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UNIVERSITY OF CALIFORNIA PUBLICATIONS 


ZOOLOGY 


WILLIAM EMERSON RITTER 
AND 


CHARLES ATWOOD KOFOID 


. EDITORS 


VOLUME XIil 


WITH 22 PLATES 


UNIVERSITY OF CALIFORNIA PRESS 
BERKELEY 
1913-1916 


SoG. FS 
eee ©. 5556 


we 


[92] 


CONTENTS 


. A Study of a Collection of Geese of the Branta canadensis Group 


from the San Joaquin Valley, California, by Harry 8. Swarth; 
aya tilt Len C OS tot 2 gemencesecee pecans statue soe astra: sec eeerec ts saree geaeecte a2 a0 snaccee memes eres 


. Nocturnal Wanderings of the California Pocket Gopher, by Harold 


C. Bryant .... 


. The Reptiles of the San Jacinto Area of Southern California, by 


SST ALM OP ETS ACSA Gi ces csc eroe taee oh eas ee eres esteeese esse cu eee saee Sonn Sa 


. An Account of the Mammals and Birds of the Lower Colorado Val- 


ley with Especial Reference to the Distributional Problems 
Presented, by Joseph Grinnell; with plates 3-13 -..................... 


. Aplodontia chryseola, a New Mountain Beaver from the Trinity 


Region of Northern California, by Louise Kellogg -..................-... 


. A Previously Undescribed Aplodontia from the Middle North Coast 


Of Galliiommitasbiyy Weller eles l abyil ON eae emecnee keene eem cee eecreeerceesea see 


. A Second Species of the Mammahan Genus Microdipodops from 


Calitorniary by vJiosepiu Grinnell) eas seca wee scence cee 


. Distribution of River Otters in California, with Description of a 


New Subspecies, by Joseph Grinnell; with plate 14 ........0........ 


. Four New Pocket Gophers from California, by Joseph Grinnell —_.. 
. Three New Races of Vespertilionid Bats from California, by Hilda 


Wood! (Grinnell) 7-2. Beso cesar aera cease eee oe ener ay cae 


. Eutamias sonomae, a New Chipmunk from the Inner Northern 


Coast Belt of California, by Joseph Grinnell — 


. Batrachoseps major and Bufo cognatus californicus, New Amphibia 


from Southern California, by Charles Lewis Camp ~.-..................- 


3. Report upon Mammals and Birds found in Portions of Trinity, Sis- 


kiyou and Shasta Counties, California, by Louise Kellogg; 
Ni /AlLe Oyo) EY eof 1S Ut eae ne er ee 


. An Analysis of the Vertebrate Fauna of the Trinity Region of 


Northern California, by Joseph Grinnell .............20.2002-22-2------- 


. The Status of the Beavers of Western North America, with a 


Consideration of the Factors in their Speciation, by Walter 
SS MTR OT; ae soe ers sass cence sasenwoacees satvac note seseatescestenoacnccsencessuesscicnepnaveszccauteeer 


. Two New Aplodontias from Western North America, by Walter 


TE TAREE AO 8 ec cee oO EPP Er e 


. Notes on the Local Distribution and Habits of the Amphibians and 


Reptiles of Southeastern California in the Vicinity of the Turtle 
Mountains, by Charles Lewis Camp; with plates 19-22 _.. 


SUTIN be oh Asa er a re Be ee ee ee a 
LDASERS ADD so Se A oa een Se ee a ce ope Ee Re IS ae See eet en Bees 


PAGES 


1-24 
25-29 


31-50 


51-294 


297-500 
301-304 


305-310 
311-316 


317-320 
321-325 


327-334 


335-398 


401-412 


413-495 
497-501 
5038-544 


545 
558 


UNIVERSITY OF. CALIFORNIA PUBLICATIONS 
IN 


ZOOLOGY 
Vol. 12; No. 1, pp. 1-24, pls. 1-2, 8 text figs. November 20, 1913 


A STUDY OF A COLLECTION OF GEESE OF 
THE BRANTA CANADENSIS GROUP FROM 
THE SAN JOAQUIN VALLEY, 
CALIFORNIA 


BY 
HARRY S. SWARTH 


UNIVERSITY OF CALIFORNIA PRESS 
BERKELEY 


UNIVERSITY OF CALIFORNIA PUBLICATIONS 


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Significance of Such Investigations, by William E. Ritter; Part II, 
Practicability of the Determinations, by Samuel E. Bailey. Pp. 1-10. 


October 1908) 75 oe See oe a ae Ne creat ape Re 10 
2. (SXIV) The Leptomedusae of the San Diego Region, by Harry Beal 

Torrey. Pp. 11-31, with 11 text-figures, February, 1909° «2.002500... 20 
3. (XXV)-~-The Ophiurans of the San Diego Region, by J. F. 


don. Pp, 33-64, plates.1-6.. July, 1909° = 2c «30 
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by Wm, E. Ritter. Pp. 65-114, plates. 7-14. November, 1909. 2.2)... 0 


5, (XXVII) Three Species of Certanthus from Southern California, by 
H. B. Torrey and F. L. Kleeberger.. Pp. 115-125, 4 text-figures, 


December, TINGE ss ak a Ae gh aS Ee Tse ete Re 10 


6. The Life History of Trypanosoma dimorphon Dutton & Todd, by 
Edward Hindle. Pp. 127-144, plates 15- 17, 1-text-figure. December, “ 
YO a See SO eR ie a ae An a een ie GUA ae ee ain - 

(XXVIII) A Quantitative Study of the Developnient of the Salpa 
Chain in Salpa fusiformis-runcinata, by. Myrtle Elizabeth Johnson. 


4 


Bd 0 pel S50 9d OAR Bi dt ines Bb ecm ae cee tap pd SST Mien ee Er .35- 


8. A Revision of the Genus Ceratocorys, Based on. Skeletal Morphology, 
by Charles Atwood Kofoid.- Pp. 177-187. May, 1910 ..200.c.a le 10 
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dranth and Holdfast, by Harry Beal Horrey. Pp. 205-221; 16 text- 
figures. 
11, (XXXI) Note on Geotrovism in Corymorpha; by Harry Beal aon 4 
Pp. 223-224;.1 text-figure. 


Nos 10 and 11 in one cover. August, RORY AS SOS Per dee craheslres 20 

12. The Cyclostomatous Bryozoa. of the West Coast of North Anierica, by 
Alice Robertson... Pp. 225-284; plates 18-25. December 1910 ......... 60 

13. Significance of White Markings in Birds of the Order Passeriformes, 
by Henry Chester-Tracy. Pp. 285-312. December, -1910° wee 
14, (XXXIII) Third Report on the Copepoda of the San Diego Region, by = 
Calvin Olin Esterly.° Pp. 313-352; plates 26-32: February, 1911... 40 


15..The Genus Gyrocotyle, and Its Significance for Problems of Cestode 
Structure and Phylogeny, by Edna Harl Watson. Pp. 353-468; plates 
BS-48.— DuMe, VOD on ease eac ach cee engceten canine neee [ees Fe 1.00 
Index, pp. 469-478. 


*Roman numbers indicate sequence of the Contributions from the pateraiaey. of the - 


Marine Binlenitas Association of San Diego: 


sien ic led etn eat dS 


UNIVERSITY OF CALIFORNIA PUBLICATIONS 
IN 


ZOOLOGY 
Vol. 12, No. 1, pp. 1-24, pls. 1-2, 8 text figs. November 20, 1913 


A STUDY OF A COLLECTION OF GEESE OF THE 
BRANTA CANADENSIS GROUP FROM 
THE SAN JOAQUIN VALLEY, 
CALIFORNIA 


BY 
HARRY 8. SWARTH 


(Contribution from the Museum of Vertebrate Zoology of the University of California) 


The status of the geese of the Branta canadensis group occurring 
in California has been heretofore in an uncertain and unsatisfactory 
condition, owing largely to a lack of specimens in collections repre- 
senting the different subspecies found within the state. During the 
winter of 1911-1912 the Museum of Vertebrate Zoology obtained a 
large series of these geese from the San Joaquin Valley, California. 
This was part of a collection of water birds from the region collected 
and prepared by Mr. R. H. Beck, and acquired through the benefac- 
tion of Miss Annie M. Alexander. This material, together with skins 
already in the Museum collection, promised to go far towards solving 
the problems involved, but it required only a cursory examination of the 
series to show that nothing could be settled without additional specimens 
representing typical B. canadensis canadensis and B. c. occidentalis. 
The published descriptions of the last named were so much at variance 
with specimens at hand which were supposed to be of the race, that 
it was necessary to examine the type in order to have a basis for any 
satisfactory conclusions. Through the courtesy of Dr. C. W. Rich- 
mond, Acting Curator, Division of Birds, of the United States National 
Museum, these wants were supplied by loan from the collection of 
that institution. 

In the following pages the four forms of Branta canadensis are 
discussed on the basis of the accumulated material, one hundred and 


2 University of California Publications in Zoology [ Vou. 12 


fifty-three skins altogether. The numbers used refer, unless otherwise 
stated, to the collection of birds in the University of California Museum 
of Vertebrate Zoology. 


Branta canadensis canadensis 


Of specimens referable to canadensis proper, there are in the 
Museum collection thirty-five winter birds from Los Banos, California 
(nos. 21944-21960, 21962-21979), and one breeding bird from Lake 
Tahoe (no. 17224), and there were available five additional skins, one 
from Los Angeles County (no. 588, Grinnell coll.), and four loaned 
by the United States National Museum, from Rhode Island (no. 
175197), North Carolina (no. 169168), District of Columbia (no. 
70996), and Ontario (no. 223137), respectively. 

The specimens composing our series of canadensis are more uniform 
in size, color, and markings than those of any other of the groups into 
which the species has been separated. The conspicuous features of 
this form are large size and pale coloration, especially of underparts, 
associated with white cheek patches continuous across throat, and 
absence of white half collar at base of black neck. The tarsus is 
usually shorter than combined length of middle toe and claw. 

In the series indicated above there is but slight variation in shade 
of gray of underparts, the winter birds from the San Joaquin Valley, 
California, being of precisely the same hue as winter birds from the 
East. As regards color patterns, the following unusually marked 
birds are the only variants in the series. Three specimens (nos. 21962, 
21949, 21952) show slight traces of a white half collar at base of neck, 
and two (nos. 21962, 21972) have a few small black spots on center 
line of throat, the white cheek patches being otherwise continuous with 
one another across throat, but in no instance are these differences 
sufficiently marked to cause hesitaney in placing the birds in the 
canadensis category. 

Of the thirty-nine specimens in which I was able to count the 
number of tail feathers, one had fourteen rectrices, nine had sixteen, 
twenty-three had eighteen and six had twenty. 


Branta canadensis hutchinsi 
The hutchinsi series at hand forms a perfect connecting link between 
B. c. canadensis and B. c. minima, the gradation between hutchinsi 
and minima in particular being so gradual that several specimens 
might with equal propriety be placed in either subspecies. 


1913 | Swarth: Geese of California 3 


Of the thirty-six specimens considered as hutchinsi (nos. 6497, 
17241, 21961, 21980-21983, 21985-22013), twenty-five are males, eight 
are females, and of three the sex was not ascertained. Six have a 
white half collar at base of neck and a black line on throat, three have 
white half collar with no black throat line, twelve have black throat 
line with no white half collar, and thirteen have neither white collar 
nor black throat line. Seventeen specimens have sixteen rectrices each, 
while seven have eighteen. Tarsus and middle toe with claw are more 
nearly the same size than in the other forms of the canadensis group. 
Minimum and maximum measurements are: tarsus, 68 and 86 mm., 
middle toe with claw 67 and 85. Usually the tarsus is a trifle the 
longer. 

The descriptions of hutchinsi, as given in literature (e.g., Coues, 
1903, p. 905; Ridgway, 1887, p. 117), characterize the form as of 
smaller size than but otherwise similar to canadensis, giving no hint 
of its intergradation with minima. Our series, as stated above, forms 
an unbroken chain between the two forms. The series averages paler 
than minima, many specimens being quite as light-colored as canaden- 
sis, but there are few birds in the lot which combine the pale color, 
and head and neck markings of canadensis with the smaller size of 
typical hutchinsi, as should be the case. 

A possible explanation of the intermediate character of so many 
of the birds wintering in California is that the breeding ground of 
these variants lies in the region between the habitats of typical minima 
and hutchinsi, and that there really is geographical continuity of range 
in the summer home of the two forms, correlated with gradual blending 
of characters in the birds occupying the middle ground. Then, if the 
individuals wintering in California came from the more western part 
of the summer home of hutchinsi as well as from all parts of the range 
of minima, we should find here, as is actually the case, vast num- 
bers of typical minima, a lesser number of intergrades, and compar- 
atively few typical hutchinsi. On this supposition we should expect 
to find at points farther east but few intergrades and the majority of 
birds typical of hutchinsi. 

The latest authority on the distribution of North American ducks 
and geese (Cooke, 1906, pp. 77, 78) describes the subspecies minima 
and hutchinsi as occupying in part the same region during the breed- 
ing season, western Alaska and parts of the Aleutian Islands, a con- 
dition that is certainly unusual unless the two are distinct species. 


4 University of California Publications in Zoology [ Vou. 12 


Such is evidently not the.case, as shown by the large number of speci- 
mens combining the characters of the two forms. 

It is possible that the asserted occurrence of occidentalis in Cali- 
fornia is based in part upon the capture of birds of such characters 
as those just mentioned, for in some particulars they would answer 
fairly well to some published descriptions of that race. Comparison 
of California birds with these intermediate characters with specimens 
of occidentalis from the northwest coast region, however, shows that 
the affinities of the former le in other directions, and my belief, as 
indicated above, is that they are intergrades between hutchinsi and 
minima. 


Branta canadensis minima 


There seems to be a greater range of variation in color and mark- 
ings in this than in any of the other forms. Sixty-seven specimens 
were examined (nos. 19456, 19457, 21984, 22014-22077). These show 
the following combinations of patterns: a well-defined white mark 
(collar or half collar) at base of neck, and white cheeks separated by 
black bar on the throat, 32 specimens; with black throat bar but with 
no white collar, 21; with white collar but with no black on throat, 6; 
with neither white collar nor black throat bar, 7. These divisions 
have been made rather arbitrarily, for taking any one of the char- 
acters separately, every conceivable gradation may be found (see 
plate 1). The white collar may be merely suggested by but a few 
white feathers on the lower neck, or it may be a solid band of white 
encircling the neck, an inch or more in width. In one ease the white 
has invaded forward so that nearly all the feathers of the fore neck 
are tipped with this color. : 

The throat may be as purely and extensively white as in canadensis, 
there may be a few flecks of black along the median line, or there may 
be a solid black bar. In one or two instances this bar is of such width 
that the white cheek markings are reduced to small, oblong patches 
(in no. 22026 they measure 44 by 15 mm.). These may be thickly 
speckled with black. 

The color of underparts ranges from a uniform dark cinnamon or 
tawny olive to as light a gray as in any specimen of typical canadensis. 
The rectrices are usually sixteen in number; of fifty-one birds in which 
they could be counted, there were forty-six with sixteen tail feathers, 
three with fourteen, and two with eighteen. 


1913 | Swarth: Geese of California 5 

The black neck usually shades gradually into the brown of the 
breast, but in some, the gray-colored birds, it is about as sharply 
defined as in canadensis. In several instances there is a well-defined 
black area on the upper breast, just below the white collar. 

As each of the above characters varies independently of the others 
there is very little uniformity of appearance in a series of these birds, 
and, except in very general terms, it is not possible to designate for 
the subspecies any character of color or pattern. 

