
surface between the median ridge and a row of spines parallel to the outer mar- 
gin is convex and rugose; the hand is narrow, the outer margin concave, lined 
with spines and fringed with pinnate tomentum; a median spinate ridge from 
proximal end to base of dactyl us is bordered by two concave surfaces; fingers 
long, half the length of the hand, contorted; pollex terminates in a sharp, slighdy 
upturned point; dactylus with upper margin toothed, a short median spinate 
ridge between this and the prehensile edge; a fringe of pinnate tomentum veils 
the cutting edge. 

Ambulatory legs long, fairly slender; merus crested with short setae, trans- 
versely rugose; propodus long, cylindrical, with a few setae; dactyli long, com- 
pressed, curved, with needle-like tips. 

Epistome heavily plicated. 

Sexual variation and color in life. — Unknown. 

Measurements.— Female holotype (left cheliped missing) : length of cara- 
pace 4.5 mm., width 4.6 mm.; length of hand 5 mm.; width at base of finger 1.5 
mm.; length of fingers 2.5 mm.; length of carpus 2.5 mm., width 1.5 mm. 

296 San Diego Society of Natural History 

Range. — Known only from type locality. 

Material examined. — Five female specimens, collected in Magdalena Bay, 
Baja California, in 12 fathoms, by the author. The ovigerous holotype for this 
proposed species is the largest of the series. 

Habitat. — Evidently among sponges and corallines. 

Remarks. — This species is allied to P. serratifrons Stimpson, 1858, but 
differs from that species in the shape of the front, the median lobe being triangu- 
lar instead of rounded, with the lateral lobes but little less in width, instead of 
subacute and scarcely less prominent. The chelipeds are more heavily spined in 
this species than on P. serratifrons and, in addition, the hands are fimbriate with 

ORTHOCHELA, new genus 

Carapace longer than wide, transversely convex; front horizontal, truncate, 
except for a prominent, median, equilateral, rostral protuberance; the front 
nearly as wide as the carapace; lateral margins of the carapace subparallel, armed 
with short, sharp, forward-pointing spines, terminating at cervical shoulder. 
Eyes large, stalks thick, short; may be semi-retracted under lateral edge of ocular 
margin. Antennae partly excluded from orbit, nearly as long as chelipeds; flagel- 
lum naked. Chelipeds subsimilar, unequal, directed forward, as in the Galathei- 
dae, length about l x h times the length of the carapace; merus extends past the 
rostrum; carpus and hand long, cylindrical; fingers short, opening vertically. 
Ambulatory legs short, propodi longer than merus, dactyli curved at tip, simple, 
not multiunguiculate. Terminal segment of abdomen with seven plates. 

This genus has a remote affinity to the genus Minyocerus Stimpson, 1858, 
which is based on Porcellana angusta Dana, 1852, in which the chelipeds are 
somewhat similar. The carapace, however, differs from all the other genera in 
this family and is more like that of the genus Uroptychns Henderson, 1888, of 
the family Galatheidae. 

Genotype. — Orthochela pumila, new species, taken at Magdalena Bay, Baja 
California, Mexico, 1 fathom, on yellow gorgonian coral (sea-fans), December 
2, 1931, by Steve A. Glassell. 

Orthochela pumila, new species 
Plate 21, figure 1 

Type. — Male, holotype, Cat. No. 71545, and female, paratype, U. S. 
National Museum: from Magdalena Bay, Baja California, Mexico, 1 fathom; 
December 2, 1931; collected by Steve A. Glassell. One paratype male, one para- 
type female, Cat. Nos. 786 and 787, S. D. S. N. H; from same series. 

Diagnosis. — Carapace subquadrilateral, margins spinous, front unidentate, 

Glassell — New Porcellanids and Pinnotherids 297 

truncate to width of carapace from base of rostrum. Chelipeds forward-pointing, 
long, cylindrical, fingers moving in a vertical plane. 

Description. — Carapace subquadrilateral, longer than wide, transversely 
convex, cervical groove ill-defined, surface polished, hard, microscopically punc- 
tate; lateral margins armed with a row of closely set, forward-pointing spines, 
terminating at the shoulder behind the cervical groove. Posterior margin 
straight, entire. Front truncate, wide, extending nearly the width of the carapace, 
where it turns backward in a curved line, thence runs tangent to its former 
direction, finally sweeping outward to form the postorbital tooth which covers 
the first antennal peduncle; the rostrum is a large, truncate-tipped, equilateral tri- 
angle whose base is the line of the front; it is armed at its truncate apex with a 
row of short, sharp, protruding teeth; the lateral margin of the truncate front, 
anterior to the eyes, is also minutely toothed. The antennae are long, naked, 
and may be pointed in a direction either straight forward or straight backward. 
The eyes are large, with stout, short stalks, and may be retracted until half of the 
cornea is visible beneath the upper ocular margins. 

The chelipeds are subsimilar, unequal, directed forward and in large speci- 
mens may be 2 1 /? times the length of the carapace; the merus extends past the tip 
of the rostrum; the length of the carpus is about equal to the width of the cara- 
pace, half as wide as long, subcylindrical, smooth and having a concavity at its 
distal inner end for the partial reception of the hand; the major hand is nearly 
twice the length of the carpus, cylindrical, with the fingers in a vertical plane; 
the fingers are short, being a trifle more than 1/5 the length of the hand; the 
pollex is serrated with teeth on its outer distal margin; the fingers of the minor 
chela are longer in proportion, and not so heavy. 

The ambulatory legs are bent underneath the body in a grasping position; 
the merus is short, compressed, on the anterior margin ending distally with a 
high, sharp outward-pointing tooth; the carpus is short; propodus long and 
slightly curved, longer than the merus; the dactyli are long, curved at tip and 
armed on the under side with a row of supplementary spines, simple, not mul- 

The outer maxillipeds have the ischium lightly crossed with transverse 

The ultimate segment of the abdomen has seven plates. 

Sexual variation. — Apparently only a difference in size, the males with the 
longer and heavier chelipeds. 

Color in life. — A rich yellow; lines of red on the hepatic regions. Hands 
with a few red blotches on outer surface; fingers with red bases and tips. 

Measurements. — Largest male (this specimen had lost its minor cheliped) : 
length of carapace 3.6 mm., width 2.8 mm.; length of major cheliped 9.8 mm.; 
length of hand 5.1 mm.; width of merus, carpus and hand approximately the 
same, 1.5 mm. Female paratype: length of carapace 3.8 mm., width 3 mm.; 
length of major cheliped 7.7 mm.; hand 3.2 mm.; carpus 2.2 mm.; merus and 

298 San Diego Society of Natural History 

ischium 1.3 mm.; length of first ambulatory leg 3.8 mm.; merus 1 mm.; carpus 
0.5 mm.; propodus 1.5 mm.; dactylus 0.8 mm. 

Range. — Known only from the type locality. 

Material examined. — A series of both sexes, 35 specimens, collected at 
Santa Margarita Island, Magdalena Bay, Baja California, December 2, 1931, 
in 1 fathom, by the author. 

Habitat. — These little crabs were found clinging to yellow gorgonian coral, 
along with Isopods and Amphipods, all of which harmonized so exactly in color 
with their host that they were to be distinguished only with difficulty. 

Remarks. — This proposed species shows a rather close relationship to the 
members of the Galatheidae in the position of its chelipeds, and by having 
spinose lateral margins on the carapace. From their size, the location of their 
eyes, and the shape of the front, the specimens collected might be thought to be 
immature or even larval forms, if it were not that nearly all the females found 
were gravid. 


Fabia unguifalcula, new species 
Plate 21, figure 2 

Type. — Female, holotype, Cat. No. 788, San Diego Society of Natural 
History: from Punta Penasco (Rocky Point), Sonora, Mexico, low tide; May 3, 
1935; collected by Steve A. Glassell. One female paratype, from same locality, 
in the collection of Steve A. Glassell, Beverly Hills, California. 

Diagnosis. — Carapace with sides subparallel. Ischium-merus of outer 
maxilliped crescentoid, palp two-jointed. Dactyls of legs falcate. Hands short, 
wide, heavy; fingers dentate. 

Description. — Carapace smooth, glossy, membranaceous, subtransparent, 
much wider than long, anteriorly arcuate, sides subparallel. Front turned abruptly 
downward. Eyes subovoid, cornea minute, not visible in a dorsal view. Basal joint 
of antennae short and wide. A narrow furrow leading backward from the buccal 
angle. Ischium-merus of outer maxillipeds crescentic, palp with two joints, ulti- 
mate segment wide, rounded distally. 

Chelipeds stout, equal; merus short, not extending far past sides of carapace; 
carpus long, wide, rounded dorsally, inner proximal margin tomentose; hands 
short, wide, heavy, thick, smooth; lower margin sinuous; pollex slightly deflexed, 
with upturned, sharp-pointed tip, armed with a triangular-shaped cutting edge, 
with a median tooth and proximal denticles; dactylus long, falcate, armed proxi- 
mally with a prominent denticulate tooth. Tips of fingers crossing. 

Ambulatory legs paired, the first pair differing from the others in that the 
upper crest of the merus, the anterior lower margin of the carpus and propodus 

Glassell — New Porcellanids and Pinnotherids 299 

are margined with tomentum. The dactyli of all legs are falcate, the fourth the 
least, the second the longest and sharpest. 

The abdomen covers the sternum, is circular and fringed with hair. 

Sexual variation and color in life. — Unknown. 

Measurements. — Female holotype: length of carapace 4 mm., width 5 mm. 

Range. — Known only from type locality. 

Material examined. — Two ovigerous females, the larger the type, collected 
by the author at Punta Peiiasco (Rocky Point) , Sonora, May 3, 1935. 

Habitat. — Collected in the inter-tidal zone. Association and host not de- 

Remarks. — This proposed species is allied to F. granti Glassell, 1933, but 
differs in that the front is nearly a continuation of the curve of the anterior mar- 
gins, instead of being advanced; by the first ambulatory leg bearing tomentum, 
instead of being smooth and naked; and by the hands being short, stout, sub- 
quadrate and compact, instead of long and increasing in width distally. In addi- 
tion, F. granti appears to be a much larger species. 

Dissodactylus xantusi, new species 
Plate 21, figure 4 

Type. — Female, holotype, Cat. No. 71546, and male, paratype, U. S. 
National Museum: from Espiritu Santo Island, Gulf of California, Mexico, low 
tide; December 8, 1931; collected by Steve A. Glassell. Paratype, female, para- 
type, male, Cat. Nos. 789 and 790, S. D. S. N. H.; from same series. 

Diagnosis. — Carapace convex fore and aft. Dorsal ridge short, oblique. 
Legs stout; dactyli of legs one to three bifurcate for almost half their length. 
Carpus with a transverse, setose, median ridge. Hand crossed on upper margin 
with three oblique, setose ridges. Terminal segment of male abdomen nearly an 
equilateral triangle. Palp of outer maxilliped three- jointed. 

Description. — Carapace distinctly broader at lateral angles than posteriorly; 
lateral margins subequal, antero-lateral arcuate, postero-lateral straight, slightly 
concave posteriorly. Dorsal surface naked and polished, lightly punctate, convex 
fore and aft, slightly from side to side, depressions on metabranchial regions and 
margins of cardiac; a raised rim on antero-lateral margin sharply turns at lateral 
angle and continues obliquely on dorsal surface for a short distance. Margin of 
front slightly convex, slightly advanced beyond curve of antero-lateral margins; 
posterior margin sinuous. 

Palp of outer maxilliped with three joints; the penultimate spatulate, truncate, 
with inner margin straight, with outer margin arcuate, widest distally; terminal 

300 San Diego Society of Natural History 

segment small, slightly advanced beyond line of penultimate segment and located 
on that segment's inner distal angle. 

Merus of chelipeds extends but little beyond margin of carapace, upper distal 
margin setose; carpus as broad as long, crossed with an interrupted, transverse, 
median, setose ridge, the distal margin arcuate, setose; hands proximally swol- 
len, upper crest straight, rounded, obliquely crossed by three distinct, setose 
ridges, the bristles pointing forward; the under margin has three light, oblique, 
setose ridges, the proximal the longest, a sparse row of long pinnate hair on inner 
surface to base of pollex; the outer surface is crossed with short, oblique, setose 
ridges, the distal ridge continuing on the base of the pollex; fingers long, close- 
fitting, with tips crossing. 

Ambulatory legs slightly compressed, margined with fine, light hair; merus 
stout; carpus of first leg, only, with a longitudinal, sub-oblique ridge; the propo- 
dus has a sharp, setose crest, the bases of the bristles being just beneath this 
crest on the anterior side and pointing at right angles to the axis of the segment; 
dactyli of the first three legs bifurcate for nearly half their length; surface of 
legs highly polished, lightly punctate. 

The abdomen of the female does not cover the sternum; the lateral margins 
of the segments, including the proximal half of the sixth, are subparallel; the 
terminal segment is broadly triangular, more than twice as wide as long. Male 
abdomen fused in third, fourth and fifth segments, widest at third, lateral mar- 
gins converging to proximal half of penultimate segment, terminal segment an 
equilateral triangle. Surface of abdomen in both sexes highly polished, lightly 

Sexual variation. — Hands of males heavier in proportion than those of 
the females. The females are the larger specimens. 

Color in life. — A chocolate brown with a design of cream-colored lines and 
spots; the gastro-cardiac region divided by a transverse arcuate line, surmounted 
anteriorly by a broad V-shaped design. Legs with terminals of the joints banded 
with cream color; dactyli light-colored, as are the fingers of the hands; hands 

Measurements. — Female holotype: length of carapace 4.5 mm., width 6.1 
mm. Male paratype: length 3.8 mm., width 4.9 mm. 

Range. — Gulf of California. 

Material examined. — Twenty specimens of both sexes were collected at 
Espiritu Santo Island, Gulf of California, December 8, 1931, by the author. 
The type specimens were selected from this series. Small series of both sexes 
were also collected by the author at the following stations: Las Animas Bay, 
Baja California, January 2, 1932; Coyote Cove, Concepcion Bay, Baja Cali- 
fornia, January 18, 1932; San Felipe, Baja California, June 1, 1934; Punta 
Periasco (Rocky Point) , Sonora, May 2, 1935. 

Glassell — New Porcellanids and Pinnotherids 301 

Habitat. — Commensal on the exterior ventral surface of Echinoids, such as 
Mellita and Encope. They are in close association with D. nitidus Smith; both 
species may at times be found on the same host. 

Remarks. — This proposed species is allied to D. nitidus Smith, 1870, which 
it closely resembles, but from which it differs in the shape of the chelipeds, by 
the hands being rougher, the fingers stouter, by being naked under the pollex, 
instead of decorated with a tuft of thick black tomentum, and by the penulti- 
mate segment of the palp of the outer maxillipeds not having its lateral margins 
straight and parallel. 

This species is dedicated to Louis John Xantus de Vesey, in appreciation 
of his character as a gentleman and of his attainments and zeal as a votary of 
natural history. 

Pinnixa richardsoni, new species 
Plate 21, figure 3 

Type. — Male, holotype, Cat. No. 791, San Diego Society of Natural 
History: from Balboa, Canal Zone, Panama, upper tidal zone; February 22, 
1936; collected by Frank Richardson. 

Diagnosis. — Carapace twice as wide as long, regions deeply furrowed. 
Hand thin, compressed; fingers acuminate. Third leg longer than body width, 
heavy. Dactyli of ambulatories horizontally compressed. Third, fourth and fifth 
segments of male abdomen fused. 

Description. — Carapace twice as wide as long, covered with very short 
setae; anterior and antero-lateral margins together forming a strongly convex 
arch, reaching to the widest part of the cardiac region and meeting the posterior 
angle at an obtuse angle; posterior margin transverse at its middle for 1/3 of 
carapace width. Gastric and cardiac regions delimited, the latter wider; three 
longitudinal, narrow gastric furrows, the median short, reaching half way back; 
branchial region crossed by four obliquely transverse furrows, the hinder one 
deep and parallel to the posterior margin. A short dorsal hepatic furrow is 
directed inward and forward. The antero-lateral margin is tuberculate distally. 
The eyes are small, dorsally placed, filling their orbits. The front is horizontal, 
truncate, entire, not extending past buccal area in dorsal view. Fronto-orbital 
width V4 width of carapace. 

Chelipeds small, longer than first leg; merus hairy, carpus and manus mar- 
gined with long brown hair; carpus wide, compressed, with a high distal crest, 
outer surface covered with microscopic hair; hands thin, sharp-edged, compressed, 
not as wide as carpus, flat on outer surface, covered with minute hair, which, 
like those on the carpus, do not obscure the view of the surface; upper margin 
arched, lower margin straight; fingers narrow, longitudinal, acuminate, gaping in 
proximal two-thirds, movable finger crested with long hair, lower margin of 

302 San Diego Society of Natural History 

pollex nude. A median longitudinal fringe of long hair on inner side of carpus 
and hand. The inner proximal end of the pollex furnishes hair at the gape. 

Ambulatory legs stout; the first is remarkable for the shape of its propodus 
and dactyl, the propodus being wide, short and heavy, its upper margin much 
the shorter; the dactyl is horizontally compressed, wide, short, crooked, up- 
turned, its margins tangent to those of the propodus, as are the margins of the 
dactyli of the other legs; the third leg is very heavy, stout, 1/6 longer than the 
width of the carapace and 1/3 longer than the second leg; the merus is equal in 
length to that of the carapace, and nearly half as wide as long; there is a tubercle 
on the distal posterior surface of the carpus and propodus. The presence of a 
dense growth of tomentum and setae, covering the entire surface posteriorly, 
makes a close observation difficult; the dactyl is stout, heavy, dull-pointed, hori- 
zontally fringed with setae; the fourth leg reaches nearly to the end of the merus 
of the third leg. 

The abdomen is widest at the third segment, very long and narrow, over- 
lapping the buccal cavity; third, fourth and fifth segments regularly tapering, 
fused, sixth long and narrow, tapering, seventh long, rounded at tip which is 
margined with hair, sides converging. Ischium-merus of outer maxilliped with 
upper distal margin arched, a transverse line of hair located centrally. 

Sexual variation and color in life. — Unknown. 

Measurements. — Male holotype: length of carapace 6.5 mm., width 12.8 
mm.; orbital width 3.2 mm.; transverse posterior margin 4 mm.; length of sec- 
ond leg 9.8 mm.; of third leg 15 mm. 

Range. — Known only from type locality. 

Material examined. — The single male specimen, the holotype. 

Habitat. — Unknown. This specimen was dug out of heavy, thick mud in 
the upper tidal zone, from a small channel margined with mangrove trees. No 
host noted. 

Remarks. — This proposed species is very closely allied to P. valerii Rathbun, 
1931, but differs in the outer surface of the hands and carpus being smooth, in- 
stead of covered with tomentum (as is a topotype of P. valerii, before me) ; by 
the outer distal margin of the outer maxilliped being arched, instead of angular; 
and by some of the segments of the abdomen being fused, instead of articulated. 
Otherwise there are a great many similarities between these two unique species, 
which would seem to set them apart from other species of this genus with which I 
am familiar. 

I take pleasure in dedicating this species to Mr. Frank Richardson, orni- 
thologist, and graduate student of the University of California, to whom I am 
indebted for this specimen. 

Glassell — New Porcellanids and Pinnotherids 

Plate 21 

Fig. 1. Ortbochela pumila, n. gen. and n. sp. Male. 

Fig. 2. Fabia unguifalcula, n. sp. Outer maxilliped. 

Fig. 3. Dissodactylus xantusi, n. sp. Outer maxilliped. 

Fig. 4. Pinnixa richardsoni, n. sp. Outer maxilliped. 



Volume VIII, No. 22, pp. 305-320, plate 22 



A. M. Strong and H. N. Lowe 

San Diego Society of Natural History 


Printed for the Society 

December 7, 1936 


U. S. Grant, IV, Chairman 

Fred Baker Clinton G. Abbott, Editor 



A. M. Strong and H. N. Lowe* 

San Diego Society of Natural History 

Included among the shells dredged by the Templeton Crocker 
Expedition of 1932 off the west coast of Central America and Mexico 
were a large number of specimens of the genus Phos. These, and the 
specimens dredged by the junior author off Taboga Island, Panama; 
Acapulco, Mexico; and in the Gulf of California, Mexico, have added 
greatly to the amount of west coast material available for study. The 
large series of specimens of some of the species and representatives of 
many of the described forms in these collections seem to warrant a 
review of the genus as applied to the west coast. 

The writers wish to express their appreciation to Mr. Templeton 
Crocker, whose generosity provided most of the specimens upon which 
this paper is based. Acknowledgment is also due Dr. G. Dallas Hanna, 
Curator of the Department of Paleontology of the California Academy 
of Sciences, for assistance and advice. The information regarding the 
records of the species occurring fossil in the Tertiary of western North 
America has been furnished to the authors by Dr. Leo G. Hertlein, of 
the Department of Paleontology, California Academy of Sciences, and 
Dr. U. S. Grant, IV, of the Department of Geology, University of 
California at Los Angeles. Dr. Alexander Wetmore, Assistant Secre- 
tary, Smithsonian Institution, kindly furnished photographs of certain 
specimens in the U. S. National Museum. These photographs were 
prepared under the supervision of Dr. Paul Bartsch, to whom west 
coast students are under obligation for many similar courtesies in the 

The type of the genus Phos Montfort, 1810, "Murex" senticosus 
Linnaeus, is an Indo-Pacific shell. Woodring 1 stated that the West 
Indian and Panamic species are probably not congeneric. He consid- 
ered the West Indian species to be descendants of the Eocene and 

* H. N. Lowe, outstanding authority on west coast shells, died on June 10, 1936. The 
San Diego Society of Natural History, to which he bequeathed his large conchological 
collection, dedicates to his memory the publication of this paper, the last to bear his name. — 

1 W. P. Woodring, Carnegie Inst. Washington, Publ. 385, 1928, p. 260. 

308 San Diego Society of Natural History 

Oligocene genus Tritaria Conrad and placed the Tertiary and living 
species in a new subgenus Antillophos, type "Cancellaria" candei 
d'Orbigny, Recent, West Indies. This species was united by Tryon 
with the west coast Phos veraguensis Hinds, which, while undoubtedly 
distinct, is closely related. For a thorough study of the entire group, 
world wide material, both fossil and living, would be required; so for 
the purpose of this paper the name Phos is used as it was by the older 

Recent species of the genus on the west coast seem to be limited 
to tropical waters, Gulf of California to Panama. Nearly all of the 
records are from dredged material in depths ranging from 20 to 100 
fathoms. Little dredging has been done in this area and the literature 
on the various species is limited. Hinds described and figured four 
species in the portion of the Zoology of the Voyage of the Sulphur 
dealing with Mollusca. In addition to these, one species described by 
Reeve, one by Carpenter, one by A. Adams, one by Powys as a Nassa, 
and one by C. B. Adams as a Triton make up the list from the older 

Dall, 2 in his "Summary of the Family Alectrionidae from the 
West Coast of America," cited under the genus Phos only one of 
these older species, Phos crassus Hinds. In this and other papers he 
described a number of new species, either as Phos or which should be 
referred to Phos. Several of these were unfigured, but the types in the 
U. S. National Museum have been examined by the junior author. 
Some of these are here placed in synonymy with species described by 
Hinds. A study of the large number of specimens now available shows 
that the character of the nuclear whorls, presence or absence of a 
columellar keel, and dentition of the outer lip are specific characters. 
The color, number of ribs, and details of sculpture are more variable. 

In the following pages all species of Phos for which we have been 
able to find west coast records are mentioned. The genus is described 
in ZittePs 3 Text-book of Palaeontology as follows : 

"Shell elongate, bucciniform, turriculate; spire sharp, elevated, 
whorls ornamented with prominent longitudinal costae, and less salient 

2 W. H. Dall, Proc. U. S. Nat. Mus., vol. 51, no. 2166, 1917, p. 578. 

3 W. H. Dall in Text Book of Palaeontology adapted from Karl A. von Zittel by 
Charles R. Eastman, vol. 1, 1913, p. 556. 

Strong & Lowe — The Genus Phos 309 

spiral threads and sulci, often varicose. Aperture oblong, outer margin 
Urate within. Columella excavated, plicate in front; canal short, slightly 

Phos articulatus Hinds 
Plate 22, figure 6 

Phos articulatus Hinds, Zool. voy. Sulphur, vol. 2, Moll. pt. 2, October 1844, 
p. 38. "Panama."— Sowerby, Thes. Conch., vol. 3, (pt. 19), 1859, p. 91, 
pi. 221, fig. 32. "Panama."— Tryon, Man. Conch., vol. 3, 1881, p. 218, pi. 
83, fig. 516. Panama; western Colombia. 

Phos turritus A. Adams, Proc. Zool. Soc. London, 1850, p. 154. "Panama, coral 
sand, 6 to 10 fathoms. "—Sowerby, Thes. Conch., vol. 3, (pt. 19), 1859, 
p. 91, pi. 221, fig. 37. Original record cited. — Tryon, Man. Conch., vol. 3, 
1881, pi. 83, fig. 517. 

Specimens were dredged by the Templeton Crocker Expedition at the fol- 
lowing localities : 

Loc. 27587 (C A. S.), off Cape San Lucas, Lower California, Mexico, near 
big rocks, in 20-25 fathoms. 

Loc. 27585 (C. A. S.) , Lat 23° 02' N., Long. 109° 32' W. A few miles off 
shore at Gorda Point, in San Jose del Cabo Bay, Lower California, Mexico, in 
20-25 fathoms. 

Loc. 27581 (C A. S.), between Isabel Island and Mazatlan, Sinaloa, 

Loc. 27571 (C. A. S.) , Lat. 16° 39' N., Long. 99° 24' 30" W., to Lat. 16° 
38' N., Long. 99° 27' 30" W. About 33 miles slightly east of Acapulco, Guerrero, 
Mexico. This is about 32 miles west of Dulce Bay. 

Loc. 27568 (C. A. S.), Lat. 14° 52' N., Long. 93° 04' W., in 35 fathoms. 
About 23 miles west of San Simon Bar, Mexico. 

Loc. 27566 (C A. S.), Lat. 14° 25' N., Long. 92° 28' W., in 35 fathoms. 
About 28 miles west of Champerico, Guatemala. 

A free translation of Hinds' original Latin description is as follows : 

"Shell elongate-ovate, white, clouded with brown, whorls rounded, ribbed; 
with spiral lines; below the suture flatly sloping to an angular shoulder; articulated 
with narrow whitish brown bands; with about 14 ribs at the periphery, sometimes 
swollen; columella smooth." 

To this can be added that the nucleus consists of five spirally threaded whorls 
and that the outer lip is dentate. P. turritus A. Adams was also described from 
Panama. Tryon stated that it is the same as P. articulatus Hinds and there is 
nothing in the descriptions or figures to indicate that they can be separated. 

310 San Diego Society of Natural History 

Phos chelonia Dall 
Plate 22, figure 3 

Phos chelonia Dall, Proc. U. S. Nat. Mus., vol. 51, 1917, p. 578. "Dredged at 
the Galapagos Islands in 40 fathoms." 

Dall's description of this species is as follows: 

"Shell very similar in general appearance and size to P. varicosus Gould, 
having similar whitish varices, about three or four to a whorl, but differing by 
having the whorls appressed to the suture, not deeply impressed, and in having a 
nuclear shell of five or six whorls, deeply spirally sulcate instead of a nearly smooth 
one of three and a half whorls. The color of the present species is pale yellowish 
with a tinge of brown, as in varicosus, which also has narrower and more numer- 
ous ribs between the varices." 

Phos varicosus Gould 4 was described from the Philippines and is said by 
Tryon 5 to be a synonym of P. roseatus Hinds from the same region. Phos chelonia 
differs from the other species described from the west coast in the presence of 
the whitish varices. No specimens that could be referred to this species were col- 
lected by either the Templeton Crocker Expedition or the junior author. 

Phos cocosensis Dall 

Plate 22, figure 7 

Phos cocosensis Dall, Proc. U. S. Nat. Mus., vol. 18, April 23, 1896, p. 11. 
"U. S. Fish Commission station in 66 fathoms, near Cocos Island, Gulf of 
Panama."— Dall, Bull. Mus. Comp. Zobl., vol. 43, no. 6, 1908, p. 306, pi. 
8, fig. 5. Near Cocos Island, Gulf of Panama. Also at U.S.S. Albatross Sta. 
3387, in 127 fathoms.— Dall, Proc. U. S. Nat. Mus., vol. 51, 1917, p. 578. 
"Gulf of Panama, Cocos Island, (Gulf of California?) ." — Zetek, Rev. 
Nueva, nos. 1 & 2, 1918, p. 22. 

Dall gave the type locality as near Cocos Island, U.S.S. Albatross Sta. 3368 
in 66 fathoms. He also listed the species from U.S.S. Albatross Sta. 3387, in 127 
fathoms and in a later paper he cited it questionably from the Gulf of California. 
His original description is as follows: 

"Shell elongate, acute, eleven-whorled, including a nucleus of four whorls; 
color yellowish white, with variable brown spiral banding; sculpture of eleven or 
twelve narrow, little elevated, distant ribs, more or less angulated at the shoulder; 
spiral sculpture of numerous rather sharp, close threads, flatter on the last whorl, 
with a few more prominent between the suture and the shoulder; sutures distinct, 
whorls moderately rounded; aperture longer than wide, with an entire outer lip, 

4 A. A. Gould, Proc. Boston Soc. Nat. Hist., vol. 3, 1849, p. 143. "Philippine Islands." 
— Gould, Otia Conch., 1862, p. 66. — Gould, U. S. Explor. Exped., (Wilkes Exped.), 
Moll. Atlas, 1856, pi. 20, figs. 360, 360a. 

5G. W. Tryon, Jr., Man. Conch., vol. 3, 1881, p. 217, pi. 83, fig. 511. 

Strong & Lowe — The Genus Phos 311 

slightly thickened and internally lirate; throat white, pillar with a groove near its 
anterior edge; canal short, deep; siphonal fasciole moderate; body with a thin 
white callus. Height, 47; last whorl, 28; diam. 19 mm." 

In size and in the shouldered whorls this species is quite similar to P. articu- 
latus Hinds, but it seems to differ in the details of the sculpture and in the appar- 
ently smooth nuclear whorls. According to Dall, P. cocosensis differs from Phos 
beaui Crosse dC Fischer, of the West Indies, in the greater size and in the strong 
regular ribs. No specimens collected by the Templeton Crocker Expedition or 
the junior author have been referred to the species. 

Phos crassus Hinds 
Plate 22, figure 10 

Phos crassus Hinds, Ann. & Mag. Nat. Hist., new ser., vol. 11, no. 70, April, 
1843, p. 257. "Panama and Gulf of Fonseca; dredged as solitary shells in 
from 3 to 14 fathoms, mud." — Hinds, Zool. Voy. Sulphur, vol. 2, Moll. pt. 
2, October 1844, p. 37, pi. 10, figs.l, 2. Original record cited. — Sowerby, 
Thes. Conch., vol. 3, (pt. 19), 1859, p. 90, pi. 221, fig. 5. Panama.— Tryon, 
Man. Conch., vol. 3, 1881, p. 218, pi. 83, fig. 521. Panama.— Dall, Proc. 
U. S. Nat. Mus., vol. 51, 1917, p. 578. "Lower California and the Gulf of 

Buccinum crassum Hinds, C. B. Adams, Ann. Lyceum Nat. Hist. New York, 
vol.5, 1852, p. 291. Panama. 

The following is a free translation of Hinds' original Latin description: 

"Shell elongate-ovate, subturrited, solid, pale brown; whorls rounded, ribbed; 
ribs strong, rather distant; crossed by prominent cords; outer lip strongly toothed; 
columella with a lamella produced directly and boldly forward." 

To this can be added that there are three smooth nuclear whorls forming a 
rather blunt apex. This is the heaviest of the west coast species, with strong, 
rather coarse sculpture. The single specimen taken by the Templeton Crocker Ex- 
pedition was dredged at Loc. 27567 (C. A. S.) , off Oaxaca, Mexico. A specimen 
in the collection of the junior author was collected at Panama. Dall gave the 
range as Lower California and Gulf of California, but it is doubtful if it occurs 
in the Gulf. 

Phos fusoides (C. B. Adams) 

Triton fusoides C. B. Adams, Ann. Lyceum Nat. Hist. New York, vol. 5, 
1852, p. 340. "Taboga," Panama. — Carpenter, Proc. Zool. Soc. London, 
1863, p. 347.— Tryon, Man. Conch., vol. 3, 1881, p. 220. 

Phos fusoides (C B. Adams), Pilsbry & Lowe, Proc. Acad. Nat. Sci. Phila- 
delphia, vol. 84, p. 116. "Dredged at 20 fathoms at Taboga Island." 

312 San Diego Society of Natural History 

Adams described this species as follows : 

"Shell ovate-fusiform, slender; whitish, stained with brown, with a white 
spiral stripe near the middle of the whorls; with prominent narrow not approxi- 
mate ribs, about nine on each whorl, crossed by numerous raised fine spiral lines, 
of which the alternate ones are mostly larger; apex acute; spire conic; whorls 
eight, convex, with a well impressed suture; aperture long subovate; canal short. 

"Mean divergence about 35°; length .76 inch; breadth .28 inch." 

A single specimen dredged in 20 fathoms at Taboga Island, Panama, by the 
junior author was compared with Adams' single type specimen. The nucleus is 
lost in both and both are dead, rather worn shells. Pilsbry and Lowe stated that it 
"is undoubtedly a Phos, in the same group as P. gaudens Hinds, from which it 
differs in weaker sculpture and greater number of axial ribs." 

Phos gaudens Hinds 
Plate 22, figures 1, 5 

Phos gaudens Hinds, Zool. Voy. Sulphur, vol. 2, Moll. pt. 2, October 1844, 
p. 38, pi. 10, figs. 5 and 6. "Gulf of Tehuantepec, west coast of Mexico. 
Dredged from thirteen fathoms." — Sowerby, Thes. Conch., vol. 3, (pt. 19), 
1859, p. 92, pi. 222, figs. 30, 31. Original record cited.— Tryon, Man. 
Conch., vol. 3, 1881, p. 218, pi. 83, fig. 518; pi. 84, fig. 527. "Gulf of Tehuan- 
tepec, W. Coast of Mexico; W. Columbia." — Tomlin, Jour. Conch., vol. 18, 
no. 6, 1927, p. 160. Dredged off Gorgona Island, Panama.— Pilsbry & Lowe, 
Proc. Acad. Nat. Sci. Philadelphia, vol. 84, 1932, p. 116. "Montijo Bay," 
Panama.— Strong, Hanna & Hertlein, Proc. Calif. Acad. Sci., ser. 4, vol. 
21, no. 10, 1933, p. 119. Acapulco, Guerrero, Mexico. 

Phos mexicanus Dall, Proc. U. S. Nat. Mus., vol. 51, 1917, p. 578. "(Cat. No. 
212111 U. S. N. M.) Ranges from Cape St. Lucas to Panama."— Zetek, 
Rev. Nueva, nos. 1 & 2, 1918, p. 22. Panama.— Woodring, Carnegie Inst. 
Washington Publ. 385, 1928, p. 260 (foot-note) .—Pilsbry & Lowe, Proc. 
Acad. Nat. Sci. Philadelphia, vol. 84, 1932, p. 116. "Dredged at 20 fathoms. 
Guaymas; Mazatlan; Manzanillo; Acapulco." 

Not Phos mexicanus BoSE, Inst. Geol. Mexico, Bol. 22, 1906, p. 38, pi. 4, figs. 
18-21. "Division Tuxtepec, Pliocene— Paso Real cerca de Tuxtepec 
Oax[aca] ." 

Mangilia? dejanira Dall, Proc. U. S. Nat. Mus., vol. 56, 1919, p. 72, pi. 20, fig. 
12. "Dredged in Santa Maria Bay, Lower California; Dr. Paul Bartsch." 

Specimens of this species were dredged by the Templeton Crocker Expedi- 
tion at the following localities: 

Loc. 27588 (C. A. S.), Lat. 24° 14' to 18' N, Long. Ill 28' to 29' W. 
About 13 miles southeast of Cape Tosco, Sta. Margarita Island, off the west 
coast of Lower California, Mexico. 

Loc. 27849 (C A. S.), Lat. 23° 12' N, Long. 106° 29' W. Dredged in 12 
fathoms off Cape San Lucas, Lower California, Mexico. 

Strong & Lowe — The Genus Phos 313 

Loc. 27581 (C. A. S.), between Isabel Island and Mazatlan, Sinaloa, 

Loc. 27580 (C. A. S.), dredged about one-half mile east of Isabel Island, 
between Isabel Island and Mazatlan, Sinaloa, Mexico. 

Loc. 27583 (C. A. S.), Lat. 22° 44' N., Long. 105° 59' W. Dredged in 
10-17 fathoms, about 38 miles southeast of Mazatlan about 8 miles off shore. 

Loc. 27574 (C. A. S.), Lat 18° 33' N., Long. 103° 45' W. Dredged near 
Manzanillo, Colima, Mexico, in 52 fathoms. 

Loc. 27573 (C. A. S.), Lat. 18° 14' N., Long. 103° 23' W. Just off shore 
at Maruata, Mexico, and about VA miles southeast of Pt. Telmo, Mexico, in 60 

Loc. 27571 (C. A. S.), Lat. 16° 39' N., Long. 99° 24' 30" W. to Lat. 16° 
38' N., Long. 99° 27' 30" W. Dredged in 20-45 fathoms, about 33 miles slightly 
east of Acapulco, Guerrero, Mexico, about 32 miles west of Dulce Bay. 

Loc. 27527 (C. A. S.) , Acapulco Bay, Guerrero, Mexico. 

Loc. 27567 (C. A. S.), dredged in the Gulf of Tehuantepec, between Aca- 
pulco and Pt. Angeles, Oaxaca, Mexico. 

The junior author secured specimens of the species at the following local- 
ities : 

San Felipe, east coast of Lower California, Mexico. 

Off Punta Penasco, Sonora, Mexico, in 10 fathoms. 

Concepcion Bay, east coast of Lower California, Mexico, in 15 fathoms. 

Off Guaymas, Sonora, Mexico, in 20 fathoms. 

Off Mazatlan, Sinaloa, Mexico, in 20 fathoms. 

Off Acapulco, Guerrero, Mexico, in 15 fathoms. 

Montijo Bay, Panama, in 15 fathoms. 

The following is a free translation of Hinds' original Latin description : 

"Shell elongate-ovate, pointed, shining, pale, with dark brown bands near 
the suture; whorls rounded, ribbed, ribs about 9 at the periphery, with white 
nodes, interspaces finely striated, toward the base banded; aperture elongate- 

To this can be added that there are four, smooth, glassy nuclear whorls and 
that the outer lip is nearly smooth. Dall compared P. mexicanns with P. articu- 
latus Hinds but the latter has more numerous axial ribs and quite different nuclear 
whorls. In the description of Mangilia dejanira, Dall questioned if it belonged in 
the genus Mangilia as there was no anal fasciole. The type is a young specimen 
of this species. 

Phos minusculus Dall 

Plate 22, figure 4 

Phos minusculus Dall, Proc. U. S. Nat. Mus., vol. 51, 1917, p. 578. "Dredged 

314 San Diego Society of Natural History 

in Panama Bay in 26 to 47 fathoms." — Zetek, Rev. Nueva, nos. 1 SC 2, 
1918, p. 22. Panama. 

Dall's description of this species is as follows: 

"Shell very small and thin, with about six whorls without the nucleus; 
whorls rounded, sutures distinct, with two undulated spiral threads in front of it, 
and in front of these six flattened threads with wider channeled interspaces be- 
tween the sutures on the penultimate whorl; these are not swollen where they 
cross the ribs, of which on the last whorl there are 14, with wider interspaces; 
there are no intercalary spirals; outer lip slightly varicose, with about 10 internal 
lines; labium smooth, with no subsutural callus and no anterior keel on the pillar. 
Length, 12; diameter, 5; length of last whorl, 8 mm." 

No specimens referable to this species were secured by the Templeton 
Crocker Expedition. The junior author dredged four specimens in Concepcion 
Bay, Gulf of California, in 15 fathoms. They were compared with the type from 
Panama and were found to be identical. The nucleus consists of four, smooth, 
glassy whorls. 

According to Dall this is the smallest species of this genus, so far as known. 

Phos veraguensis Hinds 
Plate 22, figures 2, 8, 9 

Phos veraguensis Hinds, Ann. & Mag. Nat. Hist., new ser., vol. 11, April 1843, 
p. 257. "Pueblo Nueva, coast of Veragua; dredged in some numbers from 
26 fathoms, mud."— Hinds, Zool. Voy. Sulphur, vol. 2, Moll. pt. 2, Octo- 
ber 1844, p. 37, pi. 10, figs. 13, 14. Original record cited.— Sowerby, Thes. 
Conch., vol. 3, (pt. 19) , 1859, p. 92, pi. 222, fig. 41. Original record cited.— 
Tryon, Man. Conch., vol. 3, 1881, p. 219, pi. 84, fig. 529. Veragua, W. 
Coast of Central America. [The records "West Indies; Senegal" refer to 
West Indian species.] — Tomlin, Jour. Conch., vol. 18, no. 6, 1927, p. 160. 
Coiba Island, Panama.— Strong, Hanna & Hertlein, Proc. Calif. Acad. 
Sci., ser. 4, vol. 21, no. 10, 1933, p. 119. Acapulco, Guerrero, Mexico. 

Phos biplicatus Carpenter, Proc. Zool. Soc. London, 1856, p. 166. Panama. — 
Tryon, Man. Conch., vol. 3, 1881, p. 220. 

F[usinus]. porticus Dall, Nautilus, vol. 29, no. 5, Sept. 1915, p. 56. "It is an 

inhabitant of Panama." 

Phos alternatus Dall, Proc. U. S. Nat. Mus., vol. 51, 1917, p. 578. "From the 
Gulf of California." (Cat. no. 212110, U. S. N. M.) 

Specimens of this species were dredged by the Templeton Crocker Expedi- 
tion at the following localities: 

Loc. 27585 (C. A. S.) , Lat. 23° 02' N., Long. 100° 32' W. A few miles off 
Gorda Point, in San Jose del Cabo Bay, Lower California, Mexico, in 25 fathoms. 

Loc. 27584 (C. A. S.), Lat. 23° 03' to 23° 06' N., Long. 109° 36' to 

Strong & Lowe — The Genus Phos 315 

109° 31' W. About 10 miles due east of San Jose del Cabo, Lower California, 
Mexico, in 20 to 220 fathoms. 

Loc. 27574 (C. A. S.), Lat. 18° 33' N., Long. 103° 45' W. Near Manza- 
nillo, Colima, Mexico. Just off shore at Black Head (Pta. San Juan de Lima) 
and about 20 miles northwest of Pt. Telmo, in 52 fathoms. 

Loc. 27572 (C. A. S.), a few miles south of Acapulco Bay, Guerrero, 
Mexico, in 15-20 fathoms. 

Loc. 37573 (C. A. S.), Lat. 18° 14' N., Long. 103° 23' W. Just off shore 
near Maruata, Mexico, and about 1% miles southeast of Pt. Telmo, Mexico, in 
60 fathoms. 

Loc. 27571 (C. A. S.), Lat. 16° 39' N., Long. 99° 24' 30" W., to Lat. 
16° 38' N, Long. 99° 27' 30" W. About 33 miles slightly east of Acapulco, 
Guerrero, Mexico, and about 32 miles west of Dulce Bay, in 20-45 fathoms. 

Loc. 27527 (C. A. S.), dredged in Acapulco Bay, Guerrero, Mexico. 

Loc. 27558 (C. A. S.), between Punta Arenas and Bat Island, about 5 to 
6 miles off Delmas, Costa Rica, in 50-60 fathoms. 

The junior author secured specimens of the species at the following locali- 

Punta Libertad, Sonora, Mexico. 

Off Angel de la Guardia Island in the Gulf of California, Mexico, in 20 

Off Carmen Island in the Gulf of California, Mexico, in 20 fathoms. 

Off Acapulco, Guerrero, Mexico, in 20 fathoms. 

The following is a free translation of Hinds' original Latin description: 
"Shell elongate-ovate, cancellated, brown; whorls somewhat rounded, obso- 
letely banded; transversely cancellated, subnodose at the intersections; outer lip 
crenulated; columella smooth or slightly calloused." 

To this can be added that there are three nuclear whorls of which the first 
two are smooth and the last has a very fine spiral thread, also that the columella 
has two sharp basal folds. Specimens have been compared with both Dall's and 
Hinds' types. The latter are said to have been acid-treated and so have lost the 
fine details of sculpture. The large series of specimens examined shows consider- 
able variation in these details. Phos biplicatus Carpenter is unfigured and Tryon 
stated that "the diagnosis applies fairly well to Phos veraguensis Hinds." The 
principal difference in the description seems to be that the basal fold of the 
columella, not mentioned by Hinds, is slightly bifid. In the description of 
Fusinus porticus Dall it is stated that the type may perhaps not prove to be a 
Fusinus. It is undoubtedly a young specimen of this species. 

Powys 6 described a Nassct pallida from Panama. Reeve 7 figured the shell and 
Tomlin 8 stated that it is now recognized as being a Phos. Sowerby 9 also figured 

6 L. W. Powys, Proc. Zool. Soc. London, 1835, p. 96. 

7 L. A. Reeve, Conch. Icon., vol. 8, Nassa, 1853, pi. 9, fig. 60. 

8 J. R. le B. Tomlin, Proc. Mai. Soc. London, vol. 20, pt. 2, 1932, p. 95. 

9 G. B. Sowerby, Thes. Conch., vol. 3, 1859, p. 94, pi. 222, figs. 19-21. 

316 San Diego Society of Natural History 

a shell under this name but gave the locality as the Philippines. Tryon 10 copied 
both figures and gave both localities. He included under it P. notatus Sowerby, 
described from the Philippines. 

Faustino 11 in his catalog of the marine mollusks of the Philippine Islands 
has cited Phos pallidus (Powys) from the Philippine Islands and in the syno- 
nymy of that species he included Phos notatus. It seems probable that the locality 
given by Powys was an error and that it is a Philippine shell. 

Reeve 12 figured a Phos cumingii without description or locality. Sowerby 13 
copied the figure and gave the locality as western Columbia. Tryon 14 included 
the species with others under Phos gaudens Hinds. The figure is poor, but 
resembles some of the west coast species in the genus Strombina more than those 
in the genus Phos. 

The genus Phos is very poorly represented in the literature of the Tertiary 
of Western North America. 15 "Phos?" martini Dickerson, 16 of the Eocene of 
Marysville Buttes, is probably not a Phos. Phos blakianus Anderson & Hanna, 17 
described from the type Tejon Eocene near Grapevine Creek, Kern County, 
California, is said to be an Endopachychilus Cossmann. 18 Phos dumbleana An- 
derson, 19 of the Miocene of Kern River, California, may be a Phos as stated by 
Hanna 20 but "Nassa" chehalisensis Weaver 21 from the Miocene of western 
Washington, considered by Etherington, 22 to be a variety of "Tritiaria (Antillo- 
phos) dumblei," appears closer to Nassarius of the N. perpinguis group than to 
the genus Phos. Phos cocosensis Dall has been recorded as a Pleistocene fossil 
from Albemarle Island of the Galapagos group. 23 

10 G. W. Tryon, Jr., Man. Conch., vol. 3, 1881, p. 218, pi. 83, figs. 494, 496. 

11 L. A. Faustino, Bureau of Science, Manila, Philippine Islands, Monograph 25, 
1928, p. 242. 

12 L. Reeve, Elem. Conch., vol. 1, 1860, p. 67, pi. 3, fig. 16. 

13 G. B. Sowerby, Thes. Conch., vol. 3, 1859, p. 91, pi. 222, fig. 38. 
HG. W. Tryon, Jr., Man. Conch., vol. 3, 1881, p. 218, pi. 83, fig. 519. 

15 Phos mexicanus Bose, Inst. Geol. Mexico, Bol. Nr. 22, 1906, p. 38, pi. 4, figs, 
18-21. This species reported in the Tuxtepec division of the Pliocene of Oaxaca, belongs to 
the Caribbean province, not to the Pacific. 

16 R. E. Dickerson, Univ. Calif. Publ. Bull. Dept. Geol., vol. 7, no. 12, 1913, p. 288, 
pi. 13, fig. 5. 

17 F. M. Anderson and G. D. Hanna, Calif. Acad. Sci., Occas. Papers, vol. 11, 
1925, p. 73, pi. 8, fig. 16, pi. 11, figs. 8, 9. 