The accompanying plates illustrate some of this variation. One 
(plate 2) figures six specimens showing white collar and _ black 
throat patch in different combinations. The other (plate 1) shows 
two series of selected winter specimens, illustrating, respectively, varia- 
tion in extent of white collar and of black throat patch. To our present 
knowledge these variations in color and pattern are not correlated with 
any peculiarities in the geographical distribution of the subspecies, but 
are due to individual variation, greater in this than in any other of the 
races of Branta canadensis. To ascertain postively that this is the 
ease, however, will require the collecting of an extensive series of 
minima upon the breeding grounds. 

Altogether, the small size of minima, and particularly its very small 
bill, seem to be about its best diagnostic characters; but in this an 
arbitrary line must be drawn for separation from hwtchinsi, for speci- 
mens will be found showing every degree of gradation between the two. 

Comparative length of tarsus and middle-toe-and-claw seems to 
be a fairly dependable character, for minima is a decidedly long-legged 
bird as compared with canadensis. With a few exceptions in each 
ease, it may be said that in canadensis the middle-toe-and-claw is appre- 
ciably longer than the tarsus, in hutchinsi the middle-toe-and-claw is 
about the same length as the tarsus, and in minima the tarsus is longer. 


Branta canadensis occidentalis 


Of this race there are in the Museum collection eight skins of use 
in this connection; from the Sitkan district, southeastern Alaska, four 
specimens (nos. 18, 19, 9916, 23245), two adult males, an adult female, 
and the head and neck of an immature bird; from Prince William 
Sound, Alaska, four specimens (nos. 1129, 1130, 1132, 1133), two 
adult males, an adult female, and the head and neck of another adult 
female. 


6 University of California Publications in Zoology [ Von. 12 


Baird’s type of Bernicla occidentalis (coll. U.S. Nat. Mus. no. 5994) 
from Port Townsend, Washington, has been available for comparison 
with these Alaskan specimens, and in general it may be said that all 
appear to belong to the same race. The differences are (1) that the 
type specimen has a faintly indieated trace of a white half collar at 
the base of the neck, which none of the Alaskan birds possesses; (2) it 
has a more nearly continuous. line of black spots separating the white 
cheek patches; (3) it is of a more reddish brown color ventrally. These 
are all differences which, judging from more extensive series of other 
subspecies of canadensis, may well be due to individual variation, and 
altogether the Alaskan birds appear to be sufficiently like the type of 
occidentalis to justify the application of that name to the breeding 
birds of the regions where they were secured. Of the Alaska series 
the Prince William Sound birds are smaller and darker than those 
from the Sitkan district. The type of occidentalis is rather more 
reddish brown below than any of the others, the effect being produced 
by the broad tipping of cinnamon on the broceoli brown feathers of 
the underparts. 

From typical canadensis all of these northwest coast birds differ 
in their exceedingly dark coloration, and although there is some 
variation in the specimens from different regions, as indicated above. 
it is nothing compared with the gap between the darkest canadensis 
and the palest occidentalis at hand. 

In descriptive literature treating of the subject (Ridgway, 1884, 
pp. 456-459; 1887, pp. 116, 117; Coues, 1903, pp. 902, 904-906; ete.) 
stress has been laid upon the presence of the white collar at the base 
of the black neck, and the black line on the throat dividing the white 
cheek patches, these being generally cited as distinguishing characters 
of the race occidentalis. In the type of the subspecies the white half 
collar is barely suggested, and the black on the throat is merely a 
string of disconnected black spots. Of the Alaskan specimens, not one 
shows even a single white feather at the base of the neck, and while 
the black throat bar is in three cases faintly indicated by a few black 
spots, in the remaining five there is not a mark to interrupt the con- 
tinuity of the white cheek and throat patch. .Thus these supposedly 
characteristic markings are shown to be no more constantly present 
in the race occidentalis than they are in true canadensis, where a sug- 
gestion of such markings occasionally occurs. 

Judging from our specimens and taking birds of the Sitkan dis- 
trict, Alaska, as typifying the subspecies occidentalis, the distinctive 


2913] Swarth: Geese of California 


~l 


characters of this form, as distinguished from canadensis, are: (1) 
extremely dark coloration; (2) slightly smaller size, that is, the max- 
imum of occidentalis is below the largest canadensis; (3) proportion- 
ally longer tarsus. 

Of six specimens of occidentalis, four have eighteen rectrices each, 
one has sixteen and one fourteen. 

The greater size of occidentalis serves to distinguish it from hutch- 
insi. Although occasional specimens of the latter are as darkly colored 
as occidentalis, all such specimens examined are of small size, evidently 
intererades between hutchinsi and the dark-colored minima. 

Although in general tone of coloration occidentalis approaches 
minima, the difference in size between the two is in most cases so 
ereat as to render improbable any confusion of the races. This differ- 
ence is to some extent bridged by our specimens from Prince William 
Sound, which, as mentioned above, are distinctly smaller and darker 
than southern Alaska examples of occidentalis. The region they 
inhabit is closer to the range of minima, and it is fair to believe that 
these specimens illustrate a step in the gradual transition between 
the two forms, which probably occurs. They are, however, distinctly 
closer to typical occidentalis than to minima. 

The type specimen of occidentalis, in its small size, reddish colora- 
tion, and suggestions of white collar and black throat lne, appears to 
approach minima even more closely than do the Prince William Sound 
specimens. The probability suggests itself that this bird, of inter- 
mediate character, is from some point between the habitats of typical 
occidentalis and minima, and a winter visitant only, at the point of 
capture, Port Townsend, Washington. This implies a mode of migra- 
tion that is paralleled by that of other species of birds of the north- 
west coast region, such as with the fox sparrows (Passerella iliaca 
subspp.) and the hermit thrushes (Hylocichla guttata subspp.). That 
is, the individuals inhabiting the northernmost part of the region (as 
at Prince William Sound) pass over the Sitkan district immediately to 
the southward, and spend the winter in the southernmost parts of 
the humid region, while the birds of the Sitkan district either remain 
there throughout the year, or else move but a short distance to the 
southward. It is the northernmost individuals that perform the 
longest flights. 


8 University of California Publications in Zoology | Vou. 12 


CONCLUSIONS 


In the foregoing pages several facts have been demonstrated with 
a fair degree of certainty. First, in regard to the breeding bird of 
California: It has been generally believed that Branta c. occidentalis 
breeds south into California; the various books dealing with the subject 
practically agree in this notion. In Belding’s ‘‘Geese which occur in 
California’’ (1892, pp. 96-101) occidentalis is explicitly designated 
as the common goose of this group in the state, and true canadensis 
is mentioned only as of possible occurrence, the author not having per- 
sonally met with it. It is probable that many of the later, more gen- 
eral, works have taken this as their authority. It also seems highly 
probable that the idea was originally based on a misconception, Cali- 
fornia specimens not having been actually compared with eastern ones. 

Our single summer bird, from Lake Tahoe, taken with a set of eggs 
(see Ray, 1912, p. 70), is, as mentioned above, unquestionably canaden- 
sis, furnishing a definite record for the subspecies within the state. 
There is also a record (Merrill, 1888, p. 143) of the breeding of B. c. 
canadensis at Fort Klamath, in southern Oregon, it being explicitly 
stated that occidentalis does not occur in the region, winter or summer. 
These two facts afford good grounds for the assertion that the goose 
found breeding throughout the lake region of southern Oregon and 
northeastern California is Branta c. canadensis and not B. c. occiden- 
talis, as has been generally assumed. Moreover, this is what would 
naturally be expected, the general faunal complexion of the region 
being distinetly similar to that of the Great Basin, to the eastward, 
and not at all like that of the humid northwest coast region, the home 
of occidentalis. 

Second, regarding the alleged occurrence of occidentalis in Cali- 
fornia during the winter- months: In our series of the canadensis 
eroup of geese from the San Joaquin Valley, there are, as already 
shown, many typical examples of B. c. canadensis, and none which 
at all approaches B. c. occidentalis. In addition, it may be said that 
the collector of this series obtained many of his birds from market 
hunters, whose daily bags he had opportunities of examining, and that 
he was constantly on the lookout for specimens illustrating every 
variation of color and markings. That no example of occidentalis was 
secured means that none was seen among many times the number of 
canadensis that were actually preserved. 


1913 | Swarth: Geese of California 9 


In this connection it may also be said that the present writer had 
occasion at one time, during the winter of 1903-1904, to examine 
several times a week the geese received by various Los Angeles dealers, 
and although canadensis, hutchinsi, and minima were seen in about 
the same proportional numbers as in the series of San Joaquin Valley 
birds under discussion, not a single specimen of occidentalis was ever 
discovered. 

A eritical examination of the literature wherein the white-cheeked 
goose has been ascribed to California, fails to reveal any definite or 
substantial basis for the statement. The following citations are taken 
from Mr. Grinnell’s manuscript list of the birds of California, and 
refer to the subspecies Branta canadensis occidentalis as oceurring in 
this state: 

American Ornithologists’ Union Cheek-List (1886, p. 127); Ridgway (1887, 
p. 117); Belding (1892, p. 100); American Ornithologists’ Union Check-List, 
second edition (1895, p. 68); Grinnell (1902, p. 21); Bailey (1902, p. 68); Fisher 
(1906, p. 194); Cooke (1906, p. 78); Sheldon (1907, p. 187); American Ornithol- 


ogists’ Union Check-List, third edition (1910, p. 86); and Salvadori (1895, p. 
115, as Branta occidentalis) . 


With one exception none of the writers named offers any 
explanation or justification for the appleation of the name occidentalis 
to the species treated. Belding, in his ‘‘Geese which occur in Cali- 
fornia,’’ explicitly applied the name to the large California goose. 
He did not inelude true canadensis as occurring in the state, however, 
and apparently relied upon his memory in deciding that the birds 
seen were different from eastern ones. Thus he recognized but one 
large form of Canada goose within the state, calling it occidentalis, 
but without making any comparison with eastern material. We agree 
with him in so far that we also have detected but one of the two forms, 
but, on making careful comparison of Californian and eastern skins, 
have discovered no differences; hence we use the name canadensis for 
the California bird. 

Thus it seems safe to assert that the oft-repeated statement regard- 
ing the occurrence of the white-cheeked goose in California has no basis 
in fact, or at any rate rests on no authenticated published record. 

Third, as regards the status of the breeding goose of southeastern 
Alaska: In reports upon collections of birds from the region, made for 
this Museum, the name occidentalis was employed with some hesitancy 
(see Grinnell, 1909, p. 199; 1910, p. 373; Swarth, 1911, p. 47). This 
was principally because the birds secured did not show the definite 


10 University of California Publications in Zoology | Vou. 12 


white collar and black throat patch ascribed to occidentalis in descrip- 
tive literature, markings usually likened to those of minima, in which 
they are frequently very strongly developed. 

At present the conclusion to which the accumulated facts and 
specimens seem to point, is that there is a dark-colored, northwest 
coast race of Branta canadensis, which occupies the immediate vicinity 
of the coast, northwest at least to Prince William Sound, and for an 
undetermined distance southward, though probably not occurring in 
the summer farther south than the southern boundary of British 
Columbia. The specimen which served as the type of Bernicla occi- 
dentalis Baird appears to be an example of this dark, coast race, but 
is probably not truly representative of the form. Unfortunately some 
of the most variable characters have been seized upon, and used as the 
distinguishing marks of the subspecies, and consequently confusion 
has ensued in the attempt to fit the descriptions of this bird to the 
more normal representatives of the race. 

Since so many of the water birds of the coast of southern Alaska 
and British Columbia are resident the year through in that general 
region, it is very probable that the white-cheeked goose belongs in the 
same category. In a letter recently received from Mr. Allen HE. Hassel- 
borg, a resident of Juneau, Alaska, and familiar with the native birds 
and mammals, he confirms this view, saying that the geese are about 
as abundant in the Sitkan district in winter as in summer. During the 
winter they frequent the more sheltered south- and west-facing bays 
and inlets, avoiding localities exposed to the cold land winds, while 
in summer they are of more general distribution. That this subspecies 
does not perform as extensive migrations as other members of the group 
is evident from its non-oceurrence in California. If it occurs in this 
state at all it should be found along the extreme northern coast. 

Reference to the literature treating of the geese of the Branta 
canadensis group shows very little uniformity in results arrived at by 
different writers. Among the various views expressed are the possi- 
bilities: (1) that these geese all belong to one exceedingly variable 
species; (2) that there are two species, canadensis with its race hutch- 
insi, and occidentalis with the race minima; (3) that there are four 
species, perfectly distinet, but frequently hybridizing. In this con- 
nection see Ridgway, 1884, p. 457. 

Study of the series of specimens examined in connection with the 
present paper is strongly confirmatory of the belief expressed in the 
nomenclatural treatment accorded the Branta canadensis group in the 


1913 ] Swarth: Geese of California 13L 


A. O. U. Check-List (1910), namely, that there is one widely ranging 
and exceedingly variable species, of which we can recognize four differ- 
ent subspecies, canadensis, hutchinsi, occidentalis, and minima. The 
material at hand illustrates every gradation of size, color, and pattern, 
from typical canadensis on the one hand, through hutchinsi. to typical 
minima at the opposite extreme. There is, however, a lack of material 
indicative of intergradation between occidentalis and the other forms, 
except as shown by the Prince William Sound birds, which approach 
minima in size, and by the type specimen of occidentalis which, as 
previously pointed out, also inclines toward minima in certain of its 
characters. It is highly probable that collecting in the proper regions 
in summer would be productive of specimens variously intermediate 
between occidentalis and canadensis, or between occidentalis and 
minima. : 

The presence in California during the winter months of three 
subspecies of Branta canadensis, namely, canadensis, hutchinsi and 
minima, occurring in large numbers in associated flocks, together with 
the presence of a fairly large proportion of intermediates with diffi- 
culty referred to any one form rather than another, is caleulated to 
produce an erroneous impression as to the distinctness and stability 
of the various forms. It must be borne in mind that this is the winter 
habitat of races which, except where our data is inadequate, are known 
to occupy widely different areas during the summer months, and that 
it is merely the similarity of their preferences and requirements which 
brings the various races together in their common winter home. The 
same may be said of Anser albifrons gambeli, Chen hyperboreus, and 
Chen rossi, these absolutely distinct species, together with the different 
subspecies of Branta canadensis, all occurring during the winter 
months in exactly the same localities in California. 

Birds taken upon their breeding grounds do not exhibit any such 
diversity of characters as is illustrated in series of winter birds taken 
at one locality. On the contrary, breeding birds from any one 
locality are wonderfully uniform in all particulars. Much remains 
to be learned in regard to the summer ranges of the North American 
geese, but such exact data as is available serves to bear out the above 
statement. 

There are, however, published accounts of high degree of authen- 
ticity which are not clearly explicable, such as the alleged occurrence 
of hutchinsi and minima over the same region, in part, during the 
breeding season. It should be borne in mind, though, that the confu- 


1D University of California Publications in Zoology [ Vou. 12 


sion heretofore existing in regard to the respective summer habitats 
of canadensis and occidentalis was of exactly the same nature, and it 
accordingly seems probable that if the two northern forms could also 
be given more careful and critical study than has heretofore been 
accorded them, many of the discrepancies now apparent could be satis- 
factorily explained. 

Nelson (1887, p. 84) gives the center of abundance of hutchinsi 
as along the lower Yukon and from there south to the Kuskoquim; 
of minima (1. @., p. 86), as along the Alaskan coast of Bering Sea, 
north to Point Barrow, and extending far inland up the rivers. Cooke 
(1906, p. 77) gives the summer home of hutchinsi as including the 
Alaskan shores of Bering Sea, and also the western Aleutian and the 
Near islands, exactly the same range as is aseribed to minima (I. ¢., 
p. 78). 

Such statements as these are hard to comprehend if the birds are 
to be considered as two races of the same form, but there are so many 
chances for error, or for misunderstanding of data, that it seems 
almost certain that further careful work will demonstrate that the 
two forms do actually oceupy separate and well-defined areas during 
the breeding season. It is certainly possible for migrating individuals 
of one form to have been captured upon the summer home of the 
other, while the variation in color and patterns in both forms is so 
ereat that there is chance for error in identification unless specimens 
be secured and examined with great care. 

There appears to be a dearth of definite data bearing upon the 
question. The two statements cited above are of a general nature, 
and are not supported by mention or description of particular speci- 
mens to uphold the ideas expressed. In fact there seems to be but few 
instances in which this has been done. Clark (1910, p. 47) gives 
hutchinsi as the breeding goose of Agattu and Attu islands. Although 
specimens were not saved, descriptions are given of some which were 
shot and thrown away, descriptions which, as regards color and pat- 
tern, fit minima very well, though the dimensions given are certainly 
large for that form. 

Bent (1912, p. 13) identifies a breeding goose from Attu Island 
as minima. Thus the statements in regard to the respective breeding 
ranges of hutchinsi and minima are seen to be contradictory and con- 
fusing; but, as stated above, this may well have arisen through mis- 
understanding of data, or misintérpretation of characters. Where we 
do possess more exact and abundant data, as in regard to the more 


1913] Swarth: Geese of California 13 


southern parts of the summer habitat of canadensis, or the summer 
home of occidentalis, we find no such confusion. 

Reasoning from geographical grounds, and in the belief that hutch- 
insi and minima are two subspecies of the same species, Branta cana- 
densis, the two forms should certainly not be expected to occur together 
during the breeding season on the same parts of the Alaskan coast 
of Bering Sea and on the Aleutian Islands. As hutchinsi is the more 
eastern species, every alleged occurrence of its breeding in the above 
region, where minima is known to be common, should be most fully 
substantiated. Minima is not known to occur elsewhere in summer, 
and it may be inferred that this is the breeding ground of the race. 

Following is a list of the characters which have been used in litera- 
ture in differentiating the four subspecies of Branta canadensis. These 
are compiled from the various handbooks dealing with the subject, and 
are those characters which have been supposed to be diagnostic. 


B. ¢e. canadensis 


(large ; é 
[B. e. oecidentalis 
General size medium B. ec. hutehinsi 
(small B. e. minima 


(pale grayish, fading 
gradually into white 
of crissum 


fB. ce. canadensis 
|B. e. hutehinsi 
Color of under 


puraacexct body dark brown, abrupt- 


ly defined against 


ie ce. occidentalis 
white of crissum 


B. ¢. minima 


continuous across (B. e. canadensis 


) B. e. hutehinsi 
Triangular white BERD | B. e. hutehinsi 


See aueles . @ occidentalis 


separated by black B 
B. ¢. minima 
B 
B 


bar on throat 


. @ canadensis 
e. hutehinsi 
White collar on 


usually very 
lower neck y k 


ae . @ occidentalis 
distinet; always 


B 
B. e. minima 


usually absent { 


present 
18 to 20 B. ¢e. canadensis 
Number of 18 to 20 B. e. occidentalis 
rectrices 16 B. e. hutechinsi 
14 to 16 B. ¢. minima 


14 Unversity of California Publications in Zoology [ Vou. 12 


In testing the above characters upon the large series of geese avail- 
able in the present study many proved so extremely variable as to be 
of no practical value, while others are based entirely upon miscon- 
ceptions. 

Careful examination of the series of geese at hand justifies the fol- 
lowing diagnoses as more nearly expressing the differences existing 
between the four subspecies. 


Branta canadensis canadensis 

Size large. : 

Tarsus usually shorter than middle toe and claw. 

Color of under surface of body pale gray; much lighter shade than back. 

Black of neck abruptly defined against gray of breast. 

Gray of lower surface fading gradually into white of crissum. 

White cheek patches usually confluent across throat; occasionally a few 
black spots along median line of throat; or slight encroachment of 
black indicating the beginnings of the black isthmus. 

White collar at base of black neck usually absent; occasionally faintly 
indicated. 

Number of rectrices 14 to 20 (usually 18). 


Measurements in millimeters. Wing, 418-527 (500.6); tail, 184-174 
Minimum, maximum, and (156.8); eulmen, 47-58 (52.4); tarsus, 
average of ten adult males. 76-98 (92.3); middle toe and claw, 

77-106 (97.3). 
Branta canadensis occidentalis 

Size large. 

Tarsus as long as, or slightly longer than, middle toe and claw. 

Color of under surface of body dark; broccoli brown, about same shade 
as back. 

Black of neck abruptly defined against brown of breast. 

Brown of abdomen abruptly defined against white crissum. 

White cheek patches usually confluent across throat; occasionally a few 
black spots along median line of throat, or slight encroachments 
of black indicating the beginnings of thé black isthmus. 

White collar at base of black neck usually absent; occasionally faintly 
indicated. 

Number of rectrices 14 to 18 (usually 18). 

Measurements in millimeters. Wing, 4383-495 (465.7); tail, 184-154 
Minimum, maximum, and _ (142.7); culmen, 48-51 (48.7); tarsus, 
average of four adult males. 86-99 (92.2); middle toe and claw, 

83-95 (87.2). 
Branta canadensis hutchinsi 

Size medium. 

Tarsus about the same length as middle toe and claw. 

Color of under surface variable; pale gray to dark brown. 

Pattern of head and neck markings variable. Black dividing line on 
white throat may or may not be present. White collar at base of 
neck may or may not be present. 

Number of rectrices 16 to 18 (usually 16). 


1913] Swarth: Geese of California 15 


ae a 


Fig. a—Branta canadensis canadensis, adult male, no. 21971, Los Baiios, Cali- 
fornia; lateral view of head and bill; x 0.83 +. 

Fig. B—Branta canadensis hutchinsi, adult male, no. 22004, Los Bafios, Cali- 
fornia; lateral view of head and bill; x 0.83 +. 

Fig. c.—Branta canadensis minima, adult male, no. 22018, Los Banos, Cali- 
fornia; lateral view of head and bill; x 0.83 +. 

These figures, approximately natural size, show the relative proportions of 
the bill in the three forms, one of the most reliable single characters differ- 
entiating the subspecies. 


16 University of California Publications in Zoology (Vou. 12 


Measurements in millimeters. Wing, 390-455 (419); tail, 101-140 
Minimum, maximum, and (122.1); culmen, 34-45 (39.4); tarsus, 
average of ten adult males. 68-86 (77); middle toe and claw, 

67-85 (73.5). 


Branta canadensis minima 

Size small. 

Tarsus much longer than middle toe and claw. 

Color of under surface variable; pale gray to dark brown. 

Pattern of head and neck markings extremely variable; black dividing 
line on white throat may or may not be present; white collar at base 
of neck may or may not be present. Every possible combination of 
the above patterns and colors. 

Number of reetrices 14 to 18 (usually 16). 

Measurements in millimeters. Wing, 337-421 (385.3); tail, 94-128 
Minimum, maximum, and (111.7); culmen, 26-86 (80.8); tarsus, 
average of ten adult males. 61-81 (72.9); middle toe and claw, 

55-76 (64.60). 


9 4| |4 

8 4 4| [4 

7 4 4| |4| [4 | 

6| |a|4| | 4) 4) 4) 4/4) 4 . 

S| |a\4| | 4) 4) 4) 4| 4) 4 aime |e] |a/e\e 

4| |a|ala|4|4| 4] 4/4[@) ele| |e| le [a] jeieie] | [le 

3| |a|a|a|4|4|ala[a[@a| ACIS e se ee 8 a a 

2 4|4| 4) 4\ 4) 4| 4 (PPP eleleleieiee a\e\a\5\8 |f\8 a 

1|a| 4) 4) 4| 4) 4| 4) 4 @ PP eleleieieeleieie| jeiaimiaisia| (as aia 
26 27 28 29 30 3) 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 5B 
4 Branta c. minima @ = Branta c. hutchinsi ®@ Branta c. canadensis 


Fig. p.—Length of culmen of B. c. canadensis, B. c. hutchinsi, and B. ec. minima. 
Each symbol represents a specimen. Numerals at left of diagram indicate 
number of specimens; numerals at the bottom, length of culmen in millimeters. 


82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 9B 99 100 {Ol 102 103 104 105 106 107 


4 Branta c. minima @ Branta c. hutchinsi @ = Branta c. canadensis 

Fig. r.—Ratio per cent of middle-toe-and-claw to tarsus in B. ce. canadensis, 
B. c. hutchinsi, and B. c. minima. Numerals at left of diagram indicate number 
of specimens; numerals at the bottom, ratio. 


17 


Geese of California 


Swarth: 


1913] 


a{ | a|4 @ | 4|\ a 

1\4|4| 4 © 5° 4a © r] 
61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 9! 92 93 94 95 96 97 
4 Branta c. minima e Branta c. hutchinsi @ Branta c. canadensis 


Fig. r.—Length of tarsus of B, ec, cana s, B. c. hutch and B. ce. 7 
specimens; numerals at the bottom, length of tarsus in millimeters. 


= GOS 
ne 
NAN 


NINISIN 


ima. Numerals at left of diagram indicate number of 


: 4 4 E JE) | | 

4| 4\4 4 4\4 | 

s|a| [a] |4|4 y ae |e e ° 5 a 

2| A|a|4| 4 4| 4 4 e| *,eJeJele ele| je a | [eieijele| fo | 
ifalalala|alal ala} al al [aleeeejejeje| 4) Pee, je] jm) Me |e miami ieee ajole| |e] (e |e 


55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 Bl 82 83 84 BS 86 87 BB 89 90 91 92 93 94 95 96 97 98 99 100 JQ) 102 103 104 105 106 


4 Branta c, minima e Branta c. hutchinsi = Branta c. canadensis 


Fig. G.—Length of middle-toe-and-claw of B. ¢. canadensis, B. ¢. lutchinsi, and B, e. 
at the bottom, length of middle-toe-and-claw in millimeters. 


ima, Numerals at left of diagram indicate 


number of specimens; numera 


[ Vou. 12 


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University of California Publications in Zoology 


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SISUapvUuvI I DIUDIG: 


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— 


1913] Swarth: Geese of California : 19 


SUMMARY 


Briefly summarized, the following are the conclusions reached in 
the present paper: 

(1) The geese of the Branta canadensis group are best regarded 
as one species, Branta canadensis, with four subspecies, canadensis, 
occidentalis, hutchinsi, and minima; this without attempting to go 
into the problems involved in the nomenclature of the races—the ques- 
tions arising as to the proper application of the names lewcopareia, 
hutchinsi, minima and occidentalis—but accepting the usage adopted 
in the 1910 edition of the A. O. U. Check-List. 

(2) The status of the group in California is as follows: The form 
found breeding in the state is Branta c. canadensis; the forms occur- 
ring in winter are canadensis, hutchinsi, and minima. Contrary to 
the statement repeated in practically all ornithological books dealing 
with the subject, B. c. occidentalis does not occur in California at 
any season. 

(3) Branta c. occidentalis is a well-defined subspecies occupying 
the humid, northwest coast region, where it is practically resident, 
performing only the most limited migrations, or none at all. 

(4) In differentiating the subspecies of Branta canadensis undue 
emphasis has heretofore been placed upon certain characters which are 
for the most part too variable to be depended upon. Thus the cause 
of the confusion which has existed relating to the true status of B. c. 
occidentalis is partly due to mistaken ideas as to which are the most 
nearly constant characters of the subspecies. A careful analysis of 
many of the contradictory statements on record in regard to the sum- 
mer habitats of B. c. hutchinsi and B. c. minima makes it appear 
probable that this uncertainty also is largely due to misconceptions 
as to the real characters of these forms. 


Transmitted February 6, 1913. 


20 University of California Publications in Zoology | Vou. 12 


LITERATURE CITED 


AMERICAN ORNITHOLOGISTS’ UNION COMMITTEE. 

1886. Check-list of North American birds. (New York, American Ornith- 
ologists’ Union), pp. i-viii, 1-892. 

1895. Check-list of North American birds. Ed. 2, revised (New York, 
American Ornithologists’ Union), pp. i-xi, 1-372. 

1910. Check-list of North American birds. Ed. 3, revised (New York, 
American Ornithologists’ Union), pp. 1-480, 2 maps. 

BalLey, F. M. 

1902. Handbook of birds of the western United States. (Boston and 
New York, Houghton, Mifflin and Company; Cambridge, the 
Riverside Press), pp. i-xcii, 1-512, 3 pls., 601 figs. in text. 

BEtpine, L. 
1892. Geese which occur in California. Zoe, 3, 96-101. 
Bent, A. C. 

1912. Notes on birds observed during a brief visit to the Aleutian 
Islands and Bering Sea in 1911. Smithsonian Inst., Miscellaneous 
Collections, 56, no. 32, 1-29. 

CuarK, A. H. 

1910. The birds collected and observed during the cruise of the United 
States Fisheries Steamer ‘‘ Albatross’’ in the North Pacifie Ocean, 
and in the Bering, Okhotsk, Japan, and Eastern seas, from April 
to December, 1906. Smithsonian Inst., U. S. Nation. Mus., Proe., 
38, 25-74, figs. in text. 

Cooke, W. W. 

1906. Distribution and migration of North American ducks, geese and 

swans. U.S. Dept. Agric., Bureau Biol. Sury., Bull. no. 26, 1-90. 
Cougs, E. 
1903. Key to North American birds, fifth edition, 2, i-vi, 537-1152, 1 pl., 
figs. in text, 354-747. 
FisHer, W. K. 
1906. Tame wild geese. Bird-Lore, 8, 193-195, 5 figs. 
GRINNELL, J. 

1902. Cheeck-list of California birds. Pacifie Coast Avifauna, 3, 1-98, 2 pls. 
(maps). 

1909. Birds and mammals of the 1907 Alexander expedition to south- 
eastern Alaska. The birds. Univ. Calif. Publ. Zool., 5, 181-244, 
pl. 25, 2 figs. in text. 

1910. Birds of the 1908 Alexander Alaska expedition, with a note on 
the avifaunal relationships of the Prince William Sound district. 
Univ. Calif. Publ. Zool., 5, 361-428, pls. 32-34. 

MERRILL, J. C. 

1888. Notes on the birds of Fort Klamath, Oregon. Auk, 5, 139-146, 251- 

262, 357-866. 
NELSon, E. W. 

1887. Report upon natural history collections made in Alaska between 
the years 1877 and 1881. Arctie Series of Publications issued in 
connection with the Signal Service, U. S. Army, 3, 337 pp., 21 pls. 


1913] Swarth: Geese of California 21 


Ray, M. 8. 
1912. Nesting of the Canada goose at Lake Tahoe. Condor, 14, 67-72, 
4 figs. in text. 
Ripeway, R., in Barro, S. F., Brewer, T. M., and Ripeway, R. 
1884. The water birds of North America, vol. 1. Memoirs Mus. Comp. 
Zool., 12, i-xi, 1-537, figs. in text. 
1887. A manual of North Ameriean birds. (Philadelphia, J. B. Lippincott 
Co.), pp. i-xi, 1-631, pls. [1] + i-exxiv. 
Sanvaport, T. 
1895. Catalogue of the birds in the British Museum, 27, i-xv, 1-636, 
pls. i-xix. 
SHELDON, H. H. 
1907. A collecting trip by wagon to Eagle Lake, Sierra Nevada Moun- 
tains. Condor, 9, 185-191. 
SwartH, H. 8. 
1911. Birds and mammals of the 1909 Alexander Alaska expedition. Univ. 
Calif. Publ. Zool., 7, 9-172, pls. 1-6. 