18 R. B. Stewart, Proc. Acad. Nat. Sci. Philadelphia, vol. 78, 1927, pp. 302, 391. 

19 Pleurotoma (Clathurella) dumblei Anderson, Proc. Calif. Acad. Sci., ser. 3, (Geol.), 
vol. 2, no. 2, 1905, p. 204, pi. 15, figs. 60, 61; not Harris, 1895. Renamed Phos dumbleana 
by Anderson in Hanna, Proc. Calif. Acad. Sci., ser. 4, vol. 13, no. 10, 1924, p. 183. 

20 G. D. Hanna, Proc. Calif. Acad. Sci., ser. 4, vol. 13, no. 10, 1924, p. 183. 

21 C. E. Weaver, Univ. Wash. Publ. Geol., vol. 1, no. 1, 1916, p. 46, pi. 5, figs. 69, 70. 

22 T. J. Etherington, Univ. Calif. Publ. Bull. Dept. Geol. Sci., vol. 20, no. 5, 1931, 
p. 100, pi. 12, figs. 6,21, 22. 

23 W. H. Dall and W. H. Ochsner, Proc. Calif. Acad. Sci., ser. 4, vol. 17, no. 4, 
1928, p. 96, pi. 2. fig. 14. 



Fig. 1. Phos gaudens Hinds. This is a" figure of the type specimen of Phos 
mexicanus Dall (Proc. U. S. Nat. Mus., vol. 51, 1917, p. 578, Cat. 
No. 212111, U. S. N. M.), from U. S. Bureau of Fisheries Station 
3034, off Point Fermin, Lower California, Mexico, dredged in 24 
fathoms. Length 23 mm., diameter 8.5 mm., length of last whorl 13 
mm. This is not Phos mexicanus Bose. (p- 312) 

Fig. 2. Phos veraguensis Hinds. This is a figure of the type specimen of Phos 
alternatus Dall (Proc. U. S. Nat. Mus., vol. 51, 1917, p. 578, Cat. 
No. 212110, U. S. N. M.), from U. S. Bureau of Fisheries Station 
3037, off Guaymas, Sonora, Mexico, dredged in 20 fathoms. Length 
26 mm., width 12 mm. (p- 314) 

Fig. 3. Phos chelonia Dall. This is a figure of the type specimen of Phos 
cheloma Dall (Proc. U. S. Nat. Mus., vol. 51, 1917, p. 578, Cat. 
No. 194961, U. S. N. M.), from U. S. Bureau of Fisheries Station 
2813, Galapagos Islands, in 40 fathoms. (p-310) 

Fig. 4. Phos minusculus Dall. (Proc. U. S. Nat. Mus., vol. 51, 1917, p. 578, 
Cat. No. 122775, U. S. N. M.) , from U. S. Bureau of Fisheries Sta- 
tion 2804, dredged in Panama Bay, in 47 fathoms. Length 12 mm., 
diameter 5 mm., length of last whorl 8 mm. (p. 313) 

Fig. 5. Phos gaudens Hinds. Plesiotype, No. 6997 (Calif. Acad. Sci. Paleo. 
type coll.), from Loc. 27581 (C. A. S.), dredged between Santa 
Isabel Island and Mazatlan, Sinaloa, Mexico; Templeton Crocker 
Expedition. Length 24.5 mm., diameter 9 mm., length of last whorl, 
approximately 10.4 mm. (p. 312) 

Fig. 6. Phos artiadatus Hinds. Plesiotype, No. 6998 (Calif. Acad. Sci. Paleo. 
type coll.) , from Loc. 27585 (C. A. S.) , Lat. 23° 02' N, Long. 109° 
32' W., dredged in 25 fathoms, a few miles off Gorda Point in San 
Jose del Cabo Bay, Lower California, Mexico; Templeton Crocker 
Expedition. Length 46.1 mm., diameter 18.9 mm., length of last 
whorl, 21 mm. (p. 309) 

Fig. 7. Phos cocosensis Dall. A reproduction of the figure of the type given by 
Dall (Bull. Mus. Comp. Zool., vol. 43, no. 6, 1908, pi. 8, fig. 5), 
from U. S. S. "Albatross," station 3368, near Cocos Island, Gulf of 
Panama, in 66 fathoms. Length 47 mm., diameter 19 mm., length of 
last whorl, 28 mm. (p-310) 

Fig 8. Phos veraguensis Hinds. Plesiotype, No. 6999 (Calif. Acad. Sci. Paleo. 
type coll.) , from Loc. 27584 (C. A. S.) , Lat. 23° 03' to 23° 06' N, 
Long. 109° 36' to 109° 31' W., in 20 to 220 fathoms. About 10 
miles due east of San Jose del Cabo, Lower California, Mexico; 
Templeton Crocker Expedition. Length 24.8 mm., diameter 11.7 
mm., length of last whorl 13 mm. (p. 314) 

Fig. 9. Phos veraguensis Hinds. Plesiotype, No. 7000 (Calif. Acad. Sci. 
Paleo. type coll.), from the same locality as the specimen shown in 
figure 8. Length 28.3 mm., diameter 12.8 mm., length of last whorl 
15.3 mm. This figure shows an axial section of the interior of the shell 
and reveals the presence of two small plates on the columella, (p. 3 14) 
Fig. 10. Phos crassus Hinds. Plesiotype, No. 7001 (Calif. Acad. Sci. Paleo. 
type coll.), from Loc. 27567 (G A. S.), dredged off the coast of 
Oaxaca, Mexico, in the Gulf of Tehuantepec; Templeton Crocker 
Expedition. Length 41 mm., diameter 19 mm., length of last whorl 
21mm. (p- 311) 

Strong & Lowe — The Genus Phos 

Plate 22 




Volume VIII, No. 23, pp. 321-336 





Dickey Collections, California Institute of Technology 


Printed for the Society 

June 15, 1937 



U. S. Grant, IV, Chairman 

Fred Baker Clinton G. Abbott, Editor 






Dickey Collections, California Institute of Technology 

Some years ago (Trans. San Diego Soc. Nat. Hist., 6, No. 19, 
April 30, 1931), van Rossem published a list of Sonora land birds based 
in part on specimens in the Dickey collection, in part on specimens in 
the collection of the San Diego Society of Natural History, and in part 
on still others from private collections. A further report, particularly re- 
lating to water birds, is now desirable. 

In the interim since 1931, the San Diego Society of Natural History 
has acquired from several sources a considerable amount of Sonora ma- 
terial, including a large donation of Sonora bird skins from Mr. Griffing 
Bancroft. The San Diego Society of Natural History specimens recorded 
in these notes were in almost every instance taken by Mr. Bancroft or his 
assistants. Additional material (either as specimens or notes) used in 
the preparation of this paper has also very kindly been placed at our dis- 
posal by Mr. A. W. Anthony, Dr. Louis B. Bishop, Mrs. Donald Dickey, 
Mr. Chester Lamb, Mr. J. T. Wright, the Museum of Vertebrate 
Zoology, the Museum of Comparative Zoology and the University of 

Specimens are in the Dickey collection unless otherwise noted. 
Observations, unless otherwise credited, are by van Rossem. 

Gavia arctica pacifica (Lawrence) 

The Pacific Loon was noted as a common spring migrant off San Esteban 
Island between April 17 and 20, 1930. 

Colymbus dominicus bangsi subsp. nov. 

Type. — Female adult, no. 218269, Museum of Comparative Zoology; 
Santiago, Lower California, Mexico, November 15, 1887; collected by M. Abbott 

Subspecific characters. — Resembles Colymbus dominicus brachypterus Chap- 
man of southern Texas and Middle America, but, sex for sex, bills definitely 
smaller; upper parts (including pileum) slightly grayer and paler; breeding 
plumage darker below, with spotting more prominent. 

Range. — Arid Tropical Zone of Lower California, southern Sonora, and 

324 San Diego Society of Natural History 

probably other portions of northwestern Mexico. 

Remarks. — Nearly ten years ago van Rossem noticed the small bills and 
dark summer coloration of the Lower California least grebes. At the time he 
talked the matter over with the late Outram Bangs, but there were certain points 
of disagreement and van Rossem naturally deferred to Bangs. Later Bangs wrote 
that he was convinced, after a careful restudy of the question, that the Lower 
California colony belonged to a perfectly distinct race. Chester Lamb (in 1933) 
collected a specimen at Agiabampo on the coast of extreme southern Sonora, 
and very kindly allowed van Rossem to examine it. Since the bird was in breed- 
ing plumage it would seem, taken in connection with the date (April 21), that 
the race bangsi is also a breeding bird of Sonora. 

On various occasions, ornithologists have expressed curiosity concerning 
the systematic status of the least grebes of Central America. With a fair amount 
of material, it may be stated that such El Salvador specimens as van Rossem 
has seen belong apparently with brachypterus. This is shown by the following 



Exposed Culmen 

6 $ from Texas 


• 93 



5 $ from El Salvador 




■ 24.1 


8 $ from Lower Calif. 




■ 21.2 


3 $ from Texas 


■ 92 



6 9 from El Salvador 





(2 immature) 


6 9 from Lower Calif. 




■ 18.8 

Colymbus nigricollis californicus (Heermann) 

Observed as follows: San Esteban Island, April 17, 1930, common; Tiburon 
Island, April 19, 1925, abundant; San Pedro Martir Island, April 20, 1930, 
common; San Pedro Nolasco Island, April 21, 1930, common; Guaymas Har- 
bor, April 21 to 25, 1930, common. 

At these dates most of the birds were in breeding plumage and were thus 
readily identifiable. Large rafts of scores, or even hundreds, were frequently 
observed on the open sea. 

Aechmophorus occidentalis (Lawrence) 

Price, under circumstances none too certain, has previously recorded the 
Western Grebe from the mouth of the Colorado River in winter. In the Thayer 
collection at the Museum of Comparative Zoology, is a specimen taken by W. W. 
Brown, Jr., at "Precidio" | = Guaymas] on March 27, 1905. 

Puffinus griseus (Gmelin) 

Two individuals were seen (unmistakably) 5 miles west of Tiburon Island 
on April 25, 1925. 


Oceanodroma melania melania (Bonaparte) 

Seen in considerable numbers off San Esteban Island on April 21, and off 
the entrance to Guaymas Harbor on April 27, 1930. 

Halocyptena microsoma Coues 

Observed as extremely common off San Esteban Island, April 17, 1930; 
off Tiburon Island, April 19, 1925; off Guaymas Harbor, April 21, 1930. At 
these dates it was obvious that both this species and melania were at the height 
of their migrations. Great rafts of up to several hundred individuals were fre- 
quently seen sitting on the open water. 

Phaethon aethereus mesonauta Peters 

The Red-billed Tropic-bird has been recorded at practically all times of the 
year from many Gulf points. However, it was interesting to observe it at the 
mouth of the Colorado River on April 25, 1925. 

Although there seems to be in some quarters a reluctance to recognize 
mesonauta, it appears to be a good race — at least so far as one can judge by the 
characters shown by Gulf specimens. 

Pelecanus erythrorhynchos Gmelin 

Ten individuals, none of which showed the bill knob which is indicative of 
breeding activity, were seen on an estero in Guaymas Harbor on April 22, 1930. 

Phalacrocorax auritus albociliatus Ridgway 

In addition to published data (Tiburon Island; Punta Penascosa; mouth of 
the Colorado) , there are additional records to show the Farallon Cormorant to be 
a common species in the Gulf, although there appear to be no actual breeding 
records. Lamb (MS) noted it as common at Guaymas on January 3, 1933; 
Brown (Mus. Comp. Zool.) took a specimen there on March 23, 1905; and van 
Rossem found it commonly at San Pedro Martir Island on April 18, 1925. 

Phalacrocorax penicillatus (Brandt) 

Brandt's Cormorant was noted as follows: San Pedro Martir Island, April 
18, 1925, where nesting commonly; San Esteban Island, April 17, 1930, abun- 
dant off shore; San Pedro Nolasco Island, April 21, 1930; off Guaymas, April 
30, 1930, common. 

Fregata magnificens rothschildi Mathews 

A specimen from Guaymas, June 14, 1930, and another from Lobos Island, 
May 28, 1930 (S. D. S. N. H.), are certainly of the species magnificens. We 
follow Swarth in recognizing the race rothschildi, but doubt that it will bear 
investigation when adequate material is assembled. 

Ardea herodias treganzai Court 
The Great Basin race of the great blue heron has been reported frequently 

326 San Diego Society of Natural History 

as occurring south to Guaymas, but actual specimens to verify these ascriptions 
have been conspicuous by their absence. Additional sight records are: Guaymas, 
January 3 and 6, 1933 (Lamb); and San Pedro Nolasco Island, April 21, 1930. 
Specimens (S. D. S. N. H.) taken six miles north of Guaymas, June 15, 1928, 
and at Tobari Bay, April 28, 1930, are definitely of the race treganzai. Both 
were breeding birds. 

Butorides virescens anthonyi (Mearns) 

J. T. Wright took a migrant anthonyi at Tecoripa on March 1, 1929, and a 
juvenile, still in partial down, at Saric on August 15, 1929. Pierce Brodkorb 
informs me that in the University of Michigan collections is a skin of this race 
collected by Berry Campbell at Pilares in northeastern Sonora on July 27, 1935. 
These records, combined with others previously published, appear to establish 
anthonyi as rather common in northern Sonora. 

Butorides virescens virescens (Linnaeus) 

The most careful scrutiny fails to reveal any characters by which to dis- 
tinguish the breeding green herons of the Arid Tropical Zone of Sonora from 
virescens of the eastern United States. Oberholser (Proc. U. S. Nat. Mus., 42, 
1912, 533) has shown that virescens occurs north on the Pacific coast of Mexico 
as far as Mazatlan. The presence of this race in southern Sonora is, while inter- 
esting, not particularly remarkable, save that it would seem to indicate that 
frazari of Lower California is of southern, rather than of northern, origin. 

Specimens of virescens, all of which were breeding when collected, have been 
examined from Tobari Bay (2), April 27 and June 12, 1930; Lobos Island, (1), 
May 25, 1930; Kino Bay, (1), May 16, 1930. All except the first listed are in 
the collection of the San Diego Society of Natural History. 

Florida caerulea caerulescens (Latham) 

Six specimens in the collection of the San Diego Society of Natural History 
are the dark colored tropical race. They were collected at Lobos Island (2) , May 
30, 1930; and at T6bari Bay (4) , June 12, 1930. 

Dichromanassa rufescens dickeyi van Rossem 

Lamb noted one at Guaymas on January 3, 1933; the Museum of Compara- 
tive Zoology contains a specimen taken by Brown in the same locality on March 
19, 1905; and in the collection of the San Diego Society of Natural History are 
two taken at Tobari Bay on April 28, 1930. These three examples are typical 

Casmerodius albus egretta (Gmelin) 

Records additional to those already published are that American Egrets 
were noted as common at Guaymas in late April and early May, 1930; and as 
breeding commonly at Tobari Bay from April 26 to May 1, 1930. 

Leucophoyx thuia brewsteri (Thayer and Bangs) 
Lamb noted a Snowy Egret at Guaymas on January 3, 1933. A skin from 


Lobos Island (S. D. S. N. H.) taken May 25, 1930, is just about intermediate 
in characters between brewsteri and thula. In the absence of further material this 
bird is referred to brewsteri, in keeping with the determinations of Thayer and 
Bangs, Bent, and others who have examined Sonora specimens. 

Hydranassa tricolor ruficollis (Gosse) 

Two specimens, collected at Tobari Bay on April 27 and 29, 1930, are in 
the collection of the San Diego Society of Natural History. 

Nycticorax nycticorax hoactli (Gmelin) 

Black-crowned Night Herons were noted by Lamb at Guaymas on January 
6, 1933; and the collection of the San Diego Society of Natural History con- 
tains a (presumably) breeding specimen taken in that locality on June 22, 1928. 
The species was noted as common at Tobari Bay between April 26 and May 1 
1930. 7 

Nyctanassa violacea bancrofti Huey 

A specimen (S. D. S. N. H.) taken at Tobari Bay on April 28, 1930, seems 
to be exactly like Lower California examples of this race. 

Heterocnus cabanisi (Heine) 

The Dickey collection contains a specimen taken by Wright at Tesia on 
March 17, 1930. Wright states that the Tiger Bittern is fairly common and 
breeds in the Mayo River Valley. Van Rossem found no trace of it anywhere 
along the coast. The single skin seems to be identical with El Salvador specimens 
of similar age. 

Botaurus lentiginosus (Montagu) 

Wright took a Bittern at El Doctor on January 25, 1929. 
Mycteria americana Linnaeus 

The Wood Ibis was found to be commonly and generally distributed all 
along the coast from Guaymas southward to Tobari Bay in April and early 
May, 1930. In the collection of the San Diego Society of Natural History is a 
specimen taken six miles north of Guaymas on June 15, 1928. 

Guara alba (Linnaeus) 

Three specimens in the collection of the San Diego Society of Natural His- 
tory were taken, respectively, at Tobari Bay on April 28 and June 12, 1930, and 
at Guasimas Lagoon on May 12, 1930. The White Ibis was noted as common 
in the first named locality from April 26 to May 1, 1930. 

Ajaia ajaja (Linnaeus) 

Wright states that Spoonbills are common in the Mayo River Valley. Van 
Rossem found them common at Tobari Bay between April 26 and May 1, 1930, 
and saw several at Guaymas on May 5 of the same year. Three specimens taken, 

328 San Diego Society of Natural History 

respectively, at Tobari Bay, April 28 and May 1, and at Kino Bay, May 16, 1930, 
(S. D. S. N. H.) are all in breeding plumage. However, the largest ova in the 
Tobari Bay females were only about 3 mm. in diameter and the birds were 
obviously not nearly ready to breed. 

Chen hyperborea (Pallas) 

Donald Dickey collected two specimens at San Luis on December 9, 1925, 
and Wright took another at El Doctor on February 7, 1929. Both localities are 
in the Colorado River Delta. 

Dendrocygna bicolor helva Wetmore and Peters 

A flock of at least 100 Fulvous Tree-ducks was seen on an exposed tide 
flat in Guaymas Harbor on May 5, 1930. 

Querquedula cyanoptera (Vieillot) 

A small flock of seven or eight Cinnamon Teal was seen just south of 
Nogales on December 21, 1931. The Dickey collection contains two specimens 
taken by Wright at El Doctor, January 27, and Saric, September 14, 1929. 

Nettion crecca carolinense (Gmelin) 

El Doctor (January 20 and 24, 1929); Saric (August 17, 1929); and 
Chinobampo (February 8, 1930) are localities and dates which supplement the 
few previously published records. 

Dafila acuta tzitzihoa (Vieillot) 

Two specimens were taken by Wright at El Doctor on January 21, 1929. 
There are only two previously published records, both from the northern part of 
the State. The apparent scarcity of the Pintail in Sonora is surprising. Van 
Rossem did not meet it anywhere along the coast in the winter of 1931-32, 
although ducks of several species were abundant there. 

Mareca americana (Gmelin) 

Lamb noted the Baldpate at San Jose de Guaymas on January 6, 1933. The 
Museum of Comparative Zoology contains two specimens taken by Brown at 
Aranjuez [ = San Jose de Guaymas] on March 22, 1905. 

Spatula clypeata (Linnaeus) 

In the Museum of Comparative Zoology is a specimen taken by Brown at 
"Precidio" = Guaymas] on March 25, 1905. Wright found the Shoveller to 
be a fairly common winter visitant at El Doctor in January, 1929, and took a 
specimen on January 27. 

Nyroca americana (Eyton) 

A specimen in the Museum of Comparative Zoology, taken by Brown at 
Guaymas on March 28, 1905, verifies certain sight records from the same locality. 


Nyroca affinis (Eyton) 

The Lesser Scaup is probably the commonest salt-water duck in the State. 
Lamb noted it at Guaymas on January 3, 1933, and at San Jose de Guaymas on 
January 6; Huey collected one at Kino Bay, February 23, 1935; and Brown 
(M. C. Z.) took two at Aranjuez on March 22, 1905. Many lesser scaups were 
noted at Guaymas between April 21 and 25, 1930, but it is believed that most, 
if not all of these birds were non-breeding individuals which would not have 
gone north. 

Charitonetta albeola (Linnaeus) 

One specimen, an adult male, was taken by Wright at Ciudad Obregon on 
November 17, 1930. 

Melanitta perspicillata (Linnaeus) 

Surf Scoters were seen in small numbers off San Esteban Island from April 
17 to 19, 1930. 

Erismatura jamaicensis rubida (Wilson) 

A specimen in the Museum of Comparative Zoology, collected by Brown at 
Guaymas on March 10, 1905, is apparently the first and only authentic State 
record of the Ruddy Duck. The "Pachico, Sonora" record in the Biologia Cen- 
trali Americana is really a Chihuahua citation and dates back to Allen's ambigu- 
ously worded report on the specimens collected by the Lumholtz expedition. 

Mergus merganser americanus Cassin 

Lamb noted the American Merganser at San Jose de Guaymas on January 
9, 1933. Van Rossem found it to be fairly common about San Esteban Island on 
April 17, 1930, and also noted a pair at Guaymas on April 22, 1930. 

Mergus serrator Linnaeus 

This is a very common coastwise winter duck, and there are numerous locali- 
ties on record. The latest seasonal dates noted are two "Precidio" = Guaymas] 
specimens in the Museum of Comparative Zoology, taken by Brown on March 
20 and 21, 1905. 

Porzana Carolina (Linnaeus) 

Wright took three specimens at El Doctor on January 24, 27, and 31, 1929. 

Fulica americana americana Gmelin 

Lamb noted Coots at San Jose de Guaymas on January 19, 1933; Wright 
took three specimens at El Doctor on January 21 and 24, 1929; Brown (M. C. 
Z.) collected one at "Precidio" [= Guaymas] on March 21, 1905. 

^ Squatarola squatarola (Linnaeus) 
The Black-bellied Plover was noted as a fairly common migrant at Tobari 

330 San Diego Society of Natural History 

Bay from April 26 to May 1, 1930. At this time most of the individuals ob- 
served were in breeding plumage. 

\' Charadrius hiaticula semipalmatus Bonaparte 
A common migrating species at Tobari Bay from April 26 to May 1, 1930. 

v Charadrius wilsonia beldingi (Ridgway) 

Lamb collected a specimen at Agiabampo on April 18, 1933. As previously 
recorded by many authors, Belding's Plover is a common permanent resident the 
entire length of the Sonora coast. 

v Numenius hudsonicus Latham 

A specimen taken by Brown (M. C. Z.) at "Precidio" [— Guaymas] on 
March 25, 1905, is apparently the only individual to have been taken, to date, in 
Sonora. The species was observed as migrating commonly at Tobari Bay from 
April 26 to May 1, 1930. 

v Numenius americanus americanus Bechstein 

The subspecific status of the long-billed curlews which were noted as 
common at Tobari Bay from April 26 to May 1, 1930, is in doubt. However, 
the single example of the species taken to date (van Rossem, 1933) is of the 
nominate race. 

J Limosa fedoa (Linnaeus) 

Some of the flocks which were noted at Tobari Bay between April 26 and 
May 1, 1930, totaled at least 200 individuals. At this time all, or nearly all, of 
the godwits seen were in breeding plumage. Two specimens were collected on 
April 28 and 29, respectively. 

^ Tringa flavipes (Gmelin) 

Two specimens were collected by Wright at Ciudad Obregon on Novem- 
ber 14, 1929. 

Tringa melanoleuca (Gmelin) 

Brown (M. C. Z.) took three at Guaymas on February 27, 1905; Lamb 
noted one at Hermosillo on December 22, 1932; and Wright collecetd two at 
Ciudad Obregon on November 14, 1929. 

*J Tringa solitaria cinnamomea (Brewster) 

Two specimens taken by Wright at Saric on September 14, 1929, are 
typical of the western race of the solitary sandpiper. 

"0 Actitis macularia (Linnaeus) 
The University of Michigan possesses a specimen taken by Campbell at 


Pilares, July 14, 1935; Wright took two at Saric on August 2 and September 
18, 1929. Van Rossem noted Spotted Sandpipers as common all along the central 
and southern coast and off-shore islands from April 17 to May 14, 1930. Some 
specific localities are: San Esteban Island, San Pedro Martir Island, San Pedro 
Nolasco Island, Tobari Bay, and Guaymas. 

Catoptrophorus semipalmatus inornatus (Brewster) 

The Western Willet, as previously has been noted, is seasonally abundant 
along the length of the Sonora coast. Additional records are: El Doctor from 
January 21 to 29 (4 specimens), 1929; Guaymas [ "Precidio" ] , March 27 and 
29, 1905; T6bari Bay, April 20, 1930; and San Esteban Island (noted), April 
17, 1930. 

v Heteroscelus incanus (Gmelin) 

On April 21, 1930 a single Wandering Tattler was seen in company with 
two spotted sandpipers on the rocky shores of San Pedro Nolasco Island. 


Arenaria interpres morinella (Linnaeus) 

Turnstones are evidently regular spring migrants along the Sonora coast, 
as the following records indicate: one Lamb collection, Agiabampo, April 18, 
1933; one seen at San Pedro Nolasco Island, April 21, 1930; two taken at Tobari 
Bay, April 29 and May 1, 1930. In the last named locality they were fairly com- 
mon and occurred as singles, pairs, and trios, usually in company with other 
shore birds. 

V Arenaria melanocephala (Vigors) 

Two Black Turnstones were seen at San Esteban Island on April 17, and 
another pair was noted on the rocky shores of San Pedro Nolasco Island on 
April 21, 1930. 

Limnodromus griseus scolopaceus (Say) 

Wright took seven at Ciudad Obregon on November 13, 1929; the species 
was noted as an abundant migrant at Tobari Bay from April 26 to May 1, 
1930; and in the collection of the San Diego Society of Natural History are 
two (probably non-breeding) specimens taken at Tobari Bay on June 27, 1930. 

Capella gallinago delicata (Ord) 

Wright took two at El Doctor on January 21 and 24, respectively, and 
another at Tecoripa on March 25, 1929. In the collection of the San Diego 
Society of Natural History are two taken 15 miles south of Nogales on February 
5, 1929. 

^ Calidris canutus rufus (Wilson) 

This was the most abundant shore bird present at Tobari Bay from April 
26 to May 1, 1930. A single specimen was taken on April 28. 

332 San Diego Society of Natural History 

Crocethia alba (Pallas) 

The Sanderling was found to be an abundant migrant at Tobari Bay from 
April 26 to May 1, 1930. 

** Ereunetes mauri Cabanis 

The M. C. Z. collection contains a specimen collected by Brown at Guay- 
mas on March 26, 1905. Western Sandpipers were abundant from Guaymas to 
Tobari Bay from April 22 to May 1, 1930. One flock at the last named locality 
contained at least 500 birds. 

^ Erolia minutilla (Vieillot) 

Wright took eight specimens at El Doctor on January 27, 1929, and in the 
M. C. Z. are two collected at Guaymas on February 22 and 23, 1905. The Least 
Sandpiper was noted as abundant from Guaymas to Tobari Bay from April 
22 to May 14, 1930. 

^ Erolia alpina sakhalina (Vieillot) 

The Red-backed Sandpiper was an abundant migrant at Tobari Bay be- 
tween April 26 and May 1, 1930. One specimen was collected on April 28. 

* Himantopus himantopus mexicanus (Miiller) 

Wright took five specimens at Ciudad Obregon on November 14, 1929. 
At least a dozen Stilts was observed in one flock at Tobari Bay on April 29, 

v Recurvirostra americana Gmelin 

Brown (M. C. Z.) took an Avocet at Guaymas on March 22 and another 
at Aranjuez on March 28, 1905; Wright collected one at Ciudad Obregon on 
November 14, 1929; and van Rossem saw a small flock of four at Tobari Bay 
on April 29, 1930. 

* Steganopus tricolor Vieillot 

Two birds were collected by Wright at Saric on September 10, 1929. 
Numerous flocks and individuals were seen in migration off Tiburon Island on 
April 19, 1925. 

V Lobipes lobatus (Linnaeus) 

Wright collected one specimen at Saric on September 10, 1929, and this is 
apparently the only inland record for the State. Northern Phalaropes were seen 
at sea in immense numbers off San Esteban Island on April 17, and off Tiburon 
Island on April 19, 1930. 

s. Larus delawarensis Ord 

The Ring-billed Gull was noted by Lamb at Guaymas on January 3, 1933; 
Brown (M. C. Z.) took a specimen in the same locality on March 23, 1905; van 
Rossem noted the species as common there between April 22 and May 14, 1930. 


Larus californicus Lawrence 

Noted as common off San Esteban Island, April 17 to 19, 1930, where ob- 
served drifting in large flocks in a northerly direction. California Gulls were also 
found to be common at Guaymas on various dates between April 21 and May 
14, and at Tobari Bay from April 26 to May 1, 1930. 

V Larus atricilla Linnaeus 

Laughing Gulls in breeding plumage were commonly seen by van Rossem 
at Guaymas from April 22 to May 14, 1930, at which latter date he left the 
locality. At Tobari Bay, from April 26 to May 1, 1930, they were even more com- 
mon and, more often than otherwise, were in pairs. Two specimens collected at 
Tobari Bay were in breeding condition, and the actions of many pairs observed 
strongly indicated that they were either breeding or about to do so. The number 
of laughing gulls in spring and early summer on the coast of central and southern 
Sonora makes more understandable the occurrence of occasional breeding pairs 
on Salton Sea in southern California. 

^ Larus Philadelphia (Ord) 

Brown (M. C. Z.) took specimens at Guaymas ["Precidio"] on March 
23 and 29, 1905; Lamb noted the species in the same locality on January 22, 
1933, and van Rossem also observed it there in considerable numbers from April 
22 to May 4, 1930. In the last instance, individuals were noted in every stage 
of plumage from one year old immatures to fully plumaged adults. 

* Gelochelidon nilotica vanrossemi Bancroft 

Gull-billed Terns were noted as not uncommon at Tobari Bay from April 
26 to May 1, 1930. They were usually in pairs and it was the distinct impression 
that they were preparing to breed on one or more sand islands at the entrance to 
the bay. Unfortunately no specimens were taken, but there can be little doubt 
that they were of the same race as that which breeds on Salton Sea in southern 

v Hydroprogne caspia imperator (Coues) 

One or more Caspian Terns were seen daily at Guaymas from April 21 to 
23, 1930. 

v Sterna forsteri Nuttall 

Forster's Terns were seen at the mouth of the Colorado River on April 23, 
1925; at Guaymas (common) on April 21 and May 4, 1930; and at Tobari Bay 
from April 26 to May 1, 1930. A specimen was collected at Guasimas Lagoon on 
May 12, 1930. In no instance was there any evidence of breeding, and the 
species was evidently purely migratory. 

Sterna albif rons mexicanus subsp. nov. 
Type. — Breeding male adult, no. 30,285, Dickey collection; Tobari Bay,. 

334 San Diego Society of Natural History 

Sonora, Mexico, April 29, 1930; collected by A. J. van Rossem; original number 

Subspecifc characters. — Similar to Sterna albifrons browni Mearns of 
southern California and northern Lower California, but size definitely smaller; 
coloration darker throughout and with the underparts even more strongly 
suffused with pearl gray. 

Range. — Coast of southern Sonora (Arid Tropical Zone) from Guaymas 
south to Sinaloa, and probably south along the west coast of Mexico for some 

Remarks. — The least terns of the Sonora coast were immediately recog- 
nized in the field as distinct from the familiar browni of southern California. 
At both Tobari Bay and Guaymas they were either breeding or preparing to do 
so, but were decidedly rare and only seven specimens could be collected. 

Regarding the distribution of mexicanus, one hesitates to suggest that all 
breeding least terns to the southward will be found to belong to that race. 
However, it would seem most improbable that browni, as a breeding race, reap- 
pears to the southward of mexicanus. 
Measurements. — 

Wing Exposed culmen 

167 - 177 26.7 - 28.3 

162 - 164 24.6 - 26.0 

170-174 26.1-27.0 

160 - 165 24.9 - 25.2 

v Thalasseus maximus maximus (Boddaert) 

Anthony noted the Royal Tern at Estrada de Tasiola on December 4, 1930. 
Additional records are: mouth of the Colorado River, April 22 and 23, 1925; 
Guaymas, April 21 to May 14, 1930; T6bari Bay, April 26 to May 1, 1930; 
Lobos Island (S. D. S. N. H.), May 25, 1930. Previously published data show 
this tern to be a fairly common breeder from George Island southward. 

V^ Rynchops nigra oblita Griscom 

The nine Skimmers collected at Algodones Lagoon (May 1, 1930), and 
Guasimas Lagoon (May 12, 1930) seem satisfactorily to belong to this race 
which was described by Ludlow Griscom (Ibis, 1935, p. 545) on the basis of six 
winter specimens from the Pacific coast of Guatemala. There are slight differ- 
ences to be noted between the Sonora birds and the published description, but 
these are probably due to season. Compared with typical nigra, these specimens 
are tinged with pale gray on the axillars and under wing-coverts, and also have 
more gray on the inner webs of the lateral rectrices. 

Although no nests were found it was perfectly evident that the breeding 
season was at hand. 















Vol. VIII, No. 24, pp. 337-348 June 15, 1937 





Robert W. Webb 

California Institute of Technology and 
University of California at Los Angeles 


The study of the fossils of the Point Loma Terrace deposits was 
undertaken primarily to determine the ecology of the fauna. The speci- 
mens were examined, identified, and their identifications checked by spe- 
cialists in Pleistocene Marine Paleontology. The work was done by the 
author while a graduate student in the Division of Geological Sciences of 
the California Institute of Technology. 

Location of the Area 

The material for study was obtained from the lowermost marine ter- 
race on the west side of Point Loma, a promontory of considerable length 
which extends into the Pacific Ocean in a north-south direction, protect- 
ing San Diego Harbor on the west. The entire southern half of the penin- 
sula is a government military and naval reservation; the northern and 
broader half is largely a residential district of the city of San Diego. 

Collections and Method of Study 

The material for study was collected by Dr. W. P. Popenoe, Cura- 
tor in Invertebrate Paleontology, and Mr. David Scharf, graduate 

* Contribution No. 216, Balch Graduate School of the Geological Sciences, California 
Institute of Technology, Pasadena, California. 

338 San Diego Society of Natural History 

student, of the California Institute of Technology, on a collecting trip 
in 1930. A later trip for additional collecting was made in the winter of 
1935-1936 by Dr. Popenoe and the writer. 

The fossils were identified by the author, largely according to the 
nomenclature of Grant and Gale. 1 The identifications were checked by 
Dr. Popenoe, under whose supervision the work was conducted. Dr. U. S. 
Grant, Associate Professor of Geology, University of California at Los 
Angeles, checked the fossil list, edited the manuscript, and offered help- 
ful suggestions. Mr. A. M. Strong, of Los Angeles, aided in the identifi- 
cation of the small gastropods; in addition he contributed, from his per- 
sonal experience, data on the ecology of the forms. 

Review of Pertinent Literature 

There are few papers on the geology of the Point Loma area. No 
thorough geologic study has ever been published, although Messrs. U. S. 
Grant and L. G. Hertlein are now preparing a systematic study of the 
region. The only geologic report of significance is that of Ellis and Lee, 
which contains a generalized geological map of Point Loma, indicating the 
distribution of the marine terrace deposits and their relations to under- 
lying materials. No description of them is undertaken, nor are any paleon- 
tological data recorded. 

A paper by Berry 3 includes a list of fossils from the "Coal Mine," 
on the west side of Point Loma, which contains thirty-one species. These 
"Coal Mine" fossils are identical in age with those listed in the present 

Stephens 4 briefly discusses the occurrence of fossils on the Plei- 
stocene terraces, and gives a small faunal list. 

Geologic Setting of the Terrace 
The terrace material from which the fauna was collected lies uncon- 

1 Grant, U. S., and Gale, H. R. Catalogue of the Marine Pliocene and Pleistocene 
Mollusca of California and Adjacent Regions. Memoirs, San Diego Soc. Nat. Hist., Vol. 1, 
pp. 1-1036, 1931. 

2 Ellis, Arthur L., and Lee, Charles H. The Geology and Ground Waters of the 
Western Part of San Diego County, California. U. S. Geol. Surv. Water Supply Paper 446, 
pp. 1-321, 1919. 

3 Berry, S. Stillman. Fossil Chitons of Western North America. Calif. Acad. Sci. 
Proc, 4th ser., Vol. 1 1 , pp. 399-536, 1922. 

4 Stephens, Frank. Notes on the Pleistocene Deposits of San Diego County, Califor- 
nia. Trans. San Diego Soc. Nat. Hist., Vol. 5, pp. 245-256, 1929. 

Webb — Pleistocene of Point Loma 339 

formably on sediments of Cretaceous age, called "Chico" by Ellis and 
Lee. 5 These Cretaceous rocks are mostly very fine-grained cherty and 
marly shales, rather carbonaceous in places. A small fauna has been re- 
ported from them by Fairbanks. 6 Associated with the terrace deposits are 
gravels of continental origin, which mostly overlie, but occasionally inter- 
finger with the terrace sediments. 

The terrace materials themselves are moderately fine-grained brown- 
ish to yellowish sands, with some conglomeratic material and much marl. 
The best collecting was near the base of the terrace directly overlying the 
Cretaceous shale. Here the fossils were well-preserved in marls that sur- 
round huge boulders of the Cretaceous shale. These are products of marine 
erosion on an old shoreline along which the terrace materials were de- 

The maximum height of the terrace above sea level is about 100 feet 
(to the north) and the minimum twenty-five feet (to the south) . 

Relation to Other Terraces 

The position of the Point Loma terrace indicates that its uplift was 
produced by the last diastrophic movement of the southern coast, since 
it is the lowest terrace exposed, and has, in part, at least, been destroyed 
by marine processes since the last diastrophism. This is further indicated 
by the fact that at least three older terraces have been recognized by Ellis 
and Lee, 8 and more by other workers. While no positive correlation of 
this lowermost terrace can be made with the terraces to the north and 
south, the fact that the Point Loma terrace shows a marked local increase 
in elevation from the south toward the north where it approaches the ele- 
vation of the La Jolla terrace of Hanna, 9 which he has shown to be the 
lowest exposed in the La Jolla Region, may be indicative of a similar age 
for the Point Loma and La Jolla terraces. Hanna, however, suggests corre- 
lation of his La Jolla terrace with the Chula Vista terrace of Ellis and 
Lee. 10 The Chula Vista terrace is, however, not the lowest one mapped by 

5 Op. cit., p. 51. 

6 Fairbanks, Harold W. The Validity of the so-called Wallala Beds as a Division 
of the California Cretaceous. Amer. Jour. Sci., 3rd ser., Vol. 45, pp. 473-478, 1893. 

7 Ellis and Lee (op. cit.), and others, have shown that a slight submergence, produc- 
ing the embayments of the San Diego coastal area, was the last diastrophic movement. This 
movement is not recorded in the Point Loma terrace. 

8 Op. cit., p. 25, and plate vi. 

9 Hanna, Marcus A. Geology of the La Jolla Quadrangle, California. Univ. Calif. 
Pub., Bull. Dept. Geol. Sci., Vol. 16, pp. 187-246, 1926. 

10 Op. cit., p. 26, and plate vi. 

340 San Diego Society of Natural History 

Ellis and Lee, but is the next to the lowest. From the account of Ellis and 
Lee, it seems that the Nestor terrace, said by them to have an elevation 
of from twenty-five feet to 100 feet above sea level, is more nearly the 
equivalent of the Point Loma terrace. This correlation is also suggested 
by Gale. 11 

The correlation of terraces in the region is complicated by the ina- 
bility to trace terraces directly from one geographic locality to another, 
and by local warping known to have taken place in the Point Loma and 
adjacent blocks. 

Climatic Inferences from the Fauna 

Analysis of the fauna as a whole, and of each locality collection, was 
undertaken in an effort to determine climatic and temperature variations 
which are known to have taken place in the Pleistocene in other west coast 
localities. 12 Considering all those forms living only as far south as San 
Diego as dominantly northern forms and thus indicative of cooler water, 
and those ranging only as far north as Santa Barbara as southern forms 
and thus indicative of warmer water, one finds a consistently high per- 
centage (64-65 % ) of the forms (for the whole fauna and for each lo- 
cality) which indicate neither warm nor cold water (on the basis outlined 
above) and which are of wide geographic range. The balance is almost 
equally divided among northern and southern forms. The general per- 
centages for the entire fauna are: northern, 16.6%; southern, 19.2%; 
intermediate, 64.1 % . Thus a temperate water condition is indicated. This 
is additional evidence for each fauna coming from the same horizon in 
the lowermost terrace, as well as evidence for the water temperature 
having been essentially the same as today at the time the forms were de- 

The absence from the fauna of such forms as Chione gnidia, Dosinia 
ponderosa, and Turritella goniostoma, which are found in other terraces 
of the San Diego region, and which do not live today 13 on the San Diego 

1 1 Gale, H. R., in Grant and Gale. Op. cit., p. 64. 

12 Bailey, T. L. Lateral Change of Fauna in the Lower Pleistocene. Bull. Geol. Soc. 
Amer., Vol. 46, pp. 489-502, 1935. Clark, Alex. The Coolwater Timms Point Pleistocene 
Horizon at San Pedro, California. Trans. San Diego Soc. Nat. Hist., Vol. VII, pp. 25-42, 
1931 Woodring W. P. Fossils from the Marine Pleistocene Terraces of the San Pedro 
Hills, California. Amer. Jour. Sci., Vol. XXIX, pp. 292-305, 1935. 

13 Professor Grant informs me that Dosinia ponderosa has been reported living in San 
Diego Bay today, but that this report is probably based upon fossil specimens which were 
thought to be Recent dead individuals. 

Webb— Pleistocene of Point Loma 341 

coast, but live farther south, indicates that some change of water tempera- 
ture (lowering) took place between the deposition of the upper terraces 
and those of Point Loma. Furthermore, the mean annual surface tempera- 
ture of the seas where the above forms now live is respectively 63, 63, and 
64 degrees Fahrenheit, while the mean annual surface temperature of the 
waters at San Diego is 62 degrees F., 14 in which today live most of the 
forms found in the faunal list. 

Gale 15 points out that the Nestor terrace, in which he includes the 
Point Loma terrace, contains warm water faunas like those of the Palos 
Verdes terraces. The data presented above show that the Point Loma ter- 
race contains faunas indicative of temperatures like those of today, and 
not warmer. Since the faunas from which the temperature conditions 
were inferred by Gale, lived on the bay side of Point Loma, and on the 
landward side of San Diego and Mission bays, where they would have 
been protected from open ocean influences, one might expect warmer 
water facies in equivalent faunas on the landward side of an island sepa- 
rated from the mainland by a shallow coastal lagoon or strait, such as 
Point Loma was known to have been during the Pleistocene, prior to the 
last uplift. 16 

The presence of one specimen of Tegula montereyi in the collection, 
which is typically a northern form, and which has not been reported living 
farther south than the Santa Barbara Islands, would be of great signifi- 
cance in indicating a cooler water temperature, if other supporting evi- 
dence were forthcoming. The presence of so many facts indicating present 
day temperature conditions at the time of deposition, is strongly sugges- 
tive of the fact that Tegula montereyi of this fauna is a reworked or 
washed-in form; or, that its geographic range in the Pleistocene was 
greater than today. 

In studying the geographic distribution of the fauna of the Point 
Loma terrace, Stephens 17 states that of those forms found: "Some have 
a southern distribution occurring now rarely or not at all this far north. 
More have a northern habitat." 

The present survey, involving almost three times as many species as 
listed by Stephens, does not support this interpretation, since as many 

14 Temperatures from data compiled by U. S. Grant. 

15 Gale, H. R., in Grant and Gale. Op. cit., p. 64. 

16 Stephens, Frank. Op. cit., p. 248-49. 

17 Op. cit., p. 249. 

342 San Diego Society of Natural History 

southern as northern forms were found in the fauna ; in fact, a few more 
southern than northern. 

Thus it is seen that temperatures closely similar to those found 
today prevailed in this area and that no temperature variations of warm 
and cool water facies (either lateral or vertical) are indicated by this 

Summary of the Ecologic Features of the Fauna 

The fauna contained in the collection examined by the writer com- 
prises a total of 102 species. Of these more than ninety per cent are 
living today on exposed coasts, in shallow water, near shore, generally 
between tides. None of the forms present are extinct. Ten per cent of 
the species whose habitat indicates deeper water or bay conditions are of 
large depth range in general. Assuming that the physical conditions and 
paleogeography at the time the forms lived were essentially as today, it 
seems reasonable to suppose that the bay forms were washed outside onto 
the outer coasts by the currents, and were deposited with the shore fauna. 

A few forms, such as Astrea inaequalis, Calliostoma turbinum, 
Diadora aspera, Tegula montereyi, Tritonalia interfossa, and Mitra idae, 
which appear in the faunal list, generally are thought to be of deeper 
water habitat. They may be present in the fauna because they were washed 
up from deeper water. Some of them appear to be water-worn. 

Age of the Terraces 

Since the Nestor terrace bevels the Pliocene San Diego formation, 
its post-San Diego age is unquestioned; and since the Nestor is either 
older than or equivalent to the Point Loma terrace, the Pleistocene age 
of the Point Loma terrace is strongly suggested. That this terrace is the 
result of the latest uplift of the region has already been shown. 

The very youthful stage of erosion of the Point Loma terrace, which 
is incised with few, small, V-shaped, steep gradient arroyos, entering the 
sea at discordant elevations, indicates a brief lapse of time since the 
exposure of this lowermost terrace. Terraces to the south and east of 

18 Professor Grant informs me that many of the smaller gastropods in the collections of 
Mr. Stephens were identified for him by the late Mr. Tom Oldroyd, who, being much more 
familiar with the northern microscopic fauna than the southern, unconsciously assigned several 
doubtful specimens to northern forms which were probably really west Mexican species with 
which he was unfamiliar. 

19 Ellis and Lee. Op. cit., p. 25. 

Webb — Pleistocene of Point Loma 343 

Point Loma show much later physiographic stages. Thus a late Pleistocene 
age is indicated. 

From a faunal standpoint, the Pleistocene age is supported by the 
fact that no extinct species are present in the fauna, and by the distinctly 
recent aspect of the faunal suite. 


The fauna of the Point Loma terrace in San Diego County, Califor- 
nia, consists of over 100 species. The ecology of the species indicates 
an open, exposed coast, with shallow water, largely inter-tidal habitat. 
The age of the fauna is Pleistocene, probably late Pleistocene. The cli- 
matic conditions at the time the forms lived were similar to those found 
in the area today. 

(For list of species and collecting localities, see following pages.) 


San Diego Society of Natural History 


Key to Ecologic Features of the Fauna 

(A) On rocks, or in sand, between high and low tides, on reefs and moss 
(B ) Along rocky shores and beaches, in the reach of the surf. 

(C) Bay form; brackish water; tidal flats. 

(D) Fairly deep water, on rocky bottom. 

(E) On big kelp. 

(F) On shells of Tegula funebralis. 

(G) On eel grass, lettuce, and seaweed. 
(H) Parasitic on star-fish and sea cucumbers. 
( I ) Nestler. 

(J) Borer. 

Localities Range 

Name Key20 Key21 


(A) Acanthina Iugubris (Sowerby) 

(A) Acanthina spirata (Blainville) 

Acmaea sp 

(A) Acmaea digitalis (?) textilis (Gould) 

(E) Acmaea insessa (Hinds) 

Acmaea instabilis (Gould) 

(B) Acmaea (?) limatula (Gould) _ 

(A) Acmaea mitra Eschscboltz in Rathke _ 

(B) Acmaea paleacea Gould _ 

(B) Acmaea scabra (Gould) - - 

(B) Aletes squamigerus Carpenter... 

Amphissa sp — - - 

(A) Amphissa versicolor Dall _.-- — 

(A) Astrea (Pachypoma) inaequalis (Martyn)' __ 

(A) Astrea (Pomaulax) undosa (Wood) __ 

Bittium sp - 

Calliostoma sp 

(E) Calliostoma canaliculatum (Martyn) 

(D) Calliostoma (?) turbinum Dall — 

Chiton sp — - 

(C) Cerithidea californica (Haldeman) _ 

(A) Conus californicus Hinds — 

Crepidula sp - 

(A-D) Crepidula aculeata (Gmelin) 

(F) Crepidula adunca Sowerby 

(A-D) Crepidula Ungulata Gould 

(A-D) Crepidula nummaria Gould 

(C) Crepidula onyx Sowerby 

(D) Diodora aspera (Eschscholtz in Rathke) 

Epitonium sp — - - 

(B) Epitonium (Nitidiscala) tinctum (Carpenter) 

(A) Fissurella volcano Reeve 

Fusinus sp - - -— 

(A) Fusinus kobelti (Dall) variety monksae (Dall).. 
Gadinia sp — - 

(B) Gadinia (?) reticulata (Sowerby) - 

(A) Haliotis cracherodii Leach _. - 

(B) Haliotis corrugata Gray.. 