PLATE 1 


Ventral view of heads of Branta canadensis minima, illustrating variation 
in color patterns. The Museum numbers of the specimens shown, reading from 
left to right, are as follows: upper row, 22076, 22038, 22021, 22060, 22014, 
22045, 22088; lower row, 22031, 22021, 22027, 22039, 22017, 22044, 22056. 

The specimens in the upper row are arranged to show variation in the 
breadth of the white collar, ranging from the one at the extreme left, in 
which there is more white than black on the neck, through varying degrees 
to the specimen at the extreme right, in which the white collar is totally 
lacking. 

The lower row shows variation in the black throat bar, this mark being most 
highly developed in the specimen at the extreme left, and ranging through 
lessening degrees of extensiveness to total absence in the one at the extreme 
right. 


As shown here there is no correlation between these two markings, each 
of them varying independently of the other. Another color character, the 
shade of the underparts (hardly apparent in a black and white illustration) 
also varies independently of either of these. The birds shown on the plate 
were all collected in their winter home. 


PLATE 2 


Ventral view of heads of Branta canadensis minima, illustrating variation 
in color patterns. The Museum numbers of the specimens shown, reading from 
left to right, are as follows: upper row, 22088, 22014, 22076; lower row, 22035, 
22042, 22033. 

Typical minima is supposed to be marked with a white collar at the base of 
the black neck, and with a black line on the throat dividing the white cheek 
patches, as shown in no. 22085. The specimens figured illustrate every possible 
combination of these markings, demonstrating the impossibility of properly 
diagnosing the subspecies on the basis of color or pattern. 


UNIV, CALIF. PUBL. ZOOL. VOL. 12 [SWARTH] PLATE 2 


—— 


— 


Vol, 8. 


Vol. 9, 


Se 


‘UNIVERSITY OF CALIFORNIA PUBLICATIONS— (Continued) 


(Contributions from the Museum of Vertebrate Zoology.) 

1, Two New Owls from Arizona, with Description of the Juvenal Plum- 
age of Strix occidentalis occidentalis (Xantus), by Harry 8. Swarth. 
Moy tee Oe aiaeg 2 9 Sa A 0 eae Sacer iste oa Rs sa cee iat aR We =n | Sem pen se eaten 

2. Birds’ and Mammals of the 1909 Alexander Alaska» Expedition, by 
Harry S. Swarth. Pp, 9-172; plates 1-6; 3-text-figures. January, 1911. 

3. An Apparent Hybrid in the Genus Dendroica, by Walter P. Taylor, 
PVA RaAT bg POUL ALY eh ON Beetham ee Ba cen YER ee SES 

4. The Linnet of the Hawaiian Islands: a Problem in Speciation, by 

5 


Joseph Grinnell, Pp. 179-195, February, 19110... cke nk. 
«The Modesto Song Sparrow, by Joseph Grinnell. Pp. 197-199. Feb- 
GUC B AA epg Ro 8 Bt LSE Par eh As sk erm ie ent oe cr NM Hef <a> Se RANI SRS 

6. Two. New Species of Marmots from Northwestern America, by H. S. 
Swarth. Pp. 201-204>> “February, Oa as Bar ee Sa 

7. Mammals of the Alexander Nevada Expedition of 1909, by Walter P. 
Taylox.— Pp. 205-307.osume) LOM =e Se eee Son atest keene 

8. Description of a New Spotted Towhee from the Great Basin, by J. 
Grinnell,"-Pp> 309-314 FA nenst, Lon oe ee eee 

9. Description of a New Hairy Woodpecker from Southeastern Alaska, by 
HS, Swarth,-Pp. $13-818.-October, 1911 2222.2 ne 

10, Field Notes on Amphibians, Reptiles and Birds of Northern Humboldt 
County, Nevada, with a Discussion of Some-of the Faunal Features 

of the Region, by Walter P. Lexa Pp. 319-486, plates '7-12. 
ECP UAT yee OF os ke sete oe eT Ce eS 

Index; pp. 437-446. 


1, The Vertical Distribution of Fucalanus elongatus in the San Diego 
Region during 1909, by Calvin O. Esterly. Pp. 1-7. May, 1911........ 
2, New and Rare Fishes from Southern California, by Edwin Chapin 
Starks and William M.Mann. Pp. 9-19, 2 text-figures. July, 1911. 
Classification and Vertical Distribution of the Chaetognatha of the San 
Diego Region, Including Redescriptions of Some Doubtful Species*‘of 
the Group, by Ellis L. Michael. Pp. 21-186 pls. 1-8, December, 1911, 
Dinoflagellata of the San Diego Region, IV. The Genus Gonyaulax, with 
Notes on Its Skeletal Morphology and a Discussion of Its Generic 
and Specific Characters, by Charles Atwood Kofoid. Pp, 187-286, 
plates 9-17. 
5. On the Skeletal Morphology of Gonyaulax catenata (uevander), by 
Charles Atwood Kofoid.. Pp. 287-294, plate 18, 
Dinoflagellata of the San. Diego Region, V..Cn Spiraulaz, a New Genus 
of the Peridinida, by Charles Atwood Kofoid. Pp. 295-500, plate 19. 
Nos. 4, 5, and 6 in one cover. “September, 1911>_..0- 0. 
Notes on Some Cephalopods in the Collection of the University of Cali- 
fornia, by S. S. Berry:.- Pp. 301-310, plates 20-21. September, 1911. 
Ona Self-Closing Plankton Net for Horizontal Towing, by -Charles 
Atwood Kofoid. Pp. 311-348, plates 22-25, 
On an Improved Form of Self-closing Water-bucket for Plankton In- 
vestigations, by Charles Atwood Kofoid. - Pp. 349-352, 
Nos. 8 and 9. in one cover. November, 1911.00... ee Ss 
Index, pp. 353-357. 


1,-The Homed Lizards of California and Nevada of the Genera Phryno- 

soma and Anota, by Harold C. Bryant. Pp. 1-84, plates 1-9. Decem- 

Taken Urs Ae Re ae a ae ek LI Si, EE EP NS, omy ms aL 

2, On a Lymphoid Structure Lying Over the Myelencephalon of Lepisos- 

teus, by Asa C, Chandler, Pp. 85-104, plates. 10-12. December, 1911. 

-8. Studies on Early Stages of Development in Rats and: Mice, No. 3, by 

- BE. L. Mark and J. A. Long. The Living Eggs of Rats and Mice with 

a Description of Apparatus for Obtaining and Observing Them (Pre- 

liminary paper), by J. A. Long. Pp, 105-136, plates 13-17. February, 

BAS 8 0A Siar ane Ee eee Saale Sect ieln a CO AS Or mee Ue eR Rea oR Be eae 

4. The Marine Biological Station of San Diego, Its History, Present Con- 

ditions, Achievements, and Aims, by Wm. E. Ritter, Pp. 137-248, 

platescts-24 and “2 maps. IW arch, N91 as ees 

5, Oxygen and Polarity in Tubularia, by Harry Beal Torrey. Pp. 249- 

251, May, 1912.2... 

6, The Occurrence and Vertical Distribution of the ‘Copepoda ‘of. the ‘San 

Diego Region, with particular reference to Nineteen Species, by Cal- 

vin ©. Esterly. Pp. 258-340, 7 text-figures. July, 1912 220 

7. Observations on the Suckling Period in the Guinea-Pig, by J. Marion 

Reads—Pp. 341-851.- September; 1992 33 oe 

Haeckel’s Sethocephalus éucecryphalus (Radiolaria), a Marine Ciliate, 

py Charles Atwood Kofoid. Pp. 353-357, September, 1912 _..........: 
Index, pp. 359-365, : 


% 


sa 


> 


Peay an 


a 


1.00 


10 
-10 


1.75 


1.50 
10 


40 


UNIVERSITY OF CALIFORNIA PUBLICATION: S—(Continued) Z 


Vol. 10, Spleen from the Museum of Vertebrate Zoology.) 


2. 
3. 
4 


oy 


10. 


Vol, 11. 1. 
2. 


3. 


7 


Vol. 12. 1. 


1. Report on 4 Collection of Birds and Mammals from Vancouver Island,. 
by Harry S. Swarth. Pp. 1-124, plates 1-4, February, 1912.00.03... 


A New Cony from the Vicinity of Mount Whitney, by Joseph Grinnell. 


Pp, 125-1293 January, 1912. sso 6 ne ee 


The Mole of Southern California, by J. Grinnell and H. 8. Swarth., . 


Pp.. 131-136, 2 text-figures. 
Myotis orinomus Elliott, a Bat New to California, by J. Grinnell and 
H.S. Swarth. Pp. 137-142, 2 text-figures. 
Nos. 3 and 4 in one cover. April, 1912 2222.20 tne 


. The Bighorn of the Sierra Nevada, by Joseph Grinnell, Pp, 143-153, © 


4 Lext-heourés, Way TOV ee ah tog Dea nearest ep agente aig 


. A New Perognathus from the San Joaquin Valley, Celta by. 


Walter P, Taylor. Pp. 155-166, 1 text-figure. 


. The Beaver of West Central California, by Walter P. Taylor... Pp. 


167-169, ; 
Nos. 6 and 7 in one cover, May, 1912 ........: tle a ee ate 


. The Two Pocket Gophers of the Region Contiguous to the Lower Colo- 


rado River, in California and Arizona, by Joseph Grinnell. Pp. 171- 
LUBE SACS BOGS aaa ae ESR ae cc gee it ae Se ahaa 


. The Species of the. Mammialian Genus Sorex. of West-Central Cali- 


fornia, with a note on the Vertebrate Palustrine Faunas of the 
Region, by Joseph Grinnell. Pp.-179-195, figs. 1-6. March, 19138....... a 
An Account of the Birds and Mammals of the San Jacinto Area of 
Southern California, with Remarks Upon the Behavior of Geographic 
Races on the. Margins of Their Habitats, by J, Grinnell and H. 8. 
Swarth. Pp. 197-406, pls. 6-10, October, 1913. 222.222... Se 
Index, pp. 407-417. 


Birds in Relation to a Grasshopper Outbreak in California, by Harold 


C. Bryant. Pp. 1-20: November, 1912 -2..20.2.cccccccl ec cecescce co eecccenenseene 5 


On the Structure and Relationships of Dinosphaera palustris (Lemm.), 
by Charles Atwood Kofoid and Josephine Rigden Michener. Pp. 21- 
29;-. WeCeni ber, “LOIS ee es ee Se ee eee 

A Study of Epithelioma Contagiosum of the Common Fowl, - by 
Clifford D. Sweet. Pp. 29-51. January, TOUS 3. OS Se 

The Control of Pigment Formation in Amphibian Larvae, by Myrtle 
E. Johnson. Pp. 53-88, plate 1.. March, 1913 2.2.22. cee. eteei eee 


. Sagitta californica, nu. sp., from the San Diego Region, including 


Remarks on Its Variation and Distribution, by Ellis L. Michael. 
Pp: 89-126, plate 2. Med BLS aaa sacs cncp Sasa gene 


. Pycnogonida from the Coast of California, with Description of Two 


New Species, by H. V. M. Hall, Pp. 127-142, plates 3-4. August, 1913. 
Observations on Isolated Living Pigment Cells from the Larvae of 
Amphibians by S. J. Holmes. Pp. 143-154, plates 5-6. 


. Behavior of Ectodermic Epithelium of Tadpoles when Cultivated in 


Plasma, by S. J. Holmes. Pp. 155-172, plates 7-8. 


Nos. 7 and 8 in one cover. September, 1913 -2.....0.02..c.. cots : 


A Study of a Collection of Geese of the Branta Canadensis Group from 
the San Joaquin Valley, California, by Harry S. Swarth: Pp. 1-24, 
plates 1-2, 8 text figs. November, 1913... 


. Nocturnal Wanderings of the Oalifornia Pocket Gopher, by Harold C. 


Bryant, Pp. 25-29, 1 text fig. November, 1913 00 te 


. ‘The Reptiles of the San Jacinto Area of Southern California, by Sarah 


Rogers Atsatt. Pp. 31-50; November, 1913 —.... Bu achacst-scpa spent ia 


2,00 


Py + are Pale ts 


i? 
wed 


Fragctt 3) wa 


UNIVERSITY OF CALIFORNIA PUBLICATIONS 
IN 


“ZOOLOGY 
Vol. 12, No.-2, pp. 25-29, 1 text fig. November 20, 1913 


NOCTURNAL WANDERINGS OF THE 
CALIFORNIA POCKET GOPHER 


BY 
HAROLD C. BRYANT 


UNIVERSITY OF CALIFORNIA PRESS 
BERKELEY : 


UNIVERSITY OF CALIFORNIA PUBLICATIONS 


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Vol. 6. 1. (XXIII) On the Weight of Developing Eggs. Part I, The Possible 
Significance of Such Investigations, by William E. Ritter; Part It, 
Practicability of the Determinations, by Samuel E. Bailey. Pp. 1-10. 


Octo ber; 4. 9068 ee ren ee Ts te Se -10 
2. (XXIV) The Leptomedusae of the San Diego Region, by Harry Beal 

Torrey. Pp. 11-31, with 11 text-figures. February, 1909 -............-: J. 20 
3. (XXV) The Ophiurans of the San Diego Region, by J. F. McClen- ~ 


don. . Pp. 33-64, plates 1-6. Tully, 1909 2.n cece ccen ele ece neces 30 
4, (XXVI). Halocynthia johnsoni n.sp.: A comprehensive inquiry as. to : 
the extent of law and order that prevails in a single animal species, 

by Wm.'E. Ritter. Pp. 65-114, plates 7-14. November, 1909 ~........ 50 
5, (XXVII) Three Species of Cerianthus*from Southern California, by 
H. B. Torrey and F. L. Kleeberger. Pp. 115-125, 4 text-figures. 


December 1 909 os. 23 set ee ee ee ate co caso ~. 10 ; 


6. The Life History. of Trypanosoma dimorphon Dutton & Todd, by 
Edward Hindle. Pp. 127-144, plates 15-17, 1 text-figure. December, i 
TOD Ge ee oe Se er hig BEE SS ES os See ts ago 

7. (XXVIII) A Quantitative Study of the Development of the Salpa 
Chain in Salpa fusiformis-runcinata, by Myrtle Elizabeth Johnson. 

Pp. V45-1 7625 Rae, VOTO es ac npd eccn sneer eee Fe ces tty acon 35 

‘8. A Revision of the Genus Ceratocorys, Based on Skeletal Morphology, 


by Charles Atwood Kofoid. Pp. 177-187. May, 1910. .t.2.....c.ceeessne 10 


9. (XXIX) Preliminary Report on the Hydrographic Work Carried on by 
the Marine Biological Station of San Diego, by George F. McEwen. 
Pp. 189-204; text-figure and map. May, 1910 2.2.2. ese eeedeeeeesene 15 
10. (XXX) Biological Studies on Corymorpha. III, Regeneration of Hy- 
dranth and Holdfast, by Harry Beal Torrey. Pp. 205-221; 16 text- 
figures. ‘ 
At, (EXXI) Note on Geotropism in Corymorpha, by Harry Beal Torrey. 
Pp. 223-224; 1 text-figure. 


Nos 10 and 11°in one cover. August, 1910 2n..1n epee 20 

42. The Cyclostomatous Bryozoa of the West Coast of North America, by 
Alice Robertson. . Pp. 225-284; plates 18-25.. December 1910 <2... *~ 60 

13. Significance of White Markings in Birds of the Order Passeriformes, 
by Henry Chester Tracy. Pp. 285-312. December, 1910 wansesnneecntrnene 625 

14. (XXXIII) Third Report'on the Copepoda of the San Diego Region, by 
Calvin Olin Esterly. Pp. 313-352; plates 26-32. February, 1911 ...... 40 


15; The Genus Gyrocotyle, and Its Significance for Problems of Cestode 
Structure and Phylogeny, by Edna Earl Watson. Pp. 353-468; plates 
33-480 Tarmac ngenn tmeepntetnndacne conn nens nbapmeretetenensarens 1.00 


Index, pp. 469-478. 


* Roman*numbers indicate sequence of the Contributions from the Laboratory of the 
Marine Biological Association of San Diego. ¢ : ; 


UNIVERSITY OF CALIFORNIA PUBLICATIONS 
IN 


ZOOLOGY 
Vol. 12, No. 2, pp. 25-29, 1 text fig. November 20, 1913 


NOCTURNAL WANDERINGS OF THE 
CALIFORNIA POCKET GOPHER 


BY 


HAROLD C. BRYANT 


(Contribution from the Museum of Vertebrate Zoology of the University of California) 


An incident bearing on the life-history of the pocket gopher has 
been recently reported to the writer by Mr. J. E. Light of Berkeley, 
California. He found on the morning of May 1, 1913, more than fifty 
pocket gophers (Thomomys bottae bottae) stuck in a strip of oil about 
two feet wide which had been left along the side of a street in process 
of repair in North Berkeley. These facts have been fully verified by 
the writer. All of the gophers appeared to have been traveling in the 
same direction, namely, east, and had crossed a macadam road before 
reaching the oil. The sticky oil had rendered them helpless in a short 
time, and their struggles to escape resulted only in entrapping them 
the more firmly, thus hastening their death. Some had been torn to 
pieces by cats and dogs after being caught in the oil, and others had 
doubtless been carried away altogether by these predators. Ten of the 
gophers were collected the next day within a stretch of two hundred 
feet. Two of these were apparently adult males, the rest being females 
and half-grown young. 

If this incident be taken as evidence, gophers come out of their 
burrows at night and travel above ground. Some mammals are known 
to travel about during the rutting season more than at other times. 
The time of year and the occurrence of large numbers of half-grown 
individuals, however, would seem to preclude the use of this fact in 
explanation of the phenomenon in this case. Bailey (1895, p. 16) in 
speaking of the pocket gopher states: ‘‘ Apparently only the males 


26 University of California Publications in Zoology [ Vou. 12 


leave the burrows in quest of mates, though positive information on 
the subject is difficult to obtain.”’ 

The only reference to a habit of foraging above ground that the 
writer has been able to find is the following from E. T. Seton (1909, 
p. 571): “‘Frequently, possibly every night, the gopher quits the 
burrow and sallies forth into the open air, foraging for grain and other 
foods not obtaimable underground. These it crams into its pouches, 
then retires to its burrow to consume them. The cover of night is 
essential to these expeditions ; they are seldom made in broad daylight, 
though they may be undertaken in twilight or by the light of the 
moon.’’ The present writer has been told that in California different 
species of pocket gophers have been occasionally observed above ground 
at night. 

Mr. Light in experimenting on methods of trapping in the same 
locality (North Berkeley) has found that gophers will follow ditches. 
By digging a small trench and sinking into the earth empty oil cans 
as pitfalls, a method also used in trapping moles, he has been successful 
in catching a number of pocket gophers. This is further evidence that 
gophers travel about on top of the ground at night to a greater extent 
than has hitherto been supposed. 

This habit of quitting the burrow at night would also seem to be 
substantiated by the fact that barn owls capture large numbers of 
pocket gophers. One of these birds has been recorded as capturing 
as many as fifteen gophers in a single night. It hardly seems probable 
that so large a number could be obtained were not some of them picked 
up on top of the ground. 

Unlike most of the strictly nocturnal rodents, the pocket gopher 
may often be seen feeding in the daytime, especially if the day be a 
cloudy one. A quivering plant, on the roots of which the gopher is 
feeding, or a momentary glimpse of a dark head disappearing down 
a hole, is usually the only evidence to be noted. It is very seldom, 
if ever, that a gopher is seen outside of his burrow during daylght, 
except when flooded out by irrigation. The writer’s only experience 
in this regard was in a newly ploughed field, where he found a large 
gopher wandering about in one of the furrows. It had doubtless been 
ploughed out, or its burrow had been disturbed by the ploughing. 
In the endeavor to reach a food plant a gopher will sometimes go as 
much as six inches from the mouth of the burrow, but it will dart back 
in alarm at the slightest disturbance. In the late summer a circular 
area of a radius of about the distance from the rump to the nose of a 


1913] Bryant: Wanderings of Pocket Gopher 27 


gopher can be found picked clean around the mouth of the burrow, 
showing that the food-getting has been limited largely to this eireum- 
seribed area. 

Another thing which it seems difficult to explain in connection with 
the incident first noted is the fact that if gophers do travel about at 
night they are certainly successful in finding their burrows again or 
in digging new ones before daylight appears. Their digging ability 
would seem to be sufficiently effective to afford them plenty of chance 
to conceal themselves again. Still, it is the common belief of observers 
that each one of the complex systems of tunnels constitutes the per- 
manent home of an individual gopher. 

If it be true that pocket gophers forage regularly above ground, 
we have a partial explanation of how it became possible for gophers 
to be entrapped in the oil. But why there should have been so many 
on one particular night, or why they were apparently all traveling 
in the same direction, remains unexplained. Such a migration seems 
the more remarkable in that pocket gophers are supposed to lead a 
solitary life. Bailey (loc. cit.) states that from the time the young are 
half grown and big enough to start burrows of their own each indi- 
vidual lives entirely alone, except during the short mating season in 
early spring. If each of the gophers caught in the oil had been living 
by itself, it seems remarkable that so many should have come out of 
their burrows on the same night and should have moved in the same 
direction. 

The data at hand will not allow of the conclusion that this migra- 
tion, if it can be called such, is exactly comparable with the sporadic 
migrations of lemmings, or of meadow mice. It is certain, however, 
that from some unknown eause large numbers of gophers left their 
burrows on the night of May 1, 1913, in North Berkeley. That they 
were either engaged in foraging or were seeking to improve their food 
supply, when entrapped in the oil, seems probable. 

The accompanying photograph (fig. 1) shows four half-grown 
pocket gophers, as seen by the writer on June 1, 1913, caught in oil 
which had seeped down into the gutter in the same locality where 
Mr. Light’s observations were made. One was still breathing when 
found. A meadow mouse (Microtus californicus) and three other 
gophers were similarly entrapped a few yards away the same night. 

The above incident further suggests crude oil as a possible method 
of getting rid of gophers. If so many gophers were entrapped in a 
strip of oil two feet wide in Berkeley, why would not a strip the same 


28 University of California Publications in Zoology [ Vou. 12 


Figure 1 


Photograph of four pocket gophers (Lhomomys bottae bottae) as found 
entrapped in erude oil which had seeped into the gutter from a newly made 
street in North Berkeley, California. One gopher was still breathing when 
discovered. Although firm earth extended within six inches of the animal, 
escape from the sticky oil had proven impossible. 


width prevent gophers from migrating into an orange orchard? And 
carrying the idea still farther, if a gopher may be entrapped in oil 
on top of the ground, why may it not be entrapped in the same sub- 
stance if it is placed in the burrow? The experiment should be tried. 
It might prove effectual in an attempt to rid new land of these pests, 
even if it does not prove practicable on a small seale. 

One fact, at least, would appear to minimize the value of oil for 
this purpose. Two days after application the surface of the oil hardens 
and small animals are able to cross without danger. On the other 
hand, its cheapness helps to balance this disadvantage. A thing to be 
further considered is the undesirable effect of erude oil on the soil. 

The effectiveness of crude oil, or asphaltum, as a trap for animals 
is demonstrated by the recent findings of great masses of vertebrate 


1913] Bryant: Wanderings of Pocket Gopher 29 


remains in the asphalt beds of Rancho La Brea, near Los Angeles, 
California. In these remarkable beds the bones of numerous pre- 
historic as well as present-day animals, including rodents, are found 
accumulated in masses from twenty to thirty feet deep (see Merriam, 
1911, pp. 199-213). 


Transmitted September 5, 1913. 


LITERATURE CITED 


BAILEY, V. 
1895. The pocket gophers of the United States. U. S. Dept. Agric., Div. 
Biol. Surv. Bull., 5, 1-47, 1 map, 6 figs. in text. 
MERRIAM, J. C. 
1911. The fauna of Rancho La Brea. Part I. Oceurrence. Mem. Univ. 
Calif., 1, 199-213, pls. 19-23, 1 fig. in text. 
Seton, E. T. 
1909. Life-histories of northern animals (Chas. Seribner’s Sons, N. Y.), 2 
vols., xxx + 1267, 46 pls., 38 maps, 182 figs. in text. 


Vol, 7. 


Vol. 8. 


Vol, 9. 


UNIVERSITY OF CALIFORNIA PUBLICATIONS— (Continued) 


(Contributions from the Museum of Vertebrate Zoology.) 

1, Two NéW Owls from Arizona, with Description of the Juvenal Plum- 
age of Strix occidentalis occidentalis (Xantus), by Harry S. Swarth. 
PtSi ay, pL OOS eo | a ES Se Re a 

. Birds and Mammals of the 1909 Alexander Alaska Expedition, by 
Harry S. Swarth. Pp. 9-172; plates 1-6; 3 text-figures. January, 1911. 

3. An Apparent Hybrid in the Genus Dendroica, by Walter P. Taylor. 

BOF ETS T0755 Fb ODEUBLYy 1 GLU as ve eS 5 

4, The Linnet of the Hawaiian Islands: a Problem in Speciation, by 

Joseph Grinnell, Pp. 179-195. February, 1910 o.oo. ecccece cece 

5, The Modesto Song Sparrow, by Joseph Grinnell. Pp. 197-199, Feb- 

pC 1s: oes fe Ral aes malin ome A LG Ge a Se ak op a Be SOR Ca at 

6. Two New Species of Marmots from Northwestern America, by H. S. 

Swarth.. Pp. 201-2045 February; 1912s ee 

7. Mammals of the Alexander Nevada Expedition of 1909, by Walter P. 

8 

9 


to 


Taylor. Pp, 205-307. =dune, LON to ee ne a ee 
~ Description of a New Spotted Towhee from the Great Basin, by J. 
Gtinnell,Pp..309-311,.Ausust, 1914 fo nn ee Se 
. Description of a New Hairy Woodpecker from Southeastern Alaska, by 
H. S. Swarth. Pp. 313-318. -October, 1911 200.020 


10. Field Notes on Amphibians, Reptiles and Birds of Northern Humboldt 


County, Nevada, with a Discussion of Some of the Faunal Features 

of the Region, by Walter P. Taylor. Pp, 319-486, plates 7-12. 

aad} Phe (Ch oa pte ks Gini origi oe, eae BARR ea Ae hs Beale Pei tT OR Mey hee Nes ARON br 
Index, pp. 437-446. 


1. The Vertical Distribution of Hucalanus elongatus in the San Diego 
Region during 1909, by Calvin O. Esterly. Pp. 1-7. May, 1911 ........ 
2. New and Rare Fishes from Southern California, by Edwin Chapin 
Starks and William M, Mann. Pp. 9-19, 2 text-figures. July, 1911, 
3. Classification and Vertical Distribution of the Chaetognatha of the San 
Diego Region, Including Redescriptions of Some Doubtful Species of 
the Group, by Ellis i. Michael, Pp. 21-186 pls, 1-8. December, 1911. 
4, Dinoflagellata of the San Diego Region, IV. The Genus Gonyaulax, with 
Notes on Its Skeletal Morphology and a Discussion of Its Generic 
and Specific Characters, by Charles Atwood Kofoid. - Pp, 187-286, 
plates 9-17, 
5. On the Sixeletal Morphology of Gonyaular catenata (Levander), by 
Charles Atwood Kofoid. Fp. 287-294 plate 18, 
6. Dinoflagellata of the San Diego Region, V. On Spiraulax, a New Genus 
of the Peridinida, by Charles Atwood Kofoid.. Pp. 295-300, plate 19, 
Nos. 4, 5, and 6 in one cover. September, 1911 .....:2.0..20 
7. Notes on Some Cephalopods in the Collection of the University of Cali- 
fornia, by S. S. Berry. Pp. 301-310, plates 20-21. September, 1911. 
8; On a Self-Closing Plankton Net for Horizontal Towing, by Charles 
Atwood Kofoid. Pp, 311-348, plates 22-25. 
9, On an Improved Form of Self-closing Water-bucket for Plankton In- 
vestigations, by Charles Atwood Kofoid. -Pp, 349-352. 
Nos. 8 and 9 in one cover. November, 1911] 20020. cetsci elec a 
Index, pp. 353-357. 
1, The Horned Lizards of California and Nevada of the Genera Phryno- 
soma and Anota, by Harold C, Bryant, Pp. 1-84, plates 1-9. Decem- 
POON) Peed ai) SUAS Cate Mcp iste Rakin Wh Ben one EC RRNA Hentai OPN OTS 
2, On a Lymphoid Structure Lying Over the Myelencephalon of Lepisos- 
teus, by Asa C. Chandler. Pp. 85-104, plates 10-12. December, 1911. 
3. Studies on Early Stages of Development in Rats and Mice, No. 3, by 
EB, L, Mark and J. A. Long.. The Living Eggs of Rats and Mice with 
a Description of Apparatus for Obtaining and Observing Them (Pre- 
liminary paper), by J. A, Long. Pp. 105-136, plates 13-17. February, 
AS 5 Saag —-Re Ce RUSE ap rs es DE Prk ip a hE gr Sat eS 
4, The Marine Biological Station of San Diego, Its History, Present Con- 
ditions, Achievements, and Aims, by Wm. E. Ritter, Pp. 137-248, 
3 Diates 18-24 and: 2'mans.s Marehy TOU2 sree a ac castasueee 
5. Oxygen and Polarity in Tubularia, by Harry Beal Torrey. Py. 249- 
251. May, 1912 . 
6. The Occurrence and “Vertical Distribution of the Copepoda ‘of ‘the ‘San 
Diego Region,.with particular reference to Nineteen Species, by Cal- 
vin O: Esterly. Pp. 253-340, 7 text-figures. July, 1912 20. 
7. Observations on the Suckling Period in the Guinea-Pig, by J. Marion 
Read. Pp. 341-351. September, 1912 . 


. 8. Haeckel’s Sethocephalus euceeryphalus (Radiolaria), a “Marine ‘Ciliate, 


by Charles Atwood Kofoid. Pp. 353-357.. September, 1912 -............ 
Index, pp. 359-365. 


1.75 


1.50 
10 


40 


Vol. 10. 


Vol, 11. 


Vol. 12. 


UNIVERSITY OF CALIFORNIA PUBLICATIONS— (Continued) 


(Contributions from the Museum of Vertebrate Zoology.) 

1,, Report on a Collection of Birds and Mammals from Vancouver Island, 
by Harry S. Swarth. Pp. 1-124, plates 1-4. February, 1912.20.00... 

2, A New Cony from the Vicinity of Mount Whitney, by Joseph Grinnell, 
Pp. 125-129, January, 1912 

. The Mole of Southern California, by J: Grinneli and H. 8. Swarth. 

Pp. 131-136, 2 text-figures. 

4, Myotis orinomus Elliott, a Bat New to Callfornia, by J. Grinnell and 
H. S. Swarth. Pp. 137-142, 2 text-figures. 