(A) Haliotis rufescens Swainson ._ — 

(A) Haminoea virescens (Sowerby) — 

Hipponyx sp — - - - 

(A) Hipponyx antiquatus (Linnaeus) _ .._ 

(A) Hipponyx tumens Carpenter 

768 769 

X X 

x "x 

.._. X 

X X 

X X 

X X 

._ _- 

_ X 

20p_Present; R— Rare (2-5); C— Common (5-20); A— Abundant (204"). 
21 M — Miocene; P — Pliocene; PI — Pleistocene; R — Recent. 

Webb — Pleistocene of Point Loma 


GASTROPODS {Cominuii') 

(B-A) Homalopoma carpenteri (Pilsbry)_ 

(A) Hyalina (Hyalina) califbrnica (Tomlin) 

(A) Hyalina (Cystiscus) jewettii (Carpenter) _ 

(?) Ischnochiton sp _ - — 

(G) Lacuna (?) Carpenter _ - 

(B) Littorina scutulata Gould _ ~ - 

(B) Macron lividus(A. Adams) _ - 

(A) Mangelia (Mitromorpha) filosa (Carpenter) _ 

(A) Mangelia (Mitromorpha) gracilior (Hemphill in Tryon) 

(H) Melanella (?) rutila (Carpenter) 

(D) Mitra idae Melvill _ 

(B) Mitrella carinata (Hinds) variety gausapata (Gould) 

(B-C-D) Nassarius (Schizopyga) mendicus variety cooperi (Forbes) 

(E) Norrisia norrisii (Sowerby) „ 

(A) Olivella biplicata (Sowerby) — — 

(C-D) Retusa (Acteocina) culcitella (Gould) — 

Spirotropis (Antiplanes) sp._ — 

(A) Tegula (Chlorostoma) aureotincta (Forbes) 

(B) Tegula (Chlorostoma) funebralis (A. Adams) - 

(D) Tegula montereyi (Fischer in Kiener) 

Tegula (Promartyn) pulligo Martyn — 

Tritonalia sp. 

(D) Tritonalia foveolata (Hinds) — - 

(D) Tritonalia interfossa (Carpenter) 

Trophon sp - 

(B) Truncatella californica Pfeiffer 

(B) Truncatella stimpsoni Stearns — 

Turbonilla sp._ - 


Barbatia (Acar) pernoides (Carpenter) - 

(I) Cumingia lamellosa Sowerby _ 

(D) Glans carpenteri (Lamy)_ _ _ 

(A) Hinnites multirugosus (Gale) 

(D) Irus lamellifer (Conrad) _ — 

(B) Kellia suborbicularis (Montagu) variety laperousii (Deshayes) 
Lacuna sp 

(C) Leptopecten latiaurarus (Conrad) 

(D) Lucina californica Conrad ._ _ _ _ 

(B) Mytilus (Mytilus) californianus Conrad 

(C) Ostrea lurida Carpenter 

(A-B-C) Pecten (Aequipecten) bellilamellatus Arnold _ 

(A-B-C) Pecten (Aequipecten) circularis Sowerby 

(J) Pectricola carditoides (Conrad) 

(D) Platydon cancellatus (Conrad) 

(D) Pododesmus macroschisma (Deshayes) _ _ 

(D) Psephidia lordi (Baird) variety ovalis Dall _ 

(A) Pseudochama exogyra (Conrad) 

(B) Saxicava arctica (Linnaeus). 

(B-C) Saxidomus nuttalli Conrad 

(B) Semele decisa (Conrad) - 

(B) Semele rupicola Dall 

(A) Septifer bifurcatus (Conrad) 

(B) Tellina (?) meropsis Dall.. _ 

(A-B) Venerupis (Protothaca) staminea (Conrad) 

(B) Venus (Chione) succinta Valenciennes _ _ 


Barnacle fragments 

Coral fragments- 

Echinoid fragments 

Tetraclita squamosa (Bruguiere).. 


346 San Diego Society of Natural History 


768. Yellow fossiliferous marl from marine terraces about 100 feet above sea 
level, and about 100 yards southwest of the intersection of Ladera and 
Cordova Streets, and about 4 miles N12°W of the lighthouse on Point 
Loma, San Diego County, California. Sept. 22, 1930. Popenoe and 

769. Marine terrace 100 feet above present sea level, exposed on west side of 
Point Loma peninsula and about two miles N17V2°W of the lighthouse 
on Point Loma, San Diego County, California. Pleistocene marls just 
above Eocene-Pleistocene contact. Sept. 22, 1930. Popenoe and Scharf. 

771. Marine terrace about 100 feet above present sea level along ocean front 
on west side of Point Loma peninsula, about 100 yards north of the 
north boundary of Fort Rosecrans Military Reservation and about 3.1 
miles N13°W of the lighthouse on Point Loma, San Diego County, 
California. Marls just above Eocene sandstones. Sept. 22, 1930. Popenoe 
and Scharf. 

1185. Marine terrace about fifty feet above present sea level, exposed on west 
side of Point Loma peninsula, and about 200 yards south of the south 
boundary of the Theosophical Society grounds, on the Fort Rosecrans 
Military Reservation. Nearly on the contact between the terrace deposits 
and the Cretaceous. Point Loma peninsula, San Diego County, Califor- 
nia. February 8, 1936. Popenoe and Webb. 

1186. Marine terrace about fifty feet above present sea level, exposed on west 
side of Point Loma peninsula, and about three-tenths of a mile south of 
the south boundary of the Theosophical Society grounds, on the Fort 
Rosecrans Military Reservation. Nearly on the contact between the terrace 
deposits and the Cretaceous. Point Loma peninsula, San Diego County, 
California, February 8, 1936. Popenoe and Webb. 

1187. Marine terrace about fifty feet above the present sea level, exposed on the 
west side of Point Loma peninsula, and about one-half mile south of the 
south boundary of the Theosophical Society grounds, on the Rosecrans 
Military Reservation. Nearly on the contact between the terrace deposits 
and the Cretaceous. Point Loma peninsula, San Diego County, Califor- 
nia. February 8, 1936. Popenoe and Webb. 

1 188. Marine terrace about fifty feet above the present sea level, exposed on the 
west side of Point Loma peninsula, and about eight-tenths of a mile south 
of the south boundary of the Theosophical Institute grounds, on the 
Fort Rosecrans Military Reservation. Nearly on the contact between the 
terrace deposits and the Cretaceous. Point Loma peninsula, San Diego 
County, California. February 8, 1936. Popenoe and Webb. 




Vol. VIII, No. 25, pp. 349-360, plate 23 June 15, 1937 




Laurence M. Huey 

Curator of Birds and Mammals, San Diego Society of Natural History 

During the past four years there has been accumulated in the collec- 
tion of the San Diego Society of Natural History a small series each of a 
number of mammals from southwestern Arizona and the northwestern 
coastal district of Sonora, Mexico. Study of this material has revealed the 
presence of several apparently undescribed races which are herewith 

The writer wishes to express his gratitude and appreciation to Mrs. 
Florence V. V. Dickey and Mr. A. J. van Rossem for loan of specimens 
from the Dickey collections in Pasadena, California; to Dr. E. Raymond 
Hall of the Museum of Vertebrate Zoology, Berkeley, California, for the 
loan of comparative material; and to Mr. Bernard Bailey for the use of 
specimens from his private collection now on deposit in the Natural 
History Museum in San Diego, California. The skull illustrations are 
by Mr. Allan J. Stover. 

Neotoma lepida aureotunicata subsp. nov. 
Punta Penascosa Desert Wood Rat 

Type. — From Punta Penascosa, Sonora, Mexico; no. 10907, collection of the 
San Diego Society of Natural History; adult male; collected by Laurence M. 
Huey, February 14, 1934. 

Characters. — A race of Neotoma lepida characterized by very bright buff 
color, with a suffusion of buffy over entire dorsal and belly surfaces of the 
body; tail bicolored, and feet white. The molar series of aureotunicata is slightly 
longer and heavier than that of either Neotoma lepida flava or Neotoma lepida 

350 San Diego Society of Natural History 

Measurements of 3 Neotoma lepida aureotunicata 

"o S 



-c S 






"5 c 

2 o 

C jj 





•S " 


J S 

o . 






o £ 


DC « 

£ c 

C p 


> a. 
~Z a 
< 3 













10907 1 
























Range. — Known only from the type locality. 

Comparisons. — Compared with N. I. jlava from Tinajas Altas, Gila Moun- 
tains, Yuma County, Arizona, N. I. aureotunicata is much deeper in color, 
with a bicolored tail, dark ears and slight huffy suffusion on belly. Compared 
with N. I. auripila, from Agua Dulce Mountains, Pima County, Arizona, much 
brighter in color, with less blackish cast dorsally, bicolored tail and lighter buffy 
suffusion on belly. Blossom (Occas. Papers of the Museum of Zoology, Univ. 
of Michigan, No. 315, May 29, 1935) described a smallish, dark form, N. I. 
bensoni from the Pinacate lava beds, Sonora Mexico. I have not seen specimens 
of bensoni, but from the description, it is a much darker race than aureotunicata 
and confined to the black lava of the Pinacate region. 

Remarks. — The three specimens which form the basis of this newly described 
race were taken from a small group of rocky hills that form the promontory of 
Punta Penascosa. This locality is completely isolated as far as terrain inhabitable 
by Neotoma lepida is concerned. The nearest locality this species is known to 
inhabit is the Pinacate range, which lies over twenty miles northward and is 
separated from Punta Penascosa by that distance of bleak, sandy, level desert, an 
association hostile to the species. From the experience of the writer and from 
published accounts of the general region, it would seem that Neotoma lepida 
inhabits the higher rocky slopes of all these desert ranges in southwestern Arizona 
and adjacent Sonora. In most cases this rocky type of habitat is separated by inter- 
vening valleys and wide plains of sandy desert, which are uninhabitable by this 
species. Thus biological islands are formed and the Neotoma lepida population 
is divided into isolated groups. As with islands in the sea, the smaller the area the 
more susceptible are the inhabitants to specialized development. In fact further 
study and exploration of this general desert region, both in southwestern Arizona 
and northwestern Sonora, will without doubt bring to light additional well- 
marked races of this species. 

Specimens examined. — Neotoma lepida auripila: 3 from Agua Dulce 
Mountains, 7 miles east of Papago Well, Pima County, Arizona; Neotoma 
lepida jlava: 4 from Tinajas Altas, Yuma County, Arizona; Neotoma lepida 
aureotunicata; 3 from type locality as above. 

1 Type. 

Huey— New Mammals from Arizona and Sonora 351 

Neotoma lepida harteri- subsp. nov. 
Gila Bend Desert Wood Rat 

Type. — From 10 miles south of Gila Bend (or, exactly, from the summits 
of a group of lava hills on the east side of the Ajo railroad, about 2 miles north 
of Black Gap), Maricopa County, Arizona; no. 11462, collection of the San 
Diego Society of Natural History; adult male; collected by Laurence M. Huey, 
February 8, 1936. 

Characters. — A dark, blackish-colored race of Neotoma lepida with medium 
body size, relatively long tail and small hind feet. In body size harteri is larger 
than its southern relatives, N. I. aunpila or N. I. flava, more nearly approaching 
the average body measurement given by Benson in his paper describing N. I. flava 
(Occas. Papers of the Museum of Zoology, Univ. of Michigan, No. 317, July 
1, 1935, table, pp. 4-5) for Neotoma lepida devia from the Painted Desert in 
northern Arizona, which is the region nearest the type locality of devia from 
which he had specimens. However, in color harteri is quite different from the 
three forms above mentioned, being dark (nearly black) dorsally, with the under 
color tending towards grayish rather than towards the buffy cast found in the 
southern Arizona and Sonoran forms of N. lepida. The tail of harteri is dark 
(nearly black), in about 60 f /c of the specimens examined varying but slightly on 
the ventral side to dusky, and in the remainder uniformly colored above and below. 
In this tail character harteri stands apart from all the specimens of either aunpila, 
flava or devia so far examined. The ears of harteri are deep, dusky black. The 
median line on the side is buffy, with a buffy suffusion covering the belly. A 
white pectoral patch and a white inguinal patch of relatively large size are 
present on all the specimens of this race examined. These white patches approach 
in size those of devia and are not restricted to a small area as in aunpila. Cranially, 
harteri compares closely with aunpila and flava, but differs slightly from devia 
in a few minor characters. 

Measurements. — Type: Total length, 280; tail, 135; hind foot, 29; ear, 28. 
Skull (type) : Greatest length, 36.5; condylobasal length, 34.6; zygomatic 
breadth, 19.9; interorbital constriction, 5.4; nasals, 13.0; alveolar length of upper 
molar series, 7.3. 

Range. — So far as known, the hills south of Gila Bend, Arizona, but further 
collecting may reveal N. I. harteri to have an extensive range in the desert moun- 
tains to the eastward of the type locality. 

Specimens examined. — Besides the list mentioned in the preceding descrip- 
tion: — Neotoma lepida devia: 5 from Hoover Dam Ferry, Mohave County, 
Arizona; 3 from Mud Spring, 12 miles WSW of Chloride, Mohave County, 
Arizona; 3 from foot of The Needles, Colorado River, Mohave County, Ari- 
zona; 1 from Colorado River above Bill Williams River, Mohave County, Ari- 
zona; 2 from 10 miles below Cibola, Colorado River, Yuma County, Ari- 

~ It gives the writer pleasure to dedicate this race to Samuel George Harter who as a 
youth chose natural science as his life-work and has been the writer's companion on many 
desert collecting trips. 


San Diego Society of Natural History 

zona; 4 from base of Castle Dome, Yuma County, Arizona; 2 from 10 miles 
east of Quartzsite, Yuma County, Arizona. Neotoma lepida barter:: 9 from type 
locality as above. 

Ammospermophilus harrisii kinoensis subsp. nov. 
Sonora Antelope Ground Squirrel 

Type. — From Bahia Kino, Sonora, Mexico; no. 11284, collection of the 
San Diego Society of Natural History; adult female; collected by Laurence M. 
Huey, February 22, 1935. 

Characters. — This race is characterized by being darker and more grizzled 
dorsally and having lighter-colored hind feet than either Amnios permophilus 
harrisii harrisii or Ammospermophilus harrisii saxicola. The molar teeth are 
smaller and the tooth row is slightly shorter than in either of the other races 
mentioned. A. h. kinoensis also differs from saxicola in having larger hind 

Measurements. — Type: Total length, 229; tail, 80; hind foot, 40; ear, 5. 
Skull (type) : Greatest length, 38.3; zygomatic breadth, 23.2; interorbital con- 
striction, 10.8; nasals, 10.8; alveolar length of upper molar series, 6.8; longitudinal 
length of bullae, 9.9. 

Average Measurements of Ammospermophilus harrisii 

o £ 



■£ » 




~n c 

on "■ 







-5 5 

O u 


"0 2 
2 — 







an a 


> a. 

< 3 


i-J o 

Tinajas Altas 










Camp Verde 










Bahia Kino 

234.5 6 









Range. — So far as known, coastal district of Sonora, Mexico, from Porto 
Libertad south to Bahia Kino. 

Remarks. — The fact that Audubon and Bachman named A . harrisii without 
giving a locality from which the original specimen was collected has led to some 
speculation. Merriam (N. A. Fauna, No. 2, Oct. 30, 1889, pp. 19-20) properly 
allocated this species to southern and western Arizona; and since then Mearns 
(Proc. U. S. Nat. Museum, Vol. 18, p. 444 — advanced sheets published March 
25, 1896) divided the species into two races, describing his form A. h. saxicola 
from Tinajas Altas, Yuma County, in southwestern Arizona. Comparison of 
Arizona material substantiates the fact that two tenable forms are existent in the 

3 15 specimens. 

4 Body measurements taken from table of 20 specimens in Mearns, Mam. of Mex. 
Boundary, pp. 307-308. 

5 Skull measurements from 7 specimens in collection of Bernard Bailey. 

6 7 specimens. 

Huey — New Mammals from Arizona and Sonora 353 

area south and west of the great Mogollon plateau, though there is still a ques- 
tion of how the names may, in the future, be applied. The crux of this question 
lies in the possibility of some future student being able to designate the type 
locality of A. harrisii. The writer has followed Mearns and used the measure- 
ments of specimens from Fort ( = Camp) Verde, Arizona, as most typical of 
the race barrisu. 

When adequate material is assembled "the long tail" character assigned by 
Mearns to his race saxicola vanishes. However, the specimens from southwestern 
Arizona are discernible by their lighter coloration. In fact, color is the main 
character by which the group harrisii has been divided into these races, and this 
character is more readily seen in series than individually. 

Specimens examined. — Ammospermophilus harrisii harrisii: 8 from Camp 
Verde, Yavapai County, Arizona; tabulated body measurements of 20 speci- 
mens from same locality (see Mearns, Mam. of Mexican Boundary of United 
States, Bull. U. S. Nat. Mus., No. 56, pp. 307-308); 2 from Wickenburg, 
Maricopa County, Arizona; 5 from 6 miles northeast of Paradise, Cochise County, 
Arizona; 1 from 25 miles south of Tucson, Pima County, Arizona; 1 from 5 
miles northwest of Sells, Pima County, Arizona. Ammospermophilus harrisii 
saxicola: 15 from Tinajas Altas, Gila Mountains, Yuma County, Arizona 
(type locality); 3 from 7 miles east of Papago Well, Pima County, Arizona; 3 
from Punta Pehascosa, Sonora, Mexico. Ammospermophilus harrisii kinoensis: 
6 from Porto Libertad, Sonora Mexico; 7 from Bahia Kino, Sonora, Mexico 
(type locailty). 

Thomomys bottae growlerensis subsp. nov. 
Growler Valley Pocket Gopher 

Type. — From 7 miles east of Papago Well, Pima County, Arizona (or, 
exactly, along a well wooded desert wash on the southwestern side of a range of 
hills in the southern end of Growler Valley; the Agua Dulce Mountains form the 
southern boundary of this locality and are not far distant); no. 12387, collec- 
tion of the San Diego Society of Natural History; adult male; collected by 
Laurence M. Huey, March 16, 1937. 

Characters. — In color Thomomys bottae growlerensis differs from T. b. 
phasma, its nearest western relative, by being brighter, more golden and lacking 
the pallid appearance of either T. b. phasma or T. b. depauperatus. The skull of 
growlerensis, compared with those of T. b. phasma and T. b. vanrossemi (this 
latter form from Punta Pehascosa, Sonora, but in the same general desert area) 
is narrower, the zygomatic arches are less sharply angled and the audita! bullae 
are more rounded. The interpterygoid space of growlerensis is narrower than that 
of phasma and wider than that of vanrossemi. 

Measurements. — Type: Total length, 208; tail, 62; hind foot, 30; ear, 5. 
Skull (type) : Condylobasal length, 37.4; spread of maxillary arches, 22.7; 
interorbitaJ constriction, 6.7; nasals, 13.3; alveolar length of upper molar series, 

Range. — Known only from the type locality. 

Specimens examined. — Thomomys bottae phasma: 9 from Tule Well, 
Yuma County, Arizona (near type locality). Thomomys bottae depauperatus: 

354 San Diego Society of Natural History 

15, of which 9 were from 2 miles north of Tinajas Altas ( = 7 miles south of Ra- 
ven Butte), and 6 from proximity of Tinajas Altas, Yuma County, Arizona. 
Thomomys bottae vanrossemi: 12, including type, from Punta Penascosa, So- 
nora, Mexico. Thomomys bottae growler ensis: 8 from the type locality as given, 

Thomomys bottae comobabiensis subsp. now 
Comobabi Pocket Gopher 

Type. — From 5 miles northwest of Sells, Pima County, Arizona (elevation, 
approximately 2400 feet); no. 12460, collection of the San Diego Society of 
Natural History; adult female; collected by Laurence M. Huey, March 22, 

Characters. — A medium-sized, brownish-colored gopher, more nearly re- 
sembling those found on the higher parts of the mountains of south-central 
Arizona. The skull resembles that of Thomomys bottae modicus, but is smaller, 
with rounder, more inflated bullae and relatively shorter, heavier rostrum. 

Measurements. — Type: Total length, 215; tail, 70; hind foot, 28; ear, 4. 
Skull (type) : Condylobasal length, 35.7; spread of maxillary arches, 21.4; 
interorbital constriction, 6.5; nasals, 12.1; alveolar length of upper molar series, 

Range. — Known only from the type locality. 

Remarks. — Goldman described Thomomys bottae pusillus (Journ. Wash. 
Acad. Sciences, Vol. 21, No. 17, Oct. 19, 1931, p. 422) from a single adult 
female specimen. The measurements given by him indicate pusillus to be a 
diminutive form, not approaching the size of the race here described. 

Specimens examined. — Thomomys bottae modicus: 4 from Tubac, Santa 
Cruz County, Arizona; 11 from Santa Cruz River, 2 miles south of Tumacacon 
Mission, Santa Cruz County, Arizona; 2 from Tucson, Arizona; 12 from Fort 
Lowell, Pima County, Arizona. Thomomys bottae comobabiensis: 7 from the 
type locality as given above. 

Thomomys bottae aridicola subsp. nov. 
Gila Bend Pocket Gopher 

Type. — From 10 miles south of Gila Bend (or, exactly, on Ajo railroad 
right of way, about 2 miles north of Black Gap) , Maricopa County, Arizona; no. 
11424, collection of the San Diego Society of Natural History; adult female; 
collected by Laurence M. Huey, February 1, 1936. 

Characters and Remarks.— A medium-sized, slightly tawny-appearing 
gopher, tending in color towards the preceding form described in this paper 
rather than towards the more buffy pallid types of gophers inhabiting the desert 
regions to the westward. In skull characters, the resemblance lies in the same 
direction, but aridicola has a longer, more flattened brain-case, with wider-spread- 
ing zygomatic arches and a more slender rostrum. The skull of aridicola is weak 
and light-boned and may show a relationship to T. b. subsimilis. T. b. aridicola 

Huey — New Mammals from Arizona and Sonora 355 

is not related to either T. harquahalae or T. b. cervinus, as comparison of meas- 
urements clearly demonstrates. The incisors do not project beyond the nasals. 

Measurements. — Type: Total length, 212; tail, 63; hind foot, 29; ear, 5. 
Skull (type) : Condylobasal length, 36.7; spread of maxillary arches, 23.5; inter- 
orbital constriction, 6.8; nasals, 12.9; alveolar length of upper molar series, 7.5. 

Range. — Known only from the type locality. 

Specimens examined. — Thomomys bottae aridicola: 2 from type locality as 
given above. Thomomys bottae cervinus: 11 from Phoenix, Maricopa County, 

Perognathus intermedius lithophilus subsp. now 
Porto Libertad Rock Pocket Mouse 

Type. — From Porto Libertad (or, exactly, the summit of a rocky hill IV2 
miles NNW of the fresh water spring on the beach) , Sonora, Mexico; no. 11211, 
collection of the San Diego Society of Natural History; adult male; collected by 
Laurence M. Huey, February 5, 1935. 

Characters. — Perognathus intermedius lithophilus is darker and more gray- 
ish dorsally than either Perognathus intermedius intermedius or Perognathus in- 
termedius phasma and lacks the pinkish cast found in these two northern races. 
In size lithophilus resembles phasma and is slightly smaller than intermedius. 
Cranially, the mastoid bullae are less extended and the posterior part of the skull 
is slightly more arched and deeper than in either of the other two mentioned 

Measurements. — Type: Total length, 166; tail, 91; hind foot, 19; ear, 5. 
Skull {type) : Occipitonasal length, 23.5; mastoid breadth, 12.7; interorbital 
constriction, 6.2; nasals, 9.3; alveolar length of upper molar series, 3.4. Eight 
specimens including the type averaged: Total length, 166.8 (162-170) ; tail, 91.7 
(82-97); hind foot, 20.1 (19-21); ear, 5. Skull: occipitonasal length, 23.3 
(22.4-24.2); mastoid breadth, 12.6 (12.2-12.9); interorbital constriction, 6.0 
(5.9-6.2); nasals, 9.0 (8.8-9.3); alveolar length of upper molar series, 3.2 

Range. — So far as known, the vicinity of Porto Libertad, Sonora, Mexico. 

Specimens examined. — Perognathus intermedius intermedius: 48 from Cas- 
tle Dome, Yuma County, Arizona; 1 from Ehrenberg, Yuma County, Arizona. 
Perognathus intermedius phasma: 53 from Tinajas Altas, Yuma County, Ari- 
zona. Perognathus intermedius lithophilus: 9 from Porto Libertad, Sonora, 
Mexico (type locality) . 

Perognathus longimembris pimensis subsp. nov. 
Pima Silky Pocket Mouse 

Type. — From 1 1 miles west of Casa Grande, Pinal County, Arizona; no. 
12579, collection of the San Diego Society of Natural History; adult male; 
collected by Laurence M. Huey, May 22, 1937. 

Characters. — The dorsal color of the type of P. I. pimensis is darker than 
P. 1. bombycinus, nearly matching Ridgway's Avellaneous (Nom. of Color, 

356 San Diego Society of Natural History 

1912). This ground color resembles that of P. I. panamintinus , but pimensis 
lacks the blackish tipped hairs found on panamintinus. Cranially, these two races 
differ widely. The skull of pimensis is slightly larger than that of bombycinus and 
has a more rounded brain case and deeper, more inflated mastoid bullae. The 
zygomatic arches are more widely spreading anteriorly. The interparietal is small, 
nearly equal-sided, though this character is not one that can be used in contrast 
with bombycinus, but rather one that is shared with it and used when comparing 
with all other members of the longimembris group except P. I. kinoensis from So- 
nora, Mexico. In fact it is notable that these southwestern Arizona and Sonora 
forms of longimembris may be grouped by this character. 

Measurements. — Type: Total length, 144; tail, 83; hind foot, 18; ear, 4. 
Skull (type) : Greatest length, 21.2; mastoid breadth, 12.2; interorbital constric- 
tion, 5.0; nasals, 7.4; tooth row, 2.7. 

Range. — So far as known, from the vicinity west of Phoenix, Maricopa 
County, Arizona, south to the type locality west of Casa Grande, Pinal County, 

Remarks. — Records of Perognathus longimembris from Arizona are few. 
Osgood (Proc. Bio. Soc. Wash., Vol. 20, p. 19, Feb. 23, 1907) described Perogna- 
thus bombycinus (now P. longimembris bombycinus) from Yuma, basing his 
description on one specimen from that locality and two from northwestern So- 
nora, Mexico. Grinnell subsequently recorded (Univ. Pub. in Zool., Vol. 12, 
No. 4, p. 243, March 20, 1914) eighteen specimens of P. bombycinus taken on 
the Arizona side of the Colorado River at Ehrenberg. In 193 1 Goldman described 
Perognathus longimembris arizonensis (Proc. Bio. Soc. Wash., Vol. 44, p. 134, 
Oct. 17, 1931) from 10 miles south of Jacobs Pools, Houserock Valley, north 
side of Marble Canyon of the Colorado River, Arizona. This form ranges north 
of the Grand Canyon and into southern Utah. Nine specimens from two locali- 
ties in Arizona and one in Utah were represented in this description. So far as 
the writer is aware, these are the only localities of occurrence of Perognathus 
longimembris hitherto recorded for Arizona. 

On October 17, 1930, Bernard Bailey collected a single specimen of 
Perognathus longimembris, now in the collection of the San Diego Society of 
Natural History, at Marinette, Maricopa County, Arizona; and on May 21 
and 22, 1937, the writer captured two male specimens in a sandy area 11 miles 
west of Casa Grande, Pinal County. It is upon these three specimens that the 
present description is based. The single specimen from Marinette is in winter 
pelage and is darker and grayer than the Casa Grande specimens. It also does 
not have as well developed cranial characters, but is sufficiently close to pimensis 
to be included in the new race. However, a good series might prove it to be worthy 
of subspecific separation. 

Specimens examined. — Perognathus longimembris kinoensis: 4 from Bahia 
Kino, Sonora, Mexico (including the type). Perognathus longimembris bomby- 
cinus: 2 from 6 miles east of Yuma, Arizona (type locality); 2 from 3 miles 
west of Pilot Knob, Imperial County, California; 3 from San Felipe, Lower Cali- 
fornia, Mexico. Perognathus longimembris bangsi: 3 from Palm Springs, River- 
side County, California (type locality) ; 8 from below San Felipe Narrows, San 
Diego County, California. Perognathus longimembris panamintinus: 15 from 

Huey — New Mammals from Arizona and Sonora 


Junction Ranch, Argus Mountains, Inyo County, California; 6 from Nemo 
Canyon, Panamint Mountains, Inyo County, California; 7 from Harrisburg Flat, 
Panamint Mountains, Inyo County, California; 4 from mouth of Goler Canyon, 
Panamint Valley, Inyo County, California. Perognathus longimembris pimensis: 
1 from Marinette, Maricopa County, Arizona; 2 from 11 miles west of Casa 
Grande, Pinal County, Arizona (type locality) . 

Bassariscus astutus yumanensis subsp. now 
Yuma Cacomistle 

Type. — From Tinajas Altas, Gila Mountains, Yuma County, Arizona; no. 
12272, collection of the San Diego Society of Natural History; adult male; col- 
lected by Laurence M. Huey, March 6, 1937. 

Characters and Comparisons. — A race of Bassariscus astutus differing from 
B. a. arizonensis of central and eastern Arizona in slightly smaller size, lighter 
dorsal color and having heavier proportions of black in the bands on the upper 
side of the tail. In this latter character B. a. yumanensis is similar to B. a. octavus 
of southern California, but has a more pallid color dorsally. Cranially yumanensis 
differs from both of the above mentioned races in having a smaller skull, with 
shorter, heavy rostrum, a more curving tooth row, a shallower brain case, and the 
tympanic bullae slightly shorter in length but rounder and much more deeply 
expanded when viewed in profile. (See Plate 23) . 

Measurements in Millimeters 




5 D 

5 = 

. 5 

c z 




2 5 

■z ° 

.2 c/i 







15 5 

£ o 






£ ~ 













5k « 


— o 







































Remarks. — The race B. a. yumanensis represents another form of lighter- 
colored, subspecifically different mammals from the southwestern desert section 
of Arizona. The range of Bassariscus astutus extends from Transition Zone in 
the north to Lower Sonoran or even Arid Tropical in the south. This provides a 
wide latitude of environmental conditions, and, as a result, when a good assem- 
blage of specimens is brought together from widely separated localities, the effects 
of varied habitat are quickly discernible. This is true as regards both color and 
size. For example yumanensis has size tendency towards the Lower California 
form palmarius, but stands apart in color tone, as would be expected when the 
climatic variation of their two ranges is compared. Similarly, conditions of 

7 Type. 

Huey — New Mammals from Arizona and Sonora 

Plate 23 

Bassariscus astufus anzonensis. 

Bassariscus asfufus yumanensis. (Type) 

Bassariscus asfufus ocfavus. 

Comparison of Bassariscus skulls. X3/5 

Huey — New Mammals from Arizona and Sonora 359 

habitat are reflected in the appearance of the other races of the species. 

There are two trapper-taken specimens in the Dickey collection labeled 
from Laguna Dam, Imperial County, California. Both of these specimens lack 
skull, front feet and measurements. One of them (no. 9975) is fairly representa- 
tive in color of the race yumanensis, but the other (no. 9976) is doubtful and, 
judging from tail characters, is probably referable to anzonensis. However, with- 
out a skull the identity of this latter specimen cannot be definitely determined. 
The writer has positive knowledge of the presence of Bassariscus at this Califor- 
nia locality, and he also knew the trapper and something of his travels. At the 
time these specimens were taken, he was a prospector, trapping as a side line, 
and was often away on short excursions into territory east of Tucson, Arizona, 
well within the range of anzonensis. His failure to appreciate the need of scien- 
tific accuracy, and the resemblance of no. 9976 to anzonensis , would tend to dis- 
qualify the locality data on this specimen. 

Range. — Mountains of the arid region in extreme southwestern Arizona and 
southeastern California, and probably in contiguous territory of northwestern 

Specimens examined. — Bassariscus astutus anzonensis: 2 from 14 miles east 
of Fort Lowell, Pima County, Arizona. Bassariscus astutus octavus: 2 and 1 skull 
from San Luis Rey River, altitude 1700 feet, near Escondido, San Diego County, 
California; 1 from Bear Flat, San Antonio Canyon, Los Angeles County, Cali- 
fornia. Bassariscus astutus raptor: 1 from Eel River Bridge, Mendocino County, 
California; 1 from Low Gap, Trinity County, California; 1 from Hyampom, 
Trinity County, California; 1 from Bridgeville, Humboldt County, California; 
2 from Eldridge, Sonoma County, California. Bassariscus astutus palmarius: 
6 from San Ignacio, Lower California, Mexico. Bassariscus astutus insulicola: 
2 from San Jose Island, Lower California, Mexico. Bassariscus astutus saxicola: 
2 from Espiritu Santo Island, Lower California, Mexico. Bassariscus astutus 
yumanensis: 3 from Tinajas Altas, Gila Mountains, Yuma County, Arizona 
(type locality) ; 1 from Laguna Dam, Imperial County, California. 



Vol. VIII, No. 26, pp. 361-362 June 15, 1937 



A. J. van Rossem and The Marquess Hachisuka 

Dickey Collections, California Institute of Technology 

Recently, when the writers were shown a breeding pair of Mexican 
Black Hawks collected by Mr. W. J. Sheffler in southeastern Arizona, 
we at once recognized them as something quite distinct from typical 
Buteogallus anthracinus anthracinus of southern Mexico and Central 
America. Mr. Sheffler generously allowed us to borrow these specimens 
for study and our investigations show that a well-marked race is present 
in northwestern Mexico and the contiguous parts of the United States. 
This race we name as 

Buteogallus anthracinus micronyx subsp. nov. 

Type. — Breeding male adult, no. 1477, collection of W. J. Sheffler; Arivaipa 
Creek, Graham County, Arizona, June 3, 1936; collected by W. J. Sheffler. 

Subspecific characters. — Most closely resembling Buteogallus anthracinus 
anthracinus (Lichtenstein) of southern Mexico and Central America, but general 
coloration paler and more brownish plumbeous (less blackish) ; mottling on 
under wing coverts and remiges more extensive and, in the latter instance, tend- 
ing to gray rather than buff or reddish brown; lores and sub-ocular streak pure 
white instead of buff or pale buff. In the matter of size, micronyx has a longer 
wing and tail, and shorter, more slender claws. Juvenile very much paler than the 
corresponding stage of anthracinus, the ground color being nearly white and the 
ventral streaking narrow. 

Range. — Southern Arizona, south to western Chihuahua and southern 

Remarks. — Lichtenstein's type of Falco anthracinus was collected by Deppe 
on his first (von Sack) trip to Mexico and therefore came from southeastern or 

362 San Diego Society of Natural History 

southwestern Mexico. This type is now in the Berlin Museum and was examined 
by van Rossem in 1933, but his notes and measurements concerning the speci- 
men have unaccountably disappeared. However, three specimens from the state 
of Vera Cruz (Alvarado; Pasa Nueva) and one from Tehuantepec (all in the 
Museum of Vertebrate Zoology) belong unmistakably with the southern race. 
We are unalile to perceive any significant differences between black hawks from 
southern Mexico and those from Guatemala, and El Salvador. However, two 
specimens from Limon, Costa Rica (Dickey collection) , are more intensely 
black below than any typical anthracinus examined, and in addition have notably 
larger and more powerful feet and claws. Additional material may show the 
existence of a southern Central American race, in which case the name of Buteo- 
gallus anthracinus bangsi (Swann) will probably be applicable. 

Measurements. ' — 
Adult Males Wing Tail Middle Claw Hind Claw 

2 micronyx from Arizona and Sonora 365-385 220-222 19.0-19.7 21.5-22.5 
4 anthracinus from southern 

Mexico and Central America 355-360 195-200 21.5-22.0 25.0-25.1 

Adult Females 

3 micronyx from Arizona and Sonora 395-398 222-237 20.0-23.0 24.0-26.0 
6 anthracinus from southern 

Mexico and Central America 370-390 204-220 22.5-25.5 26.0-27.8 

2 anthracinus (?) from Costa Rica 385-390 218-226 28.0-29.5 31.1-33.1 

Specimens examined. — B. a. anthracinus, Mexico, 6; Guatemala, 1; El 
Salvador, 10; Costa Rica, 2. B. a. microynx, Chihuahua, 1 (Colonio Pacheco) ; 
Sonora, 4 (Sonora; Alamos; Guirocoba) ; Arizona, 2 (Arivaipa Creek) . B. a. 
subtilis. El Salvador, 8. 

1 Immature birds approximate adults in measurements save that almost invariably they 
have longer tails. 

J ** 1i JAN 3 193! 



Vol. VIII, No. 27, pp. 363-366 December 1 5, 1937 



Laurence M. Klauber 

Curator of Reptiles and Amphibians, San Diego Society of Natural History 

Through the kindness of Robert Hoard of San Diego State Col- 
lege, I have received a Sonora from the state of Sonora, Mexico, which, 
while obviously related to Sonora occipitalis, the Shovel-nosed Ground 
Snake of southeastern California and southwestern Arizona, differs con- 
spicuously from that species in pattern. It is likewise somewhat low in 
ventral scale counts, although the significance of the latter difference 
cannot be determined until more specimens of the new form are avail- 
able. The territory intervening between the most southerly known speci- 
mens of S. occipitalis (these are from southern Arizona) , and the type 
locality of the new snake (central Sonora) is suitable in character to 
either form, consequently the two may later be shown to intergrade. 
However, the extreme southerly specimens of S. occipitalis, as compared 
with those from the center of its range, do not show directive character 
trends toward the new form. I therefore deem it advisable to consider it 
a full species, until intergradation be demonstrated, and suggest for it 
the name of 

Sonora palarostris sp. nov. 
Sonoran Shovel-nosed Ground Snake 

Holotype. — No. 26,771 in the collection of LMK. Collected 5 miles south 
of Magdalena, Sonora, Mexico, by George Lindsay, April, 1937. 

Diagnosis. — A Sonora resembling S. occipitalis in configuration, but with 
fewer dorsal blotches and with a low ventral scale count. An examination of 219 

364 San Diego Society of Natural History 

specimens of S. occipitalis reveals from 18 to 40 black dorsal rings on the body, 
the average being 26.3. The holotype of S. palarostris has 10 black rings. S. occipi- 
talis males have from 147 to 170 ventrals, the average being 155; S. palarostris 
has 144. 

Description of the Type. — Adult male.The head is somewhat wider than 
the neck, and is narrowed anteriorly. The snout is wedge-shaped in profile, with 
the lower jaw inset, the rostral projecting beyond the mental. The rostral is wider 
than high, recurved above, and deeply concave below; it extends farther backward 
below than above. The scales on the top of the head consist of a pair of inter- 
nasals, which are wider than long; a pair of prefrontals which widen laterally; a 
large hexagonal frontal; a pair of supraoculars, not conspicuously imbricate and 
shorter than the frontal; and a pair of contacting parietals, considerably larger 
than the frontal, and one-third longer than wide. The nasal is small, entire, longer 
than high, and with the nostril somewhat posterior to the center. There is a single 
loreal on each side, smaller than the nasal, and longer than high. The preoculars 
are 1 — 1; postoculars 2 — 2, the lower being the smaller. The temporals are 1+2. 
There are 7 supralabials; the 6th is the largest; the 3rd and 4th contact the eye. 
The mental is small and quadrangular. There are eight infralabials, the first pair 
contacting on the median line; the fourth is the largest. The chin shields are in 
two pairs; the first pair in contact and nearly twice as long as the second; the latter 
are separated by a single gular. There are six gulars between the posterior tips of 
the first chin shields and the first ventral. The dorsal scale rows number 
15 — 15 — 15; they are smooth and polished, with single apical scale-pits. The 
ventrals number 144; anal divided; subcaudals 39, all divided except the rather 
blunt terminal scale. 

The pattern consists of alternating black and red rings or blotches, separated 
by narrow strips of yellow ground color. 1 There are 10 black rings on the body 
and 3 on the tail, the last being at the tail tip. The black rings are about twice 
the width of the yellow separating strips, and the red rings or blotches in turn 
have about twice the longitudinal extent of the black. In terms of dorsal scales 
(end to end) the rings have approximately the following widths: yellow, If 
scales; black, 3i scales; red, 62 scales. Both the black and red rings narrow on the 
sides, so that laterally the ground color is more in evidence than dorsally. The 
anterior black ring does not contact the ventrals; the second, while not complete, 
is represented ventrally by a pair of black spots. The third and all subsequent 
black rings completely encircle the body. They widen ventrally, compared with 
their lateral extent, but are not as wide as on the mid-dorsal line. The red blotches 
fade out laterally at the first row of scales above the ventrals; the two red rings 
on the tail are complete ventrally, but none on the body crosses the ventrals. 
There are a few black dots irregularly disposed in the red areas. 

The snout is cream colored. There is a large black parietal blotch covering 
the posterior 2/3 of the frontal and extending to the posterior edge of the parietals; 
on the sides the blotch engages the eyes and the upper edges of the posterior 
supralabials. While this blotch is analogous to the crescent-shaped black blotch 

• The ring arrangement formula is that usually given as distinguishing the coral snake 
from various harmless snakes. 

Klauber — New Snake from Mexico 365 

characteristic of S. occipitalis, in this form it is more rectangular. The underside 
of the head is cream. Referring to Ridgway (Color Standards, 1912), the three 
dorsal colors of the alcoholic type are Maize Yellow, Brazil Red, and Black; 
these colors were observed shortly after preservation. The ventral shade is Cream 

The length over-all is 312 mm.; tail length 57 mm. The pupil of the eye 
is round. 

Habits. — The type specimen was found abroad crawling across the road 
at about seven o'clock in the evening. From the likeness to S. occipitalis it may be 
assumed that this snake is a burrower; however, the top of the head is slightly 
convex from frontal to rostral, whereas in S. occipitalis this digging wedge is 
flat, or may even be dished. The type contained the remains of a large spider. 

Remarks. — The number of black cross-bands on the body in S. occipitalis 
is a relatively constant character. Statistics of specimens ranging from Inyo 
County to Imperial County in California, and eastward in Arizona to Wicken- 
burg and Picacho, are as follows : 

Range 18 to 40 bands 2 

Mean 26.3 ±0.18 bands 

Standard deviation 4.03 bands 

Interquartile range 23.6 to 29.0 bands 

Coefficient of variation 15.3 per cent 

There is some tendency of the number of bands to increase as we go north 
from the southern border of California to Kern and Inyo Counties, but the Ari- 
zona specimens, nearest to the probable range of palarostris, are not low. Thus 
there is no directive tendency toward intergradation, as far as cross-bands are 
concerned. The deviation of the palarostris type, from the mean of occipitalis, 
is 4.04 times the standard deviation of the latter — a highly significant difference. 
The relative lengths of the red and black blotches in occipitalis are also quite 
different. Palarostris is a brilliantly colored little snake, and with its conspicuous 
red blotches is even more beautiful than occipitalis, which is admired even by 
persons who do not usually care for snakes. 

2 The type of occipitalis from the Mohave Desert had 33 black bands. 

At UK 

JAN 3 1938 



Volume VIII, No. 28, pp. 367-374, plate 24 December 15, 1937 




Hubert Lyman Clark 

Museum of Comparative Zoology, Cambridge, Mass. 


Several years ago Professor Hubert G. Schenck, of Stanford Uni- 
versity, sent to the Museum of Comparative Zoology a rock fragment, 
collected in Oregon in 1931, bearing the mold of most of the upper sur- 
face of a large spatangoid of which he desired identification. Besides the 
nearly complete impression, the fragment bears portions of the molds of 
at least three other individuals of the same species, but these are of prac- 
tically no assistance in determining the systematic position of the spatan- 
goid involved. Recently (1936) Professor Schenck has sent an additional 
specimen of what he thinks is the same species, but from a different locality 
— State of Washington, and has asked for a name by which these echini 
may be designated. 

There is no doubt that the molds in the earlier specimen represent 
more or less of the petaloid areas of a species of Brisaster, very similar to, 
but probably not identical with, the Recent species, B. townsendi (A. 
Ag.), which occurs along the whole western coast of North America in 
water of moderate (511-995 fathoms) depth. The apical system of the 
fossil is much more anterior than in townsendi, and the petals are straight- 
er and relatively narrower. But unfortunately townsendi is a very variable 
species and not a great deal of reliance can be placed on these differences. 

1 Abstract in Geol. Soc. Am. Proc. for 1936 (1937). 

368 San Diego Society of Natural History 

In view, however, of the very much greater size of the fossil form, it may 
perhaps be justifiable to lay some stress on the markedly anterior apical 
system and distinguish the extinct species from its Recent congener under 
the name maximum. Careful comparison with a large example of townsendi 
enables one to describe this new Brisaster as follows : 

Brisaster maximus, sp. nov. 

Plate 24, fig. 9 

Length about 84 mm., with the width approximately 76 mm.; nearly 
the whole width is shown in the large mold. If the proportions were the 
same as in townsendi, the height was about 42 mm. In the Recent species 
the apex is far back of the middle of the dorsal surface, so that its center 
is only .40 of the test length from the posterior margin of the test; in the 
fossil the apex is so near the center that the anterior margin could not have 
been more than 4 or 5 mm. further away than the rear end of the test. 
As a result of this difference in the position of the apex, the distal ends 
of petals I and V are far from the test margin in maximus, while in town- 
sendi they nearly reach it; indeed, in some specimens they would over- 
reach it if they did not diverge markedly from each other and from the 
longitudinal axis. The size of the petals and the angles which they make 
with the axis and with each other show very great diversity in townsendi, 
but it is rare for petals I and V to form as narrow an angle with each other 
as they do in maximus (about 80°) ; in townsendi the angle commonly 
exceeds 90° and may be much more. 

The tuberculation of the test in maximus was apparently very much 
as in townsendi, the larger tubercles occurring beside the petals, especially 
near the distal ends. Fragments of the peripetalous fasciole can be dis- 
tinguished here and there, most evidently around the tip of petal V and 
thence anteriorly towards petal IV. 


The holotype of maximus is in the Museum of Comparative Zoology, 
Harvard University, No. 3830; a plastotype is in the Schenck Collection, 
Stanford University, No. 2196. Collector: John T. Holman, who de- 
scribes the type locality (Holman field locality No. 44) as follows : 

Washington County, Oregon, from the center of the south line of 
Section 12, T. 3 N., R.4W.; dug well along road at C. H. Bonham farm; 
1000 feet south of coal seam exposure; Pittsburg Bluff formation, Re- 
fugian Stage; Acila shumardi zone, "Oligocene" of Pacific Slope authors. 

Clark — New Sea-urchin from Oregon 369 

The locality is shown on the map in the monograph by Schenck (1936, 
p. 43). 

Specimen from Washington 

The specimen sent by Professor Schenck in 1936 comes from "Loc. 
N. P. 55, 2 near Porter, Washington," H. Hannibal, collector. It con- 
sists of the upper and lower portions of a mold in a soft, clay-like, gray 
rock. In connection with petal IV, on the lower portion, is a small part of 
the test with many minute spines still present, while on the upper portion 
a large part of the peripetalous fasciole can be made out. The specimen 
was badly crushed before fossilization and its normal size and proportions 
are therefore uncertain. Apparently it was 60-65 mm. long and 50-55 mm. 
wide. While it seems unquestionably a Brisaster, there are some features 
of the petals I and V that are unlike maximus and make it possible to 
have some doubts as to its identity with the Oregon specimens, in spite 
of Professor Schenck's opinion (presumably on stratigraphical grounds) 
that "it must belong to the same species." But in view of the great diversity 
shown by individuals of townsendi, it seems foolish to doubt that the 
Oregon and Washington fossils represent the same species of Brisaster, 
since their geological position is essentially the same. Whether maximus 
is the direct forerunner of townsendi is a matter that admits of more un- 

The Genus Brisaster 

The genus Brisaster was named by Gray (Cat. Rec. Ech. Brit. Mus., 
p. 61, 1855) as a subgenus of Schizaster with three species; fragilis, gib- 
bendus, and cubensis. The year before, d'Orbigny (Pal. France Cret., p. 
270, 1854) had placed cubensis in the genus Periaster. In 1883, Pomel 
(Class. Meth. Ech., p. 36) made gibbendus the type of the genus Paraster. 
Thus, Schizaster fragilis Agassiz and Desor, 1847, alone is left to be the 
type of Brisaster. 

A. Agassiz, Duncan, and W. B. Clark and Twitchell never recog- 
nized Brisaster, but treated it as a synonym of Schizaster. However, Mor- 
tensen (Ingolf Ech. pt. 2, pp. 122-123, 1907) discussed the genus Schiz- 
aster, recognizing four subgenera : Paraster, Schizaster, s.s., Tripylaster, 
and Brisaster. H. L. Clark (Mem. Mus. Comp. Zobl., vol. 46, no. 2, pp. 
159 et seq., 1917) accepted Mortensen's four biologic units as valid 

2 Professor Schenck informs me that the initials stand for "North Pacific" and that the 
number is H. Hannibal field locality No. 55. 

370 San Diego Society of Natural History 

genera, and included six Recent species in Brisaster, two of which (lati- 
frons and townsendi) , occur in the eastern and northern Pacific Ocean. 
In short, there can be no doubt that Brisaster is a valid nomenclatural 

The species of Schizaster and Brisaster are distinguished by three 
characters, as follows : 

1. Test relatively high and more or less swollen in Schizaster, es- 
pecially posteriorly; more flattened and widened in Brisaster. 

2. Genital pores : two in Schizaster, three in Brisaster. 

3. In Schizaster, petals II and IV are short, wide, divergent; petal 

III deeply sunken, but usually not very broad. In Brisaster, petals II and 

IV are long, comparatively narrow, directed well forward, sometimes so 
markedly so as to be nearly parallel for a short distance; petal III some- 
what sunken, broad. 

The time range of Brisaster is given commonly as Eocene to Recent, 
as may well be. Lambert and Thiery, for example, list a number of species 
of Brisaster which they assign to the Eocene. There is not one that can 
unqualifiedly be called Brisaster, although Schizaster pyrenaicus Munier- 
Chalmas is a likely candidate. 

Clark — New Sea-urchin from Oregon 371 

Selected Bibliography 

Agassiz, Alexander 

1904. The Panamic Deep Sea Echini. Mem. Mus. Comp. Zool., vol. 31, 
pp. 1-243, 112 pis., chart, 319 text figs. 

Agassiz, Louis (and Desor, Edouard) 

1847. Catalogue raisonne des families, des genres et des especes de la classe 
des Echinodermes. Ann. Sci. Nat. ser. Ill, Tom. VI-VIII, Paris. 

Clark, Hubert Lyman 

1917. Hawaiian and other Pacific Echini, based upon collections made by 
the U. S. Fish Commission Steamer Albatross in 1902, Commander 
Chauncey Thomas, U. S. N., Commanding. The Spatangina. Mem. 
Mus. Comp. Zool., vol. 46, no. 2, p. 85-283, 22 pis. 

Gray, John E. 

1855. Catalogue of the Recent Echinida in the British Museum, pt. 1, 
Echinida Irregularia, London. 

Kew, William S. W. 

1920. Cretaceous and Cenozoic Echinoidea of the Pacific Coast of North 
America. Univ. Calif. Pub. Dept. Geol., Bull. vol. 12, no. 2, pp. 
23-236, 40 pis., 5 figs. 

Merriam, John C. 

1899. The Tertiary Sea Urchins of Middle California. California Academy 
of Sciences, Proc. (3rd Series), Geol., vol. 1, no. 5, pp. 161-174, 2 pis. 

Mortensen, Th. 

1907. Echinoidea. The Danish Ingolf- Expedition, 1895-1896. Scientific Re- 
ports, vol. 4, pt. 2, pp. 1-200, 19 pis. 

d'Orbigny, Alcide 

1853-1860. Paleontologie francaise, Description zoologique et geologique de 
tous les animaux mollusques et rayonnes fossiles de France, Terrains 
cretaces, VI Echinodermes, Atlas de pi., Paris. 

Pomel, M. A. (Auguste) 

1883. Classification Methodique et Genera des Echinides Vivants et Fossiles, 
Thesis for Dr.'s Degree, Paris, pp. 132, 1 pi., Alger. 

Schenck, Hubert G. 

1936. Nuculid Bivalves of the Genus Acila. Geol. Soc. Am., Special Paper 
4, pp. 149, 18 pis., 15 figs, in text. 

372 San Diego Society of Natural History 

Explanation of Plate 

Figs. 1-6. Brisaster jragilis (Agassiz and Desor) , x 3/4. 

Recent. Hypotype No. 6064, (Stanford University Paleo. Type 
Coll.). Off east coast of the United States, 140-216 fathoms. 
Length of specimen 49.2 mm., width 46.8, thickness 30.8. 

Figs. 7-8. Brisaster latifrons (Agassiz) , x 4/5. 

Recent. Eastern Pacific, "Albatross" station 3431, 995 fathoms. 

(Copy of protographs) . 

Fig. 9. Brisaster maximus H. L. Clark, n. sp., x 2/3. 

Holotype No. 3830, (Museum of Comparative Zoology, Har- 
vard University). From Washington County, Oregon, Sec. 12, 
T. 3 N, R. 4 W. "Oligocene." 

Figs. 10-12. Brisaster townsendi (Agassiz). Recent. Eastern Pacific. 

Figs. 10-11, x 4/5. Hypotype No. 6949, (Calif. Acad. Sci. Loc. 
28046) . Howe Sound, British Columbia. Collector, S. A. Glassell. 
Length, 50.4 mm., thickness 26.8 mm. 

Fig. 12, x 4/5. Hypotype No. 6950, (Calif. Acad. Sci. Loc. 
28046). Howe Sound, British Columbia. Length, 56.5 mm. 
(Photographs of this species by Frank L. Rogers, W. P. A. pro- 
ject) . 

Figs. 3, 8, and 12 are views of the upper surface. 
Figs. 2, 7, and 10 are views of the lower surface. 
Figs. 1 and 1 1 are views of the rear end. 
Figs. 5 and 6 are side views. 
Fig. 4 is an anterior end view. 

Clark — New Sea-urchin from Oregon 

Plate 24 

»s !/o. 


I 1 , 

.y \'v. 



>^; ; 



JAN 3 1938 



Vol. VIII, No. 29, pp. 375-378, text figs. 1, 2 December 15, 1937 



Loye Miller 

Professor of Biology, University of California at Los A ngeles 

Through the very great courtesy of Director Clinton G. Abbott and 
Mr. Frank Gander of the museum of the San Diego Society of Natural 
History, I have lately come into possession of an interesting bird femur 
from the San Diego Pliocene. The specimen was collected by Mr. Gan- 
der's son from an exposure of marine gravels on Market Street near 
Euclid Avenue in the eastern part of the city of San Diego. This forma- 
tion and this same locality yielded, five years ago, the second known 
specimen of Mancalla californiensis Lucas (Miller, L., 'Condor,' XXXV, 
pp. 34-35, Jan., 1933). Dr. U. S. Grant has made some study of the in- 
vertebrate fauna of these beds with the conclusion that they are of Upper 
Middle Pliocene age, well above the middle of the San Diegan formation. 

The matrix is fine grained, gray sand of quite uniform texture. 
Enough motion had taken place prior to the entombment of the specimen 
to wear away the more delicate contours, a fact that would indicate that 
relatively shallow and active water had deposited the stratum. 

The bone has been extensively mineralized without appreciable stain- 
ing. A clean fracture of the shaft shows a central cavity that seems, at 
first impression, rather small for a bird bone. However, the diving birds 
do not have such large shaft cavities in their bones, especially in the femur. 
In the femur of the loon, the cavity is almost rendered nil by the abundant 
cross struts of dense bony tissue. In the fossil specimens such struts are 

376 San Diego Society of Natural History 

not found at the point of fracture. In those highly specialized divers, the 
loons and grebes, that depend mainly upon the feet for underwater prog- 
ress, the femur tends toward a short, thick, and strongly curved form. 

The puffins, on the other hand, have relatively weak feet. In rapid 
underwater progress, their wings are brought into play in a true under- 
water flight. The pelvic limb is less specialized and the long, slender 
femur does not differ radically from that of their surface feeding rela- 
tives, the gulls. The fossil femur here discussed is of this latter type. It 
is about the size of that of the Herring Gull from which it differs as 
would a living puffin. The trochanter is less developed, there is a slightly 
greater dorsiventral curvature and a much greater lateral arching. Anhin- 
ga, Sula, the cormorants, and the diving anserines are entirely different. 
The fossil is most closely like the puffins of the genus Lunda, from which it 
differs in certain details as well as in its much greater size. There seems 
to be no genus of the living Alcidae to which the specimen may properly 
be referred. It is therefore deemed necessary to establish a new genus for 
which the name Pliolunda is proposed. 

Pliolunda diegense, new gen. and sp. 

Type. — Right femur, No. 33409, Museum of Paleontology, University of 
California; from the Upper Middle Pliocene, San Diegan formation. 

Most closely like Lunda cirrhata in general appearance, curvature of shaft, 
and axes of the articulations. Size much greater; trochanteric ridge less prolonged 
down the shaft; obturator ridge more pronounced; popliteal area more depressed; 
rotular groove slightly wider; external condyle more prolonged and inner condyle 
less prolonged up the anterior face of the shaft. 

These differences are sufficiently pronounced to prevent assignment to the 
genus Lunda, but they are not considered to be fundamental, nor do they appear 
to be correlated with adaptive differences. The femur of the Great Auk (Plautus 
impennis) has much the same characters of the distal end, but is not so closely 
like it proximally. Plautus might properly be considered as the extreme of speciali- 
zation among the Recent alcids, but this specialization is not reflected in the 
femur. The Lucas Auk (Mancalla californiensis) from the same formation is 
almost as large as the Great Auk, hence the specimen here discussed could not 
be properly considered of that species, the femur of which is not known. It does 
appear, however, as a contemporary of the Lucas Auk, and it may have had 
similar habits. 

Remarks- — The present day metropolis of the puffins and their nearer allies 
is well to the northward of San Diego. During the later stages of Pliocene time 
there were accumulated at a number of stations along the Southern California 
coast quite extensive molluscan faunas, some species of which serve as tempera- 
ture indicators of fairly definite nature. 

Miller — Extinct Puffin from San Diego 


According to Grant and Gale (Mem. San Diego Soc. Nat. Hist., Vol. I, 
1931, p. 35), the San Diegan formation begins with a fairly warm water fauna 
which changes definitely to a cool water fauna toward its close so that Upper 
Pliocene is found to contain "numerous specimens of species that live today 
between Alaska and Puget Sound." Into such a picture of advancing cold, this 
new species of Pliocene puffin would fit quite harmoniously. 


Measurements — Measurements of type specimen of Plwlunda are as fol- 

Length from trochanter to outer condyle 55.7 mm. 

Least shaft diameter 4.5 mm. 

Transverse diameter through condyles 10.8 mm. 

Figs. 1,2. Anterior and external views of the 
type specimen Pliolunda diegense, approximately 
natural size. 

aiin JAN 3 1938 



Volume VIII, No. 30, pp. 379-406, plates 25, 26 





George Willett 

Los Angeles Museum 


Printed for the Society 

December 15, 1937 


U. S. Grant, IV, Chairman 

Fred Baker Clinton G. Abbott, Editor 





George Willett 

Los Angeles Museum 

In 1926 Professor A. J. Tieje (Bull. Am. Assoc. Petr. Geol., Vol. 
10, p. 510), in a discussion of the Pliocene and Pleistocene history of the 
Baldwin Hills, referred to a warm water fauna uncovered in Trench 6 
of the Los Angeles Outfall Sewer, giving it the name of the Centinela 
Gravels. A much more extensive exposure of what is apparently the same 
fauna occurred a few years later during the widening of Lincoln Avenue, 
which crosses the outfall sewer about two miles northeast of Playa del 
Rey. At a point just south of the sewer, at an altitude of about fifty feet 
above sea level, excavations by steam shovels cut into the upper part of the 
fossiliferous strata, exposing large numbers of marine invertebrates. 

During the summer of 1935 a number of lots of fossil shells from 
this section were brought to me for identification, and, after studying 
them, I became sufficiently interested in certain features of the collections 
to undertake a rather careful examination of the locality from whence 
they came. 

The fossiliferous stratum, from eight to twelve inches thick in most 
places, was found to be mainly from two to four feet below the present 
surface. It was bordered, both above and below, by sand which sometimes 
contained sparsely scattered, small, water-worn stones. In some sections 
there was a thin stratum of echinoderms a few inches above the mollusk- 
bearing vein, but this was by no means constant. 

During 1935 and 1936 I made many trips to this fossil locality, and 
excavated, screened and carefully examined several tons of material. This 
resulted in an accumulation in the Los Angeles Museum of more than 
30,000 specimens. While the majority of these are mollusks, several other 
groups were well represented. No attempt was made to preserve all the 
specimens uncovered, in the case of the more common species only a good 
representation being kept, and all badly worn or broken specimens being 
discarded except in case of the rarer species. A million would probably be 

382 San Diego Society of Natural History 

a conservative estimate of the total number of specimens examined. 

In addition to the above, I have had access to the collections of John 
Q. and Tom Burch, Alex Clark, Mr. and Mrs. Philip M. Connelly, 
Miss Edna T. Cook, Mrs. Bertha M. Fuller, Steve A. Glassell, J. C. 
Marsh, Miss Alice Waterbury, and H. C. and Homer L. White, all of 
whom possess considerable material from the Del Rey deposits. To these 
friends, who have not only allowed free use of their collections, but have 
donated numerous specimens to the Los Angeles Museum, my sincere 
thanks are due. I am also indebted to the following students for classify- 
ing material in their respective fields : Steve A. Glassell, decapod crusta- 
ceans; Dr. U. S. Grant, echinoderms; Dr. Howard R. Hill, barnacles 
and bryozoans; and Dr. Hildegarde Howard, birds. Finally my gratitude 
is expressed to Dr. U. S. Grant and A. M. Strong for helpful informa- 
tion regarding literature and taxonomic questions. 

Among the more striking features of this deposit are the abundance 
of crustacean remains, the great number of specimens of small mollusks, 
such as Pyramidellids, Melanellids and Turrids, and the periodic purity 
of the entire assemblage; that is, it may be almost entirely attributed to 
one distinct and rather exact period. There has apparently been very little 
mixing of materials from different geological ages, as is so common in 
many of our coastal fossil deposits. That this horizon is the same as Pro- 
fessor Tieje's "Centinela Gravels" is indicated, not only by locality, but 
by comparison with a list of fifty-five species of mollusks in an unpublished 
manuscript of Professor Tieje. 

From a study of the nature of the marine fauna of this section, it 
would appear that its habitat was sandy ocean bottom, at a depth of from 
ten to twelve fathoms, near the mouth of a bay or slough, the latter feature 
being indicated by the presence of a few examples of marsh species such as 
Melampus, Helisoma and Gyraulus, which must have drifted down from 
coastal marshes. 

A summary of the material preserved is as follows. Of mammals 
there are the remains of two species, a seal, and a dolphin or porpoise, 
that we have not identified more closely up to the present. According to 
Dr. Hildegarde Howard, ten species of birds are represented, two being 
extinct and the other eight apparently Recent. One of the fossil species is 
Chendytes lawi, a very large diving duck, first described by Dr. Loye 
Miller from the Upper San Pedro formation of Santa Monica Canyon. 
The other fossil bird is a hitherto undescribed gannet, of the genus Moris, 

Willett — Pleistocene Fauna from Baldwin Hills 383 

this genus being previously known on the Pacific coast by only one species, 
from the Miocene of Kern County. In the 'Condor' (vol. 38, 1936, p. 
213) Dr. Howard has named this gannet Moris reyana and has given a, 
detailed account of the avian remains found in this deposit. 

There is a goodly representation of fish material, more than 700 
specimens being preserved, but we have, as yet, not found any ichthyolo- 
gist who can give the time necessary to identify all of the elements. Teeth 
of at least two species of sharks are rather common, and teeth and stingers 
of rays are even more so. There are also teeth of a sheephead, and many 
ear bones, vertebrae and other elements unidentified as yet. 

Echinoderms were abundant, but very fragile, and perfect specimens 
difficult to obtain. According to Dr. U. S. Grant, the following are repre- 
sented: Strongylocentrotus sp. ?., Lorenia cardiformis A. Agassiz, and 
Dendraster sp. nov. The latter species is known only from late Pleistocene, 
the others being found living in this same latitude today. The Round 
Bryozoa, Lichenopora radiata (Audouin), the only member of the group 
found, was abundant and usually well preserved. The barnacles, identified 
by Dr. Howard R. Hill, are of three species. The Pink Barnacle, Balanus 
tintinnabulum californicus Pilsbry, was abundant; two specimens of Tet- 
raclita squamosa rubescens Darwin were found, also a number of seg- 
ments of the Whale Barnacle, Coronula regina Darwin. 

The decapod crustaceans were studied by Steve A. Glassell, who 
reports the following species : Callianassa longimana Stimpson, Darda- 
nus arnoldi Rathbun, Dromidia larraburei Rathbun, Randallia ornata 
(Randall) (by far, the most abundant species), Hepatus [meatus Rath- 
bun, Heterocrypta occidentalis (Dana) (second in numbers), Meso- 
rhoea idae Rathbun, Pyromaia tuberculata (Lockington), Pugettia pro- 
ducta (Randall), Pugettia richii Dana, Taliepus nuttallii (Randall), 
Loxorhynchus grand is Stimpson, Callinectes bellicosus Stimpson, Por- 
tunus xantusii (Stimpson), Cancer branneri Rathbun, Cancer anten- 
narius Stimpson, Cancer gracilis Dana, Cancer anthonyi Rathbun, Cancer 
productus Randall, Lopbopanopeus frontalis (Rathbun), Lophopanopeus 
diegensis Rathbun, and Cycloxanthops novemdentatus (Lockington) ; 
also three species not yet determined, probably undescribed. 

Mr. Glassell comments on this assemblage as follows : "Of the iden- 
tified species, the greater per cent are living here today. Three (Mesorhoea 
idae and two species as yet unnamed) are only known as fossils, but may, 
like two other species (Hepatus lineatus and Callinectes bellicosus) found 

384 San Diego Society of Natural History 

in this horizon, be represented at the present time in lower latitudes. In 
this lot also have been found three species of our present day fauna which 
have not previously been reported as fossil. These are Dromidia larraburei, 
Taliepus nuttallii, and Cancer antennarius. A striking feature of the col- 
lection is the absence of the remains of strictly inter-tidal forms. So far, 
not a single specimen of these numerous species has been observed, al- 
though an occasional one might well be expected. While all of the species 
are not intrinsically shallow water forms (some having a bathymetric 
range of over fifty fathoms), still the living ones may all be taken today 
from the extreme minus tide line to at least the fifteen fathom contour. 
Due to the preponderance of fragments of one or two species (nothing 
but pieces remaining), it might be inferred that the balance of species in 
this Pleistocene deposit was not the same as the present day fauna. This, 
however, is probably not the case, for it is safe to assume that only those 
crustacean processes have survived which were structurally able to do so." 

A study of the mollusks has resulted in recognition of 296 species, 
which divide as follows: Pelecypods, 90; Scaphopods, 5; Gastropods, 
201. Five genera and forty-eight species are added to the Calif ornian fossil 
list. These genera are Ensis, Siphonodentalium, Atys, Engina and Simnia. 
The species are Mytilus adamsianus Dkr., Rochefortia reyana sp. nov., 
Bornia cooki sp. nov., Petricola tellimyalis (Cpr.), Ensis californicus 
Dall, Dentalium numerosum Dall, Siphonodentalium quadrifissatum 
Dall, Carolina trispinosa Lesueur, Atys casta Cpr., Cancellaria bullata 
Sby., Engina strongi Pils. and Lowe, Purpura carpenteri (Dall) , Purpura 
petri (Dall) , Purpura gemma (Dall) , Purpura santarosana (Dall) , Thais 
biserialis (Blain.), Simnia catalinensis (Berry), Erato vitellina Hinds, 
Alabina tenuisculpta diegensis Bartsch, Cerithiopsis cosmia Bartsch, 
Cerithiopsis halia Bartsch, Cerithiopsis oxys Bartsch, Cerithiopsis ante- 
munda Bartsch, Rissoella sp. ?, Fartulum orcutti Dall, Fartulum occiden- 
tal Bartsch, Calyptraea contorta Cpr., Acmaea cassis nacelloides Dall, 
Tricolia substriata (Cpr.), Tegula pulligo (Mart.), Calliostoma glorio- 
sum Dall, Epitonium sawinae Dall, Turbonilla sanctorum Dall and 
Bartsch, Turbonilla superba Dall and Bartsch, Turbonilla vexativa Dall 
and Bartsch, Turbonilla antestriata Dall and Bartsch, Turbonilla almo 
Dall and Bartsch, Turbonilla adusta Dall and Bartsch, Turbonilla weldi 
Dall and Bartsch, Turbonilla ista Bartsch, Turbonilla canfieldi Dall 
and Bartsch, Turbonilla regina Dall and Bartsch, Odostomia eugena Dall 
and Bartsch, Odostomia nemo Dall and Bartsch, Odostomia donilla Dall 
and Bartsch, Odostomia helena Bartsch, and Lepidopleurus nexus (Cpr.) . 

Willett — Pleistocene Fauna from Baldwin Hills 385 

That there has been a mixing of faunas, though an exceedingly lim- 
ited one, is indicated by the presence of examples of the following ten 
species of a colder water fauna: Pecten hericeus Gld., Lora fidicula 
(Gld.), Spirotropis barbarensis (Dall), Spirotropis perversa (Gabb), 
Neptunea tabulata (Baird), Exilioidea rec tiros tris (Cpr.), Trophon 
orpheus (Gld.), Ranella oregonensis (Redf.), Epitonium wroblewskyi 
(Mbrch), and Tegula pulligo (Mart.). These are all Recent species, 
occurring at the present time either further to the northward or in deeper 
water in our latitude. The total number of individuals of these ten species 
in our collections is only seventeen, so that the true ratio of their abun- 
dance would be, not ten to 289, but seventeen to several hundred thou- 
sand, as all representatives of the cold water fauna were preserved, while 
the majority of warmer water forms were discarded. Most of these seven- 
teen specimens are more or less fragmentary and all are much eroded, 
their appearance thus indicating greater age than that of the remainder of 
the fauna. There is no doubt in my mind that the few representatives of 
this older fauna were already fossils at the time the others were living. 

These deposits have been referred to as being, for the greater part, 
representatives of a warm water fauna, and that the water was even less 
cold than it is today is indicated by the following facts. Of the 286 species 
(after deleting the ten older ones) , 261 occur living in this latitude today, 
and nineteen are found, so far as we know, only further to the southward, 
many of them being confined to Mexican waters. These are: Pecten 
vogdesi Arnold, Crassinella branneri (Arnold), Crassinella varians 
(Cpr.), Aligena cerritensis Arnold, Cardium procerum Sby., Mactra 
pallida Brd.' and Sby., Dentaliam numerosum Dall, Retusa carinata 
(Cpr.), Bulla punctulata A. Adams, Pseudomelatoma penicillata semiin- 
flata Grant and Gale, Mangelia cetolaca Dall, Cancellaria bulla ta Sby., 
Mitra fultoni E. A. Smith, Cantharus fortis (Cpr.), Nassarius cerritensis 
Arnold, Purpura leeana (Dall), Thais biserialis (Blain.), Turbonilla 
sanctorum Dall and Bartsch, and Turbonilla superba Dall and Bartsch. 
Six remaining species, Rochefortia reyana sp. nov., Bornia cooki sp. nov., 
Rissoina pleistocena Bartsch, Delphinoidea coronadoensis Arnold, Epi- 
tonium clarki T. S. Oldroyd, and Ischnochiton sanctaemonicae Berry are 
listed as extinct. However, a Recent specimen of Epitonium clarki, from 
Lower Californian waters, has been examined, and it is entirely possible, if 
not probable, that the other five may eventually be found living in mod- 
erate depths off Lower California, this being a region where very little 
shallow dredging has been done to date. 

386 San Diego Society of Natural History 

Among the interesting facts brought out by a study of the genetic 
relationship of the mollusks in this deposit, two features were emphasized 
particularly. First, in no single instance, where a sufficient amount of 
comparative Recent material was available, was there any perceptible dif- 
ference in either form or sculpture between fossil and Recent examples of 
a species. Second, the accumulation of an abundant representation of 
some supposed species, previously known by very few specimens, indi- 
cates that many characters generally used in differentiating species are 
extremely inconstant, and can be considered to represent only individual 
variation within the species. 

The literature most used in this study includes : Paleontology and 
Stratigraphy of San Pedro, California, by Ralph Arnold (1903) ; A Mon- 
ograph of West American Pyramidellid Mollusks, by Dall and Bartsch 
(1909) ; Marine Shellbearing Mollusks of the Northwest Coast of Ameri- 
ca, by W. H. Dall (1921 ) ; Marine Shells of the West Coast of North 
America, by Mrs. Ida S. Oldroyd (1924-27), and Pliocene and Pleisto- 
cene Mollusca of California, by Grant and Gale (1931). No attempt has 
been made to include a complete synonymy of the species, but where the 
names in the above works differ from those used in this paper, they are 
listed as synonyms. 

The following is a list of the mollusks, with remarks on some of the 
species. A number after the name of a species refers to the number of 
specimens preserved in the Los Angeles Museum, unless otherwise stated. 

Nucula (Nucula) exigua Sowerby. — Syn., Nucula suprastriata Cpr. 
(Arnold, 1903). — Abundant; many still in pairs. (550 pairs, 1150 valves). 

Leda taphria Dall. — Syn., Nuculana taphria (Dall) (Grant and Gale, 
1931). — Common. (30 pairs, 120 valves). 

Yoldia cooperi Gabb. — 1 fragment, including hinge, found by Miss 
Edna T. Cook. 

GlycYmeris septentrionalis (Middendorff). — Syn., G. subobsoleta 
(Cpr.) : G. barbarensis Conr.: G. corteziana Dall ?. — (110 valves). I am not 
sure of the above synonymy, but believe it correct, with the possible exception of 
the last name. Shells in my collection, identified by Dr. Dall as corteziana, are 
certainly the same as the fossils, but I have not seen the type of corteziana. 
These shells differ greatly individually as regards shape, thickness and amount of 

Ostrea lurida Carpenter. — 13 valves. 

Ostrea palmula Carpenter. — 1 valve. 

Pecten (Hinnites) multirugosus Gale. — Syn., Pecten giganteus Gray 
(Arnold, 1903) : Hinnites giganteus Gray (Dall, 1921; Oldroyd, 1924).— Not 

Willett — Pleistocene Fauna from Baldwin Hills 387 

very common, probably because of sandy character of locality. (7 valves). 

Pecten (Chlamys) hericeus Gould. — Syn., "Pecten hastatus Sby." 
(Grant and Gale, 193 1 ) . — 2 valves in White collection. 

Pecten (Aequipecten) latiauritus Conrad. — Abundant. (170 
valves) . 

Pecten (Aequipecten) circularis Sowerby. — Syn., P. c. aequisulca- 
tus Cpr. (Dall, 1921; Oldroyd, 1924).— Much less common than the last. (8 

Pecten (Pecten) stearnsii diegensis Dall. — 2 right valves, 1 left 
valve, 25 fragments. 

Pecten (Pecten) vogdesi Arnold. — Syn., P. dentatus, of some authors; 
not of J. Sowerby; P. excavatus, of some authors; not of Anton: P. cataractes 
Dall (Nautilus, 27, 1914, p. 121) : P. heimi Hertlein (Proc. Calif. Acad. Sci., 
Ser. 4, 14, 1925, p. 9). — 1 right valve and 9 fragments; 2 valves in White col- 

Lima dehiscens Conrad. — 5 valves. 

Anomia peruviana d'Orbigny. — Syn., A. lampe Gray (Arnold, 1903) . 
— Upper valves common, probably having drifted in from rocky localities. 

Pododesmus macroschisma (Deshayes) . — Much less plentiful than 
Anomia. 2 valves taken by the writer and 1 by Mrs. Fuller. 

Mytilus (Mytilus) californianus Conrad. — 1 half valve (with hinge) 
found by Mrs. Fuller; 1 pair in White collection. 

Mytilus (Mytilus) adamsianus Dunker. — 1 valve. 

Volsella modiolus (Linnaeus) . — Syn., Modiolus modiolus Linn. (Dall, 
1921; Oldroyd, 1924) .— 8 valves, 12 fragments. 

Volsella capax (Conrad). — Syn., Modiolus capax Conr. (Dall, 1921; 
Oldroyd, 1924) .— 3 valves. 

Volsella flabellata (Gould) .— Syn, Modiolus flabellatus Gld. (Dall, 
1921; Oldroyd, 1924).— 9 fragments. 

Lithophaga plumula (Hanley) . — 1 pair collected by Tom Burch. 

Periploma planiuscula Sowerby. — Syn, P. argentaria Conr. (Arnold, 
1903). — Hinge teeth rather common; occasional fragments of other sections of 

Thracia (Cyathodonta) undulata (Conrad). — Syn, Cyathodonta 
dubiosa Dall, C. pedroana Dall (Dall, 1921; Oldroyd, 1924).— 10 fragments. 

Pandora punctata Conrad. — Syn, Clidiophora punctata Conr. 
(Arnold, 1903) . — 2 pairs, 6 fragments. 

Crassinella branneri (Arnold). — Syn, Astarte branneri (Arnold, 
1903).— 180 valves. 

Crassinella varians (Carpenter). — (6 pairs, 45 valves). Not listed 
by Grant and Gale, but recorded by Woodring (Am. Journ. Sci, 29, 1935, p. 
303) from San Pedro Hills. 

Glans carpenteri (Lamy) . — Syn, Lazaria subquadrata Cpr. (Arnold, 

388 San Diego Society of Natural History 

1903) : Cardita subquadrata Cpr. (Dall, 1921; Oldroyd, 1924) : Glans minuscu- 
la (Grant and Gale, 1931) . — 2 valves. 

Chama pellucida Broderip. — 34 valves. 

Lucina (Myrtea) californica Conrad. — Syn., Phacoides calif ornicus 
Conr. (Dall, 1921; Oldroyd, 1924).— 1 valve. 

Lucina (Myrtea) nuttallii Conrad. — Syn., Phacoides nuttallii Conr. 
(Dall, 1921; Oldroyd, 1924) .— Abundant. (3 pairs, 85 valves) . 

Lucina (Myrtea) tenuisculpta approximata (Dall).— Syn., L. 
tenuisculpta Cpr. (Arnold, 1903, at least part) : Phacoides approximate Dall 
(Dall, 1921; Oldroyd, 1924) .— Abundant. (375 valves). This appears to be a 
southern race of L. tenuisculpta and it is probable that Arnold's "Upper San 
Pedro" specimens are referable to it. Southern shells are smaller, with accentuated 
radial sculpture. That the two forms exist in the same latitude, as has been in- 
ferred by many authors, is perhaps doubtful. 

Lucina (Here) excavata Carpenter. — Syn., Phacoides richthojeni Gabb 
(Dall, 1921; Oldroyd, 1924).— (67 valves). Gabb's richthojeni is undoubtedly 
the adult of Carpenter's excavata. 

Taras orbellus (Gould). — Syn., Diplodonta orbella Gld. (Arnold, 
1903; Dall, 1921; Oldroyd, 1924) .— 6 valves. 

Kellia suborbicularis laperousii (Deshayes) . — Syn., Chironia sub- 
orbicularis laperousii Desh. (Grant and Gale, 1931). — This species is so fragile 
that it occurs mostly in fragments. However, 16 valves in fair condition are pre- 
served. The name laperousii is used here solely because of the statement by Grant 
and Gale that Pacific coast specimens average larger than shells of the British 
Isles. These authors used the generic name Chironia with this species on the 
grounds that Heermannson (1847) named Cardium (Lasaea) rubrum Mont, 
as type of the genus Kellia. However, Winckworth (Journ. Conch., 20, 1934, p. 
52) calls attention to the fact that Recluz (Revue Zool. Cuv., 7, 1844, p. 295) 
had previously designated Mya suborbicularis Mont, as the type of Kellia. 

Aligena cerritensis Arnold. — 7 valves. 

Rochefortia aleutica (Dall). — Common. (40 pairs, 220 valves). 

Rochefortia reyana, sp. nov. 
Plate 25, figs. 1, 2 

Similar to R. pedroana Dall, but more equilateral, and with heavier and more 
elongated hinge teeth, the hinge line in the right valve occupying almost one-half 
of the margin of the valve. Left valve with one very small lamella immediately 
below the umbone, and deflected umbonal margin. 

Types, right and left valves, No. 1046 L. A. Mus., taken by the writer, with 
thirty-six additional valves and one connected pair, in the Del Rey Pleistocene 
deposit. Type right valve measures, in millimeters: diam., 6.7; alt., 5.2; ant. 
lateral, 4; post, lateral, 3: left valve, diam., 7.6; alt., 5.9. 

Bomia retifera Dall. — Like Kellia, this shell is very fragile and seldom 
found entire. It was probably more plentiful than the specimens preserved (1 
pair and 21 valves) would indicate. 

Willett — Pleistocene Fauna from Baldwin Hills 

Plate 25 

Fig. 1. Rochejortia reyana Willett, type, right and left valves; x 4. 
Fig. 2. Bornia cooki Willett, type, left valve; x 4. 
Fig. 3. Bornia cooki Willett, type, right valve; x 4. 

Willett — Pleistocene Fauna from Baldwin Hills 389 

Bornia cooki, sp. nov. 
Plate 25, figs. 3-6 

Shell thin, white, moderately convex, oblong, inequilateral; beaks small, dis- 
tinct, situated at posterior third of shell. Surface marked by numerous concentric 
striations and growth lines of varying strength; also by several faint, rounded, 
radial ridges which start near the center of the valve and run to the ventral mar- 
gin. In the type there are four of these ridges and trace of a fifth, and in the para- 
type, in Miss Cook's collection, the ridges are fainter but more numerous (6-7) . 
The shagreened pattern, usual to members of the genus, is only perceptible near 
the margins, but it is possible that this may have been worn away on the earlier 
portions of the shell. Dentition similar to that of B. ret if era Dall, but with 
shorter laterals and wider notch. 

Type pair, No. 1047 L. A. Mus., collected, together with another right 
valve, by Miss Edna T. Cook, for whom it is named. The type measures, in 
millimeters: diam., 9.9; alt., 6.4. The paratype, in Miss Cook's collection, meas- 
ures: diam., 11.4; alt., 6.8. 

Except in dentition, this species is quite different from B. retifera Dall, the 
only other member of the genus found in this deposit. In its oblong, inequilateral 
form it is more similar to some species of the genera Erycina, Montacuta and 
Sportella, but its dentition would seem to place it with Bornia. 

Cardium (Laevicardium) elatum Sowerby. — 8 valves; many frag- 
ments noted. 

The writer prefers not to follow some recent authors who have divided this 
old familiar genus. He believes that the various divisions in the group may be 
satisfactorily indicated by using subgeneric names, as was done by Dr. Dall 

Cardium (Laevicardium) substriatum Conrad. — 1 valve found. 

Cardium (Laevicardium) procerum Sowerby. — Common. (60 
valves) . 

Cardium (Laevicardium) quadragenarium Conrad. — The most 
plentiful species of the genus. 

Cardium (Fragum) biangulatum Broderip and Sowerby. — Fairly 
common. (30 valves). 

Venus (Antigona) fordii Yates. — 1 immature valve. 

Venus (Chione) succincta Valenciennes. — Syn., Chione undatella 
Sby. (Dall, 1921; Oldroyd, 1924) : Venus neglecta Sby., V. simillima Sby. 
(Arnold, 1903).— 21 valves. 

Venus (Chione) fluctifraga Sowerby. — 3 valves. 

Venerupis (Callithaca) tenerrima (Carpenter). — Syn., Tapes 
tenerrima Cpr. (Arnold, 1903) : Paphia tenerrima Cpr. (Dall, 1921; Oldroyd, 
1924) . — Fairly common, but usually broken. (2 pairs, 4 hinges) . 

Venerupis (Protothaca) staminea (Conrad) . — Syn., Tapes staminea 
Conr. (Arnold, 1903): Paphia staminea Conr. (Dall, 1921; Oldroyd, 1924). — 
Less plentiful than the last. (3 valves, 2 fragments) . 

390 San Diego Society of Natural History 

Compsomyax subdiaphana (Carpenter) .— Syn., CalUsta subdia- 
phana Cpr. (Arnold, 1903): Marcia subdiaphana Cpr. (Dall 1921; Oldroyd, 
1924): Clementia subdiaphana Cpr. (Grant and Gale, 1931). — (2 valves). 
Dr. U. S. Grant informs me that he now considers this species generically differ- 
ent from Clementia. 

Transenella tantilla (Gould) . — Syn., Psephis tantilla Gld. (Arnold, 
1903).— 5 valves. 

Tivela (Pachydesma) stultorum (Mawe).— Syn., T. crassatelloides 
Conr. (Arnold, 1903). — (2 valves). There appears to be some doubt whether 
Mawe's Donax stultorum, stated to be from "Indian Seas," is really the same as 
this species, though his figure shows a similar shell. Possibly Conrad's name 
crassatelloides should be revived. 

Saxidomus nuttalli Conrad. — Syn., S. aratus Gld. (Arnold, 1903). — 
1 valve. 

Pitar newcombianus (Gabb) . — Syn., Callista newcombiana Gabb 
(Arnold, 1903): Pitaria newcombiana Gabb (Dall, 1921; Oldroyd, 1924).— 
1 valve. 

Amiantis callosa (Conrad) . — Syn., Callista callosa Conr. (Arnold, 
1903).— Abundant. 

Petricola tellimyalis (Carpenter) . — 70 valves. 

Petricola californiensis Pilsbry and Lowe. — Syn., P. denticulata Sby. 
(Arnold, 1903; Dall, 1921; Oldroyd, 1924; Grant and Gale, 1931).— 2 valves. 

Petricola carditoides (Conrad) . — 1 valve collected by Miss Edna 

Cooperella subdiaphana Carpenter. — 4 valves (three in Miss Cook's 

Tellina idae Dall. — Common. ( 10 pairs, 26 valves) . 

Tellina buttoni Dall. — 3 valves. 

Tellina bodegensis Hinds. — 8 valves. 

Tellina santarosae Dall. — 3 valves. 

Apolymetis biangulata (Carpenter) . — Syn., Metis alta Conr. (Arn- 
old, 1903; Dall, 1921; Oldroyd, 1924).— Common. (5 pairs, 2 valves). 

Macoma nasuta (Conrad) . — 5 valves. 

Macoma yoldiformis Carpenter. — 6 pairs, 22 valves. 

Macoma secta (Conrad). — Common. (15 valves). 

Macoma indentata Carpenter. — Syn., M. i. tenuirostris Dall (Dall, 
1921; Oldroyd, 1924) .— Abundant. (5 pairs, 40 valves) . 

Semele decisa (Conrad.) — 1 fragment found by the writer, another 
by Miss Cook. 

Semele pulchra (Sowerby) . — 7 valves. 

Donax califomicus Conrad. — 1 valve; 2 valves in Miss Cook's collec- 

Donax gouldii Dall. — Syn., D. laevigata Desh. (Arnold, 1903). — 

Willett — Pleistocene Fauna from Baldwin Hills 391 

Abundant. (30 valves). 

Gari edentula (Gabb). — Syn., Psammobia edentula Gabb (Arnold, 
1903; Dall, 1921; Oldroyd, 1924) .— 3 fragments, with hinges. 

Tagelus californianus (Conrad) . — 1 fragment, with hinge. 

Tagelus subteres (Conrad) . — 1 valve. 

Solen sicarius Gould. — 20 fragments, with hinges. 

Ensis califomicus Dall. — 24 valves. 

Siliqua lucida (Conrad) . — 1 valve, 7 hinges. 

Mactra (Mactra) califomica Conrad. — Common. (23 valves). 

Mactra (Spisula) planulata Conrad. — Syn., "Mactra falcata Gld." 
(Arnold, 1903): Spisula planulata Conr. (Dall, 1921; Oldroyd, 1924).— 
Abundant. (1 pair, 50 valves). 

Mactra (Spisula) hemphilli Dall. — Common. (3 pairs, 21 valves). 

Mactra (Spisula) catilliformis (Conrad) . — 1 valve. 

Mactra (Mulinia) pallida modesta (Dall). — Syn., "Mactra exo 
leta Gray" (Arnold, 1903) . — Rather common. (3 pairs, 45 valves) . 

Schizothaerus nuttallii (Conrad). — Syn., Tresus nuttalli Conr 
(Arnold, 1903) . — Not rare, but mostly fragmentary. (2 pairs) . 

CrYptomya califomica (Conrad) . — Abundant. (8 pairs, 60 valves) 

Corbula (Lentidium) luteola Carpenter. — Abundant; pairs of con 
nected valves common. (250 pairs, 100 valves) . 

Panope (Panope) generosa Gould. — Syn., Panopea generosa Gld 
(Arnold, 1903; Oldroyd, 1924). — Fairly common. (1 pair, 6 valves, 5 hinges) 

Saxicava arctica (Linnaeus). — 1 valve. 

Pholas pilsbryi Lowe.— Syn., TJnphaea gabbi Tryon (Arnold, 1903 
Dall, 1921; Oldroyd, 1924): Pholas gabbi Tryon (Grant and Gale, 1931). — 
1 valve, 1 fragment. 

Pholadidea (Pholadidea) penita (Conrad). — 1 pair. 

Dentalium neohexagonum Sharp and Pilsbry. — Syn., Dentalium 
pseudohexagonum Dall (Arnold, 1903). — Abundant. (700). 

Dentalium numerosum Dall. — Six specimens seem referable to this 
species. In addition to these are numerous examples, referred to neohexagonum, 
that have more ribs than the typical of that form and, although complete inter- 
gradation between neohexagonum and numerosum is not shown in our series, 
rather close relationship between the two appears to be indicated. 

Dentalium semipolitum Broderip and Sowerby. — 58 specimens, mostly 
more or less fragmentary. 

Siphonodentalium quadrifissatum Dall. — 17 specimens. 

Cadulus fusiformis Pilsbry and Sharp. — Abundant. (1300). Consider- 
ing the abundance of this species in a deposit so near those worked by Arnold, 
it is difficult to understand why he did not find it. Pilsbry (Nautilus, 17, 1904, 
p. 108) believed Arnold's figure of "Cadulus nitentior Cpr." to be "probably of a 

392 San Diego Society of Natural History 

serpuloid annelid," but, if possible, this should be checked by a study of Arnold's 
material. There is some variation in this species in both shape and diameter, and 
Arnold's figure may represent a worn Cadulus. 

Cavolina telemus tricuspida (Rivers). — Syn. Carolina occidentals 
Dall (Dall, 1921; Oldroyd, 1927).— 1 specimen collected by Miss Edna T. 

Cavolina trispinosa Lesueur. — 1 in Museum collection and 3 in col- 
lection of Miss Cook. 

Acteon (Acteon) traski Stearns. — Rather common, though usually 
broken. (225). 

Acteon (Rictaxis) punctocaelatus (Carpenter) . — 38. 
Retusa (Acteocina) culcitella (Gould). — Syn., Tornatina culcitella 
Gld., T. cerealis Gld. (Arnold, 1903) : Acteocina culcitella Gld. (Dall, 1921; 
Oldroyd, 1927) : Acteocina pedroensis T. S. Oldroyd (Proc. U. S. Nat. Mus., 
65, 1925, art. 22, pp. 23, 24) .— Common. (600) . 
Retusa (Acteocina) carinata (Carpenter) . 

Retusa (Acteocina) inculta (Gould) . — More than 600 specimens of 
the short, blunt-spired Retusas were preserved. While the majority of these appear 
referable to carinata, a few are indistinguishable from inculta and others are vari- 
ously intermediate between the two. A sufficient number of Recent specimens 
will probably show that carinata and inculta are not more than subspecifically 
distinct, the former being a southern form and the latter a more northern one 
of the same species. 

Volvulella cylindrica (Carpenter) . — Syn., Volvula cylindrica Cpr. 
(Arnold, 1903). — (700). A careful examination of this splendid series shows 
much variation in size, length of spire, and amount of spiral sculpture. It is prob- 
able that some other named species are only variants of cylindrica. 

Atys casta Carpenter. — 1 juvenile specimen. 

Cylichna attonsa Carpenter. — Syn., Cylichna alba Brown (Arnold, 
1903, at least part) : Cylichnella attonsa Cpr. (Dall, 1921; Oldroyd, 1927). — 
(1000) . Although there is considerable variation in this series, it seems advisable 
to refer them all to the above species, of which many are typical. Some specimens 
approach C. diegensis Dall, which may be the same as C. propinqua Smith. 
None appears referable to C. alba Brown, which name, in the past, has been used 
for most southern Californian fossils. This latter species is probably confined to 
northern waters and, if it has appeared at all as a fossil in southern California, it 
should be only in a cold water fauna. 

Bulla punctulata A. Adams. — Syn., Bullus punctulatus (A. Ad.) 
(Grant and Gale, 1931). — (62). Our specimens assigned to this species differ 
from available Lower Californian examples in larger size (largest, 48x33 mm.), 
slightly more globular form, and fewer (3-6) spirals in the umbilicus. This is 
probably the shell that Pilsbry (Man. Conch., 15, 1893, p. 341) refers doubt- 
fully to B. aspersa A. Adams. The difference in number of spirals in the um- 
bilicus does not appear to coincide with different localities, as both types are 
present in specimens from the west coast of South America in the H. N. Lowe 

Willett — Pleistocene Fauna from Baldwin Hills 393 

collection. For use of Bulla instead of Bullus, see Pilsbry, Nautilus, 44, 1931, 
p. 98. 

Haminoea vesicula (Gould) . — 1 juvenile. 

Melampus olivaceous Carpenter. — (14). These undoubtedly washed 
down from coastal marshes. 

Williamia peltoides (Carpenter). — (26). The species represented is 
the one with elevated apex. Whether the above name is correctly applied here may 
be open to question (see Grant and Gale, 1931, p. 464). 

Terebra (Strioterebrum) pedroana Dall. — Syn., T. simplex Cpr. 
(Arnold, 1903) : T. pedroana philippiana Dall (Dall, 1921; Oldroyd, 1927). — 
(260) . Abundant. The typical and the variant named philippiana both present. 

Conus calif ornicus Hinds. — Rather common. (16) . 

Megasurcula remondii (Gabb). — Syn., Cryptoconus stearnsianus 
Raymond (Dall, 1921; Oldroyd, 1927) : Surculites remondii (Gabb) (Grant 
and Gale, 1931).— 41. 

Megasurcula carpenteriana (Gabb). — Syn., Pleurotoma carpen- 
teriana Gabb, P. tryoniana Gabb (Arnold, 1903) : Cryptoconus car penterianus 
Gabb, C. tryonianus Gabb, C. tremperianus Dall (Dall, 1921; Oldroyd, 1927) : 
Surculites car penterianus (Gabb) (Grant and Gale, 1931) . — (130) . A common 
and very variable species. 

Lora fidicula (Gould) . — Syn., Bela fidicula GId. (Arnold, 1903, part) : 
"Lora viridula Fabr." (Grant and Gale, 1931). — 1 specimen collected by Miss 
Edna T. Cook. 

Spirotropis (Borsonella) barbarensis (Dall) . — Syn., Borsonella 
barbarensis Dall (Dall, 1921; Oldroyd, 1927): "Borsonella dalli Arnold" (Dall, 
1921, part; Oldroyd, 1927, part). — 1 specimen in Museum collection and an- 
other in collection of Mrs. E. M. Clark. 

Spirotropis ( Antiplanes) perversa (Gabb) . — Syn., Pleurotoma per- 
versa Gabb (Arnold, 1903) : Antiplanes perversa Gabb (Dall, 1921; Oldroyd, 
1927) . — 8 specimens, all much worn and few entire. Evidently of an older fauna 
than the bulk of the deposit. 

The writer cannot follow Grant and Gale in relegating such species as 
rotida, santarosana and catalinae to the synonymy of perversa. They appear to 
have not only different forms, but different ranges. 

Moniliopsis incisa fancherae (Dall) . — Syn., "Drillia inermis Hds." 
(Arnold, 1903) : Clathrodrillia halcyoms Dall (Dall, 1921; Oldroyd, 1927). — 
Abundant. (150). 

Moniliopsis incisa ophioderma (Dall) . — Syn., "Drillia inermis peni- 
cillata Cpr." (Arnold, 1903) : "Moniliopsis incisa Cpr." (Dall, Proc. U. S. Nat. 
Mus., 56, 1919, pi. 12, fig. 7; Oldroyd, 1927, pi. 18, fig. 3).— (10). Much less 
common than the last. 