Nos, 3 and 4 in-one cover. April, 1912 2.00.2. cl cect eeneeeenrcences 


5, The Bighorn of the Sierra Nevada, by Joseph Grinnell.- Pp. 148-153, : 


4 text-fisuress; May, LOS i AS Oe en Pee eee ee : 


6. A New Perognathus from the San Joaquin Valley, California, by 
Walter P. Taylor.- Pp. 155-166, 1 text-figure. 

7. The Beaver of West Central California, by Walter P. Taylor. ~ Pp. 
167-169. 4 


Nos. 6 and 7 in one cover. May; 1912 -...c0....ccecil ce eecsdecceeeneecten serene é 


8. The Two Pocket Gophers of the Region Contiguous to the Lower Colo- 
rado River, in California and Arizona, by Joseph Grinnell. Pp, -171- 
1785. JUNO; LOD ee ee a a ee eee 

9. The Species of the Mammalian Genus Sorex of West-Central Cali- 


fornia, with a note on the Vertebrate Palustrine Faunas of the — 


Region, by Joseph Grinnell. Pp. 179-195, figs. 1-6. . March, 1913 ..... ~ 
10. An Account*of the Birds and Mammals of the San Jacinto Area. of 


Southern California, with Remarks Upon the Behavior of Geographic 


Races on the Margins of Their Habitats, by J. Grinnell and H. 8. 
Swarth. Pp. 197-406, pls, 6-10. October, 1913 2.002.000 
Index, pp. 407-417. 


1. Birds'in Relation to a Grasshopper Outbreak in California, by Harold 
C. Bryant. Pp. 1-20. November, 1912 ....0.2...o. ee ecccce ected eden 
2. On the Structure and Relationships of Dinosphaera palustris (Lemm.), 
by Charles-Atwood Kofoid and Josephine Rigden Michener. Pp. 21- 
QSOS MD OCEMDOR NE OED ss hey Be reecn atta ak punansagenbaseascieseanaveaeeias 


3. A Study of Epithelioma Contagiosum of the Common Fowl, by 


Clifford D. Sweet. Pp. 29-51. January, 1913 0.22222. 
4, The Control of Pigment Formation in Amphibian Larvae, by Myrtle 
E. Johnson. Pp. 53-88; plate 1. March, 1913 22.2.0. ccecccsee cette 
5. Sagitta californica, n.sp., from the San Diego Region, including 
Remarks on Its Variation and-Distribution, by Ellis L. Michael. 
Pp. 89-126, plate 2> Dune; 1913 once ees nccecce een acersnnensennennneveceranuaneecsvee 
6. Pycnogonida from the Coast of California, with Description of Two 


New Species, by H. V. M. Hall. Pp. 127-142, plates 3-4. August, 1913, - 


7. Observations on Isolated Living Pigment Cells from the Larvae of 
Amphibians: by S. J. Holmes.» Pp, 143-154, plates 5-6. 


8. Behavior of Ectodermic Epithelium of Tadpoles when Cultivated in 


Plasma, by S. J. Holmes. Pp. 155-172, plates 7-8. 
Nos. 7 and 8 in one cover.. September, 1913 2.2..0..00.-c tice tence 


1. A Study of a Collection of Geese of the Branta Canadensis Group from 
the San Joaquin Valley, California, by Harry S. Swarth. Pp. 1-24, 
plates 1-2; 8 text figs. -November,-1913 2. oar cS a ena 

2. Nocturnal Wanderings of the California Pocket Gopher, by Harold ©. 
Bryant. Pp. 25-29, 1 text fig. November, 1913 20.0... etcceees 

3. The Reptiles of the San Jacinto Area of Southern California, by Sarah 
Rogers Atsatt.. Pp. 31-50. November, 1913 22... .cos.c pioneers 


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UNIVERSITY OF CALIFORNIA PUBLICATIONS 
IN 


ZOOLOGY 
Vol. 12, No. 3, pp. 31-50 A November 20, 1913 


THE REPTILES OF THE SAN JACINTO AREA 
OF SOUTHERN CALIFORNIA 


BY 


SARAH ROGERS ATSATT 


INTRODUCTION 


During the summer of 1908 a collection of reptiles of the San 
Jacinto area was made by two parties sent out from the Museum of 
Vertebrate Zoology of the University of California. The material 
thus obtained, together with the field records of Messrs. Grinnell, 
Swarth, Taylor, Richardson, and Camp, has constituted the major por- 
tion of the basis of this report. Several specimens collected by Mr. 
Bridwell in July, 1912, are also in the Museum collection. In July, 
1912, the writer spent two weeks collecting in the San Gorgonio Pass 
and on the Pacific side of the mountain as far south as Strawberry 
Valley. The fresh material obtained then has been of use in deter- 
mining coloration, inasmuch as the material of 1908, preserved in both 
alcohol and formalin, is discolored in some eases. 

A full account of the localities is given in a report on the 
birds and mammals of the San Jacinto area in a_ preceding 
paper (Grinnell and Swarth, 1913). In brief the regions in which 
collecting was carried on are as follows: Whitewater, Snow Creek, 
Cabezon and Banning are in the San Gorgonio Pass, which extends 
nearly east and west from the desert toward the coastal valley, from 
Lower Sonoran to dilute Upper Sonoran at Banning. Directly south 
of Cabezon the ‘‘Hall Grade’’ ascends the ridge to Hurley Flat in 
the Upper Sonoran zone; above Banning is Poppet Flat, also Upper 
Sonoran. Higher on the mountain at about 5000 feet is Schain’s 
Ranch on the border line between Upper Sonoran and Transition. 


32 University of California Publications in Zoology [ Vou. 12 


Still higher and to the southeast is the region designated as Fuller’s 
Mill in true Transition country. A well-marked valley, Strawberry 
Valley, of low Transition character, is south of Fuller’s Mill and 
southeast of and heading up towards San Jacinto Peak. Tahquitz 
Valley, high up in the Transition zone, is directly south of the Peak. 
Southeast of the Peak is Round Valley, which marked the upper limit 
of observed reptiles. Kenworthy and Hemet Lake are near the head 
of the south fork of the San Jacinto River, south of the San Jacinto 
Peak and northwest of the Santa Rosa region. They are Upper 
Sonoran with oceasional Transition ‘‘islands.’’ West of them is 
Thomas Mountain. Eastward and on the desert side of the range 
are Pinon Flat and Asbestos Spring in Upper Sonoran. A little far- 
ther east is the extreme limit of the San Jacinto mountains, where 
Dos Palmos Spring, with Carrizo Creek flowing from it to the desert, 
and the neighboring parallel canon, Deep Canon, show the Lower Son- 
oran and desert aspects. From the ridge between Kenworthy and Dos 
Palmos heads Palm Canon, leading almost directly north and emptying 
out upon the desert around the corner from Whitewater. It is Lower 
Sonoran. South of Pinon Flat are the localities higher in the Santa 
Rosa region, namely, Garnet Queen Mine, Santa Rosa Mountain and 
Toro Peak. The two latter are in high Transition. On the western 
side from Hemet Lake the San Jacinto River road leads to Vallevista, 
an ‘“‘island’’ of Lower Sonoran. In 1912 the road from the town of 
San Jacinto to Beaumont, east of Banning, was covered also. After 
encircling the base of the hills for a few miles the road leads through 
Lamb Canon, passing from the hills covered with dead grass into 
thick chaparral at the head of the canon (2500 feet). 

The reptilian population of the San Jacinto area varies in abund- 
ance of species and individuals according to locality. The number of 
specimens recorded cannot indicate exactly the relative abundance 
beeause of the inequalities of collecting, both in time and weather 
conditions. Yet a general impression of population results which prob- 
ably approximates the correct relative abundance. 

Material collected in 1908, or in 1912 by Mr. Bridwell, is quoted 
under Museum of Vertebrate Zoology numbers; that collected in 1912 
by myself is indicated by numbers bearing an asterisk. 

To Professor Charles A. Kofoid, under whose direction the work 
has been done, and to Mr. Joseph Grinnell of the Museum of Verte- 
brate Zoology, the writer desires to express her indebtedness for their 
most valuable suggestions and criticism. 


1913 | Atsatt: Reptiles of the San Jacinto Area 33 


DISCUSSION OF SPECIES 


Callisaurus ventralis (Hallowell) 
Gridivon-tailed Lizard 

Distribution: Recorded as follows: Vallevista, 1800 feet, nos. 502— 
3; Cabezon. 1700 feet, nos. 176, 187; Whitewater, 1130 feet, no. 147, 
nos. *27-32; Dos Palmos Spring, 3000-3500 feet, nos. 242, 238-9, 
476-7, 479-482, 484, 561-2; Pinon Flat, 4000 feet, nos. 478, 483; Deep 
Canon, 3000 feet, no. 241; Palm Cafion, 2500 feet, no. 240. 

The gridiron-tailed lizard is generally a Lower Sonoran form, but 
on the desert side of the Santa Rosa mountains it ranges up into 
the Upper Sonoran area on Pinon Flat. Here, however, the pifion 
belt occurs on a ridge between two Lower Sonoran areas. The species 
was most common in Whitewater, Deep Cation and Dos Palmos, that 
is, In regions having access to the true desert. Van Denburgh (1897, 
p. 50) cites the occurrence of this form near Banning, although it 
was not found there either in 1908 or 1912. With the addition of 
the sandy wash of Vallevista as a new station to the previous records 
of Cajon Pass, San Bernardino County, and Oak Springs, San Diego 
County, three places are known where this lizard occurs west of the 
mountain range. In none of these places does the lizard seem to be 
abundant. From Oak Springs eight have been recorded, from Cajon 
Pass two, while at Vallevista not over half a dozen were seen. In all 
eases a fairly direct connection with the desert is possible, but these 
stations are now isolated. These western examples are identical with 
the eastern desert form. 

At Dos Palmos at half past six in the morning several of these 
lizards were kicked out of the sand of the trail by the horses’ hoofs. 
At Whitewater in 1912 an interesting color change was noted in an 
individual which was very light on the white sand but on coming under 
the shade of a scraggly bush developed a gray pattern. 


Crotaphytus collaris baileyi Stejneger 
Bailey Lizard 

Distribution: Snow Creek, 1500-2000 feet, nos. 210-1; ‘‘ Hall 
Grade,’’ 2000 feet, no. 1; Palm Canon, 800 feet, nos. 231, 243. 

This species is generally Upper Sonoran or in the Grayia belt of 
Lower Sonoran. In the San Jacinto area it is on the lower edge of 
Upper Sonoran and in Palm Cafion is found in Lower Sonoran. 

The lizards were found inhabiting both rocky and sandy spots. 


34 University of Califorma Publications in Zoology [ Vou. 12 


Crotaphytus wislizenii Baird and Girard 
Leopard Lizard 

Distribution: Vallevista, 1800 feet, no. 496; Cabezon, 1700 feet, nos. 
174-5, no. *6; Snow Creek, 1500 feet, no. 153; Dos Palmos Spring, 
3000 feet, no. 232. On Pinon Flat several were noted. 

This species usually oceurs in Lower Sonoran and arid Upper Son- 
oran. In the San Jacinto area it is found in Lower Sonoran and on 
the edge of Upper Sonoran. 

These lizards were found on the ground in the brush. Several 
were observed to be swift runners. 


Sauromelas ater Dumeril 
Chuck-a-walla 
Two typical individuals of large size were taken. At an altitude 
of 3500 feet near Dos Palmos Spring no. 560 was shot as it was 
sitting on the topmost ledge of a boulder pile. Several others were 
seen at a distance, usually with only their heads in sight. In their 
habits they were shy. 
At Snow Creek, 1500 feet, the other, no. 193, was found alive, 
caught in a trap in the morning. It offered to bite when approached 
and puffed up its body until its skin was taut. 


Uta mearnsi Stejneger 
Mearns Lizard 

Distribution: Banning, 2200 feet, nos. 87, 121-3, 149, 190; Cabezon, 
1700-2000 feet, nos. 4, 137, 178-9, 200, no. *11; Dos Palmos Spring, 
3500 feet, no. 571; Palm Cafion, 3000 feet, no. 234; Lower Palm 
Cafion, 800 feet, nos. 235-6. 

Previous records of this species are all from the eastern slope of 
the Coast Range of San Diego County near the Mexican boundary line. 
In the San Jacinto region it is found rather generally in the Lower 
Sonoran and dilute Upper Sonoran areas on the desert (eastern) side 
of the range and through San Gorgonio Pass. In coloration the most 
noticeable feature is the generally darker and more uniform appear- 
ance of our specimens as compared with those from the San Diego 
Range. 

Throughout all their range these lizards are dwellers on the rocks, 
usually on the vertical sides. At Palm Cation they are noted as being 
shy, darting about with great rapidity, scarcely trusting themselves 


1913] Atsatt: Reptiles of the San Jacinto Area 35 


in an exposed spot. They remained out of sight during the day, seem- 
ingly to avoid the direct hot sunlight, coming out after the sun was 
below the western wall of the canon and while the air and rocks were 
still hot. At Cabezon on the ‘‘Hall Grade’? many were seen which 
were rather easy of approach. 


Uta stansburiana Baird and Girard 
Brown-shouldered Lizard 


Distribution: The wide distribution and full representation of this 
species 1s shown in the following record: Vallevista, 1800 feet, no. 
505; Snow Creek, 1500-2000 feet, nos. 88-92, 150, 192, 218-9; nos. 
*13, 19-23; Cabezon, 1700-2000 feet, nos. 15-6, 169-70, 208, no. *8; 
Schain’s Ranch, 5000 feet, nos. 332-4, 338-40, 355-62, 366-7, nos. 
*35-7; Fuller’s Mill, 5900 feet, no. 310, no. *41; Strawberry Valley, 
6000 feet, nos. 525, 3802; Kenworthy, 4500 feet, no. 578; Dos Palmos 
Spring, 3500 feet, no. 368; Palm Canon, 3000 feet, no. 252. It was 
noted frequently from Banning to Strawberry Valley and from there 
towards Hemet; it also occurred in Deep Canon. 

This form was especially abundant at ‘‘Hall Grade,’’ Snow Creek, 
Banning, Schain’s Ranch, and Dos Palmos Spring. On the road 
between Schain’s and Fuller’s Mill in one morning about three dozen 
were observed in sunny places below the pine belt. The area of greatest 
population was between 4000 and 5000 feet. This species occurs in 
Upper and Lower Sonoran zones and in small numbers in the Tran- 
sition zone. 

The individuals present a wide range of variation in color and 
pattern which corresponds to Van Denburgh’s description (1897, p. 
67). 


Sceloporus biseriatus Hallowell 
Fenee Lizard 


Distribution: This common species was collected as follows: Valle- 
vista, 1700 feet, no. 506; Cabezon, 1700-2000 feet, nos. 13-4, 134-6, 
152, 201-5, no. *10; Banning, 2200 feet, nos. 93-4, 99, 116-20; 
Schain’s Ranch, 4000-5100 feet, nos. 290-4, 328-31, 337, 351-2, 363-5; 
Fuller’s Mill, 5300 feet, nos. 100-1, no. *42; Strawberry Valley, 6000 
feet, nos. 6-9, 520-1, 559, 585, 590-1; Hemet Lake, 4400 feet, nos. 
492, 494, 509-11; Kenworthy, 4500 feet, nos. 564-5; Thomas Mt., 
6800 feet, nos. 493, 512-3; Garnet Queen Mine, 6000 feet, nos. 522-3 ; 


36 University of California Publications in Zoology [ Vor. 12 


two were seen near Relief Hot Springs, 1500 feet, on the road from 
Hemet to Beaumont. 

Apparently the only regions where this species was not present 
are the highest peaks of the area and the desert region of the Santa 
Rosa mountains. At 5900 feet and above on the road from Schain’s 
to Fuller’s Mill this form seems to be displaced by Sceloporus graci- 
osus. 

At Cabezon on May 7, 1908, a female (no. 202) was taken which 
contained an egg, yellow in color and irregularly ovate in shape. <A 
large proportion of the specimens were found on rocks, but some were 
on trees or stumps in such higher localities as Schain’s and Straw- 
berry Valley. 