There has been much confusion among authors regarding the names to be 
applied to the varieties of this well known species. It would seem that the correct 
application of names depends entirely upon the identity of Carpenter's type of 
incisa, which does not appear to be definitely established. The arrangement here 

394 San Diego Society of Natural History 

used is based on the assumption that typical incisa is the northern form with 
"grooved" spirals and axial sculpture confined to faint growth lines. This is the 
shell figured by Grant and Gale (1931, pi. 26, fig. 21) as the "typical variety," 
but is not the same as some of the forms included in their synonymy. 

The fact is that we have in southern California, both fossil and Recent, two 
common varieties of M. incisa, each of which has been referred to by several 
names. One type (jancherae, as used here) is more slender, with rounder body 
whorl, and sharper spiral sculpture, and is a dredged shell. The other (ophio- 
derma), frequently collected at low tide, is characterized by greater diameter, 
more or less flattened body whorl, less sharp spiral sculpture, and (in life) verti- 
cal reddish lines. Dall's figures of both incisa and ophioderma (Proc. U. S. Nat. 
Mus., 56, 1919, pi. 12, figs. 5 and 7) appear to be of this latter form. M. rhines 
Dall (cancellata Cpr.) is probably a color form of jancherae, Carpenter's de- 
scription calling for a white shell. Such specimens are in the writer's collection 
from Catalina Island. 

Clavus (Cymatosyrinx) empyrosia (Dall). — Syn., "C. pallidas 
Sby." (Grant and Gale, 1931, part).— 1. 

Clavus (Cymatosyrinx) halocydne (Dall). — Syn., "C. pallidas 
Sby." (Grant and Gale, 1931, part).— (3). Although Grant and Gale place 
this and the last species in the synonymy of C. pallidus (Sby.), the writer does 
not consider such action justified. A comparison of halocydne and pallidus shows 
that the latter is larger and relatively wider, and has a much heavier callus on the 
inner lip and a narrower constricted area at the suture. In halocydne this constric- 
tion, on all whorls but the last, is almost as wide as the remainder of the whorl. 
Furthermore, the color of halocydne is not white like pallidus, but light brown, 
darker in the aperture. C. empyrosia differs from halocydne and pallidus in both 
size and sculpture. 

Clavus (Cymatosyrinx) hemphilli (Stearns). — Syn., Dnllia hemp- 
hilli Sts. (Arnold, 1903): Cymatosyrinx hemphilli Sts. (Dall, 1921; Oldroyd, 
1927) : C. aeolia Dall (Proc. U. S. Nat. Mus., 56, 1919, p. 11).— (16). All 
are of the ribbed form called aeolia by Dr. Dall. A fine series of topotypes of 
hemphilli collected by Mr. and Mrs. P. M. Connelly at Todos Santos Bay, 
Lower California, are mostly quite different, both in sculpture and color, from 
Los Angeles County specimens of "aeolia," but enough intergrades have been 
examined to show that the two are conspecific. It is possible that aeolia may be a 
geographical race of hemphilli, but this remains to be demonstrated. 

Clavus (Crassispira) montereyensis (Stearns) . — Syn., Crassispira 
arsino'e Dall (Proc. U. S. Nat. Mus., 56, 1919, p. 26).— 1 specimen found by 
Miss Alice Waterbury and donated to the Museum. 

Mangelia (Mangelia) hexagona Gabb. — Syn., Mangdia branneri 
Arnold (Arnold, 1903; Dall, 1921).— 7. 

Mangelia (Mangelia) merita (Hinds). — 1. 

Mangelia (Bela) variegata Carpenter. — Syn., M. angulata Cpr., not 
Reeve (Arnold, 1903; Dall, 1921) : M. oenoa Dall, M. pulchnor Dall, M. beta 
Dall (Dall, 1921; Oldroyd, 1927) : M. barbarensis Oldroyd (1927) : "M. hece- 
tae Dall and Bartsch" (Grant and Gale, 1931). — (1800). Our specimens of 

Willett — Pleistocene Fauna from Baldwin Hills 395 

this species exhibit every variation between typical variegata and the other forms 
listed in the above synonymy. Plate 26, fig. 1, shows intergradation between the 
two extremes, typical variegata on the one hand, and the angulated variety on the 
other. That this intergradation also occurs at the present time is indicated by 
specimens in the writer's collection. While, as Grant and Gale point out (1931, 
p. 593), the shell of M. hecetae Dall is indistinguishable from some specimens 
of the angulated form of variegata, an example of hecetae in the writer's collec- 
tion, taken in southeastern Alaska, possesses an operculum, which, according to 
our present understanding, would place it in the genus Lora. 

Mangelia (Bela) cetolaca Dall. — Syn., Columbella (Aesopus) old- 
roydi (Arnold, 1903, p. 238), not Mangilia oldroydi (Arnold, 1903, p. 213): 
"Mangelia perattenuata Dall" (Grant and Gale, 1931). — (720). Grant and 
Gale considered M. perattenuata Dall, Pbilbertia phylira Dall, and P. amyela 
Dall identical with this species. However, an examination of our large series does 
not appear to substantiate their views. Perattenuata seems more tapering than 
cetolaca, with sutures far too narrow, and with the last whorl longer than the rest 
of the shell, which is not the case in specimens of cetolaca the same size as the 
type of perattenuata. Philbertia phylira has fewer and more regularly spaced 
spiral cords than M. cetolaca, and Philbertia amyela has too few axials. 

Mangelia (Bela) arteaga roperi Dall. — Syn., "Mangilia sculpturata 
Dall" (Arnold, 1903). — (360). Our specimens are uniformly more slender 
than examples of the typical form. It is probable that roperi is a southern race 
and that typical arteaga does not range as far southward as has been generally 

Cancellaria bullata Sowerby. — Rather rare. 1 specimen in the Museum 
collection, and 3 in the White collection. 

Cancellaria crawfordiana Dall. — 2 specimens found, one by the 
writer and the other by Mrs. E. M. Clark. 

Cancellaria cooperi Gabb. — 1 broken specimen. 

Olivella biplicata (Sowerby). — Common. (76). 

Olivella baetica Carpenter. — Abundant. (170). 

Hyalina (Cypraeolina) pyriformis (Carpenter) . — Syn., Merovia 
pyrijormis Cpr. (Dall, 1921): Cypraeolina pyriformis Cpr. (Oldroyd, 1927). — 
1 specimen collected by Mrs. Clark. 

Mitra idae Melvill. — Syn., "Mitra maura Swain." (Arnold, 1903): 
Stngatella idae Mel. (Dall, 1921) . — 1 specimen in White collection. 

Mitra fultoni E. A. Smith.— 11. 

Mitra catalinae (Dall). — 10. 

Fusinus barbarensis (Trask). — Syn., Fusus barbarensis Trask 
(Arnold, 1903).— 1. 

Fusinus arnoldi (Cossman) . — Syn., Fusus rugosus Trask (Arnold, 
1903) : Fusinus trash Dall (Dall, 1921; Oldroyd, 1927).— 22. 

Fusinus kobelti (Dall) . — 6. 

Fusinus monksae (Dall). — Syn., "Fusus robustus Trask" (Arnold, 

396 San Diego Society of Natural History 

1903).— 1. 

Fusinus luteopictus (Dall) . — (130) . By far, the most common species 
of the genus. 

Kelletia (Kelletia) kelletii (Forbes). — Syn., Siphonalia kellettii Fbs. 
(Arnold, 1903). Rather common. (9). 

Cantharus fortis (Carpenter). — Syn., Pisania fortis Cpr. (Arnold, 
1903) . — 1 specimen; 2 additional in White collection. 

Neptunea (Sulcosipho) tabulator (Baird). — Syn., Chrysodomus 
tabulatus Baird (Arnold, 1903; Dall, 1921; Oldroyd, 1927).— 3 fragments. 

Exilioidea rectirostris (Carpenter). — Syn., Chrysodomus rectirostris 
Cpr. (Arnold, 1903): Exilia rectirostris Cpr. (Dall, 1921; Oldroyd, 1927).— 
1 specimen. 

Engina strongi Pilsbry and Lowe. — Syn., "Engina carbonaria Rve." 
(Dall, 1921; Oldroyd, 1927). — 2 specimens; an additional one in Miss Cook's 

Nassarius (Zeuxis) tegula (Reeve) . — Syn., Nassa tegula Rve. (Arn- 
old, 1903) : Alectrion tegula Rve. (Dall, 1921; Oldroyd, 1927) .— 55. 

Nassarius (Schizopyga) califomianus (Conrad) . — Syn., Nassa 
californiana (Conr.) (Arnold, 1903) : Alectrion calij orniana Conr. (Dall, 1921; 
Oldroyd, 1927).— 290. 

Nassarius (Schizopyga) cerritensis (Arnold) . — Syn., Nassa cerri- 
tensis Arn. (Arnold, 1903) : Alectrion cerritensis Arn. (Dall, 1921; Oldroyd, 
1927). — (160). Our series appears to show that this species grades into N. 
califomianus at one end, and approaches very near to N. mendicus cooperi at the 
other, though none have as few axial ribs as cooperi. 

Nassarius (Schizopyga) mendicus cooperi (Forbes). — Syn., 
Nassa mendica cooperi Fbs. (Arnold, 1903): Alectrion cooperi Fbs. (Dall, 
1921; Oldroyd, 1927).— 135. 

Nassarius (Schizopyga) perpinguis (Hinds). — Syn., Nassa per- 
pinguis Hds. (Arnold, 1903) : Alectrion perpinguis Hds. (Dall, 1921; Old- 
royd, 1927) .— 260. 

Nassarius (Schizopyga) fossatus (Gould) . — Syn., Nassa fossata 
(Gld.) (Arnold, 1903): Alectrion fossata Gld. (Dall, 1921; Oldroyd, 1927).— 

Nassarius (Schizopyga) insculptus (Carpenter) . — Syn., Nassa in- 
sculpta Cpr. (Arnold, 1903) : Alectrion insculptus Cpr. (Dall, 1921; Oldroyd, 
1927).— 2. 

Mitrella carinata (Hinds). — Syn., Columbella carinata Hds. (Arnold, 
1903; Dall, 1921; Oldroyd, 1927) : C. carinata hind si (Gask.) Rve. (Dall, 1921; 
Oldroyd, 1927). 

Mitrella carinata gausapata (Gould) . — Syn., Columbella gausapata 
Gld. (Arnold, 1903; Dall, 1921; Oldroyd, 1927) : C. californiana Gask. (Arn- 
old, 1903): C. carinata californiana Gask. (Dall, 1921; Oldroyd, 1927). — 
(400) . In this series are examples typical of each of the two above forms and 

Willett — Pleistocene Fauna from Baldwin Hills 397 

many intergrades between them. If only southern Californian specimens were con- 
sidered, there would seem to be no justification in recognition of more than one 
race, as our shells, both fossil and Recent, show no point of division. From what 
is known at the present time, however, it appears that the range of the form 
gausapata extends considerably farther north than typical carinata. In a sense, 
therefore, they may be considered geographical races. 

Mitrella tuberosa (Carpenter). — Syn., Columbella tuberosa Cpr. (Arn- 
old, 1903; Dall, 1921; Oldroyd, 1927).— 130. 

Amphissa reticulata Dall. — 3. 

Amphissa versicolor Dall. — (40) . Because of the great amount of 
variation in these species, I find it difficult to separate reticulata from versicolor, 
especially in the case of immature specimens. Three are referred to reticulata 
largely on account of their greater size. 

Amphissa undata Carpenter. — 7. 

Purpura (Pteropurpura) carpenteri (Dall). — Syn., Murex carpen- 
feri Dall (Dall, 1921; Oldroyd, 1927).— 8. 

Purpura (Pteropurpura) petri (Dall).— Syn., Murex petri Dall 
(Dall, 1921; Oldroyd, 1927).— 46. 

Purpura (Centrifuga) leeana (Dall). — Syn., Murex leeanus Dall 
(Arnold, 1903). — (70). A rather common species, a fine growth series being 
preserved. An interesting feature is the similarity of the young of this species to 
half-grown Tritonalia barbarensis (Gabb) (see Plate 26, figs. 2, 3) . 

Purpura (Jaton) festiva (Hinds).— Syn., Murex festivus Hds. (Arn- 
old, 1903; Dall, 1921; Oldroyd, 1927) .— Abundant. (100). 

Purpura (Jaton) gemma (Sowerby). — Syn., Murex gemma Sby. 
(Dall, 1921; Oldroyd, 1927) .— 6. 

Purpura (Jaton) santarosana (Dall). — Syn., Murex santarosana 
Dall (Dall, 1921; Oldroyd, 1927).— 1. 

Tritonalia foveolata (Hinds). — Syn., Ocinebra foveolata Hds. (Arn- 
old, 19C3). — (40). Arnold's record of "Ocinebra perita Hds." may refer to 
this species. 

Tritonalia interfossa (Carpenter) . — Syn., Ocinebra interfossa Cpr. 
(Arnold, 1903). — 2, one typical and the other near to the form beta Dall. 

Tritonalia poulsoni (Nuttall in Carpenter) . — Syn., Ocinebra poulsoni 
Nutt. (Arnold, 1903). — (85). The most common member of the genus. 
Thais biserialis (Blainville). — 38. 

Thais emarginata (Deshayes).— Syn., Purpura saxicola Val. (Arnold, 
1903) .— 2 in White collection 

Acanthina spirata (Blainville). — Syn., Monoceros engonatum Conr. 
(Arnold, 1903). — (85). This series varies from the high spired form, with 
rounded whorls, to the short, carinated one. 

Trophon (Boreotrophon) orpheus (Gould). — 1. 

Forreria belcheri (Hinds).— Syn., Chorus belcberi Hds. (Arnold, 

398 San Diego Society of Natural History 

1903) . — Common. 

Bursa califomica (Hinds) . — Syn., Ranella California Hds. (Arnold, 
1903).— Abundant. 

Ranella (Priene) oregonensis (Redfield).— Syn., Tritonium ore- 
gonensis Redf. (Arnold, 1903): Argobuccinum oregonensis Redf. (Dall, 1921; 
Oldroyd, 1927).— 1 collected by Miss Edna Cook and another by J. C. Marsh. 

Simnia (Neosimnia) catalinensis (Berry) .— Syn., Neosimnia cata- 
linensis (Berry, Nautilus, 30, 1916, p. 21).— 2 specimens in Museum collection 
and 2 more in White collection. Our largest measures 24x9 millimeters, and a 
Recent specimen in the writer's collection measures 32.5x12 millimeters. When 
these measurements are considered, the statement of F. A. Schilder (Proc. Mai. 
Soc. London, 20, 1932, p. 54) that catalinensis is "evidently the young of 
loebbeckeana Weinkauff" would seem palpably erroneous. 

Cypraea spadicea Swainson. — 4. 

Trivia califomiana (Gray).— Syn., T. califomica Gray (Arnold, 
1903).— 3. 

Trivia solandri (Gray in Sowerby) . — 1. 

Erato vitellina Hinds. — 1. 

Erato columbella Menke. — 4. 

Alabina tenuisculpta diegensis Bartsch.— 5. 

Bittium (Lirobittium) omatissimum Bartsch. — 1 collected by Tom 

Bittium (Semibittium) rugatum Carpenter. — 2 collected by the writer 
and another by Miss Edna Cook. One of the features of this deposit was the 
scarcity of Bittiums which are usually so abundant in our Pleistocene localities. 

Cerithidea califomica (Haldeman).— (29). Undoubtedly washed 
down from salt marshes. 

Seila montereyensis Bartsch. — Syn., "Sella asslmilata C. B. Ad." 
(Arnold, 1903) . — (100) . The commonest species of the family. 

Cerithiopsis antefilosa Bartsch. — 2. 

Cerithiopsis cosmia Bartsch. — 29. 

Cerithiopsis oxys Bartsch. — 16. 

Cerithiopsis antemunda Bartsch. — 8. 

Cerithiopsis halia Bartsch. — 3. 

Triphora pedroana (Bartsch). — Syn., Trlfora pedroana Bart. (Dall, 
1921).— 1 in White collection. 

Rissoella sp. ? — 18 specimens tentatively referred to this genus, but 
absence of opercula and soft parts makes the assignment uncertain. These re- 
semble somewhat elongated specimens of Syncera translucens (Cpr.), but they 
are narrowly umbilicated and spirally striated. This may be the species described 
by Bartsch (Proc. U. S. Nat. Mus., 70, 1927, p. 31) as Rissoella ? califomica, 
but it appears to differ from the figure of that species in much rounder body whorl 
and less open umbilicus. 

Willett — Pleistocene Fauna from Baldwin Hills 399 

Rissoina kelseyi (Dall and Bartsch). — Syn., Alaba oldroydi Dall 
(Nautilus, 19, 1905, p. 15).— 3. 

Rissoina pleistocena Bartsch. — 1 specimen collected by Mrs. E. M. 

Turritella jewettii Carpenter. — 1 very worn specimen found by J. C. 

Turritella cooperi Carpenter. — 38. 

Vermicularia eburnea (Reeve). — 4. 

Aletes squamigerus Carpenter. — Syn., Serpidorbis squamigerus Cpr. 
(Arnold, 1903). — 19 specimens, one of which is probably referable to A. s. 
pennatus (Morch). Whether this latter form is of any ecological significance is 

Spiroglyphus lituellus (Morch). — 1. 

Micranellum crebricinctum (Carpenter) . — Syn., Caecum crebricinc- 
tum Cpr., "Caecum magnum Stearns" (Arnold, 1903) : Micranellum pedroense 
Bartsch (Dall, 1921; Oldroyd, 1927; Grant and Gale, 1931) .— Common. 

It appears to the writer that M. pedroense Bartsch, and "Caecum magnum 
Sts." as figured by Arnold, are the young of M. crebricinctum Cpr. This species, 
during juvenility, is slender, rather strongly curved, and the plug is longer and 
narrower; as it becomes older, the slender part of the shell is discarded; some of 
the curve being lost, and the plug becomes thicker and more blunt. 

Fartulum orcutti (Dall). — 3. 

Fartulum occidentale Bartsch. — Common. (920). 

Littorina scutulata Gould. — 2 immature specimens. 

Lacuna unifasciata Carpenter.— 65. 

Iselica fenestrata (Carpenter) . — Syn., Fossarus jenestrata Cpr. (Arn- 
old, 1903) .— 1 collected by Miss Edna T. Cook. 

Hipponix antiquatus cranioides Carpenter. — 2. 

Hipponix tumens Carpenter. — 2. 

Crepidula onyx Sowerby. — Common. 

Crepidula excavata (Broderip). — Abundant. 

Crepidula lingulata Gould. — Syn., Crepidula dorsata Brod. (Arnold, 
1903 ) . — Common. 

Crepidula nummaria Gould. — Syn., C. navicelloides Nutt. (Arnold, 
1903) : "C. mvea C B. Ad." (Oldroyd, 1927) .— Common. 

Crepidula nummaria glottidiarum Dall. — (30). An interesting 
feature in the presence of this race is that no trace was found of the Brachiopod, 
Glottidia, upon which it undoubtedly lived. 

Crucibulum spinosum (Sowerby). — 7. 

Calyptraea contorta Carpenter. — Syn., Galerus mammillaris Brod. 
(Arnold, 1903, at least part) : Calyptraea mammillaris Brod. (Grant and Gale, 
1931, part) . — (380) . This is probably the only species of the genus to be found 

400 San Diego Society of Natural History 

in the Upper San Pedro formation, but it is possible that the more northern C. 
fastigiata Gld. occurred in earlier periods. Contorta may be a stunted, southern 
form of fastigiata, but both differ from mammillaris, of southern waters, in their 
thinner shell. 

Polinices (Neverita) reclusianus (Deshayes). — Common. (45). 

Polinices (Neverita) alius Dall. — Abundant. (60) . 

Polinices (Euspira) lewisii (Gould) . — Only 1 specimen is referred 
to this species, although a number of individuals in our series of Polinices have 
an open umbilicus. Neither this feature nor the presence or absence of a funicle 
seem to be good characters in differentiating between lewisii and reclusianus. A 
series of specimens before me at this writing proceeds without a perceptible break 
from a completely closed umbilicus to a wide open one. The funicle is often 
present in the juvenile shell and absent in the adult. The shoulder of the whorls 
is not a constant feature in lewisii, but is usually present; it is, also, sometimes indi- 
cated in reclusianus. Lewisii grows to a much greater size than reclusianus and 
ranges considerably farther north, specimens having been taken by the writer in 
southeastern Alaska. 

Sinum scopulosum (Conrad) . — Syn., S. debile, of some authors; not 
of Gould, 1853: S. califomicum Oldroyd (Dall, 1921; Oldroyd, 1927).— Com- 
mon. (130). 

Acmaea cassis Eschscholtz subsp. ? — 2 juveniles. 

Acmaea cassis nacelloides Dall. — 1. 

Acmaea insessa (Hinds). — (30). The fact that this species, which 
lives on kelp, is the only member of the genus that is at all common in the deposit, 
is added evidence of scarcity of rocks. 

Tricolia pulloides (Carpenter) . — Syn., Phasianella pulloidea Cpr. 
(Dall, 1921): P. pulloides Cpr. (Oldroyd, 1927) .— (300). Arnold's speci- 
mens of "Phasianella compta Gld." should be checked with this species. 

Tricolia substriata (Carpenter) . — Syn., Phasianella substriata Cpr. 
(Dall, 1921; Oldroyd, 1927).— 135. 

Astraea (Pomaulax) undosa (Wood). — Syn., Pomaulax undosus 
Wood (Arnold, 1903).— 12. 

Leptothyra carpenteri Pilsbry. — Syn., Homalopoma carpenteri Pils. 
(Grant and Gale, 1931).— 1. 

Norrisia norrisi (Sowerby) . — 10. 

Halistylus pupoideus (Carpenter) . — Syn., H. subpupoideus Tryon 
(Dall, 1921; Oldroyd, 1927).— 8. 

Tegula (Chlorostoma) gallina (Forbes). — Syn., Chlorostoma gal- 
lina Fbs. (Arnold, 1903) .— 17. 

Tegula (Chlorostoma) gallina multifilosa (Stearns). — 1. 

Tegula (Chlorostoma) aureotincta (Forbes). — Syn., Chlorostoma 
aureotinctum Fbs. (Arnold, 1903). — 26. 

Tegula (? Chlorostoma) ligulata (Menke). — Syn., Chlorostoma 
vindulum ligulatum Mke. (Arnold, 1903).— 19. 

Willett — Pleistocene Fauna from Baldwin Hills 401 

Tegula (Promartynia) pulligo (Martyn) . — 1 specimen in White 

Calliostoma canaliculatum (Martyn). — Common. (70). 

Calliostoma gemmulatum Carpenter. — 18. 

Calliostoma tricolor (Gabb). — Abundant. (190). 

Calliostoma gloriosum Dall. — 1 collected by the writer, 2 by Miss 
Edna T. Cook. 

Calliostoma supragranosum Carpenter. — 2. 

Calliostoma splendens Carpenter. — 6. 

Turcica caffea Gabb. — Syn., Thalotia caffea Gabb (Arnold, 1903) . — 2. 

Margarites (Lirularia) optabilis (Carpenter) . — Syn., M. o. knechti 
Arn., M. o. nodosa Arn. (Arnold, 1903; Grant and Gale, 1931).— (30). This 
series includes both the typical form and the variety acuticostatus Carpenter. 

Vitrinella williamsoni Dall. — 29. 

Vitrinella eshnauri Bartsch. — 13. 

Vitrinella stearnsi Bartsch. — 17. 

Delphinoidea coronadoensis Arnold. — 1. 

Haliotis cracherodii Leach. — 1 fragment. 

Fissurella volcano Reeve. — Syn., F. v. crucifera Dall (Dall, 1921; 
Oldroyd, 1927) . — 3 specimens (Museum, 2; Miss Cook, 1) . 

Epitonium (Opalia) wroblewskyi (Morch) . — Syn., Opalia borealis 
Gld. (Arnold, 1903).— 1. 

Epitonium (Opalia) retiporosum (Carpenter). — 2. 

Epitonium (Asperiscala) bellastriatum (Carpenter). — Syn., Scala 
bellastnata Cpr. (Arnold, 1903) .— Common. (240). 

Epitonium (Asperiscala) clarki T. S. Oldroyd. — Abundant. (490). 

Epitonium (Nitidiscala) acrostephanum Dall. — 25. 

Epitonium (Nitidiscala) indianorum (Carpenter) . — Syn., Scala in- 
dianorum Cpr. (Arnold, 1903) . — (32) . This is a puzzling series, varying greatly 
in number and form of varices. It may not be true indianorum, though I am not 
able to distinguish it from young of that species. None approaches the size of 
adults of indianorum from the north. 

Epitonium (Nitidiscala) tinctum (Carpenter). — (35). Lacking the 
color band, this species is difficult to identify in the fossil, and it is probable that 
mistakes have been made. 150 juveniles of this group remain undetermined. 

Epitonium (Nitidiscala) cooperi Strong. — Syn., "Scala tincta Cpr." 
(Arnold, 1903): Epitonium hindsu Cpr. (Packard, Univ. Calif. Publ. Zool., 
14, 1918, p. 319) : E. fallaaosum Dall (Dall, 1921; Oldroyd, 1927).— 48. 

Epitonium (Nitidiscala) sawinae Dall. — Syn., E. catalinensis Dall 
(Dall, 1921; Oldroyd, 1927).— 64. 

Melanella micans (Carpenter). — Syn., Eulima micans Cpr. (Arnold, 
1903 ) .— Abundant. ( 1850) . 

402 San Diego Society of Natural History 

Melanella oldroydi Bartsch. — Rather uncommon. (24). 

Melanella rutila (Carpenter). — Abundant. (1100). 

Melanella sp. ? — 3 specimens, the size of rutila, but less slender and 
with higher body whorl. 

Strombiformis raymondi (Rivers). — Syn., S. riversi Bartsch (Proc. 
U. S. Nat. Mus., 53, 1917, p. 339).— (17). S. calif omica Bartsch is very simi- 
lar to this species and may be the same, but none of our specimens of the Recent 
form is as large as adults of the fossil. 

Turbonilla (Turbonilla) hypolispa Dall and Bartsch. — 21. 

It is with much hesitation that the writer employs here a division of sub- 
genera different from that in general use. The easier method would be to follow, 
without comment, the arrangement used by Dall and Bartsch in their great "Mon- 
ograph of West American Pyramidellid Mollusks," which appeared in 1909. 
However, after intensive study of west American Turbonillas, the writer is not 
convinced that the generally accepted division of the group as regards subgenera 
is not more arbitrary than natural. In fact, a number of excellent conchologists, 
known as keen students of Californian shells, have expressed their inability to 
separate the various subgenera of Turbonilla by their supposed characters. The 
natural inference drawn by an average student from such a condition might well 
be that there is no difference in value between a subgenus and a section. 

It appears to the writer that Californian members of the genus Turbonilla 
fall into five natural groups, as follows: Turbonilla (including Chemmtzia and 
Strioturbonilla) , Pyrgolampros, Pyrgiscus (including Pyrgisculus) , Bartschella 
(Dunkena), and Mormula. 

Cbemnitzia and Strioturbonilla have been differentiated from the subgenus 
Turbonilla because their axial sculpture does not extend onto the base and, in 
case of Strioturbonilla, because of spiral striations. In the species usually assigned 
to the subgenus Turbonilla the strength of the basal sculpture varies greatly; in 
some species, such as centrota and gilli, it is very weak, while in others like acra 
and diegensis, it is strong. At least one species, cayucosensis Willett (Nautilus, 
43, 1929, p. 26) , lacks basal sculpture in the young and shows it in the adult. As 
to Strioturbonilla: Although Dall and Bartsch state that the "spiral sculpture 
is always stronger than microscopic striations," in the majority of specimens 
examined by the writer this sculpture was not perceptible under a magnification 
of thirty diameters. The characters cited as a basis of separation of Pyrgisculus 
from Pyrgiscus appear to the writer to be only of sectional value, rather than sub- 
generic. Of the known Californian species, only laminata is here assigned to 
the subgenus Bartschella. Arata, which was included in this subgenus by Dall 
and Bartsch, when further material is available, may prove to be conspecific with 
weldi, generally included in Pyrgiscus. 

Turbonilla (Turbonilla) asser Dall and Bartsch. — 500. 

Turbonilla (Turbonilla) torquata (Gould). — 145. 

Turbonilla (Turbonilla) stylina (Carpenter). — 111. 

Turbonilla (Turbonilla) buttoni Dall and Bartsch. — 4. 

Turbonilla (Turbonilla) ralphi Dall and Bartsch. — Syn., "T. torquata 

Willett — Pleistocene Fauna from Baldwin Hills 403 

GH." (ArnoU, 1903).— 160. 

Turbonilla (Turbonilla) simpsoni Dall and Bartsch. — 22. 

Turbonilla (Pyrgolampros) lowei Dall and Bartsch. — 200. 

Turbonilla (Pyrgolampros) pedroana Dall and Bartsch. — 175. 

The last two species have been divided solely on the difference in number of 
ribs on the early whorls, no other stable differences being perceptible to me. 
There are also some specimens that appear to bridge the gap between pedroana 
and the following species. 

Turbonilla (Pyrgolampros) arnoldi Dall and Bartsch. 

Turbonilla (Pyrgolampros) halia Dall and Bartsch.. 

Turbonilla (Pyrgolampros) keepi Dall and Bartsch. — 1500. This 
splendid scries appears to demonstrate conclusively that the three above named 
were conspecific in late Pleistocene. They exhibit a surprising amount of variation 
and, in addition to ranging through the three already described species, there are 
numerous variants that, if found under some conditions, would undoubtedly be 
considered worthy of naming. Whether these three species are still connected, or 
whether the connecting links have disappeared since late Pleistocene, will remain 
uncertain until a larger number of Recent specimens are available for study. 

Turbonilla (Pyrgiscus) sanctorum Dall and Bartsch. — 3 specimens 
are referred to this species, although they have a few more incised spirals than 
the type has. The largest of our specimens has fifteen whorls and measures: alt., 
10 mm.; diam., 2.2 mm. 

Turbonilla (Pyrgiscus) cf. superba Da!l and Bartsch. — 1 specimen, 
taken by Mrs. E. M. Clark and donated to the Museum, appears nearest to this 
species, but differs from the type in position of median series of pits, which is a 
Ii:tle anterior to the middle of the whorl instead of posterior to it; furthermore, 
the ribs do not terminate as abruptly at the suture as is shown in the figure 
of superba. Our series of about 1000 specimens of the subgenus Pyrgiscus clearly 
demonstrates that many of the features generally used in differentiation of the 
species in the group are of little value. Variation in number and strength of both 
spirals and axials are endless. In many groups, undoubtedly of the same species, 
it is difficult to find two specimens exactly alike. These facts have caused the 
writer to adopt an entirely different view of the definition of species in the genus 
Turbonilla, with the direct result that no new ones are named in this paper, al- 
though there are numerous specimens that are different in appearance from any- 
thing hitherto described. 

Turbonilla (Pyrgiscus) vexativa Dall and Bartsch. — 2. 

Turbonilla (Pyrgiscus) antestriata Dall and Bartsch. — 290. 

Turbonilla (Pyrgiscus) almo Dall and Bartsch. — (460). A study of 
our series leads to the conclusion that the type of almo was not adult. For varia- 
tion in the species, see Plate 26, fig. 4. 

Turbonilla (Pyrgiscus) adusta Dall and Bartsch. — 1. 

Turbonilla (Pyrgiscus) weldi Dall and Bartsch. — Our 80 specimens 
exhibit great variation and appear to show intergradation between weldi, wick- 
hami and arata. 

404 San Diego Society of Natural History 

Turbonilla (Pyrgiscus) cf . ista Bartsch. — 2. 

Turbonilla (Pyrgiscus) canfieldi Dall and Bartsch. — (193). A varia- 
ble series, extending from typical canfieldi to histias; some individuals indicating 
close relationship to macbridei and almejasensis. 

Turbonilla (Bartschella) laminata (Carpenter). — 43. 

Turbonilla (Mormula) tridentata (Carpenter). — Syn., T. ambusta 
Dall and Bartsch.— 565. 

Turbonilla (Mormula) regina Dall and Bartsch. — Syn., T. catalinen- 
sis Dall and Bartsch. — ■ (18). Throughout the subgenus Mormula the number 
of incised lines and axial ribs, and basal sculpture vary greatly within the species, 
in both fossil and Recent specimens. The writer is unable to find any constant 
characters separating regina from catalinensis, or ambusta from tridentata. 

Turbonilla (Mormula) pentalopha Dall and Bartsch. — (23). The 
adult of this species is quite different in appearance from the figure of the type 
given by Dall and Bartsch; in fact, it is much more like their figure of the type of 
castanea. The last whorl is long and rounded and, in some examples, possesses 
more than forty axials, and the internal lirations are so far back as to be hardly 
perceptible without breaking away the outer lip. 

Odostomia (Chrysallida) eugena Dall and Bartsch. — 2 specimens; 
an additional one in Miss Cook's collection. 

Odostomia (Evalea) nemo Dall and Bartsch. — (2100). By far the 
most abundant Odostomia. 

Odostomia (Evalea) donilla Dall and Bartsch. — 9. 

Odostomia (Evalea) cf. phanea Dall and Bartsch. — 1. 

Odostomia (Amaura) helena Bartsch. — (188). This species varies 
considerably in diameter and in amount of tabulation of whorls. Some specimens 
show spiral sculpture. 

Lepidopleurus nexus (Carpenter) . — Syn., L. heathi Berry, L. ambus- 
tus Dall (Dall, 1921; Oldroyd, 1927) .— 2 head and 2 median valves. 

Mopalia acuta (Carpenter) . — 1 head, 2 tail and 7 median valves. 

Ischnochiton sanctaemonicae Berry. — 1 median valve found by 
Miss Cook and donated to the Museum. 

Helisoma cf. trivolvis (Say). — 2 juveniles. 

Gyraulus vermicularis (Gould). — 2. 

Zonitoides arboreus Say. — 1. 

Willett — Pleistocene Fauna from Baldwin Hills 

Plate 26 

Fig. 1. Mangelia variegata Cpr., showing intergradation between two ex- 
tremes; x 2. 

Figs. 2. 3. Purpura leeana (Dall), juv., (fig. 2), showing similarity of the 
young of this species to half-grown Tritonalia barbarensis (Gabb) (fig. 3) ; x 2. 

Fig. 4. Turbonilla almo Dall and Bartsch, showing variation in the species; 

JAN 27 1938 



Vol. VIII, No. 31, pp. 407-408 January 18, 1938 



Dickey Collections, California Institute of Technology 

During the past spring and summer van Rossem and Hannum 
continued field work in southern Sonora in order to gather data supple- 
mentary to the work initiated by van Rossem in 1930 and subsequent 
years. Many interesting facts were discovered, some of which have been 
published and others are in press. 

The presence of a strong Arid Tropical element in southern Sonora 
is now so well established that any further discoveries of tropical species 
or races may be considered as evidence purely additional to that already 
produced. One of the tropical genera of hummingbirds encountered was 
Saucerottia, whose presence was already known through the species 
beryllina (dubiously of the race viola Miller). An additional species of 
this genus was discovered during the recent field work. 

Two representatives of Saucerottia, namely sumichrasti (Salvin) 
of Oaxaca and ocai (Gould) of Vera Cruz are each known from 
(unique?) specimens in the British Museum. They differ from other 
closely related members of the genus in having bronzy instead of chest- 
nut, purplish, or blue rectrices and upper tail coverts. It is to this bronze- 
tailed group that the new species from Sonora belongs. The single speci- 
men was sent, as an additional check, to J. L. Peters and Ludlow Gris- 
com at the Museum of Comparative Zoology and neither has been able 

408 San Diego Society of Natural History 

to place it with any known hummingbird; neither can they associate it 
as a hybrid. We therefore name it as 

Saucerottia florenceae sp. nov. 

Type. — Female adult, no. 31,888, Dickey collections at the California In- 
stitute of Technology; Rarcho Santa Barbara, 20 miles northeast of Guirocoba, 
southeastern Sonora, Mexico; altitude 5000 feet in the oak-pine association; June 
9, 1937; collected by A. j. van Rossem and Robert Hannum. 

Specific characters. — Belonging, obviously, to the sumichrasti-ocai group of 
the genus Saucerottia in possessing golden, bronzy-green rump, upper tail cov- 
erts, and rectrices. Above, bright, semi-iridescent, metallic grass green, darker 
on pileum and forehead and brighter, more coppery, on rump, upper tail coverts, 
and rectrices. Under parts grayish white, heavily spangled with green save on the 
abdominal region and under tail coverts; the spots on chin and throat smaller 
and more bluish green, those on the pectoral region and sides larger and more 
grass green. Under tail coverts chiefly grayish white, but buffy gray centrally. 
Auriculars dusky, surmounted by an indistinct grayish white post-ocular streak. 
Wings, dull black with a strong purplish hue, and white concealed bases of the 
flight feathers pale buff. Rectrices, (from above) golden, bronzy green, less 
obviously so on the outer pairs whose terminal portions are more or less dusky 
and whose shafts are pale chestnut; from below the coloration of the tail is simi- 
lar, but the lateral rectrices are obsoletely mottled with steely blue on the subter- 
minal portions. Bill, black, with the basal three-fourths of the mandible flesh 
color; iris, dark brown; feet, dull black. The measurements are: wing, 58; tail, 
32; exposed oilmen, 22 mm. 

flange. — So far as known at present, the pine-oak association in the Sierra 
Madre in extreme southeastern Sonora. 

Remarks. — The unique specimen was shot at late dusk as it was flying 
actively about the topmost branches of a leafless oak. The light, in fact, was so 
dim that the fallen bird was found only by the aid of a flashlight. Two other 
hummingbirds of unknown species were seen under similar circumstances and 
one is naturally tempted to speculate on the possibility of nocturnal, or at least 
crepuscular activity in the case of certain species. 

This species is named for Florence van Rossem, the wife of the 
senior author. 

JAN 27 1938 
q Sr * 1 V 



Vol. VIII, No. 32, pp. 409-410 January 18, 1938 



Laurence M. Huey 

Curator of Birds and Mammals, San Diego Society of Natural History 

Among the mammals collected by the writer on an expedition for 
the San Diego Society of Natural History in southwestern Utah during 
the summer of 1937 is an apparently unnamed form of muskrat. 

In the absence of sufficient material at hand for adequate compari- 
son, the skulls of the muskrats secured were forwarded to Major E. A. 
Goldman of the Bureau of Biological Survey, Washington, D. C, where 
needed comparative specimens from the national collections were avail- 
able. Through his kindness a number of comparisons were made and 
reported to the writer. 

It is with pleasure that the animal is named in Major Goldman's 
honor as 

Ondatra zibethica goldmani subsp. nov. 
Virgin Valley Muskrat 

Type. — From Saint George, Washington County, Utah; no. 12915, collec- 
tion of the San Diego Society of Natural History; adult female; collected by 
Laurence M. Huey, August 11, 1937. 

Characters. — A race of Ondatra zibethica larger in size than either Ondatra 
zibethica pallida or O. z- bernardi of the Colorado River drainage system in 
Arizona, and smaller than O. z- mergens of the Great Basin region in northwest- 
ern Nevada and northeastern California. In color, O. z- goldmani is somewhat 
similar to O. Z- bernardi but has a heavier coating of guard hairs dorsally. Com- 
pared with O. z- pallida, goldmani is lighter in color; in fact pallida, despite its 
name, is the darkest of the three forms found along the lower Colorado River 

410 San Diego Society of Natural History 

drainage system. Compared with O. z- mergens, goldmani is somewhat paler 
and more uniformly light brown. Cranially, goldmani differs from mergens in 
having a relatively narrower, less massive skull; braincase decidedly narrower; 
lambdoid crest narrower, less flaring and upturned; interparietal narrower; pre- 
maxillae wider at fronto-maxillary suture; basi-occipital narrower; auditory bullae 
more inflated laterally. Compared with bernardi, goldmani has a relatively nar- 
rower, mor? elongated skull, with interparietal larger and longer, that is to say 
more extended antero-posteriorly. The bullae are distinctly larger and more fully 
inflated. Compared with pallida, goldmani has a relatively more elongated skull 
with a more slender rostrum. The interparietal differs as it does from that of 
bernardi, the zygomatic arches are more arched and the audital bullae more 

Color and Measurements of Type. — Dorsally uniform Dresden Brown, 1 
slightly darker on nose and rump, shading to lighter on sides and underparts. 
Tail thinly fringed on dorsal and ventral ridges, with dark, nearly black, hairs. 
Feet thinly covered dorsally with lighter-colored hairs. Vibrissae black. Total 
length, 502; tail, 215; hind foot, 76; ear, 17. Skull: greatest length, 59.6; 
zygomatic breadth, 36.9; nasals, 19.6; tooth row, 14.8. 

Range. — Probably limited to the riparian association along the Virgin 
River in southwestern Utah, from near Zion National Park westward at least 
to Saint George and perhaps farther westward along the course of the Virgin 
River into the extreme northwestern tip of Arizona and southeastern Nevada. 

Specimens examined by the writer. — Ondatra zibethica mergens: 1 from 
Eagle Lake, Lassen County, California. Ondatra zibethica pallida-?- 24 from 
Camp Verde, Yavapai County, Arizona (type locality). Ondatra zibethica 
bernardi: 1 9 from 4 miles south of Gadsden, Yuma County, Arizona (type 
locality). Ondatra zibethica goldmani: 7 from Saint George, Washington 
County, Utah (type locality) . 

1 Ridgway, Color Standards and Color Nomenclature, 1912. 

2 From collection of Bernard Bailey. 



Volume VIII, No. 33, pp. 41 1-454, plates 27-36 

% ** % 



Steve A. Glassell 

Research Associate in Crustacea, San Diego Society of Natural History 


Printed for the Society 

May 31, 1938 


Introduction 413 

Family Crangonidae 

Genus Homoriscus Rathbun 

Homoriscus macginitiei, sp. nov 414 

Genus Betaeus Dana 

Betaeus ensenadensis, sp. nov 416 

Family Paguridae 

Genus Paguristes Dana 

Paguristes sanguimmanus, sp. nov 419 

Paguristes anahuacus, sp. nov 421 

Family Porcellanidae 

Genus Euceramus Stimpson 

Euceramus panatelus, sp. nov 423 

Euceramus transversilineatus (Loclcington), new combination 426 

Genus Minyocerus Stimpson 

Minyocerus fork, sp. nov 430 

Genus Porcellana Lamarck, restricted 

Porcellana magdalcnensis Glassell, 431 

Genus Ulloaia, gen. nov 434 

Ulloaia perpusillia, sp. nov 434 

Genus Pisonella, gen. nov 436 

Key to the Species of Pisonella 437 

Pisonella sinuimanus (Loclcington) , new combination 437 

Pisonella tuberculipes (Loclcington), new combination 440 

Pisonella smithi (Glassell), new combination 442 

Pisonella erosa (Glassell), new combination 442 

Key to the West North American Species of Petrolisthes 442 

Key to the West North American Species of Pachycheles 444 

Family Goneplacidae 

Genus Hexapus de Haan 

Hexapus wdliamsi, sp. nov 445 

Family Pinnotheridae 

Genus Alarconia, gen. nov 446 

Alarconia seaholmi, sp. nov 448 

Genus Pinnotheres Latreille 

Pinnotheres orcutti Rathbun, 451 

Genus Fabia Dana 

Fabia granti Glassell, 452 



qJS^ Steve A. Glassell 

Research Associate in Crustacea, San Diego Society of Natural History 


The material for this paper was collected from several sources, with 
localities ranging from La Jolla, California, to La Libertad, Ecuador. 
Twelve genera in five families are represented. Of these, three genera are 
proposed as new. Four genera are newly introduced to the west American 
fauna, three of these having been monotypic genera from west Atlantic 
waters, one, a monotype, from the Indo-Pacific region. Keys are given 
for three genera of west North American porcellanids, Petrolisthes, 
Pachycheles and the proposed genus Pisonella, split off from the genus 
Pisosoma. The holotypes for all the new species are deposited with the 
San Diego Society of Natural History. 

For specimens I am indebted to Professor George E. MacGinitie of 
the California Institute of Technology, and Captain W. J. Seaholm and 
Mr. Woodbridge Williams, who brought in a splendid collection of well 
preserved material from southern waters, taken on a cruise of the yacht 
"Stranger," under the command of Captain Fred E. Lewis, of Balboa, 

For the loan and gift of comparative material, photographs of obscure 
publications, literature and advice, I am under obligations to Dr. Waldo 
L. Schmitt, of the U. S. National Museum, Dr. Fenner A. Chace, Jr., of 
the Museum of Comparative Zoology, Mr. G. Robert Lunz, Jr., of the 
Charleston Museum, Dr. K. H. Barnard, of the South African Museum, 
and Mr. Melbourne Ward, of the Australian Museum. 

To Mr. Anker Petersen, of Beverly Hills, California, who has given 
his own time and means to the drawing of the plates, I can but offer my 
thanks, with the full knowledge that his contribution is greater than my 

414 San Diego Society of Natural History 

Genus Homoriscus Rathbun 

Homoriscus macginitiei, sp. nov. 
Plate 27, figures 1-4 

Type. — Female, holotype, Cat. No. 1120, San Diego Society of Natural 
History; from La Jolla, California, low tide; March 4, 1935; collected by George 
E. MacGinitie. 

Diagnosis. — Rostrum anteriorly subovate, armed. Antennal scale armed 
with 7 or 8 teeth. Chelipeds subchelate. Telson armed with 3 pairs of lateral 
spines. Outer margin of dactyl of 3rd ambulatory leg armed with 11 spines. 

Description. — Carapace lightly pubescent, with 7 sharp longitudinal crests; 
the median occupies the posterior 4/5 of carapace and is interrupted by the cervi- 
cal groove; the submedian and supero-lateral crests begin near base of rostrum, 
the former posterior to this point. Both of these crests are less than half the length 
of the carapace, the supero-lateral being the shorter of the two. The infero- 
lateral crest begins at antennal sinus of anterior margin of carapace and is about 
half as long as submedian crest. Orbit semicircular, a little narrower than the 
black cornea beneath; outer orbital angle small and blunt-pointed. Rostrum 
semioval, armed anteriorly with fine, sharp teeth; upper surface lightly granulous, 
slightly concave. Third antennular peduncle extending its entire length past 
tip of rostrum. Antennular flagellum nearly 1/2 the length of the median crest 
of the carapace. Antennal peduncles subequal in length to that of the antennular. 
Antennal acicle subovate, armed on its outer margin with a row of 7 or 8 fine, 
sharp teeth, inner margin setose. Flagellum nearly as long as the body. 

Chelipeds subchelate, somewhat resembling those of the genus Crangon. 
Manus subquadrate, upper margin carinate, straight; lower margin with a median, 
fixed tooth, which acts as the pollex, the distal lower edge armed with 3 sharp, 
triangular teeth, the proximal the largest; the dactyl is arcuate, sharp-pointed, 
setose on its upper crest and unarmed on its inner edge. The 1st ambulatory leg 
in short and stout, in comparison with the others; the lower margin of the merus 
and the upper margin of the carpus, propodus and dactylus are setose; the dactyl 
is simple and heavier than the others; of the ambulatory legs the 2nd is the 
longest, followed by the 3rd, 4th and 1st. The dactlyi of the 2nd, 3rd and 4th 
are slender, lanceolate, with their lengths as in the order given. 

The outer maxillipeds have their ischium armed on the inner side with a row 
of spinules; the merus on the distal outer end with a spine; the propodus and 
dactylus subequal in length and longer than the carpus. Telson longer than 
broad, rounded at extremity; sides obscurely spinulous, with 3 pairs of lateral 
spines. Lateral lamina longer than telson; inner lamina longer than outer; outer 
margin of inner lamina smooth, of outer lamina spinulous. 

Sexual variation and Color in life. — Unknown. 

Measurements. — Female holotype: length from rostrum to tip of telson 
18.3 mm., of carapace 7 mm., width of carapace 3.1 mm. 

Material examined. — The two ovigerous type specimens. 

Glassell — West American Decapod Crustacea 

Plate 27 





^£/"~ ^*v 




1 ' [Va 




yr ~- 


Ik. \ 



^ ; 

n. m . 

>■" Eg 

5}fes Mttl J ?. 



Fig. 1. Homoriscus macginitiei Glassell, sp. nov. Right cheliped. 

Fig. 2. Homoriscus macginitiei Glassell, sp. nov. Telson. 

Fig. 3. Homoriscus macginitiei Glassell, sp. nov. Carapace. 

Fig. 4. Homoriscus macginitiei Glassell, sp. nov. Ambulatory legs. 

416 San Diego Society of Natural History 

Habitat. — Professor MacGinitie reports finding these two specimens in a 
small pool at extreme low water, after he had turned some stones in search of 
Typhlogobius californiensis Steindacher (the Blind Goby) . 

Remarks. — This proposed species is closely allied to H. portoricensis Rath- 
bun, 1902. For the differences between these two species I cannot do better 
than quote a letter received from Dr. Fenner A. Chace, Jr., of the Museum of 
Comparative Zoology, at Harvard College, Cambridge, Massachusetts, who 
was kind enough to compare photographs of my drawings with a specimen of 
H. portoricensis, sent him from Havana, Cuba. Dr. Chace's findings are as 
follows : 

"There is no doubt in my mind but that your form is specifically distinct 
from the Atlantic species. In comparing the two, the following differences are 
the most apparent: (1) Rostrum rounded rather than bluntly acute, (2) ros- 
trum margined with a few distinct spines instead of being merely finely serrate, 
(3) rostrum reaching nearly to end of second antennular article rather than to 
middle of third segment, (4) submedian and supero-lateral crests of carapace 
entire instead of being minutely serrate, (5) antennal scale armed with 7 or 8 
spines rather than 4 or 5, (6) the dactyl of the chelipeds is slightly shorter, the 
ratio of the length of the dactyl to the length of the palm being as 1 : 0.78 
instead of as 1 : 0.73; consequently the apposable surface of the palm is longer, 
the large spine being placed more proximal, although the ratio of length to 
breadth of the palm is essentially the same in the two species — perhaps it is 
slightly broader in the Pacific species — and the armature is almost identically 
the same, (7) outer margin of palm of cheliped entire rather than armed with 
3 or 4 small spines at distal fourth, (8) outer margin of dactyl of 3rd ambu- 
latory leg armed with 11 spines rather than 2 or 3, (9) telson slightly longer 
and narrower at distal end; ratio of length to breadth as 1 : 0.60 instead of as 
1 : 0.70, (10) telson armed with three pairs of lateral spines instead of a single 
pair at the junction of the smooth lateral margin with the setose terminal mar- 
gin, and (11) outer margins of outer uropods much straighter; in H. portoricen- 
sis they are very convex. I might add that the inner margins of the dactyls of 
all the ambulatory legs in H. portoricensis are pectinate, irregularly so in the 
central portion. In the above comparisons, the characteristics of your species 
are given first in each case." 

This proposed species is named for my good friend Professor George E. 
MacGinitie, of the California Institute of Technology, who collected this aber- 
rant form and allowed me the privilege of describing it. 

Genus Betaeus Dana 

Betaeus ensenadensis, sp. nov. 
Plate 28, figures 1-3 

Type. — Male, holotype, Cat. No. 1121, San Diego Society of Natural 
History; from Estero de la Punta Banda, Ensenada, Baja California, Mexico, 
low tide; December 19, 1930; collected by George E. MacGinitie. 

Diagnosis. — Front evenly rounded, not emarginate between the eyes. 
Hands similar, oblong, compressed; propodus subtruncate at apex; dactyl falcate 

Glassell — West American Decapod Crustacea 

Plate 28 



Fig. 1. Betaeus ensenadensis Glassell, sp. nov. 

Fig. 2. Betaeus ensenadensis Glassell, sp. nov. Cheliped. 

Fig. 3. Betaeus ensenadensis Glassell, sp. nov. Telson. 

418 San Diego Society of Natural History 

at tip, armed with three well-formed teeth. 

Description.- — Front evenly rounded, not emarginate. Carapace smooth, in 
life transparent, so that vital organs may be plainly seen, opaque in preservative. 
Second peduncle of antennule nearly twice the length of the third. Flagellum of 
antennae as long as chelipeds. Antennal acicle reaches past the proximal end of 
third peduncle of antennule, entire on outer margin, terminating in a spine; 
inner margin of acicle terminates near base of spine, evenly rounded at distal 
end, margin setose. 

Chelipeds similar, 1/3 longer than length of carapace; merus lightly den- 
tate on inner margins, outer margin with a broad oblique sinus, transverse sub- 
distal groove deep; carpus with a vertical, lamellar projection on inner face 
which fits into a recess of the merus, when arm is flexed; hand oblong, com- 
pressed, lightly granulated when viewed under a lens; the length of the palm is 
greater than the length of the dactyl; the dactyl is strongly falcate at the apex, 
is armed with three strong teeth, the median the largest, the proximal the small- 
est; the propodus is lightly granulated, proximally armed with a single, small 
tooth in the gape. The propodus terminates in a single, sharp, up-turned spine, 
at the base of which the propodus is truncate; the fingers are crossed at their 
tips, and gape from base to apex. 

Color in life. — The carapace, abdomen and chelipeds are covered with light 
tinted chromatophores in reds and blues, the fingers and telson are tinted a light 

Measurements. — Male holotype (not the largest specimen, but the most 
perfect) : length from rostrum to tip of telson 19.5 mm., of carapace 6.8 mm., of 
cheliped 10 mm., of manus 5.6 mm., of dactyl 3 mm. Female paratype: length 
from rostrum to tip of telson 21.2 mm., of carapace 6.5 mm. (the hands were 
missing) . 

Range. — So far only known from the type-locality. 

Material examined. — A series of 12 specimens, from Estero de la Punta 
Banda, Ensenada, Baja California, Mexico; collected by George E. MacGinitie, 
December 19, 1930. The types were selected from this series. 

A male specimen collected by the author at the same locality, December 
25, 1936. 

Habitat. — These specimens were found at low water in the burrows of 
Callianassa and Upogebia. Professor MacGinitie reports finding them in pairs. 

Remarks. — This proposed species is allied to B. longidactylus Lockington, 
1877, but differs from that species by the hands being more uniform in both the 
sexes, and in the individual showing little if any variation, instead of varying 
from long slim fingers with little gape, to those as figured by Schmitt in Univ. 
Calif. Publ. Zool., vol. 23, 1921, pi. 12, fig. 2, where the fingers are widely 
separated throughout their length. In addition, B. ensenadensis has the apex of 
its propodus subtruncate and spine-tipped, instead of being rather blunt-pointed 
as in B. longidactylus; it differs also by the dactyli being armed with 3 well- 
formed teeth, and being falcate distally where the fingers cross each other, in- 
stead of being unarmed, or having, at best, a small proximal tooth; the tips of 
the fingers crossed slightly. It also differs by the peduncles of the antennules 

Glassell — West American Decapod Crustacea 419 

being of different length, the second nearly twice the length of the third, instead 
of being about equal length. In addition, B. ensenadensis is a much smaller 
species, a mature specimen of B. longidactylus measuring from rostrum to tip 
of telson 40.5 mm. 

B. ensenadensis resembles the other known Californian representative of 
the genus, B. harfordi (Kingsley), 1878, in size, shape and relative lengths of 
the antennular peduncles (the 2nd being nearly twice the length of the 3rd), 
but differs in that the front is evenly rounded, instead of emarginate, by the 
hands being similar and suboblong, instead of dissimilar and oval, by the ter- 
minal spine of the antennal article extending far past the rounded inner margin, 
instead of extending a little past the rounded inner edge. 

To Professor George E. MacGinitie of the California Institute of Tech- 
nology, belongs the credit for collecting and recognizing this proposed species. 

Note. — Since the above notes were written, an important extension of range 
for this species has been made by Professor MacGinitie, who collected two males 
and one ovigerous female, in Upogebia tubes, at False Bay, San Diego, Califor- 
nia, on November 18, 1937. These specimens are in the author's collection. 


Paguristes sanguinimanus, sp. nov. 

Type.— Male, holotype, and female, paratype, Cat. Nos. 1122 and 1123, 
San Diego Society of Natural History; from Punta Penasco, Sonora, Mexico, 
low tide; May 2, 1935; collected by Steve A. Glassell. 

Diagnosis. — Precervical portion of carapace longer than wide, areolate, 
laterally punctate, a gastric median groove; rostral tooth long, exceeding laterals. 
Chelipeds subsimilar, heavy, wide; inner margin of carpus not regularly, though 
distinctly, spined; hand with 4 spines on margin of palm. Flagellum reaching 
palm of hand, lightly ciliate. Eye-stalks long, cylindrical. Tip of antennal acicle 
extends but slightly past the median length of the eye-stalk, and is subequal in 
length to the distal end of the 3rd antennal peduncle. The flagellum of the 
antennule extends past the cornea. 

Description. — Carapace with precervical portion longer than wide, punctate 
laterally and on the anterior portion of the gastric region, lightly setose laterally, 
smooth centrally, a median protogastric groove. The median tooth is long and 
slightly depressed, subtruncate at the tip, extending midway between the eye- 
scales, and is 1/6 the length of the eye-stalk. The laterals extend 1/3 the distance 
of the median, are obtuse, with a short terminal spine. The margin between the 
teeth is revolute and granulous. 

Eye-stalks long, cylindrical, slightly outward turned, with tufts of setae on 
upper surface; in length they equal the length of the carpus. Ophthalmic scales 
triangular, sharp-pointed, margins entire. 

Antennal acicle bifid at tip, a strong, proximal spine on upper surface at 
proximal 1/3, two outer marginal, distal spines; in length it extends a little past 
the median portion of the eye-stalk. The 3rd antennal peduncle has 3 spines on 

420 San Diego Society of Natural History 

the upper proximal surface, in length it slightly exceeds the tip of the acicle. The 
flagellum reaches the palm of the hand and is longer than the precervical portion 
of the carapace, is lightly ciliated. 

Chelipeds subsimilar, stout, wide, heavy, the upper surface covered with 
short, sharp-pointed tubercles, interspersed with short setae and some pubescence, 
the setae not much longer than the tubercle; merus with upper distal surface 
triangular, inner surface smooth, its lower margin spined, outer surface rough- 
ened with granules, lower distal margin spined; carpus about as wide as long, 
spines more prominent on and near inner margin, outer margin not distinct, 
surface covered with short, sharp-pointed tubercles; manus semiovate, longer 
than wide, 4 conical, sharp-pointed spines on inner margin of palm, outer mar- 
gin with small spines distally, entire surface set with sharp-pointed tubercles; 
fingers close set, tips corneous. 

Ambulatory legs stout, rugose and setaceous, the first pair margined on the 
upper crest of the carpus, propodus and dactylus with spines, the second pair 
with a distal carpal spine only; the dactyli are slightly twisted. 

The distal edge of the telson is armed with 4 well-spaced teeth on each 
segment; the right is the largest. 

Color in alcohol. — The carapace, merus and carpus have a buff ground color 
with numerous circular red spots, the perimeter only being colored; the mem- 
branous covering of the branchials is reddish-purple. The hands are blood-red. 
The eye-stalks are orange-red, the base purple. 

Measurements. — Male holotype (the largest specimen) : length from ros- 
trum to tip of telson 103 mm., of carapace 25 mm., of precervical portion of 
carapace 13 mm., width 11 mm., length of cheliped 38 mm., of merus 11 mm., 
of carpus 9 mm., width of carpus 8 mm., length of manus 13 mm., width of 
manus 10 mm., length of eye-stalk 9 mm. 

Range. — Gulf of California. 

Material examined. — A large series of 20 or more males, and 20 or more 
females, non-ovigerous, from Punta Penasco, Sonora, Mexico, low tide; May 2, 
1935; the types were selected from this series. 

A series of 10 or more males, and 10 or more females, non-ovigerous, from 
the same location; April 12, 1937. Both series collected by the author. 

Habitat. — This hermit crab, contrary to the majority of species in the genus, 
is apparently a littoral form, being very numerous at the type-locality, from 
mean low water down. The carcinoecia was a species of Turritella. 

Remarks. — This proposed species is closely allied to P. digueti, Bouvier, 
1892, which it resembles in many particulars, such as the shape of the front and 
the form of the chelipeds, particularly the hands. It differs from that species, 
however, by lacking distinct heavy spines on the inner carpal margin, instead of 
having 3 large conical spines, by the antennal acicle extending only a little past 
the middle of the eye-stalk, instead of 2/3 the length of the eye-stalk, by the 
upper proximal surface of the 3rd antennal peduncle being armed with 3 spines, 
instead of with 2 spines, and by the ophthalmic scales being sharp-pointed, with 

Glassell — West American Decapod Crustacea 421 

margins entire, instead of having a bifid tip. 

In addition to these structural differences, there is one of anatomical dis- 
tortion, for P. sanguinimanus occupies a shell with a circular aperture, the cara- 
pace remaining normal, while P. digueti favors a dwelling such as Strombus, 
which distorts the carapace, depressing the precervical portion and distending 
the branchial regions. This may be taken as a general statement. 

While both species live in the same waters, P. digueti has been recorded 
at depths ranging from 10 to 40 fathoms, and P. sanguinimanus has so far only 
been taken as a shore form. 

Paguristes anahuacus, sp. nov. 

Type. — Male, holotype, and female, paratype, Cat. Nos. 1124 and 1125, 
San Diego Society of Natural History; from Punta Penasco, Sonora, Mexico, 
low tide; May 2, 1935; collected by Steve A. Glassell. 

Diagnosis. — Rostral tooth long, sharp-pointed, margined, concave on upper 
surface, extending well between the ocular scales for more than half their length. 
Eye-stalks extending past merus. Flagellum not reaching distal end of carpus, 
lightly ciliated. Chelipeds densely tomentose; carpus with 5 inner-marginal 
spines; palm of hand with 3; hand nearly twice as long as wide. 

Description. — Precervical portion of carapace nearly 1/3 longer than wide, 
tuberculate and tomentose laterally. Median tooth long, sharp, pointed, heavily 
margined, upper surface concave, the apex extending slightly past the center of 
the eye-scales; lateral teeth short, their outer margins convex, the inner concave. 
The margin between the laterals and the median is deep and revolute. 

Eye-stalks long, nearly as long as the width of the carapace, heavy at the 
base, cylindrical distally, extending past the merus of the chelipeds and to the 
tip of the 3rd antennular peduncle. The ophthalmic scales are bifid, toothed 
on the outer margin, entire on the inner, and are tomentose distally. 

The antennal acicle extends 2/3 the length of the eye-stalk, and is armed 
on its proximal inner edge with a single, sharp-pointed tooth; the outer margin 
has 1 or 2 teeth at the distal 1/3; the tip is bifid. The outer distal portion of the 
2nd antennal peduncle extends 1/3 the length of the acicle, is bifid and spined 
on the outer edge. The acicle is covered with long pinnate tomentum. The distal 
end of the 3rd antennal peduncle just reaches past the acicle. The flagellum ex- 
tends past the middle of the carpus, is subequal in length to that of the hand, 
and is lightly ciliated. 

The chelipeds are subequal; the upper crest of the merus and the upper 
surfaces of both the carpus and manus are densely tomentose, though not 
entirely covering the fingers; merus trigonal, distally spined, a transverse sub- 
distal groove extending down both sides, the outer surface rectangular, lightly 
rugose, the inner lower margin spined; carpus subequal in length to the merus, 
widest distally, the inner margin armed with 5 upward- and forward-pointing, 
corneous-tipped, conical spines, the outer margin with 8 or more smaller spines, 
the upper surface with numerous well-spaced tubercles; a prominent spine over 

422 San Diego Society of Natural History 

the upper hinge joint of the hand, the inner face smooth. The hands are nearly 
1/2 longer than wide, the inner margin of the palm with 3 spines, upper surface 
flat, covered with sharp-tipped tubercles, not quite in a distinct pattern; the 
outer distal margin of the pollex is spined, as is the upper proximal edge of the 
dactyl. The tips of the fingers are corneous, spooned. The hand is densely cov- 
ered with tomentum except for the inner edges of the fingers. 

The ambulatory legs are thickly margined with tomentum down to the 
corneous tips of the dactyl i; the 1st pair extend past the chelipeds by the length 
of their dactyli; the carpi, propodi and dactyli of the 1st pair are crested with 
spines, the 2nd pair with a distal carpal spine only. The distal end of the telson 
is wider than its base, broadly V-shaped and armed with small teeth. 

Color in alcohol. — Carapace reddish-brown with light blue spots. Chelipeds 
orange with cream-colored spots. Ground color of ambulatory legs orange over- 
cast with blue, which gives the appearance of a dull lavender. The eye-stalks at 
their distended bases are light orange; the stalk is white with a submedian band 
of dusky violet-blue which distinguishes the species. The antennules are light 
blue. The antennal flagellum has it joints light blue on their proximal end and 
white distally. The tomentum is cream color. 

Measurements. — Male holotype: length from rostrum to tip of telson 
46.6 mm., of carapace 12.6 mm., of precervical portion 8.2 mm., width 5.5 mm., 
length of cheliped 22 mm., of merus 6 mm., of carpus 6 mm., of manus 7.5 mm., 
width 4 mm., length of eye-stalk 4 mm. 

Range. — So far only known from the upper end of the Gulf of California. 

Material examined. — A series of at least 100 specimens of both sexes, 
collected at Punta Pehasco, Sonora, Mexico, low tide; May 2, 1935; and a 
smaller series taken at the same locality, April 12, 1937; both series collected by 
the author. 

Habitat. — Found from extreme low water to a depth of 10 fathoms. The 
carcinoecia of the type specimens was a species of Turritella. They are abundant 
at the type-locality. 

Remarks. — This proposed species is allied to P. aztatlanensis Glassell, 
1937, in that the chelipeds are of similar shape, especially the hands, the 
flagellum being short and lightly ciliated; but it differs in that the median spine 
is long, extending well between the eye-scales, instead of being triangular and 
extending between the bases of the eye-scales, by the carpus of the chelipeds 
having 5 inner-marginal spines, the hand 3, instead of the carpus having 4 
spines, the hand 3, and by the chelipeds being densely clothed with tomentum, 
instead of being lightly tufted. The front of the carapace in this proposed species 
somewhat resembles that of P. spinipes A. Milne Edwards, 1880. 

This is another of the few species in this primitive genus of Paguridae, 
which may be termed littoral. 

The name of this species is taken from a Nahuatl word signifying "within 
the water." 

Glassell — West American Decapod Crustacea 423 

Genus Euceramus Stimpson 

Euceramus panatelus, sp. now 
Plate 29 

Type. — Male, holotype, Cat. No. 1126, San Diego Society of Natural 
History; from La Libertad, Ecuador, 6-9 fathoms; March 24, 1937; collected by 
Woodbridge Williams, on Captain Fred E. Lewis' yacht "Stranger." 

Diagnosis. — Carapace slightly longer than twice the width, nude, transverse, 
minute striations; frontal region tridentate, the median slightly longer than the 
laterals. Antennal flagellum 1/3 longer than carapace, ciliated as in the genus 
Lepidopa. Chelipeds subequal in length to carapace, fingers gaping. Maxillipeds 
attached to a broadly truncate sternal segment. 

Description. — Carapace slightly longer than twice the width, regularly 
curved like a segment of a cylinder, nude, cervical groove defined, minutely 
striated transversely, the lateral marginal carina only broken near posterior 
margin, precervical portion anteriorly granulate. Posterior margin widely V- 
shaped, edge revolute. Front horizontal, tridentate, teeth sharp, triangular, the 
median slightly the longest, separated from the laterals by a U-shaped sinus. 
Orbits incomplete, with concave superior margins, a microscopic spinule at its 
outer margin. A single lateral spine at the shoulder, behind the cervical groove. 
The eyes are retractile, stalks cylindrical, slightly contracted below cornea. When 
extended, the eyes extend past the frontal teeth and are equally as far advanced 
as the base of the flagellum; when retracted, they are not visible in a dorsal view. 
The antennules extend half their length past the eyes. The antennae are massive; 
the width of the first peduncle is nearly 1/4 the width of 'he carapace, and 
equal to the length of the 2nd peduncle; the 3rd is short. The flagellum is nearly 
1/3 longer than the carapace, lined on its inner surface with two rows of inward 
directed cilia, so that when the antennae are brought together a hairy tube is 
formed, as in the antennae of the genus Lepidopa. 

The chelipeds are subsimilar, unequal, slightly shorter than the length of 
the carapace; merus 1/2 the width of the carapace and nearly as wide as long, 
unarmed, rugose; the carpus is equal in length to the merus, cylindrical, widest 
distally, unarmed, rugose on entire outer surface; the major hand is slightly 
more than twice as long as wide, slightly flattened on the outer surface of the 
palm, distended on the inner, with a wide rugose crest on the upper margin, the 
outer surface is transversely rugose, the rugae anteriorly bordered with short 
setae. The lower margin is proximally median between the articulations, formed 
of an obliquely longitudinal ridge of interrupted rugae which becomes beading 
on the lower margin of the pollex. From the inner side of the palm, and ending 
at the lower marginal crest, is a row of oblique rugae. The hand is widest at the 
base of the pollex. The pollex is nearly horizontal, slightly upturned at the tip 
and armed with a short distal cutting edge. The dactyl is falcate, with the upper 
margin armed with a few spinules and setae, on the under side with a median 
blunt tooth. The fingers gape widely from base to tips. The minor hand is simi- 

424 San Diego Society of Natural History 

lar to the major, except that it is not so stout, the fingers more slender and 
curved at their tips; in addition they are armed on their cutting edges with a 
row of small teeth. These fingers also gape from base to tips. The dactyli cross 
their respective pollices on opposite sides. 

The ambulatory legs differ from each other, not alone in length, the 2nd 
and 3rd being longer than the 1st, but also in the relative shape of their carpi, 
propodi and dactyli: all three legs are margined with fine setae. The carpi and 
meri of the 2nd and 3rd legs are subequal in length to their meri, while the car- 
pus is shorter than the merus in the 1st leg. The propodi of the 1st and 2nd 
legs are distorted, subequal in length to the dactyl in the 1st, shorter than the 
dactyl in the 2nd. The dactyli of the 1st and 2nd are slightly curved at the tip, 
rounded on their upper surfaces and slightly flattened beneath. In the 3rd leg, 
which is carried up over the back, the propodus is compressed, is as wide as long, 
and is shorter than the dactyl; the dactyl is sub-equal in length to its carpus, is 
vertically compressed, curved, and with a blunt tip. The lateral edges of all three 
pairs of dactyli are lined with setae. The fingers of the manus in the chelate last 
leg are half the length of the hand. The hand is 1 mm. in length. 

The sternal piece, to which the maxillipeds are attached, is broadly trun- 
cate in front. The telson is composed of 7 plates. 

Sexual variation. — In the female the hands are subequal, subsimilar and 
slighter than those in the male. The male abdomen is narrower than that of the 

Color in alcohol. — Carapace cream, with front red and a transverse red 
median band. Fingers of chelipeds flecked with red. 

Measurements. — Male holotype: length of carapace 8.7 mm. width 4 
mm., length of antennal flagellum 12.5 mm., length of major cheliped 7.5 mm., 
of merus 2 mm., of carpus 2 mm., of manus 3.5 mm., of dactyl 1.6 mm., width 
of hand 1.6 mm., length of 1st ambulatory propodus 1 mm., of dactyl 1 mm., 
length of 2nd ambulatory propodus 1.3 mm., of dactyl 1.5 mm., length of 3rd 
ambulatory propodus 0.9 mm., of dactyl 1.6 mm. 

Range. — From Tenacatita Bay, Mexico, to La Libertad, Ecuador. 

Material examined. — The following specimens were all collected by Wood- 
bridge Williams, on Captain Fred E. Lewis' yacht "Stranger." 2 males and 2 
females, La Libertad, Ecuador, in 6-9 fathoms, sand with mud bottom; March 
24, 1937. One ovigerous female, San Jose, Guatemala, in 10 fathoms, sand bot- 
tom; April 1, 1937. One female, Isle Grande, Mexico, in 10-13 fathoms, sand 
bottom; April 8, 1937. One male and one female, Tenacatita Bay, Mexico, in 
5 fathoms, fine grain sand with shell; April 11, 1937. 

Habitat. — Found on a sand and mud, or sand and shell, bottom, in from 
5 to 13 fathoms. Without doubt, to judge from the structure of the ambulatory 
legs and the disposition of the peculiarly ciliated antennae, this species is a 
burrowing form which remains concealed just under the surface of the sand. 

Remarks. — This proposed species is closely allied to E. praelongus Stimp- 
son, 1860, of the Atlantic coast, of which it is the Pacific analogue. It differs 
in that the dactyli of the ambulatory legs, with the exception of the first pair, 
which are equal to the length of their propodi, are longer than their propodi, 

Glassell — West American Decapod Crustacea 

Plate 29 

<* X-f 

—VS5Z3SP*'' " v">^~w*K 

Euceramus panatelus Glassell, sp. nov. Male. 

426 San Diego Society of Natural History 

instead of "nearly as long as penult joint;" by the chelipeds being nearly as long 
as the carapace, instead of much shorter than the carapace; and by the maxilli- 
peds being attached to a broadly truncate sternal plate, instead of a triangular 
sternal piece. In addition, the antennae in E. panatelus are much longer than 
those of the Atlantic species. 

An examination of two specimens, a male and a female, of E. praelongus, 
furnished me by G. Robert Lunz, Jr., of the Charleston Museum, collected at 
Charleston, South Carolina, January 7, 1936, and a male specimen loaned me 
through the kindness of Dr. Waldo L. Schmitt, by the U. S. National Museum, 
collected by the U. S. Fish Commission Str. "Fish Hawk," off the west coast 
of Florida, in 3 fathoms, January 8, 1902, shows that in all three specimens the 
fingers of the chelipeds gape from base to apices, and are not as Stimpson de- 
scribed them "not gaping." 

This proposed species, on account of its elongated shape and red median 
band, suggested a shape of cigar sold under the trade-name "Panatela." 

Euceraraus transversilineatus (Lockington) , new combination 

Plate 30 

Porcellana transversilineata Lockington, Ann. Mag. Nat. Hist., ser. 5, vol. 2, 
1878, p. 405 (type-locality, Boca de las Piedras, Sinaloa, Mexico; types 
not extant) . 

Lockington's type-locality falls well within the Gulf of California, if I am 
correct in assuming that Boca de las Piedras, Sinaloa (name not in the Coast 
Pilot) , is the entrance to the Estero de las Piedras. The estero opens on the gulf 
6 miles southward of the mouth of the Rio del Fuerte or Santa Maria de Ahome, 
and is in lat. 25° 50' N. and long. 109° 25' W. His second locality, Angeles 
Bay, Baja California, is in the upper part of the Gulf of California. This series, 
like all of Lockington's type material, was destroyed by fire in the San Francisco 
disaster of 1906. 

On April 13, 1937, at Punta Penasco, Sonora, Mexico, I took a small series 
of 3 specimens, at extreme low water, from the sand and shell material at the 
base of gorgonian corals. On May 8, 1937, at San Felipe, Baja Calfiornia, 
Mexico, I secured another specimen under similar collecting conditions. In addi- 
tion to the above, I took, on January 1, 1932, off the north end of Tiburon 
Island, Gulf of California, a female in 12 fathoms. From this material I am 
designating one female the neotype and one male the allotype. 

Neotype.— Female; Cat. No. 1127, San Diego Society of Natural His- 
tory; from off the north end of Tiburon Island, Gulf of California, Mexico; 
January 1, 1932, 12 fathoms; collected by Steve A. Glassell. 

Allotype.— Male; Cat. No. 1128, San Diego Society of Natural History; 
from Punta Penasco, Sonora, Mexico; April 13, 1937, low tide; collected by 
Steve A. Glassell. 

Diagnosis. — Carapace 1/3 longer than wide, transversely striate. Antennae 
subequal in length to width of carapace; basal peduncle short; flagellum naked. 

Glassell — West American Decapod Crustacea 

Plate 30 


Z rrv m . 

Enceramus transversilineatus (Lockington) . Male. 

428 San Diego Society of Natural History 

Chelipeds in females subequal, similar; in males unequal, dissimilar; merus with 
an inner distal spine; carpus with a submedian inner spine. Outer maxillipeds 
attached to strongly arched sternal segment. Dactyli of ambulatories subequal, 
stout, falcate. Telson of abodmen much larger in female, triangular, with 7 

Description. — Carapace elongate, broadest posteriorly, about 1/3 longer 
than wide, regions well-marked; surface transversely crossed by well-spaced, 
asymmetric, distinct striations which begin at the inner edge of the obliquely pli- 
cated, thin, lateral margins. Distally the lateral margins end in a sharp, forward- 
pointing spine, separated from the outer antennal-marginal tooth by a V-shaped 
notch. Posterior margin a concave obtuse angle. Protogastric ridge distinct, 
toothed, divided by a median sulcus. Front tridentate, the median slightly ad- 
vanced beyond the laterals, which are on a slightly lower plane. The eyes are 
retractile, as in the genus. The antennae are short, the 2nd peduncle the longest; 
flagellum naked. 

The chelipeds in the female are equal, subsimilar; in the male they are dis- 
similar and much stouter. The merus is distally armed on the inner margin with 
a sharp spine, rugose on upper and lower surfaces as is the carpus and manus; 
carpus on upper surface rectangular, flattened, with a single, submedian, inner 
marginal spine, and in addition there may be one or more spinules distally. The 
major hand in the male is stout, thick, with rugose upper carina and a spined 
lower margin; the fingers gape from base to blunted tips; the dactyl is crested 
with smooth granules, and is armed on the under edge with 3 or more large 
teeth; the proximal is double; the pollex is horizontal, armed with a row of low 
irregular lobes. The minor hand is narrow, with a median rugate ridge which 
divides the outer surface of the hand into two subconcave surfaces; the lower 
margin is spined, the fingers long, not gaping, the tips sharp. In the female the 
hands are like the minor hand in the male. In both sexes the outer surface is 
sparsely covered with clavate setae. 

The ambulatory legs are stout, rugose and lightly covered with setae; the 
dactyli are subsimilar, long, stout, falcate, tips corneous. 

The male abdomen is much smaller than that of the female, as is the telson. 
The outer maxillipeds are attached to a broadly arched first sternal segment. 

Color in alcohol. — Deep cream with markings of orange-red. 

Measurements. — Female neotype: length of carapace 8.3 mm., width 
6.1 mm., length of hand 5 mm., width 1.5 mm., length of telson 4.1 mm., width 
at base 2.1 mm. Male allotype: length of carapace 6.2 mm., width 4.2 mm., 
length of major hand 5 mm., width 2.6 mm., thickness 1.6 mm., length of telson 
1.9 mm., width at base 1 mm. 

Range. — So far known only from the Gulf of California, Mexico. 

Material examined. — The female neotype (see neotype) . A series of 2 
males and 1 female from Punta Penasco, Sonora, Mexico, taken April 13, 
1937, one of which is the allotype (see allotype) . A single female from San 
Felipe, Baja California, Mexico, taken May 8, 1937. All collected by the author. 

Habitat. — Found from extreme low water, partly covered with sand and 
shell fragments, to a depth of 12 fathoms. 

Glassell — West American Decapod Crustacea 

Plate 3 1 

Fig. 1. Minyocerus for fa Glassell, sp. nov. Antennae. 

Fig. 2. Minyocerus farfa Glassell, sp. nov. Left 3rd ambulatory le£ 

Fig. 3. Minyocerus farfa Glassell, sp. nov. Male holotype. 

430 San Diego Society of Natural History 

Remarks.- — This species of Lockington's is more closely allied to E. praelon- 
gus Stimpson, 1860, than it is to E. panatelus Glassell, although it differs in 
many respects from both of these species: (1) the carapace is shorter for its 
breadth, (2) the dactyli of the ambulatory legs are more nearly uniform and 
falcate, (3) the antennae are shorter and naked, (4) the major hand in the 
male is heavier in proportion to its length, (5) the striations on the carapace are 
more distinct, (6) the telson is longer in proportion to its width. 

Lockington in his description of this species failed to mention the sex of 
his specimens, but it is evident that these were all females — "females with ova," 
for, had he also had adult males he would have noted the dimorphic character 
of the hands. 

Genus Minyocerus Stimpson 

Minyocerus kirki, sp. nov. 
Plate 31, figures 1-3 

Type. — Male, holotype, Cat. No. 1129, San Diego Society of Natural 
History; female, paratype, Cat. No. 1130, S. D. S. N. H.; from San Felipe, 
Baja, California, Mexico, low tide; May 11, 1937; collected by Steve A. Glassell. 

Diagnosis. — Carapace convex in both directions, oblong. Chelipeds stout, 
rugose; carpus armed on inner margin. Ambulatory legs lightly ciliate; merus 
stout, rugose. 

Description. — Carapace nearly 1/3 longer than wide, suboblong, convex in 
both directions, highest longitudinally along median line, lightly rugose, more 
distinct posterior to the faintly outlined cervical groove. Front with three sub- 
equal teeth, the median slightly advanced, if any, past the laterals. A sharp 
forward- and upward-pointing spine at the shoulder, posterior to which the 
sides are subparallel. Posterior margin slightly concave. The antennules extend 
past the apices of the frontal spines. The antennae are extremely minute and 
difficult to locate without staining. They are placed posterior to the outer orbital 
spine, have 3 movable joints and a rudimentary flagellum; their total length 
is not equal to the width of the cornea of the eye. The eyes are retractile, on 
cylindrical white stalks, and in life the cornea is extended forward as far as the 
tips of the lateral spines. 

Chelipeds stout in the male, more slender in the female, subsimilar, slightly 
unequal, more pronounced in the male; merus stout, rugose on upper surface, 
armed on inner distal margin with a sharp spine; carpus rugose and flattened 
on upper surface, armed on inner margin with a large median spine, followed 
distally by several spinules; the length of the major hand, in the male, including 
the fingers, is equal to the width of the carapace, and is nearly 1/3 as wide as 
long; the hand in the female is shorter, but has the same length to width ratio; 
the outer margins are subparallel, the lower fringed with cilia; the fingers are 
close-fitting, their tips obtuse. 

Ambulatory legs stout, margined with microscopic cilia; merus stout, nearly 
as wide as long, unarmed, rugose; the dactyli are lanceolate, sharp, curved at 
tip, nearly as long as their carpi. The telson has 7 segments. 

Glassell — West American Decapod Crustacea 431 

Color in life. — Carapace with median longitudinal area white with a yellow 
cast, branchial areas brown with a greenish cast. Antennules blue, flagellum 
yellow. Palp of maxillipeds light green. Chelipeds and ambulatory legs with a 
whitish ground banded with brown. The carapace colors extend onto the first 
two abdominal segments. ( Wm. A. Kirk, from field sketch) . 

Measurements. — Male holotype: length of carapace 3.5 mm., width 2.5 
mm., length of hand including fingers 2.5 mm., width 1 mm. Female paratype: 
length of carapace 4 mm., width 2.8 mm., length of hand 2 mm., width 0.7 mm. 

Range. — Known only from type-locality. 

Material examined. — A series of 4 males and 4 gravid females, collected 
at San Felipe, Baja California, Mexico, May 11, 1937, by the author. 

Habitat. — Found at extreme low water commensal on the sand starfish 
Luidia Columbia (Gray). A pair of crabs was usually found on a single starfish, 
one on the dorsal, the other on the ventral side. 

Remarks. — This proposed species is closely allied to M. angustus (Dana), 
1852, but differs from that species in the following respects: the upper surface 
of the carpus of the chelipeds is nearly as wide as long, depressed and armed 
with a large inner marginal tooth, instead of being oblong, nearly entire, and by 
the meri of the ambulatory legs being stout, instead of slender. 

Carlos Moriera, 1901, places Fritz Miiller's Porcellana stellicola, in syno- 
nymy for M. angustus (Dana), and the same differences exist between my pro- 
posed species and that of Miiller's, with the additional difference that in M. 
kirfa, the antennae are composed of the usual three movable segments and a 
rudimental flagellum, instead of having six segments and a rudimentary flagel- 
lum, as figured in Ann. Mag. Nat. Hist., ser. 3, vol. 11, 1863, pi. 1, fig. 2. 

This proposed species is named for my worthy friend Mr. William A. 
Kirk, of Los Angeles, California, who accompanied me to the Gulf of Cali- 
fornia, and made the discovery of this obscure little anomuran. 

Porcellana magdalenensis Glassell 
Plate 32, figures 1, 2 

Porcellana magdalenensis Glassell. Trans. San Diego Soc. Nat. Hist., vol. 8, 

no. 21, 1936, p. 295. 

At the time this species was described, the only specimens at hand were a 
series of five females, two of which were juvenile. Since then, a series of five 
males and one dismembered, ovigerous female, was collected in Acapulco Bay. 
State of Guerrero, Mexico, by Mr. Woodbridge Williams, on Captain Fred 
E. Lewis' yacht "Stranger," April 6, 1937. The adult males in this series 
differed in so many respects from the already described females, that they were 
at first considered to be a separate species. However, a close study of the juve- 
nile and adolescent specimens proved their dimorphic character. 

Description of male. — Carapace nearly smooth, except for light pubescence 
anteriorly, cervical groove well defined. Front broad, slightly less than half the 
length of the carapace, tridentate, the median twice the size of the laterals. 

432 San Diego Society of Natural History 

triangular, with a median longitudinal sulcus, microscopically margined with 
spinules, depressed and obtuse at the tip. The laterals are half the length of 
the median tooth, from which they are separated by a V-shaped sinus. Their 
outer spinous margins form the upper ocular margins. The lateral margin of 
the carapace is bordered with a row of sharp, upward- and forward-pointing 
spines, and is continued onto the carapace, behind the cervical groove, forming 
an unarmed though slightly granulose shoulder. The posterior margin is nearly 
straight. The antennal flagellum exceeds the length of the chelipeds. 

The chelipeds are long, unequal, dissimilar; merus nearly smooth, unarmed 
on carpal articulation, with a wide, anteriorly produced, compressed, inner, distal 
lobe, well dentated on the margin in adolescents and females, nearly obsolete 
in adult males, carpus microscopically rugose on the slightly rounded upper 
surface, 1/3 longer than wide, and the upper, inner margin may or may not be 
armed with two small spines in the adults. The major manus is naked, stout, 
unarmed on margins or surfaces; the upper margin of the palm has a slight 
carina; a blunt longitudinal median ridge extends from the proximal end to a 
point near the gape; from this ridge to the outer, slightly beaded or simply round- 
ed margin, the surface is slightly concave, as is the outer surface of the pollex. The 
pollex is short, blunt, stout, and armed with a single low lobe. The dactyl is 
smooth, slightly curved, stout and armed with a median lobe. The fingers gape 
from their bases to their blunt apices. There is a trace of pubescence in the gape. 
The minor cheliped has the two inner marginal spines of the carpus more dis- 
tinct, the entire margin roughened with smaller spines; in adolescents and 
females the outer carpal margin has a row of upturned spines which are lacking 
in adult males; the manus is narrow, contorted, armed on its outer margin with 
a row of spines, partly concealed in pubescence, inner margin smooth; a median 
ridge, armed distally with sharp spines, divides the outer surface of the palm into 
concave surfaces, the outer pubescent. The arched, sharp-pointed dactyl is longer 
than the palm, crested with a row of spinules, and, on the scooped-out under 
surface, is setose and pubescent. The pollex is distorted, sharp-tipped, slender, 
and, like the dactyl, is setose and pubescent on its cutting edge. 

Ambulatory legs long, as in the female; dactyl of the first pair extending 
past the distal end of the carpus of the chelipeds. The telson is composed of 7 

Color in alcohol.— Carapace cream. Chelipeds orange-red. Ambulatory legs 
cream, banded with red or orange. 

Measurements. — Adult male: length of carapace 3.6 mm., width 3.8 mm., 
length of carpus 3 mm., width 2 mm., length of major manus 5.5 mm., width 
2.2 mm., length of minor manus 4.5 mm., width 1.3 mm. 

Range. — From Magdalena Bay, Baja California, Mexico, to Panama. 

Material examined. — In addition to the type-series from Magdalena Bay, 
I examined an adolescent male, from Perico Island, Panama, collected by the 
U. S. Fish Commission, S. S. "Albatross," October 26, 1904. This specimen 
was sent me for identification by Dr. Waldo L. Schmitt of the U. S. National 
Museum, and has been returned to that institution. 

Remarks. — The juveniles of both sexes are quite similar to the adult female 

Glassell — West American Decapod Crustacea 

Plate 32 


jt/j : ■■ i 

Fig. 1. Porcellana magdalenensis Glassell. Female. 
Fig. 2. Porcellana magdalenensis Glassell. Male. 

434 San Diego Society of Natural History 

Ulloaia, gen. nov. 

Carapace oblong-ovate, slightly longer than broad, convex, regions defined, 
surface squamo-tuberculate, lateral margins carinate, toothed. Front in dorsal 
view with a deep, wide, V-shaped, median notch, on each side of which are 2 
short, multi-spined spinules, separated from each other by a notch. The median 
or rostral process in frontal view is subvertical, truncate and serrate on the lower 
edge. Eyes small, not retractile. First antennal peduncle removed from the eye, 
not joining the margin of the carapace; nagellum short, slightly more than 1/2 
the width of the carapace. Chelipeds short; carpus cylindrical, slightly longer than 
wide; hands compressed, weak. Ambulatory legs short, compressed, bent; dactyli 
simple, not multiunguiculate. 

This proposed genus is rather distantly related to Minyocerus, Stimpson, 
1858, in which the carapace is concave, the ambulatory legs short, the dactyli 
not multiunguiculate. In the shape of the carapace and its peculiar front, this 
genus differs from all the other genera in the family; in fact, in the general 
appearance of the carapace, it somewhat resembles species of the genus Mithrax, 
Latreille, 1817, of the family Majidae. 

Genotype. — Ulloaia perpusillia, new species, taken at Punta Penasco (Rocky 
Point), Sonora, Mexico, low tide, April 12, 1937, by Steve A. Glassell. 

Remarks- — This proposed genus is named for Francisco de Ulloa, conquis- 
tador, explorer and navigator, who was the first to prove that Baja California 
was not an island, by directing his ship into the treacherous upper reaches of 
the Gulf of California, after which he traversed the eastern coast of the penin- 
sula, doubled Cabo San Lucas, at the lower end, and sailed westward into the 
setting sun. 

Ulloaia perpusillia, sp. nov. 
Plate 33, figure 1 

Type. — Male, holotype, Cat. No. 1131, San Diego Society of Natural 
History; from Punta Penasco, Sonora, Mexico, low tide; April 12, 1937; col- 
lected by Steve A. Glassell. 

Diagnosis. — Carapace oblong-ovate, longer than broad, convex, with a 
raised, longitudinal, median groove dividing the carapace into two halves. Gas- 
tric and cardiac regions raised. Posterior margin convex, entire. Branchial region 
with raised, flat-top tubercles. Telson with 7 plates. 

Description. — Carapace oblong-ovate, longer than broad, convex, regions 
well defined; a longitudinal groove over the gastric and cardiac regions extend- 
ing from the front to the posterior border divides the carapace into halves. 
The gastric and cardiac regions are raised, separated from each other by a well 
defined sulci; the median groove on the gastric region is bordered by longitudinal 
rugose ridges, on the cardiac region by rounded excrescences. The branchial 
regions have numerous wart-like, truncate-tipped, excrescences arising from the 
punctate surface, these granulose ridges and warts being more prominent on the 
posterior half of the carapace. The intestinal region is free from tubercles except 

Glassell — West American Decapod Crustacea 

Plate 33 

Fig. 1. Ulloaia perpusillia Glassell, sp. nov. Male holotype. 
Fig. 2. Fabia granti Glassell. Male. 

436 San Diego Society of Natural History 

for small granules bordering the median groove. The front in a dorsal view 
has a deep, granulous, median notch, on each side of which are two forward- 
and upward-pointing spinules, separated from each other by a small V-shaped 
notch. In a frontal view the rostral process is subvertical, truncate and serrate 
at the tip, a sort of apron between the antennules. There is a tubercle on the 
upper ocular margin. The lateral margin is a carinate row of granulose lobes, 
diminishing in size anteriorly. The antennae are short; the joints of the flagellum 
rather long, with sparse cilia. 

Chelipeds stout, short (only one chela remaining on the two specimens) ; 
merus short, stout, armed with an inner distal, lamellar lobe, width extremely 
narrow on the anterior carpal articulation, triangular and wide on the upper 
posterior face; carpus slightly longer than wide, inner margin with a median, 
serrate lobe; upper surface very rough, with an uneven median ridge bordered by 
sulci, a twisted row of tumid excrescences anterior to the median ridge, a row of 
unequal serrate lobes on the outer margin. The lower surface of the merus and 
carpus are on one plane, and flat. The hand is weak, compressed, shorter than 
the combined length of the lower surface of the carpus and merus, flexed, the 
arch of travel in the carpal articulation being small. The under surface of the 
hand is tomentose, granulate, and near the inner side are several longitudinal 
rows of beading. The upper surface is flattened, with a median ridge, the outer 
margin spinose; the inner margin of the palm has an upward-turned crest, and 
between this crest and the median ridge the surface is concave. Fingers short, 
thin, close-fitting, 1 /3 the length of the hand, tips crossed. 

Ambulatory legs are short, compressed, rugose, spined and lightly margined 
with tomentum; merus wide, postero-distal lamellar process shields 1/2 the 
length of the carpus on its posterior face; the carpus has a like projection over 
1/3 the propodal length; propodus fluted with ridges of spinules; dactyli fal- 
cate, corneous tipped. The telson is composed of 7 plates. 

Color in alcohol.- — Cream tipped with orange-red. 

Measurements. — Male holotype: length of carapace 3.5 mm., width 3.1 
mm., length of carpus 1.5 mm., width 1.1 mm., length of hand 2.2 mm., width 
1.2 mm. 

Range. — Known only from the type-locality. Gulf of California. 

Material examined. — One male and one ovigerous female (see type) . 

Habitat. — Found among gorgonian corals, sponges and bryozoan growths, 
at extreme low tide. 

Remarks. — The aberrant type, herein described, somewhat resembles, in 
the shape of the carapace, Ethusa sexdentata (Stimpson), as figured in Smith- 
sonian Misc. Coll., vol. 49, no. 1717, 1907, pi. 19, fig. 4, with due allowance 
for family differences. 

Pisonella, gen. nov. 

Carapace suboval, orbicular, slightly convex laterally, lateral margins high, 
ridged. Front depressed, arched and subentire in dorsal view, with median alone 
or with median lateral projections when viewed from the front. Eyes very small. 

Glassell — West American Decapod Crustacea 437 

First article of outer antennae produced, joining margin of carapace, as in the 
genus Porcellana; flagellum longer than carapace. Chelipeds short, stout; carpus 
with inner margin armed or unarmed; hands thick. Ambulatory legs stout; 
dactyli simple, not multiunguiculate. 

This proposed genus has a close affinity to the genus Pisosoma Stimpson, 
1858, which is based on P. pi sum (M. Edw.), 1837, but differs in that the eyes 
are smaller and that the basal segment of the antennae is removed from the 
ocular hiatus, instead of the eyes being large, and the first article of the outer 
antennae short, not reaching upper margin of the carapace and occupying a 
portion of the ocular hiatus, as in the genera Pisosoma and Petrolisthes Stimp- 
son, 1858. It differs, also, from the genus Porcellana. Lamarck, restricted Stimp- 
son, 1858, by the carapace not being generally longer than broad, the front not 
tridentate and prominent. 

Genotype. — Pisonella sinuimanus (Lockington) , ( = Pisosoma sinuimanus 
Lockington) . 

Remarks. — This genus is proposed for the reception of the following 

Pisonella sinuimanus (Lockington), (—Petrolisthes {Pisosoma) sinui- 
manus Lockington) , the genotype. 

Pisonella tuberculipes (Lockington), (—Pachychcles tuberculipes Locking- 
ton, = Polyonyx tuberculipes (Lockington) Nobili) . 

Pisonella smithi (Glassell) , (^Pisosoma smithi Glassell) . 

Pisonella erosa (Glassell), (^Pisosoma erosa Glassell). 

Key to the Species of Pisonella 

A 1 . Telson of abdomen with 7 plates. Carapace without lateral spines. 

B 1 . Carapace with light transverse plications or nearly smooth. Chelipeds 
smooth or lightly granulated on under surface. 
C. Carapace nearly smooth. Chelipeds granulate; carpus armed 

with an inner marginal lobe; hands unequal sinuimanus 

C 2 . Carapace lightly rugose. Chelipeds granulate on hands; carpus 

unarmed, lightly rugose; hands subsimilar smithi 

B 2 . Carapace heavily eroded. Chelipeds eroded on upper surface, rugose 
and roughened on under surface; carpus armed, eroded; hands 

unequal. Ambulatory legs eroded erosa 

A 2 . Telson of abdomen with 5 plates. Carapace with lateral spines. Chelipeds 
heavily tuberculated on upper surface, smooth on under side; carpus 
armed; hands in male dissimilar tuberculipes 

Pisonella sinuimanus (Lockington), new combination 

Plate 34, figure 2 

Petrolisthes (Pisosoma) sinuimanus Lockington, Ann. Mag. Nat. Hist., ser. 5, 
vol. 2, 1878, p. 401 (type-locality [?], La Paz and Port Escondido, Baja 

438 San Diego Society of Natural History 

California, Mexico; types not extant) . 
Petrolisthes sinuimanns (Lockington) Nobili, Boll. Mus. Zool. Anat. comp. R. 