Sceloporus graciosus Baird and Girard 


Mountain Lizard 


Distribution: Above Sehain’s Ranch, 5300 feet, nos. 296, 301, nos. 
*39, 40, 43; Fuller’s Mill, 5850-7000 feet, nos. 103, 297-309; Straw- 
berry Valley, 6000 feet, no. 592; Tahquitz Valley, 8000 feet, no. 495; 
canon east of Round Valley, 8500 feet, no. 526; Thomas Mt., 6800 
feet, nos. 514-5; Santa Rosa Peak, 7500 feet, nos. 527-533. 

The range of this lizard is in the Transition zone; but it begins 
at the very lowest margin and also reaches the upper limit of this zone, 
as for instance in the neighborhood of Hidden Lake, east of Round 
Valley. Along Fuller’s Mill ridge this species replaces Sceloporus 
biseriatus as.the altitude increases, until Sceloporus graciosus reaches 
its maximum of population between 5800 and 6000 feet. In the Tran- 
sition zone over the higher parts of the Santa Rosa region from the 
peak to Toro this was the only reptile observed. Here it was abundant 
everywhere, about logs as well as rock piles. 

The coloration of these San Jacinto specimens is much grayer in 
tone than in those from Nevada and the Sierra Nevada, which are 
brownish. It is also very much darker than that specimen figured 
(as S. gracilis) by Girard (1858, pl. xx, fig. 1). The amount of blotch- 
ing in even the most pronounced ease is less than in the type and from 
this a series can be made with gradual decrease of blotching and 
lightness of longitudinal lines until there is little pattern visible. 

The adult lizards are found on rocks, pine or cedar trunks or 
stumps. The juvenals were found more often in the shade in dead 
erass. Around Fuller’s Mill they were not shy and were reported as 
even allowing themselves to be taken with the hand. At Santa Rosa 


1913 | Atsatt: Reptiles of the San Jacinto Area 37 


Peak, however, they were too lively to be noosed. The ones observed 
in the valley by Hidden Lake were surprisingly wild, even on a cold 
gray morning darting immediately under the rocks. 


Sceloporus magister Hallowell 
Rough-sealed Lizard 

Distribution: At Cabezon, 1700 feet, two adult males, nos. 180, 
188, were taken; at Dos Palmos Spring, 3000 feet, one adult and one 
juvenal, nos. 488, 490, were secured. One individual was seen at 
Snow Creek. 

In general coloration these forms are more grayish than material 
from other localities. The juvenal color is of a light yellowish ante- 
riorly and bluish posteriorly. 

This species is a typical desert form and is found under yucea 
or cactus plants. 


Sceloporus orcutti Stejneger 
Dusky Sealed Lizard 


Distribution: Snow Creek, 1500-2000 feet, nos. 80-3, 160-4, nos. 
*14-7, 24-6; Cabezon, 1700-2000 feet, nos. 8-11, 127-30, 171, 206, nos. 
*5. 9; Banning, 2200 feet, nos. 12, 113-5, 165, 189; Poppet Flat, 4100 
feet, no. 325; Schain’s Ranch, 4800-5100 feet, nos. 289, 324; Fuller’s 
Mill, 5900 feet, no. 288; Lamb Cafion, 2500 feet, no. *45; Hemet Lake, 
4400 feet, no. 508; Dos Palmos Spring, 3000-3500 feet, nos. 251, 489, 
570; Kenworthy, 4500 feet, nos. 566-9. 

Sceloporus orcutti is found in Lower and Upper Sonoran and in 
the Transition zones, but its center of population seems to be in 
Upper Sonoran. It migrates into the upper edge of Lower Sonoran 
and appears in Transition on the Fuller’s Mill ridge. It is associated 
with the Sonoran chaparral characterized by sagebrush, manzanita, 
and serub-oak. 

In the sealation of Sceloporus orcutti one notices that the eaudal 
seales are strongly keeled, although Van Denburgh (1897, p. 87) calls 
them nearly smooth. Strong keels were found in all the specimens of 
S. orcutti in the Museum. The dorsal seales are described as varying 
from no keels to very obtuse ones, with points scarcely protruding 
beyond the seale. The S. orcutti of the San Jacinto region all show 
obtuse keeling of the dorsals. In several the tendency to sharper 
keeling and longer points is very noticeable. 


38 University of Califorma Publications in Zoology [ Vou. 12 


The most characteristic lizard of the San Jacinto region and the 
one most noticeably missed as one crosses the San Gorgonio Pass and 
enters the San Bernardino region is Sceloporus orcutti. The 
erotesque large black males with their bull-dog-like pose, the gaudily 
colored males of medium size, the paler cross-barred females and 
juvenals are inseparably associated with the foot-hills and lower areas 
of San Jacinto. Their wildness or shyness seems to vary with localities. 
Generally in the late afternoon the males are very bold and will 
calmly await approach within a few feet. 


Phrynosoma blainvillei blainvillei Gray 
Blainville Horned Lizard 

Distribution: Vallevista, 1800 feet, no. 498; Cabezon, 1700-2000 
feet, nos. 146, 167, 184-6, 277; Banning, 2200 feet, nos. 76-7, 102, 212, 
348; Poppet Flat, 2800-4300 feet, nos. 345-7; Schain’s Ranch, 4900 
feet, nos. 272-5, 322-4, 336; Fuller’s Mill, 5900 feet, no. 276; Ken- 
worthy, 4500 feet, nos. 573-5; Vandeventer Flat, 4500 feet, nos. 576— 
7; Oak Spring, 4900 feet, no. 233. 

These forms as determined by Mr. Bryant are all typical P. b. blain- 
villei. Whereas they show well the color of the ground on which they 
lived, they show no variation as to altitude or slope of the mountain. 
For typical measurements see Bryant (1911, p. 37). 

This species occurs in Lower and Upper Sonoran and low Transi- 
tion zones, but the bulk of the population is in Upper Sonoran. 


Phrynosoma platyrhinos Girard 
Desert Horned Lizard 

At Whitewater, 1130 feet, a small specimen, no. * 33, was taken 
in 1912. It was on the white sand, not far from several individuals of 
Callisaurus ventralis. In coloration it is a light gray above, marked 
with greenish gray spots. On either side of the neck the dark spots 
show plainly. Ventrally it is pure white except for a few black flecks 
in the post-anal region, on one leg, and on the tip of the tail. 


Gerrhonotus scincicauda ignavus Van Denburgh 
San Diegan Alligator Lizard 
Distribution: Vallevista, 1800 feet, nos. 500-1; Babezon, 1700 feet, 
nos. 133, 181-3; Schain’s Ranch, 4900 feet, nos. 269, 319-21; 335; 
Strawberry Valley, 6000 feet, nos. 557-8; Kenworthy, 4500 feet, no. 
224; Garnet Queen Mine, 6000 feet, no. 524. 


1913 | Atsatt: Reptiles of the San Jacinto Arca 39 


All are typical, having keeled temporal, arm, and forearm scales 
and keeled ridges in the fifth or sixth seale from the mid-ventral 
row behind the anus. 

In characteristics observed these lizards were like the rest of the 
genus. They are fond of shelter, such as thick ferns along the bottom 
of a gulch, in a grapevine under a cottonwood tree, in oak brush, under 
a lilac bush, or in a rose tangle. Although usually slow of movement 
when under no fear, after they are captured they will fight, biting a 
stick and even themselves. Occasionally they climb into bushes in 
efforts to escape pursuit. 


Cnemidophorus stejnegeri Van Denburgh 
Stejneger Whip-tailed Lizard 

Distribution: Snow Creek, 1500 feet, nos. 84-6, 154, 156-9, 213-6, 
no. *18; Cabezon, 1700-2000 feet, nos. 5, 6, 1381-2, 148, 168, 172-3, 
191, 198, nos. *1-4, 7, 12; Banning, 2200 feet, nos. 7, 95-6, 155; Pop- 
pet Flat, 3700-4500 feet, nos. 268, 349-50, nos. *34, 38; Hemet Lake, 
4400 feet, no. 516; Vallevista, 1800 feet, no. 497; Asbestos Spring, 
3500 feet, no. 247; Dos Palmos Spring, 3500 feet, nos. 230, 246, 369, 
485-7; Palm Canon, 800 feet, no. 245. At Deep Canon a few individ- 
uals were noted. This species ranges in Upper and Lower Sonoran. 

One of the distinctions between C. figris and C. stejnegeri is based 
on sealation, on the relative size of dorsal granules and the relative 
size of central gular and collar seales. Since these appear to vary 
with age, comparison has to be made on specimens accurately agreeing 
in size, as there is no structural criterion of age or relative develop- 
ment. Under such circumstances within any given group of material 
of either species one finds variation in relative proportions—such wide 
variations as to make the specific determination extremely difficult. 
The other distinction is based on the gray suffusion and the clearness 
of pattern on sides of head, neek, and gular region. C. tigris from 
Nevada in the Museum collection presents both distinet and indistinet 
markings on the sides of the head, while the specimens from Mecea, 
California, have distinet markmes and those from Colorado River 
have generally indistinct markings. C. stejnegert from the Pacific 
slope has almost no gray suffusion, but specimens from regions bor- 
dering on the desert show some gray suffusion. No. *18 taken at 
Whitewater in 1912 has the flatter and smaller gular seales charac- 
teristic of C. tigris, but it has also the slight gray suffusion, distinct 
black markings on the sides of the head, and very evident cross-band 


40 University of California Publications in Zoology [ Vor. 12 


gular spots of C. stejnegeri. A general survey of material of both 
eroups raises the question of possible intergradation. A study of 
the habits and reaction to environment of each of these species ought 
to be carried out before the question can be satisfactorily settled. In 
a general survey of masses of material one feels that the effect of 
the environment upon the individual has been great. 

This species is a ground-loving form. At Cabezon it was noted 
frequently on the mesa among the cactus; at Snow Creek amid the 
leaves under the cottonwoods or among the rocks. It seems to be able 
to run through grass no matter how close set. At Banning and Dos 
Palmos Spring several were caught during the day in unbaited mouse 
traps in the sandy wash or in the brush. At Cabezon several were 
caught in traps baited with oatmeal. The same was true in previous 
collecting at Mecea, California, of lizards of this genus. Since unbaited 
traps, as well as baited ones, seem to be effective, it is most probable 
that the lizards were not eating oatmeal. Perhaps the odor of the 
dried blood or flesh of animals previously caught in those traps 
attracts either the lizards or the insects upon which the lizards feed, 
or perhaps the novelty of the trap in the neighborhood attracts the 
lizards. In escaping pursuit at Cabezon these lizards ran along the 
ground and burrowed in the soft sand or crawled under cactus. At 
Snow Creek they took refuge between or under the many rocks with- 
out running along on the ground for any distance. 


Verticaria hyperythra beldingi Stejneger 
Belding Orange-throat Lizard 

At Vallevista, 1800 feet, in a stubble field was taken no. 504. It 
is small in size, 9.6 mm. in length, of which 6.6 mm. is tail. The tail 
has the characteristic blue color of juvenals. On the outskirts of 
the town of San Jacinto on the road to Beaumont, 1500 feet, no. *44 
was found rustling among dead cottonwood leaves. Another was seen 
but not captured. 

At Reche Canon, near Colton, during the summer of 1908 several 
were collected in such habitats as a sandy wash, on a hill sparsely 
covered with vegetation, and in the dust by the roadside. Reeord 
was made of two found in the act of copulation on July 22. 


Eumeces skiltonianus (Baird and Girard) 
Western Skink 
One specimen of this species, no. 295, was secured several miles 
above Schain’s Ranch, 5300 feet. It is small, probably a juvenal, 


1913 J Atsatt: Reptiles of the San Jacinto Area 41 


with one-third of its tail regenerated. As is characteristic, it was 
rusthng among dry leaves in a shady place. Another specimen, no. 
3803, was taken at Strawberry Valley, 5500 feet, in 1912. 


Lichanura roseofusca Cope 
California Boa 

Distribution: Cabezon, 1700 feet, no. 207; Palm Canon, 3000 feet, 
no. 223. Both specimens are deep drab and no. 223 shows three red- 
dish longitudinal bands. 

So far as observed in this region this snake is an inhabitant of 
the Lower Sonoran zone only. 

No. 207 was found trying to swallow one of the large Peromyscus 
which had been caught in mouse traps. It had wrapped its body 
about the mammal whose head it had taken into its mouth. When 
disturbed it crawled slowly away and upon being gently struck on 
the head, coiled up in a ball, completely obscuring its head. 


Diadophis amabilis Baird and Girard 
Western Ring-necked Snake 
A specimen of this snake, no. 3801, was collected at Strawberry 
Valley, 5500 feet, in July, 1912. It was crossing the road when eap- 
tured, in the afternoon. 


Lampropeltis boylei Baird and Girard 
Boyle Milk Snake 

Distribution: Cabezon, 1700 feet, no. 143; Banning, 2200 feet, nos. 
97, 221; Hemet Lake, 4400 feet, no. 507; and in 1912 seen at Cabezon 
and on the ‘‘Hall Grade.”’ 

The dorsal scales are arranged in longitudinal rows numbering as 
follows for the whole length between the head and anus: nos. 97, 221, 
507 all have 23-21-19; no. 173 has 23-21. 

This snake is associated with the chaparral of Lower and Upper 
Sonoran. 


Lampropeltis pyrrhomelaena multicincta (Yarrow) 
Coral King Snake 
Distribution: Strawberry Valley, 6000 feet, no. 729. In this 
specimen the black rings are frequently more or less joined on the 
mid-dorsal area. The snout, labials, and two-thirds of the parietals 


42 University of Califorma Publications in Zoology [ Vou. 12 


are black, the cross-bar of black touches the parietals, the yellow 
cross-bars number thirty-six. The dorsal scale rows are 21-19-17. 
This species is limited to the Transition zone. 


Rhinocheilus lecontei Baird and Girard 
Long-nosed Snake 

Distribution: Cabezon, 1700 feet, no. 144; Dos Palmos Spring, 
3500 feet, no. 228. 

No. 144 is typical in the red coloration with much black and white 
spotting. It has 207 gastrosteges and 36 single urosteges followed by 
2 sets of alternately divided and single and by 13 divided ones. Dor- 
sal scale rows are 23-21-19. No. 228 is large in size and of dark 
brown and white coloration with no red and no markings on the 
sides between the brown patches. It has 214 gastrosteges and 42 
urosteges followed by 10 double ones. Dorsal scale rows are 25—23— 
21-19. > 

This species seems to be restricted to Lower Sonoran. 


Hypsiglena ochrorhynchus Cope 
Spotted Night Snake 
While this form was not found in 1908 or 1912, it is reported (Van 
Denburgh, 1897, p. 180) from Strawberry Valley, 5000 feet, and 
from San Jacinto. 


Salvadora grahamiae Baird and Girard 
Patched-nosed Snake 
Distribution: Banning, 2200 feet, no. 145. This specimen is rather 
dark with broad dark bands. Dorsal scale rows are 17-15-13. 
From previous records this species seems associated with arid 
areas in Lower and possibly Upper Sonoran zones. 


Bascanion flagellum frenatum Stejneger 
Red Racer 

Distribution: Cabezon, 1700 feet, nos. 2, 3; Palm Canon, 800 feet, 
no. 237. Another was also seen at Cabezon and another seen with 
no. 237 near the stream which comes from Murray Canon into Palm 
Canon. 

Dorsal seale rows of nos. 2, 3 are 17—-15-13-12 and in the ease 
of no. 237, 17—15-13-12-11. 

The red racer is a desert snake. 