Univ. Torino, vol. 16, no. 415, 1901, p. 15 (Isle of Flamenco, Ecuador). 

— Rathbun, Proc. U. S. Nat. Mus., vol. 38, 1910, p. 599. 
Pisosoma sinuimanus Lockington. Glassell, Zoological N. Y. Zoological Soc, 

vol. 22 (part 1) , no. 4, 1937, p. 83. 

This species was found at two localities on the Gulf coast of Baja Califor- 
nia, Mexico; La Paz and Puerto Escondido, and was described by Lockington 
in 1878. In 1906 the type series was destroyed in the San Francisco disaster. 

In 1931 and in subsequent years, I have collected this species throughout 
the Gulf of California, and have examined many other specimens collected in the 
same locality. From a small series collected at Puerto Escondido (Hidden Har- 
bor) , I am designating one male the neotype, and one female the allotype. 

Neotype. — Male; Cat. No. 1132, San Diego Society of Natural History; 
from Puerto Escondido, Baja California, Mexico; December 19, 1931; collected 
by Steve A. Glassell. 

Allotype. — Female; Cat. No. 1133, San Diego Society of Natural History; 
Puerto Escondido, Baja California, Mexico; December 19, 1931; collected by 
Steve A. Glassell. 

Diagnosis. — Carapace suboval, anteriorly depressed, lateral margins granu- 
lar. Front with a median, triangular, depressed lobe in front view. Carpus unilo- 
bate, ridged. 

Description. — Carapace suboval, convex fore and aft, depressed anteriorly, 
lightly punctate, regions lightly denned, lateral margins granular, lightly serrate; 
posterior margin a concave obtuse angle; front entire in dorsal view, arched, 
viewed from the front, with a median, triangular, depressed lobe. 

Chelipeds unequal, similar; merus with a blunt, granulate, low, longitudinal 
lobe upon the inner distal margin; carpus more than half as wide as long, armed 
with a single, granulate, blunt tooth on proximal half of anterior margin, upper 
surface granulate, with three longitudinal, rolling ridges, divided by furrows, 
median ridge the most elevated; hands unequal, subsimilar, thick, with four 
longitudinal rolling ridges, divided by furrows, entire outer face granulate; 
outer margin thick, granulate, to upturned thick tip of pollex; inner margin 
sharply oblique, forming a flattened, triangular surface in a vertical plane whose 
base is proximal, from near proximal end to a low lobe behind the upper base of 
the dactylus, the line from the base of the hand to the base of the finger thus 
forming an obtuse angle; the dactyli are sinuous, stout, cylindrical, with blunt, 
curved, lobular tips; dactyl of major hand armed with two blunt teeth, the 
pollex with one. 

Ambulatory legs stout, granulate; carpus and propodus more granulate than 
merus; carpus of 1st and 2nd legs produced backward at posterior distal end; 
Dactyli stout, curved at corneous tip, setaceous, armed on under margin with 
a row of spines; propodus of 1st leg armed with spines on posterior margin. 
Epimera and abdomen fimbriate. Legs with a few setae. 

Color in life. — The color varies from light cream to buff; the ventral side 
is slightly iridescent. 

Glassell — West American Decapod Crustacea 

Plate 34 

.*" \ • , . -'V • 


myt u 



mm x A 41P&L 


•: «'* : 





2mm , 

j&? : ' 

Fig. 1. Pisonella tuberculipes (Lockington) . Male neotype. 
Fig. 2. Pisonella sinuimanus (Lockington). Male neotype. 

440 San Diego Society of Natural History 

Measurements. — Male neotype: length of carapace 6 mm., width 6 mm. 
Female allotype: length of carapace 4.5 mm., width 5 mm. 

Range. — From the Gulf of California to Ecuador (Nobili) . 

Material examined. — The types were selected from a series of 7 males and 
2 females, collected by the author, at Puerto Escondido (Hidden Harbor), 
Baja, California, Mexico; December 19, 1931; at low tide under rocks. 

A series of 20 males and 20 females, from the NE end of Tiburon Island, 
Gulf of California; January 2, 1932; collected by the author. 

Habitat. — Found on the under side of rocks in the lower inter-tidal zone 
to a depth of 3 fathoms. 

Remarks. — The sexes may be instantly determined by the shape of the 
terminal segments of the abdomen: in the male the ultimate plates are short and 
wide, while in the female they are subquadrate; the penultimate lateral plates 
in the male are long and narrow, the margins subparallel, while in the female 
these plates are subtriangular, widest distally. The telson is composed of seven 

Contrary to Lockington's description, in which he states that this is a 
variable species, I have found very little variation, except as to size and sex. 
A few specimens may show considerable roughness on the inner side of the 
hands, the carapace may be more punctate or lightly pubescent in others, but the 
carpus of the chelipeds is always armed with the proximal lobe on the anterior 
margin, even though distally the margin may be produced almost as far forward 
as the apex of the lobe. I have examined several hundred specimens of this 
species, all from the Gulf of California. 

Pisonella tuberculipes (Lockington) , new combination 
Plate 34, figure 1 

Pachycheles tuberculipes Lockington, Ann. Mag. Nat. Hist., ser. 5, vol. 2, 

1878, p. 404 (type-locality, La Paz, Baja California, Mexico; type not 

extant) . 
Polyonyx tuberculipes (Lockington) Nobili, Boll. Mus. Zool. Anat. comp. R. 

Univ. Torino, vol. 16, no. 415, 1901, p. 21 (Bay of S. Elena, Ecuador). 

— Rathbun, Proc. U. S. Nat. Mus., vol. 38, 1910, p. 601. 

Lockington described this species from 5 specimens (sex not noted, though 
undoubtedly females) taken at La Paz, and other ports on the Gulf of Cali- 
fornia. This type series was destroyed in the same manner and at the same time 
as the types of P. sinuimanus. 

In 1935 I collected a large series of this species at Punta Pehasco, Sonora, 
Mexico. From this material I am designating one male, the neotype, and one 
female, the allotype. 

Neotype. — Male; Cat. No. 1134, San Diego Society of Natural History; 
from Punta Pehasco, Sonora, Mexico; May 2, 1935; collected by Steve A. 

Allotype. — Female; Cat. No. 1135, San Diego Society of Natural History; 

Glassell — West American Decapod Crustacea 441 

from Punta Penasco, Sonora, Mexico; May 2, 1935; collected by Steve A. 

Diagnosis. — Carapace slightly convex; lateral margins dentate; regions 
defined, front projecting, subentire in dorsal view. Chelipeds in male, unequal, 
dissimilar, covered with granulate tubercles. Ambulatory legs tuberculate. 
Flagellum lightly ciliate. Telson with 5 plates. 

Description. — Carapace slightly longer than wide, measured from tips of 
spines, convex, with scattered tufts of pubescence, regions defined, areolate. 
Lateral margins with 6 or 7 teeth, their upper surface covered with spinules, the 
anterior tooth a spine-tipped lobe, nearly twice the width of the base of the 2nd 
tooth; the posterior tooth is the extension of a short ridge on the carapace; 
inside of the lateral spines, on the carapace, are several small granulate tubercles. 
The protogastric ridge is sharply defined, tomentose, extending much higher 
than the horizontal frontal region, and separated from a small, serrate-margined, 
hepatic lobe, by a shallow sinus. The front projects forward, is broadly arched 
or subtriangular, with a median furrow; in a dorsal view the outer margin is 
subentire, and granulous; in a front view the median tooth is sharply depressed, 
triangular, acute; the lateral lobes are separated from the median by a high 
arched sinus, and are turned down and slightly under. The upper ocular margin 
has a median, granulate tubercle, as has the upper border over the basal article 
of the antennae. There is a sharp spine on the epistome below the basal antennal 
article. The flagellum of the antennae is lightly ciliate. 

The chelipeds in the male are stout, unequal, dissimilar and covered on 
their upper surface with numerous granulous tubercles; the under surface is 
smooth; in the females the chelipeds are more nearly equal. Merus short on 
upper surface, broad, armed on inner margin with a short, lamellar, granular, 
distal lobe; carpus longer than broad, armed on inner margin with a proximal 
subhorizontal, granulous spine, nearly half as long as the inner carpal margin; 
from the distal base of this tooth to the distal end of the carpus, the margin is 
outwardly oblique and armed with several, short, stout spines. The upper surface 
of the carpus is covered with spinose tubercles, with upturned spines on the 
outer margin. The hands are thick, contorted, grotesque, unequal in the males, 
dissimilar, and are 1/3 longer than their carpi. The fingers of the major hand, 
in the male, are widely gaping, strongly curved, blunt tipped; the dactyl is fal- 
cate, armed with a large median and proximal tooth; the pollex with a smaller 
distal tooth. The fingers of the minor hand, which resemble those of both hands 
in the female, gape in a lesser degree; the dactyl is armed with a row of well 
formed teeth, the proximal the largest. The outer margins of the palms are 
bordered with spines and setae. The carpus and hands, on their upper, outer 
surfaces, are pubescent. 

The ambulatory legs are stout, roughened with rugae and granulous tuber- 
cles, and are covered with tomentum. The telson of the abdomen is composed 
of 5 plates. 

Sexual variation. — In the female the hands are more nearly alike than in 
the male. As in P. sinuimanus, the ultimate plates in the male telson are short 
and wide, the penultimate lateral plates long and narrow, the margins subpar- 

442 San Diego Society of Natural History' 

allel; in the female the ultimate plates are subquadrate, the laterals widest 

Color in life. — Muddy grey, with a dark patch on the central regions. In 
alcohol the carapace and chelipeds are light pink. 

Measurements. — Male neotype: length of carapace 4.1 mm., width 3.9 
mm., length of carpus of major cheliped 3 mm., of hand 4 mm. Female allotype 
(ovigerous) : length 3.1 mm., width 3.1 mm. 

Range. — From the Gulf of California, Mexico, to Ecuador (Nobili). 

Material examined. — Several series of both sexes, numbering more than 25 
specimens, collected by the author at San Felipe, Baja California, Mexico, 
June 5, 1933; and Punta Penasco, Sonora, Mexico, May 2, 1935, and April 
12, 1937. 

Habitat. — This little crab is found on sponge incrusted sea-fans, but more 
frequently on the rough sponges themselves, at extreme low water. They are 
quite numerous, though obscure. 

Remarks. — Lockington's description of this species is clear and unmis- 
takable. That he placed it in the genus Pachycheles, was due, more to the im- 
portance placed on the conformation of the chelipeds than its anatomical struc- 
ture, for while the chelipeds might very easily belong to Pachycheles, the epimera 
is entire, a peculiarity that removes it from that genus. 

Nobili placed this species in the genus Polyonyx, with reservations, as he 
was satisfied that it did not meet with all the requirements of that genus. He 
was influenced by noting a small spinule on the dactyli of the ambulatory legs. 

To obviate any chance of hidden characters remaining obscured by tomen- 
tum, I depilated the largest male in my series (the neotype) , by using a weak 
solution of sodium hypochlorite. When thus cleaned, the dactyli show only the 
usual small spines on the under margin, which are to be found on nearly all the 
uniunguiculate dactyli in this family. 

Pisonella smithi (Glassell), new combination 

Pisosoma smithi Glassell, Trans. San Diego Soc. Nat. Hist., vol. 8, no. 21, 
1936, p. 286 (type-locality, Miramar Beach, near Guaymas, Sonora, 
Mexico) . 

Pisonella erosa (Glassell), new combination 

Pisosoma erosa Glassell, Trans. San Diego Soc. Nat. Hist., vol. 8, no. 21, 
1936, p. 289 (type-locality, Magdalena Bay, Baja California, Mexico). 
Types of the above two species are located in the U. S. National Museum, 

and with the San Diego Society of Natural History. 

Key to the West North American Species of Petrolisthes 

A 1 . Hands bordered with setae, margined with spines. Chelipeds with carpus 
armed. Ambulatory legs pubescent or setose; upper margin of meri 

Glassell — West American Decapod Crustacea 443 

B 1 . Front trilobate. Chelipeds pubescent; hands dissimilar; fingers with 
pubescence in gape. 
C 1 . Carapace pubescent, with distinct striations. Carpus twice as 
long as wide. Meri of ambulatory legs with postero- 

distal end spined. Abdomen pubescent hirtipes 

C 2 . Carapace naked without distinct striations. Carpus less than 
twice as long as wide, armed with 3 or more spines. Meri 
of ambulatory legs at postero-distal end unspined. 

Abdomen naked nigrunguiculatus 

B 2 . Front triangular. Carapace with distinct striations. Under side of 

hands roughened; fingers with a short pile of pubescence; carpus 

with 3 or more spines. Ambulatory legs setose. 

C 1 . Carapace pubescent. Chelipeds pubescent; hands similar, carpus 

with 6 spines, twice as long as wide. Meri of ambulatory 

legs at postero-distal end spined. 

Abdomen pubescent sanfelipensis 

C 2 . Carapace naked. Chelipeds naked; hands dissimilar; carpus 
with 4 or 5 spines, less than twice as long as wide. Meri 
of ambulatory legs at postero-distal end unspined. 

Abdomen naked polymitus 

A 2 . Hands unmargined with setae. 

B 1 . Hands unmargined with spines, similar. 

C 1 . Fingers pubescent in gape. Meri of ambulatory legs at postero- 
distal end unspined, upper margin unarmed. Chelipeds 
naked; under side of hands smooth. Carapace naked. 
D 1 . Carapace smooth. Carpus unarmed; margins parallel. 

E 1 . Front triangular. Carpus twice as long as wide. Am- 
bulatory legs pubescent eriomerus 

E 2 . Front trilobate. Carpus more than twice as long as 

wide. Meri of ambulatories naked gracilis 

D 2 . Carapace roughened, regions well marked. Front trian- 
gular. Carpus less than twice as long as wide, armed with 

a lamellar lobe cinctipes 

C 2 . Meri of ambulatory legs armed on upper margin. Chelipeds 

with under side of hand roughened. 

D 1 . Fingers pubescent in gape. Meri of ambulatory legs at 

postero-distal end unspined. Carapace pubescent, areolate, 

with distinct striations. Chelipeds pubescent, with carpi 

unarmed rathbunae 

D 2 . Movable finger with a short pile of pubescence only. Meri 
of ambulatory legs at postero-distal end spined. Carapace 
heavily striate. Chelipeds naked; carpi armed with 3 or 

more spines edwardsii 

B 2 . Hands unmargined with spines, under side smooth, dissimilar; fingers 
with a short pile of pubescence; carpus less than twice as long 
as wide. Carapace pubescent. Ambulatory legs pubescent; meri 
unarmed on upper margin, at postero-distal end unspined. 

444 San Diego Society of Natural History 

C 1 . Carapace with regions well marked; frontal trilobate. Chelipeds 

pubescent; carpus armed crenulatus 

C 2 . Carapace with regions indistinct; front triangular. Chelipeds 

naked; carpus unarmed schmitti 

B 3 . Hands margined with spines, under side roughened, similar; fingers 
with a short pile of pubescence. Chelipeds pubescent; carpus 
twice as long as wide, armed with 5 or 6 spines. Carapace 
pubescent, surface smooth; front triangular. Ambulatory legs 

setose; meri armed, postero-distal end spined hirtispinosus 

A 3 . Hands unmargined with setae, margined or unmargined with spines, under 
side smooth. Chelipeds naked or pubescent. Meri of ambulatory legs 
B 1 . Carapace naked or pubescent, surface smooth. Front triangular. 
Chelipeds naked or pubescent; carpus twice as long as wide, 
armed with 3 spines; hands dissimilar, unmargined with 
spines; fingers with a short pile of pubescence. Meri of ambu- 
latory legs at postero-distal end spined. Not dimorphic. .armatus 
B 2 . Carapace naked, areolate, regions well marked. Front trilobate. Cheli- 
peds naked; carpus in female armed, 3 times as long as wide in 
male. Hands similar, margined with spines in female; fingers 
pubescent in gape. Meri of ambulatory legs at postero-distal 
end unspined. Dimorphic tiburonensis 

Key to the West North American Species of Pachycheles 

A 1 . Telson of abdomen with 5 plates. 

B 1 . Front prominent, subtriangular. 
C 1 . Gape of fingers naked. 

D 1 . Chela with setae only. Carpus with a single serrated lobe. 

Carapace pubescent rudis 

D 2 . Chela with pubescence only. Carpus with 5 teeth. Cara- 
pace lightly setose marcortezensis 

C 2 . Gape of fingers with setae and pubescence. Chela with pubes- 
cence only. Carpus with 2 or 3 teeth. Carapace naked 

except rostrum - holosencus 

B 2 . Front not prominent, subarcuate. Gape of fingers naked. Chela 

naked. Carpus unarmed. Carapace naked biocellatus 

A 2 . Telson of abdomen with 7 plates. Carpus with 3 teeth. 

B 1 . Front prominent, subtriangular. Gape of fingers pubescent. Chela 
with pubescence and setae. Carapace naked 

except rostrum pubescens 

B 2 . Front not prominent, subarcuate. 

C 1 . Gape of fingers with setae. Chela with setae only. Carapace 

lightly setose sonorensis 

C 2 . Gape of fingers naked. Chela with pubescence only. Carapace 
naked except rostrum setimanus 

Glassell — West American Decapod Crustacea 445 

Hexapus williamsi, sp. now 

Plate 35, figures 1-4 

Type.— Male, holotype; Cat. No. 1158, San Diego Society of Natural 
History; from San Jose, Guatemala, 10-13 fathoms; April 1, 1937; collected by 
Woodbridge Williams on Captain Fred E. Lewis' yacht "Stranger." 

Diagnosis. — Carapace punctate, regions lightly defined. Front 1/4 the 
width of carapace, trilobed, the median not as advanced as the laterals. Cheli- 
peds unequal; hands with a tubercle on inner, upper distal side of palm to 
engage with oblique stridulating ridge extending from the lateral margins of 
the buccal area. Sixth segment of male abdomen 1/2 longer than wide at distal 
end, 2-3-4-5-6 segments coalesced. 

Description. — Carapace nearly 1/3 wider than long, convex fore and aft, 
transversely flattened, punctate, lateral margins granulated; cervical groove de- 
fines the lateral regions; gastric and cardiac regions smoother than remaining 
surfaces, separated by a light sulcus; a light fold over the intestinal region. The 
postero-lateral is short, subequal in length to the width of the ocular hiatus, with 
two concave margins meeting at the proximal 1/3 in forming a sharp projection. 
The posterior margin is slightly convex, 1/6 wider than the length of the cara- 
pace. The upper ocular margin is raised and granulose. The front is depressed, 
wider at the tip than between the eyes; in a front view the lateral ends project 
farther than the broadly triangular median lobe; the margin between the median 
and the laterals is granulose and concave. The eye-stalks are heavy at the base, 
constricted in the middle, the cornea small in proportion to the base. A narrow, 
oblique row of stridulations on the epistome meets the buccal cavity opposite 
the distal end of the merus of the 3rd maxillipeds. The abdomen of the male is 
slender; only the 1st and 7th segments are articulated; the five interior segments 
are coalesced; the penultimate segment is nearly as wide as long at distal end; 
the ultimate segment is as high as wide, subovate. 

Chelipeds dissimilar, unequal, tomentose; the minor hand is held in a 
normal position, the major hand is carried perpendicular to the minor; major 
hand inflated, the palm nearly as wide as long, upper margin thick and, like the 
minor hand, with a tubercle on the inner distal face which engages with the 
stridulations of the epimera. The outer surface of the hands is granulose under 
a thick pile of tomentum. The lower margin of the major hand is sinuous, of 
the minor straight and beaded. The pollex in both hands is straight and the inner 
edge armed with 3 or 4 large blunt teeth. The dactyli are compressed, thin, 
fluted and armed with 3 large teeth on the major, 2 on the minor; while the 
dactyli slightly cross their pollices at the tip, they do not completely close from 
gape to apices. 

The ambulatory legs are densely margined with tomentum, punctate, the 
2nd the longest. Of the 1st leg the merus is trihedral, twisted, with a row of well 
spaced tubercles on the posterior outer margin; of the 2nd and 3rd (last) legs 
the merus is compressed, 3 times as long as broad, narrowing distally, and sub- 
equal in length to the carpus and propodus combined; the dactyli are as long or 

446 San Diego Society of Natural History 

longer than their propodi, in the 1st pair slightly twisted, in the 2nd and 3rd 
straight, long, tapering, fluted, with the crests tomentose. 

Color in alcohol. — Cream underlaid with light pink on the branchial and 
cardiac regions. Tomentum earthy brown. 

Measurements. — Male, holotype: length of carapace 5.8 mm., width 8.6 
mm., of posterior margin 6.8 mm., of front 2 mm., length of major hand 4.8 
mm., width 2.5 mm., length of 2nd ambulatory leg 10.8 mm., of 6th abdominal 
segment 2 mm., width of base 1.7 mm., width distally 1 mm., height of 7th 
segment 1 mm. 

Range. — Known only from the type-locality (see type) . 

Material examined. — Only the holotype (see type) . 

Habitat. — Taken on a fine black sand and mud bottom in from 10 to 13 
fathoms, the sand mixed with clinkers and volcanic rock. 

Remarks. — This proposed species is closely allied to H. sexpes (Fabricius) , 
1798, but differs from that species: (1) by the regions of the carapace being 
outlined by shallow sulci, instead of being usually not perceptible, (2) by the 
front being partly deflexed, the lateral lobes advanced further outward and 
downward than the median, the width about 1/4 the width of the carapace, 
instead of being vertically deflexed, truncate, and about 1/5 or 1/6 the width 
of the carapace, (3) by the dactyl of the major hand being armed with 3 large 
teeth, instead of with 2 truncate teeth near the base of the inner margin, (4) 
by the merus of the 3rd (last) leg being 3 times as long as broad, instead of 
twice as long as broad, (5) by the propodus of the 3rd leg being about 11/2 
times as long as broad, instead of semi-circular, (6) by the dactyli being as long 
as the propodi, sharp-pointed, fluted, straight, tapered, instead of short and 

This species is named for Mr. Woodbridge Williams, student, of Pomona 
College, Claremont, California, who made a splendid collection of Crustacea 
along the coasts of South and Central America, while on Captain Fred E. Lewis' 
yacht "Stranger," during the Spring of 1937. I am indebted to him for bringing 
to my attention this and many other obscure forms, which link the eastern 
Pacific fauna with those of the western Atlantic and Indo-Pacific regions. 


Subfamily Pinnotherlinae 

Alarconia, gen. nov. 

Carapace much wider than long; integument firm, regions strongly marked; 
front narrow, nearly transverse, with a median groove. Orbit broadly ovate or 
triangular, with a wide inner hiatus, which is partly occupied by the basal 
antennal joint. Antennules transversely or obliquely plicated in wide fossettes 
which communicate with each other beneath the front. Eye-stalks very short. 
Epistome linear-transverse. Ischium of maxillipeds shorter or but slightly less 
in length than merus; merus with distal margin slightly concave; palp jointed to 

Glassell — West American Decapod Crustacea 

Plate 35 

Fig. 1. Hexapus williamsi Glassell, sp. nov. Outer maxilliped. 

Fig. 2. Hexapus williamsi Glassell, sp. nov. Buccal area. 

Fig. 3. Hexapus williamsi Glassell, sp. nov. Dorsal view. 

Fig. 4. Hexapus williamsi Glassell, sp. nov. Ventral view. 

448 San Diego Society of Natural History 

summit of merus; third joint articulated on inner side of the preceding one 
near base. 

Chelipeds of moderate size; merus trigonous; hand large, compressed. Sec- 
ond ambulatory leg larger than the first; third largest of all; fourth the smallest; 
propodus of first leg subcircular, compressed. Abdomen in both sexes usually 
7-jointed and narrower at base than width of last sternal segment. In the male, 
the abdominal appendages protrude from the sternal trench opposite the lateral 
margins of the ultimate segment, bending upward and forward in a semicircle 
toward the buccal opening. 

This proposed genus is allied to the genus Pinnixa White, 1846, by the 
general shape of the carapace and the relative sizes and shapes of the ambulatory 
legs, but differs from that genus in that the ischium and merus of the outer 
maxillipeds are not fused or coalesced, but articulated, and by the ischium being 
much longer in proportion. This proposed genus is also allied to the genera 
Tritodynamia Ortman, 1894, and Asthenognatbus Stimpson, 1858, of the sub- 
family Asthenognathinae. It resembles the former in that the outer maxillipeds 
are somewhat similar, but it differs in that the eyes are not large, the carapace 
not smooth, and the 2nd ambulatory leg is not the largest. It resembles the latter 
in that the ambulatory legs, as shown by Stimpson's figure, are quite similar, but 
it differs in the disposition of the segments of the palp. In addition to the 
above, there is also a relationship to the genus Lambdophallus Alcock, 1900, 
of the subfamily Hexapodinae, as in both genera the abdominal appendages of 
the male are not distally confined under the abdomen. 

Genotype. — Alarconia seaholmi, new species, taken at Acapulco, State of 
Guerrero, Mexico, 6 to 10 fathoms; April 6, 1937; collected by W. J. Seaholm. 

Remarks. — This proposed genus is named for Hernando de Alarcon, navi- 
gator, who, under the direction of the viceroy of New Spain, was sent to support 
by sea the expedition of Francisco Vasquez de Coronado, to the Seven Cities of 
Cibola. During this adventure he discovered and explored the mouth of the 
Colorado River, in the year 1540. 

Alarconia seaholmi, sp. nov. 
Plate 36, figures 1-5 

Type. — Male, holotype, Cat. No. 1159, San Diego Society of Natural 
History; from Acapulco, State of Guerrero, Mexico, 6-10 fathoms; April 6, 
1937; collected by Captain W. J. Seaholm, on Captain Fred E. Lewis' yacht 

Diagnosis. — Carapace broadly ovate, regions well marked, branchial 
regions tuberculate and granulose, a transverse cardiac ridge, opposite the ends 
of which, on the branchial region, is a large tubercle. The 4th ambulatory leg 
extends past the merus of the 3rd by the length of the dactyl. The ischium and 
merus of the outer maxillipeds are long and narrow, the merus the longest. 

Description. — Carapace nearly 2/3 as long as wide, convex, broadly ovate; 
regions well defined with sulci; branchials granulated and tuberculated, tubercles 
more prominent opposite the cardiac region; cardiac region with a transverse, 

Glassell — West American Decapod Crustacea 

Plate 36 












Alar coma seaholmi Glassell, sp. nov. Right chela. 
Alar coma seaholmi Glassell, sp. nov. Ventral surface. 
Alarcoma seaholmi Glassell, sp. nov. Outer maxilliped. 
Alarcoma seaholmi Glassell, sp. nov. Male abdominal appendage. 
Alarcoma seaholmi Glassell, sp. nov. Dorsal view. 

450 San Diego Society of Natural History 

granulated ridge, posterior to the ridge the surface falls sharply to the posterior 
margin; on the branchial regions at each end of this ridge is a large tubercle. 
The anterolateral margin is spinulose toward the lateral angle, granular anterior- 
ly; the postero-lateral margin is sinuous, nearly 1/4 the width of the carapace; 
at the proximal end is an upright tubercle, the last of 3 tubercles which deco- 
rate the distal ends of the convex posterior margin. Front truncate, entire, slightly 
produced, with the outer angles rounded, and with a median groove on the 
upper surface; in width it is 1/7 the width of the carapace. The eye-stalks are 
stout at their bases, tapering sharply to the minute cornea, and with a submedian 
constriction; the width of the base is slightly less than the length. The length of 
the antennae is nearly twice the width of the front. Buccal cavity with parallel 
sides. Maxillipeds standing wide apart, the merus and ischium longer than broad, 
the merus longer than the ischium. 

Chelipeds similar, equal, lightly margined with fine setae; merus trigonate, 
the outer lower margin armed with small tubercles; carpus subrhomboidal 
viewed from above, the distal, triangular end covering the upper articulation of 
the hand; the lower articulation of the hand is considerably posterior to that of 
the upper, and on the inner side of the palm; the manus from the lower proximal 
end to the tip of the pollex is more than twice as long as wide; the upper crested 
margin of the palm is a little more than half as long as the thin, granulated 
lower margin. The hand is compressed; pollex horizontal, thin, with a bifid tip, 
armed on the inner margin with 3 well-spaced teeth, the median bifid, the fingers 
gape from base to apices; the dactyl is armed with a large, median, subtriangular 

Of the ambulatory legs the 3rd pair is the largest and longest, followed 
by the 2nd, 1st and 4th, all are lightly margined with setae. The 1st and 2nd 
pair have their carpal joints crested, more prominently on the 1st pair; the propo- 
dus of the 1st leg is compressed, subcircular, with a double, flattened, upper 
crest; the dactyl is horizontally compressed, lanceolate, twisted and bent upward, 
about as long as the propodus; the dactyl of the 2nd leg is compressed, straight, 
and longer than the upper margin of the propodus; the merus of the 3rd leg 
is 1/2 as wide as long, with a granulated upper margin and a granulated and 
spinous lower margin; the ischium has a strong median spine on its posterior 
margin; carpus nearly 1/3 longer than the propodus; dactyl is slightly shorter 
than the propodus, tapered, straight; of the 4th leg, whose upturned dactyl 
reaches past the distal end of the merus of the 3rd leg, the lower margin of the 
ischium is granulated, with a single subdistal tubercle; merus 3 times as long as 
wide, margins parallel, the lower granulated; the propodus is as wide as the 
merus and subequal in length to the dactyl. 

The abdomen of the male is widest at the junction of the 2nd and 3rd 
segments, the 4th and 5th segments are coalesced, the margins of the 5th and 6th 
segments are parallel, there is a line of light tomentum at the articulation which 
extends across the sternum at this point, the base of the 7th segment is less in 
width than that of the truncate distal end of the 6th, its height is 1/2 its base 
and is semiovate. The abdominal appendages of the male protrude from a sub- 
circular groove opposite the lateral margins of the terminal abdominal segment, 

Glassell — West American Decapod Crustacea 451 

and curve upward and forward toward the buccal area. At their apices they in- 
cline toward each other. 

Color in alcohol. — Light cream. Setae and tomentum red-brown. 

Measurements. — Male holotype: length of carapace 5.8 mm., width 9 mm., 
of posterior margin 6.5 mm., width of outer orbital margins 3 mm., of front 
1.3 mm., length of 1st leg 7.7 mm., of 2nd leg 9.4 mm., of 3rd leg 11.7 mm., 
of 4th leg 6.8 mm., of hand 4.5 mm. 

Range. — Known only from type-locality (see type) . 

Material examined. — A single male specimen lacking the ambulatory legs 
on the left side. 

Habitat. — Dredged on a sand and shell bottom in 6-10 fathoms. Commen- 
sal host, if any, unknown. 

Remarks. — The references in the generic description to the female abdomen 
do not refer to this proposed species, but to a different species found on this 
coast, which will be described at a later date. 

This proposed species is named for Captain W. J. Seaholm, who collected 
this and many other specimens while dredging for shells, for his sustained in- 
terest in the fields of natural science. 

Pinnotheres orcutti Rathbun 

Pinnotheres orcutti Rathbun, Bull. U. S. Nat. Mus., no. 97, 1918, p. 98, pi. 22, 
figs. 5-6, text fig. 50 (type-locality, Manzanillo, Mexico) . 
During the year 1936, Dr. Waldo L. Schmitt, of the U. S. National 
Museum, sent me for identification, among other material, a small series of 
Pinnotheres from the Tres Marias Islands, Mexico, collected by H. N. Lowe 
in March, 1930. These specimens I recognize as being Pinnotheres orcutti 
Rathbun, heretofore known only from the type specimen, a male. The following 
is a description of the female: 

Diagnosis. — Carapace calcareous, suboctagonal, high, antero-lateral mar- 
gins ridged. Front in dorsal view horizontal, bilobed, projecting. Dactyli of 4th 
pair of legs nearly 1/3 longer than the others, and longer than their propodi. 

Description. — Carapace calcareous, high, convex, suboctagonal, slightly 
longer than broad, broadest in posterior half, uneven, branchial regions with 
irregular lobes; dorsal surface pubescent and bordered by a raised rim; cardiac 
region surrounded by a furrow except anteriorly, no median tubercle near its 
posterior end, as in a smaller male. Front with two advanced, blunt-pointed 
lobes, separated by a wide U-shaped notch, behind which runs a broad median 
furrow. Lateral margin long, angled, convex; postero-lateral margin short, 
splayed out over propodite of 3rd leg; posterior margin convex. Basal segment 
of antennae elongate and obliquely placed. 

Merus of outer maxilliped wide and angled; the propodus differs from that 
of the male by being obtuse at its apex, instead of subtriangular, and by having 
the dactyl extending nearly to the extremity of the propodus, instead of only 
part way, as in the male. 

Chelipeds similar, stout, manus short, increasing greatly in width toward 
distal end, where it is slightly less in height than superior length; lower margin 

452 San Diego Society of Natural History 

concave under gape; pollex sharply turned up at apex, armed with a blunt proxi- 
mal lobe and a row of small teeth, the distal the larger, while the median tooth 
is the largest; dactylus wide at base, strongly arched and armed with a wide, 
angular tooth in front of a deep proximal notch for the reception of the proxi- 
mal lobe of the pollex; the tips of the fingers are sharp-pointed and cross each 

The ambulatory legs are narrow; the 2nd leg the longest; the dactyli of the 
first three pairs are subequal in length, slightly pubescent and with spine-like 
tips; the dactyli of the 4th pair are nearly 1/3 longer than the others, nearly 
straight, longer than their propodi, and with a fringe of pubescence on their 
lower margins. 

The abdomen is circular; its terminal segment within the perimeter, its 
posterior margins oblique, its tip with a slight median emargination. 
Color in alcohol. — Buff. Pubescence earthy brown. 

Measurements. — Length of carapace: 8.5 mm., width 8.1 mm. Length of 
dactyli of ambulatory legs: 1st 1.6 mm., 2nd 1.7 mm., 3rd 1.4 mm., 4th 2.4 mm. 
Range. — West coast of Mexico. 

Material examined. — Two females, ovigerous, and one male; from Maria 
Madre Island, Tres Marias Islands, Mexico; March 1930; collected by H. N. 
Lowe. Collection of the U. S. National Museum. 

One female, ovigerous, from Tenacatita Bay, Mexico; April 11, 1937; 5 
fathoms; collected by Woodbridge Williams, on Captain Fred E. Lewis' yacht 

Habitat. — Unknown. The Tenacatita Bay specimen had a calcareous worm 
tube attached to the carapace. 

Remarks. — The use of the outer maxilliped as an infallible method of 
determination would have been rather difficult in the Tenacatita Bay specimen, 
had that one been the only specimen examined, for its outer maxillipeds were 
different from each other, in that the dactyli of the right and left sides were 
of different lengths, that of the right extending considerably past its propodus, 
while that of the left was short of its propodal apex. In addition, there is a 
marked variation in the shape of the propodus of the outer maxillipeds in the 
sexes, that of the males being subtriangular, with its dactyl short, while in 
the female the propodus is obtuse, the dactyl reaching to a point near its tip. 
The shape of the merus in the maxillipeds of both sexes is identical. 

The only difference between the Maria Madre Island male and Rathbun's 
smaller type specimen is that the specimen I examined did not have the shallow 
right-angled indentations on either side of the 6th abdominal segment. 

The eggs are abundant, globular, and slightly less than 1/3 mm. in 

Fabia granti Glassell 
Plate 33, figure 2 

Fabia granti Glassell, Trans. San Diego Soc. Nat. Hist., vol. 7, no. 28, 1933, 
p. 342, pi. 26 (type-locality, Magdalena Bay, Baja California, Mexico). 
While collecting at San Felipe, a small fishing village near the head of the 

Gulf of California, in Baja California, Mexico, I took a large series of this 

Glassell — West American Decapod Crustacea 453 

species which were found in a tide pool, commensal with Crucibulum spinosum 
(Sowerby) . A series of more than 75 females and 15 males was collected at this 
locality on May 9, 1937. A description of the heretofore unknown male follows: 

Description. — Carapace calcareous oviform or urnal, surface flat, depressed, 
with widely separated short hairs; anterior margins raised with pubescence; 
regions not defined; antero-lateral margins strongly converging posteriorly; 
posterior margin straight, deflexed, entire, as wide as the outer ocular width. 
Front broadly arched, entire, with upper median surface depressed and pubescent. 
The cervical groove is shallow, leading back from the upper margins of orbits. 
Eyes small, pigmented. Antennae minute, though long and slender. 

Chelipeds similar, stout, short; merus crested with short pubescence; carpus 
depressed on upper surface, lightly pubescent on margins; hands short and wide, 
proximally inflated on inner side of palm; palm as wide as long, with an upper, 
broadly arched, pubescent carina; lower margin horizontal, an indistinct or 
obsolete, longitudinal, median ridge on outer surface. Pollex short, horizontal, 
except for sharply upturned, spine-like tip, armed on inner edge with an oblique, 
granulose cutting edge. Dactyl strongly curved at tip, and armed on the inner 
edge with a single submedian tooth. The fingers are close fitting, their tips 

Ambulatory legs compressed, with spatulate propodi; the dactyli com- 
pressed, slightly curved, with needle-like, corneous tips. The 2nd and 3rd legs 
with plumose natatory hairs on the carpus and propodus. The meri are mar- 
gined with a close pile of pubescence. 

The sides of the abdomen converge from the 1st and 2nd to the 7th seg- 
ment, the latter being semioval; the 1st and 2nd segments are nearly as wide as 
the sternum at this point. 

Color in alcohol. — Buff. Pubescence dirty yellow. 

Measurements. — Of largest male: length of carapace 3.7 mm., width 3.5 
mm., width of posterior margin 1.6 mm., of front between the inner ocular 
margins 1.2 mm., length of hand 1.8 mm., height of palm 1.3 mm. Of smallest 
breeding specimen: length of carapace 2.3 mm., width 2.1 mm. 

Range. — Throughout the Gulf of California, Mexico. 

Habitat. — Found commensal in Crucibulum, Acmaea and Crepidula. The 
type specimen having been taken in a worm tube does not truly indicate its host, 
but rather that the holotype was disturbed in the dredge material. 

Remarks. — This miniature male is undoubtedly a free swimmer, as the 
natatory hairs on the ambulatory legs would indicate. That it spends its time, 
other than in the breeding season, in a free state may be questioned, as a num- 
ber of the males taken in this series were alone with their host. They may be 

The genus Fabia, which is apparently restricted to American waters, has 
six recognized species at the present time, and of these species little is known 
of the males, a circumstance due to their size. Wells, 1928, described the male 
of Fabia subquadrata Dana, 1851, which, to judge from the text and figures, 
remarkably resembles Pinnotheres concharum (Rathbun), 1893, a species that 
I have frequently found in Vol sella capax (Conrad) , along with the female of 
Fabia lowei Rathbun. 



Volume VIII, No. 34, pp. 455-462, plate 37 







Loye Miller 

University of California at Los Angeles 


Printed for the Society 

May 31, 1938 


U. S. Grant, IV, Chairman 

L. M. Klauber Clinton G. Abbott, Editor 






%$*7 Loye Miller 

University of California at Los Angeles 

Dr. Chester Stock of California Institute of Technology has been 
good enough to place in my hands, for study, a fairly complete skull of 
the Asphalt Stork (Ciconia maltha) from the McKittrick Pleistocene 
asphalt. The rostral portion of the specimen has suffered several frac- 
tures which render the profile of the beak uncertain, but the cranial 
portion warrants discussion. 

Previous knowledge of the species was summed up in a special 
paper by the present writer in 1932 (Condor, vol. 34, Sept., 1932, pp. 
212-216). At that time the skull was known only from a very unimpor- 
tant fragment of the upper mandible and from a fairly well preserved 
lower mandible. 

Study of these and the more abundant trunk and limb remains led 
to the following statement : "This material differs from all living Ameri- 
can storks and is included in the species Ciconia maltha originally de- 
scribed from Rancho La Brea. . . . Generic distinction between Euxen- 
ura and Ciconia is based largely on external features and even these fea- 
tures are considered by some students to exhibit insufficient differences 
to warrant recognition of the separate genus Euxenura. 

"Conceding that the differences between existing forms are of gene- 
ric value, the form under discussion would not agree with either genus 
and a new genus would be necessary. While there is little question that, 
were the asphalt stork restored to us in its entirety, it would likely exhibit 
characters sufficient for its generic distinction, yet for the sake of sim- 
plicity it is referred, in the absence of those superficial characters, to the 
genus Ciconia." 

The last sentence of this quotation indicates a degree of conserva- 
tiveness to which I freely confess and which has been adhered to fairly 
consistently for the several years during which the fossil birds have been 
a major interest. Ciconia is the typical genus of Ciconiformes and since 
the skeletal parts heretofore examined differed from Ciconia in no radical 

458 San Diego Society of Natural History 

fashion, the fossil bird was assigned to that genus. 

Examination of the cranial parts of the stork skulls available brings 
to light a number of differences which can be applied in the study of 
fossil birds, i. e. characters not lost with the entombment of the specimen. 

Whether or not these osteologic characters are more significant than 
those of the exo-skeleton is not pertinent— the fact remains that they 
are more available to the paleontologist. 

Six rather widely separated genera of storks were examined in this 
study, Xenorhynchus from Africa, Leptoptilns from India, Ciconia from 
Europe, Euxenura from South America, and Ajaia and Mycteria from 
Central America being available. The cranial differences appear mainly 
in the occipital and basisphenoidal regions, although the general propor- 
tions of the actual brain capsule appear to be significant. Xenorhynchus, 
Leptoptilus, and Ajaia are so far divergent from the fossil form that 
they may be set aside in the present study. The closest affinities appear 
to lie with Mycteria, Ciconia, and Euxenura. 

Mycteria has a broad, flat skull when viewed in either profile. With 
the sphenoidal rostrum horizontal, the interorbital region is nearly the 
highest point of the profile. The naso-frontal area is almost as high as 
that of the cerebrum. There is no appreciable lift of the skull over the 
cerebral hemispheres. Quite in contrast are the other two genera. The 
highest point in the profile is posterior to the orbits where the skull rises 
in two swellings to accommodate the hemispheres, with a distinct sagittal 
depression between. The naso-frontal area is very low. In all these re- 
spects, the fossil cranium agrees with Ciconia and Euxenura. 

When viewed from the rear with the sphenoidal rostrum horizontal, 
there appear three major differences: (l) the ratio of transverse to ver- 
tical axis; (2) the size of the occipital area of muscle attachment in rela- 
tion to cranium; and (3) the pattern that the intermuscular lines trace 
upon the occipital plane. 

The following table of cranial diameters illustrates point number 
one with a fair degree of satisfaction. 

Table of Measurements of the Brain Case 

Maximum Depth Maximum Width Ratio 

Ciconia maltha 397 mm. 55.5 mm. 72% 

Ciconia alba 34.1mm. 47.2 mm. 72% 

Euxenura maguari 34.5 mm. 51.3 mm. 65% 

Mycteria americana 39.4 mm. 50.3 mm. 78% 

of Occiput 

of Cranium 

41.8 mm. 

55.5 mm. 


35.5 mm. 

47. mm. 


40. 1 mm. 

51.3 mm. 


42. mm. 

50. mm. 

82 % 

Miller — Pleistocene Stork 459 

It is quite impossible to get the exact area of the irregular occipital 
surface of muscle attachment. A single measurement, the maximum 
transverse diameter, is therefore taken as a rough index. This quantity 
is obtained by measuring the extreme distance between the right and left 
extremities of the occipital area as delineated by the lambdoidal crest. 

The following table again shows the fossil stork to be closely allied 
to the typical species of Ciconia. 

Table of Measurements, Comparing the Occipital Area with the 

Cranial Width 

Maximum Width Maximum Width Ratio 

Ciconia maltha 
Ciconia alba 
Euxenura maguari 
Mycteria americana 

Table of Measurements, Comparing the Occipital Width with the 

Occipital Height 

Maximum Width Maximum Height Ratio 

Ciconia maltha 41.8 mm. 23.7 mm. 59 % 

Ciconia alba 35.5 mm. 20.3 mm. 57.4% 

Euxenura maguari 40.1mm. 25.4 mm. 49 % 

Mycteria americana 42. mm. 24.7 mm. 59 % 

With regard to point number three, the pattern traced by the inter- 
muscular lines upon the occipital area is not one that can be expressed in 
numerical terms, but the general form of the area in the Asphalt Stork is 
more closely like that in the typical Ciconia than it is like that of the 
South American Euxenura. 

The rostral fragment that accompanies the cranium shows very lit- 
tle except that the beak was straight, without either the upturn of Jabiru 
or the ibis-like hook of Mycteria. The bony nares are long and poorly 
defined slits, in contrast to the more rounded and well defined nares of 
Mycteria. Both Euxenura and Ciconia have the slit-like type seen in the 
fossil beak. 

The storks are very poorly represented in the collections from 
Rancho La Brea, but they occupy a prominent place in more limited 
collections from McKittrick, owing apparently to a difference in the local 
terrain. The matrix also at McKittrick seems to have been less perfecdy 
preservative, and specimens crumbled more easily in the differential move- 

460 San Diego Society of Natural History 

ment of the matrix. We have therefore a great paucity of the more fragile 
bird skulls. 

This specimen is the only one that I have been able to examine 
that can be assigned to the true storks. The result of this and other 
studies that I have made is to confirm the original assignment to the 
genus Ciconia. 

Miller — Pleistocene Stork 

Plate 37 

Occipital aspect of cranium, (x l/l approx.) 

A. Ciconia alba. 

B. Ciconia maltha. 

C. Euxenura maguari. 

Drawings by Gerhard Bakker. 



Transactions of the San Diego Society of Natural History, Volume VIII. 
Titles of papers and new systematic names are in heavy-faced type. 

A Further Report on Birds from So- 
nora, Mexico, with Descriptions of 
Two New Races, 321-3 36 

A Key to the Rattlesnakes with Sum- 
mary of Characteristics, 185-276 

A New Genus and Species of Pigmy 
Goose from the McKittrick Pleisto- 
cene, 107-114 

A New Hummingbird of the Genus 
Saucerottia from Sonora, Mexico, 


A New Muskrat from Utah, 409-410 

A New Pelecypod Genus of the Family 
Cardiidae, 119-120 

A New Puff-bird from El Salvador, 


A New Race of Brown Towhee from 
the Inyo Region of California, 

A New Sea-Urchin from the "Oligo- 
cene" of Oregon, 367-374 

A New Silky Pocket Mouse from So- 
nora, Mexico, 73-74 

A New Snake of the Genus Sonora 
from Mexico, 363-366 

A New Subspecies of Crotalus con- 
fluentus, the Prairie Rattlesnake, 


A New Subspecies of Pocket Gopher 
from Sonora, Mexico, 1-2 

A Northwestern Race of the Mexican 
Black Hawk, 361-362 

A Study of the Skull of the Pleisto- 
cene Stork, Ciconia maltha, Miller, 

Abbott, Clinton G., 17, 39, 375 

Acanthina angelica, 31 
lugubris, 344 
muricata, 3 1 

spirata, 344, 397 
Acanthochites diegensis, 32 
Accipitcr, 127 

atricapillus striatulus, 126, 127 

cooperii mexicanus, 127 
Acila shumardi, 368 
Acmaea, 344, 45 3 

cassis, 400 

cassis nacelloides, 3 84, 400 

digitalis tcxtilis, 344 

insessa, 344, 400 

instabilis, 344 

limatula, 344 

mesoleuca, 3 2, 105 

mitella, 32 

mitra, 344 

paleacea, 344 

scabra, 3 44 

Acrididae, 130 
Acteocina culcitella, 3 92 

infrequens, 29 

pedroensis, 3 92 
Acteon (Rictaxis) punctocaelatus, 392 

(Acteon) traski, 392 
Actitis macularia, 3 30 
Adams, A., 308 

C. B., 308, 312 
Aechmophorus occidentalis, 324 
Agaronia testacea, 3 
Agassiz, A., 3 69 
Agclaius phocniceus fortis, 143 

phoeniceus nevadensis, 12 5, 143 

phoeniccus sonoriensis, 142, 143 
Agkistrodon, 188 

bilineatus, 190 

mokascn laticinctus, 189 

mokasen mokasen, 189 

piscivorus, 189, 197 
Aimophila, 144 

ruficeps scottii, 144 
Ajaia, 45 8 

ajaja, 327 
Alaba oldroydi, 3 99 
Alabina diomedeac, 31 

tcnuisculpta diegensis, 384, 398 
Alarcon, Hernando de, 448 
Alarconia, 412, 446 

seaholmi, 412, 448, 449 


San Diego Society of Natural History 

Alcid, Recent, 376 

Alcidae, 376 

Alder, 138 

Alectrion californiana, 396 

cerritensis, 396 

cooperi, 3 96 

fossata, 3 96 

insculptus, 396 

perpinguis, 3 96 

tegula, 396 
Alectrionidae, 308 
Aletes squamigerus, 344, 3 99 

squamigerus pennatus, 3 99 

Aligena cerritensis, 385, 388 

Allen, J. A., 329 

Amaral, A. do, 152 

Amiantis callosa, 3 90 

Ammospermophilus harrisii, 3 5 2, 353 
harrisii harrisii, 352, 353 
harrisii kinoensis, 3 5 2, 35 3 
harrisii saxicola, 3 5 2, 353 

Amphipod, 298 

Amphispiza bilineata, 144 

Amphissa, 344 

reticulata, 3 97 
undata, 397 
versicolor, 344, 397 

An Annotated List of Shells Collected 
at Punta Peiiasco, Sonora, Mexico, 
in February, 1934, 27-34 

An Extinct Puffin from the Pliocene of 
San Diego, California, 375-378 

An Upper Pleistocene Fauna from the 
Baldwin Hills, Los Angeles County, 
California, 379-406 

Anabernicula, 1 1 1 

gracilenta, 107, 109, 110, 111, 
112, 113 
Anachis coronata, 30 

fulva, 21 

hilli, 30 

sanfelipensis, 20 

varia, 30 

vexillum, 21,30 
Anatinae, 108, 109, 1 10 
Anderson, F. M., 3 1 6 
Angel, F., 15 5, 178 
Anhinga, 376 
Anomia, 3 87 

lampe, 3 87 

peruviana, 27, 3 87 
Anomuran, 279, 431 

Anserinae, 107, 108, 109 
Anthony, A. W., 323, 334 
Antillophos, 308 
Antiplanes perversa, 393 
Aphelocoma sordida arizonae, 140 
Aplysia, 32 

californicus, 32 
Apolymetis biangulata, 390 
Area alternata, 27 

aviculoides, 1 6 

delgada, 16 

gordita, 1 6 

gradata, 27 

illota, 27 

multicostata, 16, 17 

pacifica, 27 

reeveana, 27 

reinharti, 16, 27 
(Anadara) reinharti, 16 

solida, 27 
Arcidae, 17 
Ardea herodias treganzai, 325, 326 

Arenaria interpres morinella, 331 

melanocephala, 331 
Arenicola, 103 

Argobuccinum oregonensis, 3 98 
Armstrong, F. B., 1 1 
Arnold, Ralph, 62, 386, 388, 391, 392, 

397, 399, 400 
Arrington, O. N., 178 
Arrowweed, 1 3 5 
Artran, A. P., 178 
Asio otus wilsonianus, 134 
Aspen, Quaking, 141 
Astarte branneri, 387 
Asthenognathinae, 448 
Asthenognathus, 448 
Astraea (Pomaulax) undosa, 400 

Astrea inaequalis, 342 

(Pachypoma) inaequalis, 344 

(Pomaulax) undosa, 344 
Astrodapsis, 61 

antiselli, 61, 62, 63, 64 

arnoldi, 63 

arnoldi arnoldi, 63 

arnoldi crassus, 63 

arnoldi depressus, 63 

arnoldi fresnoensis, 63 

arnoldi peltoides, 63 

arnoldi spatiosus, 63 

crassus, 63 

depressus, 63 

Index to Volume VIII 


fresnoensis, 63 
jacalitosensis, 63 
peltoides, 63 

salinasensis, 61, 62, 63, 64 
spatiosus, 63 
tumidus, 63 
whitneyi, 63 
Astur, 127 

Atys, 3 84 

casta, 3 84, 3 92 
Audubon, J. J., 3 52 

Auk, Great, 376 
Lucas, 376 

Auriparus flaviceps acaciarum, 140 
flaviceps ornatus, 13 9, 140 

Avocet, 3 32 


Bachman, J., 3 52 

Bailey, Bernard, 349, 352, 356 

Florence M., 129, 134 
Baker, Fred, 2 5, 3 5-46 
Bakker, Gerhard, 461 
Balanus concavus, 63 

tintinnabulum californicus, 383 

Baldpate, 328 
Bancroft, Griffing, 323 
Bangs, Outram, 324, 327 
Barbatia (Acar) pernoides, 345 
Barbour, T., 178 
Barnacle, 345, 382, 383 

Pink, 383 

Whale, 383 
Barnard, K. H., 413 
Bartsch, Paul, 307, 386, 398, 402, 404 
Bartschella, 402 

Bassariscus, 3 5 8, 35 9 

astutus, 3 57 

astutus arizonensis, 357, 358, 359 

astutus insulicola, 3 59 

astutus octavus, 357, 358, 359 

astutus palmarius, 357, 359 

astutus raptor, 3 59 

astutus saxicola, 35 9 

astutus yumanensis, 3 57, 358, 359 
Batchelder, Dean E., 168 
Bela tidicula, 393 
Benchley, Belle, 178 
Benson, Seth, 3 5 1 

E. E., 181 

Bent, A. C., 327 
Berkeley, Edith, 93 
Berry, S. Stillman, 338 

Betaeus, 412, 416 

ensenadensis, 412, 416, 417, 418, 

harfordi, 419 
longidactylus, 418, 419 

Biggs, T. C., 178 
Bilderback, Norman, 178, 226 
Bischoff, Ferdinand, 143 
Bishop, L. B., 131, 132, 323 
Bittern, Tiger, 327 

Bittium, 344, 398 

(Lirobittium) ornatissimum, 3 98 
(Semibittium) rugatum, 3 98 

Blackbird, Red-winged, 142, 143 

Yellow-headed, 142 
Blossom, P. M., 3 50 
Bogert, Charles M., 178 
Bonaparte, Charles L., 5 
Bornia, 3 89 

cooki, 384, 385, 389 

retifera, 3 88, 3 89 
Borsonella barbarensis, 3 93 

dalli, 393 
Botaurus lentiginosus, 327 
Bothrops, 188, 189 

atrox, 189 
Brachiopod, 3 99 
Brant, 110 
Branta, 1 1 1 

bernicla, 110, 111, 113 

bernicla hrota, 112 

canadensis minima, 110, 112, 113 

minuscula, 111 
Brisaster, 367, 368, 369, 370 

fragilis, 369, 372 

latifrons, 370, 372 

maximus, 368, 369, 372 

townsendi, 367, 368, 369, 370, 372 

Brodkorb, Pierce, 3 26 

Brown, W. W., Jr., 324, 325, 326, 328, 

329, 330, 332, 333 
Bruncr, Stephen G., 128, 136, 139, 143 
Bryozoa, 38 3 
Bryozoan, 382, 436 
Buccinum crassum, 311 
Bucco dysoni, 4 

Bulla, 393 

aspcrsa, 3 92 


San Diego Society of Natural History 

Bulla gouldiana, 29 

punctulata, 385, 392 

Bullus, 3 93 

punctulatus, 392 

Bulpitt, E. L., 178 
Burch, John Q., 3 82 

Tom, 382, 387, 398 
Bursa californica, 398 
Burt, C. E., 178 

W. H., 123, 131 
Bushmaster, 188, 190 
Buteo albonotatus, 127 

Buteogallus anthracinus anthracinus, 361, 

anthracinus bangsi, 362 

anthracinus micronyx, 361, 3 62 

anthracinus subtilis, 362 
Butorides virescens anthonyi, 326 

virescens frazari, 326 

virescens virescens, 326 

Cabanis, Jean, 10 
Cacomistle, Yuma, 3 57 
Cactus, 124, 133, 144, 201, 106 

Giant, 124, 132, 133, 136 
Cadulus, 3 92 

fusiformis, 391 

nitentior, 3 91 

panamensis, 29 
Caecum crebricinctum, 399 

firmatum, 32 

liratocinctum, 32 

magnum, 399 
Calidris canutus rufus, 331 
Callianassa, 41 8 

longimana, 383 
Callinectes bellicosus, 38 3 
Calliostoma, 344 

angelenum, 19 

canaliculatum, 344, 401 

gemmulatum, 19. 401 

gemmuloides, 1 9 

gloriosum, 3 84, 401 

marshalli, 19, 3 2 

palmeri, 32 

splendcns, 401 

supragranosum, 401 

tricolor, 401 

turbinum, 342, 344 
Callipepla squamata pallida, 125, 127, 
128, 146 

Callista callosa, 3 90 

newcombiana, 390 

subdiaphana, 3 90 
Callistochiton, 32 

infortunatus, 32 
Calyptraea conica, 32 

contorta, 384, 399, 400 

fastigiata, 400 

mamillaris, 32 

mammillaris, 399, 400 
Campbell, Berry, 128, 133, 137, 326, 330 
Campephilus leucopterylus, 1 1 

leucorhamphus, 10 

lineatus leucopterylus, 1 1 
Camptostoma imberbe ridgwayi, 137, 139 
Cancellaria bullata, 3 84, 3 8 5, 3 95 

candei, 308 

cassidiformis, 30 

cooperi, 3 95 

crawfordiana, 3 95 

funiculata, 30 

obesa, 30 
Cancer antennarius, 383, 384 

anthonyi, 383 

branneri, 383 

gracilis, 3 83 

productus, 383 
Cantharus fortis, 385, 3 96 
Capella gallinago delicata, 331 
Caprimulgus vociferus arizonae, 134 

vociferus vociferus, 134, 135 
Cardiid, 119 
Cardiidae, 1 1 9 
Cardinal, 143 
Cardita arfinis californica, 28 

subquadrata, 3 88 
Cardium (Papyridea) aspersum, 28 

(Fragum) biangulatum, 28, "89 

corbis, 120 

(Laevicardium) elatum, 28, 389 

(Laevicardium) elenensis, 28 

(Trigonicardia) graniferum, 28 

nuttallii, 1 19, 120 

procerum, 38 5 

(Bingicardium) procerum, 28 

(Laevicardium) procerum, 3 89 

(Laevicardium) quadragenarium, 

(Lasaea) rubrum, 388 

(Laevicardium) substriatum, 389 
Carmody, Eileen, 178 
Carpenter, P. P., 17, 308, 388, 393 
Casmerodius albus egretta, 326 
Cassis abbreviatus, 32 

Index to Volume VIII 


Catoptrophorus semipalmatus inornatus, 

Caudisona Mitchellii, 15 1, 154 

pyrrha, 15 1, 157 
Cavolina occidcntalis, 3 92 

Celemus tricuspida, 3 92 

trispinosa, 3 84, 3 92 
Cemophora, 189 

Ceophloeus crythrops, 1 1 

lineatus, 11,12 

lineatus lcucoptcrylus, 1 1 

lineatus obsoletus, 12 

lineatus petersi, 1 1 

lineatus scapularis, 12 

mesorhynchus, 9, 10 
Cerastoderma, 119, 120 

Cerithidea californica, 344, 398 
mazatlanica, 3 1 

Cerithiopsis, 3 1 

antefilosa, 3 98 
antemunda, 3 84, 3 98 
cosmia, 384, 398 
halia, 384, 398 
oxys, 3 84, 3 98 

Cerithium, 3 1 

incisum, 3 1 
maculosum, 31 
stercus-muscarum, 3 1 

Chace, Fenner A., Jr., 413, 416 
Chama buddiana, 28 

echinata, 28 

pellucida, 388 
Cliaradrius hiaticula semipalmatus, 3 30 

wilsonia beldingi, 3 30 
Charitonetta albeola, 329 
Chemnitzia, 402 
Chen hyperborea, 328 

hyperboreus, 111 

rossii, 110, 112, 113 
Ghendytes lawi, 382 
Chenonetta, 1 10 
Chetopterus variopedatus, 95 
Chilomeniscus, 189 
Chione fluctifraga, 28 

gnidia, 340 

mariae, 28 

purpurascens, 28 

succincta, 28 

undatella, 389 

Chironia, 388 

suborbicularis laperousii, 3 88 

Chiton, 344 

virgulatus, 32 

Chloephaga, 1 10 

Chlorostoma aureotinctum, 400 
gallina, 400 
viridulum ligulatum, 400 

Chorus belcheri, 3 97 

Chrysodomus rectirostris, 3 96 
tabulatus, 396 

Ciconia, 457, 458, 459, 460 
alba, 458, 459, 461 
maltha, 457, 458, 459, 461 

Ciconiformes, 45 7 

Circulus annulatus, 32 
tricirinatus, 32 

Claravis mondetoura inca, 6 

mondetoura mondetoura, 5, 6 
mondetoura ochoterena, 7 
mondetoura pulchra, 6 
mondetoura salvini, 6, 7 
mondetoura umbrina, 6 

Clark, Alex, 382 

Mrs. E. M., 393, 395, 399,403 

Hubert Lyman, 367-374 

W. B., 61, 62, 63, 369 
Clathrodrillia alcestis, 30 

callianira, 30 

halcyonis, 3 93 

halis, 3 

pilsbryi, 23, 3 

rosea, 30 

thestia, 30 
Clavus empyrosia, 3 94 

(Cymatosyrinx) empyrosia, 394 

(Cymatosyrinx) halocydne, 3 94 

(Cymatosyrinx) hemphilli, 394 

(Crassispira) montereyensis, 394 

pallidus, 3 94 

pembertoni, 22,23 

Clementia, 3 90 

subdiaphana, 390 

Clidiophora punctata, 3 87 

Clinocardium, 119, 120 
blandum, 120 
bulowi, 120 

californiense, 120 
ciliatum, 120 
comoxense, 120 
coosense, 120 
decoratum, 120 
fucanum, 120 
meckianum, 120 
nuttallii, 120 
yakatagense, 120 

Clymenella, 102 
Cnemidophorus tessellatus, 168 


San Diego Society of Natural History 

Cobra, 188 

King, 205 
Cochran, Doris M., 178 
Codakia chiquita, 2 8 

distinguenda, 28 

mexicana, 2 8 
Colinus virginianus ridgwayi, 125 

Columbella californiana, 396 
carinata, 3 96 
carinata californiana, 3 96 
carinata hindsi, 396 
fuscata, 30 
gausapata, 396 
major, 30 

(Aesopus) oldroydi, 395 
tuberosa, 3 97 

Colymbus dominicus bangsi, 323, 324 

dominicus brachypterus, 323, 324 
nigricollis californicus, 324 

Compsomyax subdiaphana, 3 90 

Conant, R., 178 

Connelly, Philip M., 3 82, 3 94 

Mrs. Philip M., 382, 394 
Conrad, T. A., 61, 120, 3 90 
Conus californicus, 344, 3 93 

interruptus, 29, 30 

puncticulatus, 29 

regularis, 29 
Cook, Edna T., 382, 386, 389, 390, 392, 
393, 398, 399, 401, 404 

L. H., 178 
Cooke, Jeannette M., 3 5, 42 
Cooperella subdiaphana, 3 90 
Coot, 32 9 

Cope, E. D., 83, 15 1, 152, 154, 174, 177 
Copperhead, 189 

Broad-banded, 189 

Eastern, 189 
Coral, 345 

Gorgonian, 436 

Yellow Gorgonian, 296, 298 
Corbula, 29 

bicarinata, 29 

(Lentidium) luteola, 391 

marmorata, 29 

nasuta, 29 
Cormorant, 376 

Brandt, 325 

Farallon, 325 
Coronado, Francisco Vasquez de, 448 
Coronula regina, 38 3 
Corvus cryptoleucus, 125, 13 9, 146 

Cosmioconcha palmeri, 31 
Cottonwood, 132, 133, 136, 137, 138, 140 
Coues, Elliott, 151, 157 
Cowbird, Bronzed, 143 

Nevada, 143 
Cowles, R. B., 178 
Crab, Hermit, 294, 420 
Crangonidae, 412, 414 
Crassatellites gibbosus, 28 
Crassinella branneri, 385, 387 

varians, 29, 385, 387 
Crassispira arsinoe, 394 

bottae, 29 

nigerrima, 29 

nymphia, 29 

pluto, 29 
Crenella divaricata, 27 
Crepidula, 344, 45 3 

aculeata, 344 

adunca, 344 

arenosa, 32 

dorsata, 3 99 

excavata, 3 99 

lingulata, 344, 399 

navicelloides, 3 99 

nivea, 32, 105, 294, 399 

nummaria, 344, 399 

nummaria glottidiarum, 399 

onyx, 32, 344, 399 
Crocethia alba, 332 
Crocker, Templeton, 307 
Croneis, Carey, 5 5 
Crotalidae, 187, 188, 207 
Crotalinus viridis, 194 

Crotalus, 168, 169, 188, 190, 195, 216, 
220, 222, 223, 225, 226, 227, 230, 
231, 235, 237, 238, 239, 240 

adamanteus, 169, 176, 190, 195, 

197, 200, 205, 243, 252, 259, 

americana,. 195 
atrox, 77, 157, 162, 166, 167, 169, 

174, 176, 190, 194, 195 
barbouri, 276 
basiliscus, 190, 200, 247, 249, 25 1, 

258, 276 
boiquira, 195 
cascavella, 195 
catenatus, 195, 196, 276 
catesbaei, 195 
cerastes, 166, 168, 171, 175, 176, 

177, 192, 201, 204, 210, 230, 

231, 255, 267, 276 
cerberus, 196 

Index to Volume VIII 


cinereous, 190, 194, 195, 197, 198, 
200, 201, 205, 209, 210, 215, 
227, 230, 235, 236, 238, 239, 
248, 252, 260, 274, 275, 276 

collilineatus, 196 

collirhombeatus, 196 

concolor, 196, 276 

confluentus, 76, 77, 78, 79, 82, 83, 
152, 153, 168, 169, 171, 172, 
173, 174, 176, 194, 196 

confluentus abyssus, 76, 77, 83, 86, 

87, 173 

confluentus concolor, 77, 8 3,86 
confluentus confluentus, 76, 77, 78, 
79, 81, 82, 83, 84, 85, 86, 87, 

88, 191, 

confluentus lutosus, 76, 77, 83, 86, 

152, 171, 172, 173, 175, 176, 

confluentus mitchellii, 152, 154, 

confluentus nuntius, 76, 78, 81, 

82, 83, 84, 85, 86, 87, 88 
confluentus oreganus, 76, 77, 81, 

84, 85, 86, 87, 152, 154, 167, 

168, 170, 171, 173, 176, 177 
confluentus pyrrhus, 157 
confluentus stephensi, 152, 153, 

consors, 196 
cumanensis, 196 
cyanurus, 195, 196 
decolor, 196 
dryinas, 196 
durissus, 171, 194, 195, 196, 197, 

204, 205, 210, 215, 222, 227, 

durissus durissus, 190, 197, 200, 

233, 251, 257, 276 
durissus terrificus, 190, 194, 195, 

196, 233, 250, 251, 257, 276 
edwardsii, 194, 195, 196 
elegans, 196 
enyo, 165, 167, 171, 190, 244, 

245, 251, 258, 276 
exalbidus, 196 

exsul, 167, 191, 239, 252, 276 
fasciatus, 196 
goldmani, 157, 196 
gronovii, 196 
hallowelli, 196 
helleri, 196 

horridus, 195, 196, 245, 246, 247 
horridus atricaudatus, 192, 195, 

196, 247, 253, 268, 276 
horridus horridus, 192, 200, 246, 

247, 253, 268, 276 
immaculatus, 196 

intcrmedius, 196 

jimenezii, 1 96 

kcllyi, 196 

kirtlandi, 196 

lecontei, 194, 196 

lepidus, 168, 197, 204, 234 

lepidus klauberi, 192, 2 34, 2 54, 

269, 276 
lepidus lepidus, 192, 2 34, 2 5 4, 269, 

loetlingii, 196 
lucascnsis, 167, 176, 191, 197, 240, 

252, 261, 276 
lucifer, 196 
lugubris, 196 
melanurus, 196 
messasaugus, 196 
mexicana, 196 
minor, 196 
mitcheli, 1 54 
mitchelli, 15 4, 157 
mitchellii, 150, 151, 153, 154, 155, 

156, 157, 162, 163, 165, 166, 

167, 168, 169, 171, 172, 173, 

174, 176, 177, 179, 233 
mitchellii mitchellii, 150, 152, 153, 

154, 155, 164, 166, 167, 168, 

169, 170, 171, 172, 173, 174, 
172, 173, 174, 175, 176, 177, 
178, 181, 183, 191, 196, 198, 
210, 235, 236, 244, 254, 266, 

mitchellii pyrrhus, 150, 151, 152, 
153, 155, 157, 158, 163, 164, 
166, 167, 168, 169, 170, 171, 
172, 173, 174, 175, 176, 177, 
178, 181, 183, 191, 196, 198, 
210, 235, 236, 244, 254, 266, 

mitchellii stephensi, 150, 15 3, 154, 
162, 163, 166, 167, 168, 169, 

170, 171, 172, 173, 174, 175, 
176, 177, 183, 191, 201, 236, 

244, 254, 266, 276 

Crotalus mitchellii, the Speckled Rat- 
tlesnake, 149-184 
mitchillii, 154 
molossus, 165, 167, 169, 176, 196, 

245, 248 

molossus molossus, 190, 200, 249, 

255, 259, 276 
molossus nigrescens, 190, 249, 25 5, 

multimaculata, 196 
mutus, 196 
omiltemanus, 196 
oreganus mitchelli, 154 
orientalis, 196 


San Diego Society of Natural History 




198 : 






200 : 



230, 255. 



Crotalus ornatus, 196 
pallidus, 196 
palmeri, 197 
piscivorus, 197 
polystictus, 192, 196, 
231, 232, 254, 276 
pulverulentus, 83, 174, 
pulvis, 197 
pyrrhus, 157 
rhombifer, 197 
ruber, 167, 168, 191, 
201, 208, 210, 216, 

252, 261, 276 
salvini, 197 
scutulatus, 79, 157, 162. 

171, 191, 196, 197, 

201, 236, 237, 248, 

simus, 197 
sonoraensis, 197 
stejnegeri, 192, 205, 226, 

strepitans, 197 
terrificus, 169 
tesselatus, 197 
tigris, 78, 152, 153, 162. 

174, 175, 176, 177, 

246, 254, 267, 276 
tigris mitchellii, 154, 15 
tigris tigris, 163 
tortugensis, 190, 2 3 8, 

triseriatus, 174, 196, 197 

233, 235, 247 
triseriatus pricei, 192, 

270, 276 

triseriatus triseriatus, 

201, 247, 248, 255, 
unicolor, 190, 197, 232, 

viridis, 194, 195, 196, 

222, 232, 240 
viridis abyssus, 191, 199, 

263, 276 
viridis cerberus, 243 
viridis concolor, 191, 

242, 253, 264, 276 
viridis lutosus, 191, 199, 

253, 264, 276 
viridis nuntius, 191, 199 

263, 276 

viridis oreganus, 191, 
199, 201, 210, 241, 
253, 265, 274, 276 

viridis viridis 191, 199, 205, 209, 
241, 242, 253, 262, 275, 276 



, 201, 






, 270, 


, 245, 




, 242, 




, 242, 


, 242, 






willardi, 192, 205, 231, 255, 271. 
Crucibulum, 45 3 

spinosum, 32, 105, 399, 453 
Cryptoconus carpenterianus, 3 93 

stearnsianus, 393 

tremperianus, 3 93 

tryonianus, 393 
Cryptomya californica, 391 
Cucumber, Sea, 103, 344 
Cumingia lamellosa, 345 
Curlew, Long-billed, 3 30 
Cuspidaria didyma, 28 

dulcis, 28 
Cyathodonta dubiosa, 387 

pedroana, 3 87 
Cyclinella singleyi, 28 
Cycloxanthops novemdentatus, 38 3 
Cygninae, 109, 110 
Cygnus columbianus, 110 
Cylichna alba, 392 

attonso, 3 92 

diegensis, 392 

propinqua, 3 92 
Cymatium adairensis, 31 
Cymatosyrinx aeolia, 3 94 

hemphilli, 394 
Cypraea annettae, 3 1 

sowerbyi, 3 1 

spadicea, 3 98 
Cypraeolina pyriformis, 3 95 
Cyrtonyx montezumae mearnsi, 128 
Cytharella phaethusa, 30 


Dafila acuta tzitzihoa, 328 

Dall, W. H., 24, 26, 308, 310, 311, 313, 

314, 315, 386, 389, 394, 402, 404 
Dardanus arnoldi, 383 
Davis, C. L., 178 
Deer, 206 

Delphinoidea coronadoensis, 3 8 5, 401 
Dendraster, 3 83 
Dendrochiton, 32 

thamnophora, 32 
Dendrocygna bicolor helva, 328 
Dendrocygninae, 108, 109, 110 
Dendroica erithachorides castaneiceps, 67, 


erithachorides rhizaphorae, 67, 68 
erithachorides xanthotera, 67, 68 

Index to Volume VIII 


Dentalium fisheri, 29 

inversum, 29 

neohexagonum, 391 

numerosum, 29, 384, 385, 391 

pseudohexagonum, 391 

semipolitum, 391 

splendidum, 29 
Deppe, Ferdinand, 10, 361 
Derby, Lord, 4 

Description of a Race of Myiarchus 
cinerascens from El Salvador, 115- 


Descriptions of New Mammals from 
Arizona and Sonora, Mexico, 349- 

Deshayes, G. P., 36 
Diadora aha, 3 2 

aspera, 342, 344 

inaequalis, 32 
Dichromanassa rufescens dickeyi, 326 
Dickey, Donald, 123, 328 

Florence V. V., 67, 323, 349 
Diplodonta orbella, 388 

subquadrata, 28 

Diplotaxis, 130, 131 

Dipodomys, 168 

Dissodactylus lockingtoni, 100, 101 

nitidus, 101, 301 

xantusi, 299, 303 

Divaricella eburnea, 28 
Dolphin, 382 
Donax calif ornicus, 3 90 
gouldii, 3 90 
gracilis, 29 
laevigata, 3 90 
navicula, 29 
stultorum, 390 
Dosinia dunkeri, 28 

ponderosa, 28, 31, 340 
Dove, 5 

Drillia hemphilli, 3 94 
inermis, 3 93 
inermis penicillata, 3 93 
Dromidia larraburei, 383, 384 
Dryobates arizonae arizonae, 136 

villosus icastus, 1 3 6 
Dryocopus scapularis, 10 
Duck, 108, 109, 329, 382 
Muscovy, 109 
Ruddy, 329 
Duncan, P. M., 369 
Dunkeria, 402 

Echinoderm, 381, 382, 383 
Echinoid, 101, 301, 345 
Egret, American, 326 

Snowy, 326 
Elaeocyma, 30 

acapulcana, 2 3 

aeolia, 30 

aerope, 2 3 

ianthe, 30 

palmeri, 30 

unimaculata, 30 
Ellis, Arthur L., 338, 339, 340 
Empidonax difficilis difficilis, 137 

Encope, 101, 301 

californica, 101 

grandis, 101 

micropora, 101 
Endopachychilus, 316 

Engina, 3 84 

carbonaria, 396 
strongi, 384, 396 

Ensis, 3 84 

californicus, 3 84, 3 91 

Epitonium, 31, 344 

(Nitidiscala) acrostephanum, 401 

(Asperiscala) bellastriatum, 401 

bialatum, 3 1 

( Asperoscala) canna, 31 

catalinensis, 401 

clarki, 385 

(Asperiscala) clarki, 401 

(Nitidiscala) cooperi, 401 

crenatoides, 3 1 

crenimarginata, 3 1 

fallaciosum, 401 

hindsii, 401 

(Nitidiscala) indianorum, 401 

(Opalia) retiporosum, 401 

sawinae, 3 84 

(Nitidiscala) sawinae, 401 
(Nitidiscala) tinctum, 344, 401 

wroblcwski, 385 

(Opalia) wroblewskyi, 401 

Erato columbella, 398 

vitellina, 384, 398 
Freunetes mauri, 332 
Frismatura jamaicensis rubida, 3 29 

Erolia alpina sakhalina, 3 32 

minutilla, 332 
Erycina, 389 
Etherington, T. J., 316 
Ethusa scxdentata, 436 


San Diego Society of Natural History 

Euceramus, 412, 423 

panatelus, 412, 423, 425, 426, 430 
praelongus, 424, 426, 430 
transversilineatus, 412, 426, 427 

Fulima micans, 401 

Eumeces skiltonianus, 168 

Eupleura muriciformis, 31 
triquetra, 3 1 

Euxenura, 457, 458, 459 

maguari, 458, 459, 461 

Evans, C. L., 178 

Exilia rectirostris, 396 

Exilioidea rectirostris, 385, 396 

Fabia, 412, 453 

granti, 105 294, 299, 412, 435, 

lowei, 45 3 

subquadrata, 453 

unguifalcula, 298, 303 
Fairbanks, H. W., 49, 3 39 
Falco anthracinus, 361 
Fartulum occidentale, 3 84, 3 99 

orcutti, 384, 399 
Faustino, L. A., 316 
Felaniella serricata, 28 
Fer-de-lance, 189 
Ficus decussatus, 32 
Fisher, A. K., 70 
Fissurella volcano, 344,401 

volcano crucifera, 401 
Fitzmorris, Thomas, 178 
Flicker, 130 
Flycatcher, 115, 138 

Arizona Crested, 136, 137 

Ash-throated, 137 

Beardless, 137, 138 

Olivaceous, 137 

Western, 137 
Formicidae, 1 30 
Fossarus fenestrata, 399 
Fowler, H. W., 178 
Frazar, M. A., 12, 323 
Fregata magnificens, 325 

magnificens rothschildi, 32 5 
Florida caerulea caerulescens, 326 
Forreria belcheri, 397 
Fulica americana americana, 329 
Fuller, Bertha M., 382, 387 
Fusinus, 26, 315, 344 

ambustus, 25,26 
arnoldi, 395 
barbarensis, 3 95 
cinereus, 26 

cinereus coronadoensis, 26 
cinereus sonoraensis, 26 
dupetithouarsi, 30 
felipensis, 2 5, 30 
fredbakeri, 24 
hertleini, 2 5 
hertleini albescens, 2 5 
hertleini bruneocincta, 2 5 
kobelti, 395 
kobelti monksae, 344 
luteopictus, 25, 396 
monksae, 395 
porticus, 314, 315 
taylorianus, 26 
traski, 395 
Fusus, 26 

barbarensis, 395 
cinereus, 26 
robustus, 3 95 
rugosus, 3 95 

Gabb, W. M., 388 
Gadinia, 344 

reticulata, 344 
Gaige, Mrs. H. T., 178 
Galatheidae, 296, 298 
Gale, Hoyt Rodney, 388, 340, 341, 377, 

386, 387, 388, 393, 394, 395 
Galeodes patula, 3 
Galerus mammillaris, 399 
Gander, Frank, 375 
Gannet, 382, 383 
Gari edentula, 391 
Gastropod, 338, 342, 344, 345, 384 
Gavia arctica pacifica, 323 
Gelochelidon nilotica vanrossemi, 33 3 
Geophloeus Iineatus, 1 1 
Geothlypis trichas chryseola, 125, 142 

trichas scirpicola, 142 
Gibbula, 24 

americana, 24 
Gidley, J. W., Ill 
Gigantella, 5 1 
Githens, Thomas, 169, 176 
Glans carpenteri, 345, 387 

minuscula, 388 

Glassell, Steve A., 13-14, 91-106, 277-304, 
372, 382, 383, 411-454 

Index to Volume VIII 


Glaucidium brasilianum ridgwayi, 132 
minutissimum, 132 
minutissimum californicum, 131, 

minutissimum cobanense, 132 
minutissimum gnoma, 131, 132 
minutissimum grinnelli, 131, 132 
minutissimum griseiceps, 132 
minutissimum minutissimum, 132 
minutissimum pinicola, 131, 132 
minutissimum swarthi, 132 

Glottidia, 3 99 

Gloyd, H. K., 178, 195 

Glycimeris gigantea, 27 

maculata, 27, 29, 3 2 

multicostata, 27 
Glycymeris barbarensis, 386 

corteziana, 3 86 

septentrionalis, 386 

subobsoleta, 3 86 
Glyphostoma adria, 30 
Glyptoxanthus meandricus, 105 
Goat, 206 
Goby, Blind, 416 
Godman, F. D., 6,115 
Godwit, 330 
Goldfinch, 168 

Goldman, E. A., 2, 157, 3 54, 3 5 6, 409 
Goneplacidae, 96, 412, 445 
Goose, 107, 108, 109, 110, 111 

Brea Pigmy, 1 1 1 

Pigmy, 109, 110, 111 
Gopher, 202, 204 

Comobabi Pocket, 3 54 

Gila Bend Pocket, 3 54 

Growler Valley Pocket, 35 3 

Punta Penascosa Pocket, 1 
Gordon, Isabella, 279 
Gorsuch, David, 12 3, 136 
Goshawk, 126 

American, 126 
Grant, Chapman, 178 

U. S., 15, 20, 40, 63, 307, 338, 
340, 341, 342, 375, 377, 382, 
383, 386, 387, 388, 390, 393, 
394, 395 
Grass, Eel, 95, 344 
Gray, John E., 369 
Greasewood, 124 

Grebe, Least, 324 

Western, 324 
Gregariella denticulata, 27 

Grimthides, 5 1, 5 3, 5 5 

acanthiceps, 52 

conwayensis, 5 3, 54, 5 5, 5 6 
Griffithides conwayensis, a New Name 
for a Trilobite Species from the 
Atoka Formation of Arkansas, 5 3- 

Iongiceps, 5 1,53 

nosoniensis, 5 1, 5 2, 5 6 

olsoni, 54 

ornata, 5 3,55 

ornatus, 54 

parvulus, 5 5 

scitula, 54, 5 5 
Grinnell, Joseph, 70, 126, 129, 163, 178, 

Griscom, Ludlow, 5, 6, 115, 116, 13 2, 

334, 407 
Gryllus assimilis, 130, 131 
Guara alba, 327 
Gull, 376 

California, 333 

Herring, 376 

Laughing, 3 33 

Ring-billed, 332 
Gyraulus, 382 

vermicularis, 404 


Hachisuka, The Marquess, 321-336, 361- 

362, 407-408 
Haliotis, 37 

corrugata, 344 

cracherodii, 344, 401 

rufescens, 344 
Halistylus pupoideus, 400 

subpupoideus, 400 
Hall, E. Raymond, 349 
Hallowell, E., 194 
Halocyptena microsoma, 32? 
Haminea virescens, 2 9 
Haminoea vesicula, 3 93 

virescens, 344 
Hand, Frank, 126 
Handley, A. N., 178 
Hanetia pallida, 30 
Hanna, G. Dallas, 307, 316 

Marcus A., 3 39 
Hannibal, H., 369 
Hannum, Robert, 407, 408 
Harris, Wray, 3 5, 36, 38, 39, 40, 41 
Harter, Samuel George, 35 1 


San Diego Society of Natural History 

Hawk, 127, 206 

Black, 362 

Harris, 127 

Mexican Black, 361 

Western Cooper, 127 

Zone-tailed, 127 
Hedley, C, 3 9 
Heermansonn, A. N., 388 
Heine, F., 10 
Heliacus radiatus, 32 
Helisoma, 3 82 

trivolvis, 404 
Hellmayr, C, 115 
Hemitonia hermosa, 24 
Henne, Christopher, 69 
Hepatus lineatus, 3 83 
Heron, Black-crowned Night, 327 

Great Blue, 32 5 

Green, 326 
Herre, A. W., 178 
Hertlein, Leo G., 26, 307, 3 3S 
Hesperiphona vespertina brooksi, 144 

vespertina californica, 143, 144 

vespertina montana, 143 
Heterocnus cabanisi, 327 
Heterocrypta occidentalis, 383 
Heterodonax bimaculatus, 29 
Heteroscelus incanus, 331 
Hexapodinae, 448 
Hexapus, 412 

sexpes, 446 

williamsi, 412, 445, 447 
Hill, H. R., 178, 382, 383 
Himantopus himantopus mexicanus, 3 32 
Hinds, R. B., 308, 309, 311, 313, 315 

Hinnites giganteus, 386 

multirugosus, 345 

Hipponix antiquatus cranioides, 399 
tumens, 3 99 

Hipponyx, 344 

antiquatus, 344 
barbatus, 32 
serratus, 3 2 
tumens, 344 

Hoard, Robert, 178, 3 63 
Hoffmann, Ralph, 9, 10 
Holman, John T., 3 68 
Homalopoma, 24 

carpenteri, 34 5, 400 

concepcionensis, 24 

Homoriscus, 412, 414 

macginitiei, 412, 414, 415 

portoricensis, 416 
Howard, Hildegarde, 382, 383 
Huey, Laurence M., 1-2, 73-74, 329, 349- 

360, 409-410 
Huffman, Earl C, 103 
Hummingbird, 407, 408 

Rivoli, 131 

Rufous, 135 
Humphrey, R. R., 178 
LI) ;.lina (Hyalina) californica, 345 

(Cystiscus) jewettii, 345 

(Cypraeolina) pyriformis, 395 
Hydranassa tricolor ruficollis, 327 
Hydroprogne caspia imperator, 3 33 
Hylocichla guttata polionota, 141, 142 


Ibis, White, 327 

Wood, 3 27 
Irus lamellifcr, 345 
Ischnochiton, 345 

acrior, 32 

clathratus, 32 

(Stenoplax) limaciformis, 32 

sanctaemonicae, 3 8 5, 404 
Iselica fenestrata, 3 99 
Isopod, 298 
Ividella, 22 

Jabiru, 45 9 
Jackley, A. M., 20 5 
Jackson, Dudley, 215 
Jay, Arizona, 140 


Keen, A. Myra, 119-120 
Kelletia (Kelletia) kelletii, 396 
Kellia, 3 88 

suborbicularis laperousii, 345, 388 

Kelp, 344, 400 
Kennicott, R., 152 
Kew, W. S. W., 61, 62, 63 
Kimball, H. H., 126 
Kirk, William A., 431 

Klauber, Laurence M., 75-90, 149-184, 
185-276, 363-366 
P. M., 178 
Kobler, L. C, 178, 226 
Kuns, G. W., 178 

Index to Volume VIII 


Lachesis, 188 

muta, 190 
Lacuna, 345 

unifasciata, 345, 399 
Laevicardium, 120 
Lamb, Chester, 178, 181, 323, 324, 325, 

326, 327, 328, 329, 330, 332, 333 
Lambdophallus, 448 
Lambert, J., 370 
Lamellaria diegensis, 32 
Lampropeltis, 189 
Lark, Horned, 139 

Scorched Horned, 139 
Larus atricilla, 333 

californicus, 333 

delawarensis, 3 32 

Philadelphia, 333 
Latirus lugubris, 30 
Lazaria subquadrata, 3 87 
LeConte, John L., 194 
Leda impar, 27 

leviradius, 27 

(Adrana) penascoensis, 18, 27 

taphria, 3 86 
Lee, Charles H., 338, 339, 340 
Lepidopa, 423 
Lepidopleurus ambustus, 404 

heathi, 404 

nexus, 3 84, 404 
Lepidoptera, 130 
Leptodeira, 189 
Leptopecten latiauratus, 345 
Leptoptilus, 45 8 
Leptothyra carpenteri, 400 

concepcionensus, 3 2 
Leslie, Elizabeth, 178 
Lettuce, 344 
Leucophoyx thula brewsteri, 326, 327 

thula thula, 327 
Leucosidae, 95 
Lewis, Fred O., 178, 287, 288, 289, 413, 

431, 445, 446, 448, 452 
Lichenopora radiata, 3 83 
Lichtenstein, H., 10, 361 
Lima dehiscens, 387 

orbignyi, 27 

pacinca, 27 
Limnodromus griseus scolopaceus, 3 3 1 
Limosa fedoa, 3 30 

Lindsay, George, 363 
Linsdale, J., 178 
Liotia, 24 

carinata, 32 
Lithophaga, 17 

abbotti, 17 

aristata, 17 

attenuata, 17, 98 

plumula, 387 
Lithophagus aristata, 27 

attenuata, 27 
Littorina scutulata, 345, 399 
Lizard, 168, 189, 204 
Lloyd, \V., 126 
Lobipes lobatus, 3 32 
Lockington, W. N., 101, 104, 279, 426, 

430, 438, 440 
Loon, 375 

Pacific, 323 
Lophopanopcus bellus, 105 

diegensis, 3 83 

frontalis, 383 
Lophortyx gambelii gambelii, 128 

Lora, 395 

fidicula, 385, 393 

viridula, 393 
Lovenia cardiformis, 3 83 
Loveridge, A., 178 
Lowe, Herbert N., 15-34, 305-320, 392, 

451, 452 
Loxorhynchus grandis, 383 
Lucina californica, 345 

(Myrtea) californica, 388 

(Here) excavata, 388 

(Myrtea) nuttallii, 388 

tenuisculpta, 388 

( Myrtea) tenuisculpta approxi- 
mata, 388 
Luidia Columbia, 431 
Lunda, 376 

cirrhata, 376 
I.unz, G. Robert, Jr., 413, 42 6 
Lyonsia, 28 

inflata, 28 


MacGinitic, George E., 41 3, 414, 416, 41 8, 

Macoma indentata, 28, 390 

indentata tenuirostris, 390 

nasuta, 390 

(Cymatoica) occidentalis, 28 


San Diego Society of Natural History 

Macoma panamcnsis, 28 

secta, 3 90 

(Cymatoica) undulata, 2 8 

yoldiformis, 390 
Macrocallista squalida, 2 8 
Macron lividus, 345 
Mactra californica, 29 

(Mactra) californica, 391 

(Spisula) catilliformis, 391 

dolabriformis, 29 

exoleta, 391 

falcata, 391 

(Spisula) hemphilli, 391 

pallida, 385 

(Mulinia) pallida modesta, 391 

(Spisula) planulata, 391 
Majidae, 105, 434 
Mamba, 188 
Mancalla californiensis, 375, 376 

Mangelia angulata, 394 

antipyrgus, 3 

arteaga roperi, 30 

(Bela) arteaga roperi, 395 

barbarensis, 394 

beta, 3 94 

cetolaca, 385, 395 

(Bela) cetolaca, 395 

cymatias, 30 

(Mitromorpha) filosa, 34 5 

(Mitromorpha) gracilior, 345 

hecetae, 394, 395 

(Mangelia) hexagona, 394 

(Mangelia) merita, 3 94 

oenoa, 394 

perattenuata, 395 

pulchrior, 394 

(Bela) variegata, 394, 395 
Mangilia, 313 

branneri, 394 

dejanira, 312, 313 

oldroydi, 395 

sculpturata, 395 

Mangrove, 68, 100 
Manzanita, 1 3 5 
Marcia subdiaphana, 3 90 
Mareca americana, 328 

Margarites (Lirularia) optabilis, 401 
optabilis acuticostatus, 401 
optabilis knechti, 401 
optabilis nodosa, 401 

Marginella californica, 30 

Marl, 339, 346 

Pleistocene, 346 

Marsh, J. C, 382, 398, 399 
Marshall, W. H., 19 

William B., 63 
Martin, Purple, 140 
Martyn, Thomas, 120 
Massasauga, 195 

Eastern, 193, 273 

Western, 193, 194, 195,273 
Mawe, John, 3 90 
Meadowlark, 142 
Mearns, E. A., 123, 124, 125, 126, 352, 

Meek, F. B., 54 
Megasurcula carpenteriana, 393 

remondii, 3 93 
Meise, Wilhelm, 1 1 
Melampus, 3 82 

olivaceous, 3 93 
Melanella, 402 

mexicana, 3 1 

micans, 401 

oldroydi, 402 

rutila, 31, 345, 402 
Melanellid, 3 82 
Melanitta perspicillata, 329 
Melina (Pedalion) anomioides, 27 

(Pedalion) chemnitziana, 27 

(Pedalion) janus, 27 
Meling, Ada, 178 
Melipotis, 130 

indomita, 1 30 

Mellita, 301 

longifissa, 101 
Merganser, American, 129 
Mergus merganser americanus, 329 

serrator, 329 
Merovia pyriformis, 395 
Merriam, C. H., 352 
Mesorhoea idae, 3 83 

Mesquite, 124, 133, 134, 135, 137, 138, 

143, 144 

Metis alta, 3 90 

excavata, 28 

Micranellum crebricinctum, 399 
pedroense, 399 

Micropallas whitneyi whitneyi, 132 

Micruroides euryxanthus, 189 

Micrurus, 189 

fulvius barbouri, 189 
fulvius fulvius, 1 88 

Index to Volume VIII 


Miller, Loyc Holmes, 107, 111, 126, 128, 
129, 134, 138, 142, 375-378, 382, 
Milne Edwards, A., 104 
Minyocerus, 296, 412, 430, 434 

angustus, 43 1 

kirki, 412, 429, 430, 431 
Mistletoe, 133, 138 
Mitchell, S. Weir, 1 5 1 
Mithrax, 434 
Mitra attenuata, 30 

catalinac, 395 

dolorosa, 30 

fultoni, 38 5, 3 95 

idae, 342, 345, 395 

maura, 395 
Mitrella carinata, 3 96, 3 97 

carinata gausapata, 345, 396, 397 

diminuta, 30 

granti, 20 

ocellata guttata, 30 

tuberosa, 3 97 
Moccasin, 188, 189, 190 

Water, 189 
Modiolus braziliensis, 27 

capax, 27, 100. 387 

flabellatus, 3 87 

guyanensis, 27 

modiolus, 387 
Molothrus ater artemisiae, 143 
Moniliopsis cancellata, 3 94 

incisa, 393, 394 

incisa fancherae, 393, 394 

incisa ophioderma, 393, 394 

rhines, 394 
Monoceros engonatum, 3 97 
Montacuta, 389 

Moore, Robert T., 12, 123, 139, 144 
Mopalia acuta, 404 
Moriera, Carlos, 431 
Moris, 3 82 

reyana, 383 
Mormula, 402, 404 
Mortensen, T., 369 
Mouse, 168, 202, 204 

Kino Silky Pocket, 73 

Pima Silky Pocket, 35 5 

Porto Libertad Rock Pocket, 355 
Muller, Fritz, 43 1 

S. W., 49, 51, 52 
Murex carpenteri, 397 

elenensis, 3 1 

festivus, 397 

gemma, 3 97 

leeanus, 3 97 

petri, 3 97 

plicatus, 3 1 

santarosana, 3 97 

senticosus, 307 

squamulatus, 20 
Muricopsis erynaceoidcs, 31 
Muskrat, 409 

Virgin Valley, 409 
Mussel, 100 

Mya suborbicularis, 3 88 
MvctL-ria, 45 8, 45 9 

americana, 327, 458, 459 
Myiarclius cinerascens, 116 

cinerascens cinerascens, 115, 116 

cinerascens flavidior, 116, 117 

crinitus, 1 1 6 

inquietus, 115, 116 

nuttingi, 115, 116 

tuberculifer olivascens, 1 37 

tyrannulus magistcr, 136 
Mytilus adamsianus, 27, 384 

(Mytilus) adamsianus, 387 

(Mytilus) californianus, 345, 387 

multiformis, 27 


Nassa, 308 

angulicostis, 3 

californiana, 3 96 

cerritensis, 3 96 

chelialisensis, 3 1 6 

fossata, 3 96 

insculpta, 396 

iodes, 30 

leucops, 30 

mendica cooperi, 3 96 

pagoda, 30 

pallida, 315 

perpinguis, 396 

tegula, 396 

tiarula, 30 

versicolor, 30 

versicolor striatula, 30 
Nassarius, 3 16 

californianus, 396 

(Schizopyga) californianus, 3 96 

cerritensis, 38 5 

(Schizopyga) cerritensis, 396 

(Schizopyga) fossatus, 3 96 

(Schizopyga) insculptus, 396 

mendicus cooperi, 3 96 

(Schizopyga) mendicus cooperi, 
345, 396 

perpinguis, 3 1 6 


San Diego Society of Natural History 

Nassarius (Schizopyga) perpinguis, 396 
(Schizopyga) perpinguis, 396 
(Zeuxis) tegula, 396 

Natica marochiensis, 32 

Natrix, 189 

Nelson, E. W., 2, 131, 157 

Neosimnia catalinensis, 398 

Neotoma lepida, 349, 359, 350 

lepida aureotunicata, 349, 3 50 

lepida auripila, 349, 3 5 0, 351 

lepida bensoni, 3 50 

lepida devia, 3 51 

lepida flava, 349, 3 5 0, 35 1 

lepida harteri, 3 5 1, 3 52 

Neptunea tabulata, 385 

(Sulcosipho) tabulata, 396 
Nerita bernhardi, 32 

scabricosta, 3 2 
Neritina picta, 32 
Nettion crecca carolinense, 32 8 

New and Obscure Decapod Crustacea 
from the West American Coasts, 


New Marine Mollusca from West 
Mexico, T