1913] Atsatt: Reptiles of the San Jacinto Area 43 


Bascanion laterale (Hallowell) 
California Racer 

Distribution: Cabezon, 1800 feet, no. 166; Banning, 2200 feet, 
no. 220; Schain’s Ranch, 4900 feet, nos. 315, 316; Strawberry Valley, 
6000 feet, no. 554; Kenworthy, 4500 feet, nos. 227, 572. 

Dorsal scale rows number 17—16—15-14-13. 

This species ranges in Lower and Upper Sonoran and possibly a 
little in low Transition. It is west of the desert region in which B. f. 
frenatum oceurs. 

No. 315 was captured in a grassy meadow; no. 316 was caught 
in a mouse trap. 


Arizona elegans Kennicott 
Faded Snake 
None was taken in 1908 or 1912, but it is previously recorded at 
San Jacinto by Van Denburgh (1897, p. 194). 


Pituophis catenifer (Blainville) 
Western Gopher Snake 

Distribution: Schain’s Ranch, 4900 feet, nos. 104, 343; Strawberry 
Valley, 5500-6300 feet, nos. 551-3. Two small individuals were also 
seen in 1912 on the road from Schain’s Ranch to Strawberry Valley 
and on the road from Strawberry Valley to Hemet. 

The specimens are all typical with four prefrontals. Dorsal scale 
rows are 29-31 to 23. 

As collected in the San Jacinto area this species is limited to 
Upper Sonoran and Transition, but from other reeords it is shown 
to occur in Lower Sonoran. 


Thamnophis hammondi (Kennicott) 
California Garter Snake 

Distribution: Cabezon, 1700 feet, no. 138; Tahquitz Valley, 8000 
feet, no. 555; Kenworthy, 4500 feet, no. 226; Palm Canon, 800 feet, 
no. 244. 

Dorsal seales rows are as follows: 21-19-17. 

These loealities show this species extending from Lower Sonoran 
to high Transition. 


44 University of California Publications in Zoology [ Vor. 12 


Crotalus lucifer Baird and Girard 
Pacifie Rattlesnake 

Distribution: Banning, 2200 feet, nos. 112, 126; Schain’s Ranch, 
4900 feet, nos. 270-1, 317-8, 342 (heads only of 270-1, 318); Straw- 
berry Valley, 6000 feet, nos. 518, 556, 580; Tahquitz Valley, 8000 feet, 
no. 579; Santa Rosa Peak, 7000 feet, no. 517; Thomas Mt., 6800 feet, 
no. 581; Vallevista, 1800 feet, no. 519. 

The length of no. 579, including the rattles, was forty-five inches. 

This species ranges in both Sonoran zones and throughout the 
Transition. The alimentary tract of no. 579 contained four undigested 
half-grown ground squirrels. No. 318 contained an adult Neotoma. 


Crotalus cerastes Hallowell 
Sidewinder 
While none was taken or seen, at Whitewater in 1912 reliable 
information was given that sidewinders were frequent in that region. 


Crotalus mitchelli Cope 
Bleached Rattlesnake 

Distribution: Poppet Flat, 3300 feet, no. 341; Asbestos Spring, 
4500 feet, no. 222. The length of no. 222 was given as thirty-five inches 
and the girth as five inches. 

In the San Jacinto region this species was found on the lower 
limits of Upper Sonoran. In its general range, however, it is associated 
with desert conditions. 


Crotalus ruber (Cope) 
Red Rattlesnake 

Distribution: Cabezon, 1700 feet, no. 139; Dos Palmos Spring, 
3900 feet, nos. 225, 582-3. Reports were received in 1912 of red 
rattlers being killed at Cabezon and Hurley Flat above Cabezon. 

No. 225 measured forty-three inches without the rattles. 

This species apparently inhabits the Lower Sonoran zone and the 
lower margin of Upper Sonoran. 

No. 582 was reported as being extremely active when found at day- 
light in the morning of August 23, 1908, at Dos Palmos Spring. 
Evidently it was not chilled by the night temperature. Another very 
active and demonstrative rattler was seen, but it retreated into a eleft 
in the rocks, from which it could not be dislodged. 


1913] Atsatt: Reptiles of the San Jacinto Area 45 


CONCLUSIONS 


In the study of this collection of reptiles from the San Jacinto 
area I find a fairly definite relation of their distribution to hfe zones. 
In the Transition zone, which is the highest one in which reptiles were 
found, both the species and individuals are few in number. The 
Transition zone in general runs along the tops of the ridges, being 
superseded by the Boreal only on the highest peaks. Parts of the 
Transition are isolated from each other and still more so from the 
Transition areas of other mountains of southern California. How- 
ever, the two species Sceloporus graciosus and Lampropeltis pyrrho- 
melaena multicincta, which are characteristic of the Transition zone 
only, are found also in the Transition area of the San Bernardino 
mountains about twenty miles to the north across a depression of 5000 
feet between the two zones. 

The Sonoran zones, both Upper and Lower, encircle this area. On 
the northeastern and eastern sides of San Jacinto Peak the Upper 
Sonoran is very restricted in area because of the great steepness of 
the ridges, but on the western and the southern sides it spreads out 
to the coastal valley and the Santa Rosa mountains. It really forms 
the bulk of the actual area covered in collecting and it may be because 
of that reason that the number of individuals seems largest in Upper 
Sonoran. The Lower Sonoran occurs mainly pushing in from the 
desert through the canons on the eastern side and running up through 
San Gorgonio Pass, but appears also in “‘‘islands’’ on the western 
base of the San Jacinto area. In the Lower Sonoran we find the 
createst representation of species and a large number of individuals. 

The relative distribution in these zones is detailed in Table I. 
Where gaps resulting from insufficient representation in the collec- 
tion have occurred I have filled in with the distribution as recorded 
in the entire range of the various species by other writers, principally 
Van Denburgh (1897). Species occurring occasionally, but probably 
not in any considerable numbers, are placed in parentheses. 

There are two marked faunas in the San Jacinto area, one in 
extremely arid country, known as the Colorado desert fauna, and the 
other in a semi-humid region, known as a part of the Pacifie coast 
fauna, namely, the San Diegan fauna. While, roughly speaking, 
one of the faunas lies to the east and north of, and the other to the 
west and south of the main ridge of peaks, there are certain localities, 


[ Vou. 12 


y of California Publications in Zoology 


niversity 


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] 


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46 


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snsoreis sniodopaag 

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1913 | Atsatt: Reptiles of the San Jacinto Area 47 


especially through the San Gorgonio Pass and at the opposite end of 
the area around Dos Palmos, where the two come together. The inter- 
relations of these faunas in the cases of birds and mammals and some 
of the laws governing behavior of races on the margins of two faunas 
have been discussed in a preceding paper by Grinnell and Swarth 
(Is), 

In a tabulation of the reptiles of the San Jacinto area in regard 
to faunal distribution I find somewhat the same groups. (1) Some 
species occur in both faunas, (2) some occur in one or the other fauna 
only, and (3) some occur mainly in one fauna but invade the other 
faunal area. In this third group the species invading one fauna 
from the other fall into two sections. Im one section (a) the species 
invade the other faunal area for a short distance only, extending 
about to Cabezon from either end of the San Gorgonio Pass. The 
other section (b) contains species which invade for longer distances 
and at both ends of the area, passing from either direction through 
the entire length of San Gorgonio Pass and also pushing in from 
either direction in the region around Dos Palmos. The larger part 
of the invasion in this third group occurs in the San Diegan forms, 
which go over the ridge and range down upon the desert side. The 
invasion of all the species is along continuous associations of vegeta- 
tion, but the two desert forms, Callisaurus ventralis and Crotaphytus 
wislizenu, occur also in the isolated association or desert ‘‘island’’ at 
Vallevista. Table II details the faunal distribution of species. 

The representation in the two faunas by a species in one and a 
species, sub-species, or race, in the other does not attract one’s atten- 
tion in considering the reptiles of this area except in two cases. 
Sceloporus magister and Sceloporus orcutti are closely related species, 
the one in the Colorado and the other in the San Diegan fauna. 
Around Snow Creek and Dos Palmos the two inhabit the same geo- 
graphical area but maintain here their specific identity. The expla- 
nation of this would seem to me to rest in the fact that they inhabit 
different associations of vegetation. However, the Sceloporus orcutti 
of the San Jacinto Region in general shows a greater degree of keeling 
on the dorsal seales, thereby approaching S. magister, than does S. 
orcutti of other regions. The second ease is that of Cnemidophorus 
tigris and C. stejnegeri. C. tigris is reported from Palm Springs 
(Stone, 1911, p. 231) and oceurs eastward on the desert. C. stejnegeri 
was found on the western side and through the Pass to Snow Creek. 
The ranges of the two species probably meet near Snow Creek. At 


+ 


co 


University of California Publications in Zoology [ Von. 12 


Snow Creek were captured specimens referred by me to C. stejnegeri 
but which varied httle from C. tigris. It seems probable to me that in 
these two species we may have an illustration of the development 
through the modifying influences of light, temperature, and humidity 
of characteristics which have been described in specimens from widely 
separated areas and given the standing of specific characters. Where, 
however, these two have come into a common environment these specific 
differences have become less pronounced. 


TABLE II 
1, Species occurring in both Colorado and San Diegan faunas. 


Uta stansburiana Salvadora grahamiae 
Basecanion flagellum frenatum (?) Rhinocheilus leeontei 


2. Species occurring in one fauna and not invading the other: 


Colorado Fauna San Diegan Fauna 
Sauromelas ater Verticaria hyperythra beldingi 
Sceloporus magister Eumeces skiltonianus 
Phrynosoma platyrhinos Pituophis ecatenifer 
Crotalus cerastes Diadophis amabilis 


3. Species passing from one fauna into the other: 


From Colorado into San Diegan From San Diegan into Colorado 
(a. Invading San Gorgonio Pass a short distance) : 
Callisaurus ventralis Phrynosoma blainvillei blainvillei 
Crotaphytus collaris baileyi Gerrhonotus scincicauda ignavus 
Crotaphytus wislizenii Lampropeltis boylei 
Uta mearnsi Bascanion laterale 


Crotalus lucifer 


(b. Invading entire length of San Gorgonio Pass and also around Dos Palmos): 
Crotalus mitchelli Sceloporus biseriatus 
Sceloporus oreutti 
Cnemidophorus stejnegeri 
Lichanura roseofusea 
Thamnophis hammondi 
Crotalus ruber 


The San Jacinto mountains are separated from the San Bernar- 
dino mountains merely by a narrow pass of Lower Sonoran character, 
from three to five miles in width. Most of the San Jacinto and San 
Bernardino species, as might be expected, are identical, yet we find 
in the San Jacintos as characteristic of that region and not appearing 
in the San Bernardinos (Grinnell, 1908, pp. 160-170) the following 


1913] Atsatt: Reptiles of the San Jacinto Arca 49 


species, all of which range in Lower Sonoran: Uta mearnsi, Sceloporus 
orcuth, Verticaria hyperythra beldings and Crotalus ruber. A report 
has been made by Dr. Van Denburgh (1912, p. 149) of S. orcutti 
from material collected in 1887 or 1889 in Waterman’s Canon, San 
Bernardino mountains. Strictly speaking, on the basis of this record, 
S. orcutti would be excluded from the above list, but inasmuch as no 
collection of S. orcutti since 1889 in the San Bernardino mountains 
has been recorded, this species does not appear to be an inhabitant 
of that area at the present day. Certain of the desert forms of the 
San Jacinto region, namely, Callisaurus ventralis, Sauromelas ater, 
Rhinocheilus lecontei, and Salvadora grahamiae were not found in the 
San Bernardino area, but it is entirely possible that they may still 
be in that part of the San Bernardinos of desert character, which is 
just across the pass from Whitewater. In the San Bernardino moun- 
tains at higher levels were found forms not yet known in the San 
Jacintos, namely, Thamnophis elegans and Eumeces (sp. ?). This 
skink was recorded as Ewmeces gilberti (Grinnell, 1908, p. 163). Re- 
examination makes the previous determination questionable, but the 
true status of the species remains to be worked out. 

In a field observation of the reptiles of San Jacinto one can not 
help noticing certain general features in coloration. On the glaring 
white sand of the desert the extremely light background color of the 
reptiles and the main dorsal patterns of black and shades of gray 
and brown give a high degree of protective coloration to such species 
as Callisaurus ventralis, Phrynosoma platyrhinos and Crotalus cerastes. 
Associated with the dryness and intense light of the desert environ- 
ment seems to be the sharp contrast of extremes of light and dark 
in the color patterns of the reptiles. With the colors of the yellow, 
buff and brown sands of the higher levels and of the Pacific area the 
ground colors of Phrynosoma blainvillet blainvillei, Cnemidophorus 
stejnegert, Lampropeltis boylei and Crotalus ruber harmonize, while 
the patterns themselves have the darker shades of that group of 
colors. In the yet higher altitudes in the Transition zone, where the 
light is subdued by the large amount of shade and the dark colors of 
the foliage and where the air is more humid, the darkening of color 
and softening of pattern outlines is observed in Sceloporus biseriatus, 
Sceloporus graciosus and Crotalus lucifer. Of course in some localities 
there are exceptions to these general statements, both in individuals 
and species. Again, in Sceloporus orcutti the females and juvenals in 
their browns and greens are effectively concealed against the trunk 


50 University of California Publications in Zoology [ Vou. 12 


of a tree, yet the large black males on the light rocks are exceedingly 
conspicuous. On the whole, however, the general background colors 
of the country seems to be reflected in the general coloration of the 
reptiles. 


Zoological Laboratory, Mills College, California. 


Transmitted March 7, 1918. 


LITERATURE CITED 


Bryant, H. C. 

1911. The horned lizards of California and Nevada of the genera Phry- 
nosoma and Anota. Univ. Calif. Publ. Zool., 9, 1-84, pls. 1-9, 6 figs. 
in text. 

GIRARD, C. 
1858. United States exploring expedition, Herpetology. xvii + 496, 32 pls. 
GRINNELL, J. 

1908. The biota of the San Bernardino mountains. Univ. Calif. Publ. Zool., 

5, 1-170, pls. 1-24. 
GRINNELL, J., and Swart, H. 8S. 

1913. An account of the birds and mammals of the San Jacinto area of 
Southern California, with remarks upon the behavior of geographic 
races on the margins of their habitats. Univ. Calif. Publ. Zool., 
10, 197-406, pls. 6-10. 

STONE, WITMER. 

1911. On some collections of reptiles and batrachians from the Western 

United States. Proc. Acad. Nat. Sci., Philadelphia, 63, 222-232. 
VAN DENBURGH, J. 

1897. The reptiles of the Pacifie coast and Great Basin. Occ. Papers Calif. 
Acad. Sci., 5, 1-236, 67 figs. in text. 

1912. Notes on a collection of reptiles from Southern California and 
Arizona. Proe. Calif. Acad. Sei., (4), 3, 147-154. 


Vol. 7. 


Vol. 8. 


a Vol. 9. 


UNIVERSITY OF CALIFORNIA PUBLICATIONS— (Continued) 


(Contributions from the Museum of Vertebrate Zoology.) 
1, Two New Owls from Arizona, with Description of the Juvenal Plum- 
age of Strix occidentalis occidentalis (Xantus), by Harry S. Swarth. 
Bos Ske iam 0 ag ILS Oa Ganka a aC pe Me OO a tim Saeco eR Iu SEN 
2. Birds and Mammals of the 1909 Alexander Alaska Expedition, by 
Harry S. Swarth. Pp. 9-172; plates 1-6; 3 text-figures, January, 1911. 
3. An Apparent Hybrid in the Genus Dendroica, by Walter P. Taylor. 
PY: SL CSL Tne EF CDRUAYY AGI. cael spo  Sngehs eer es Pn 
4. The Linnet of the-Hawaiian Islands: a Problem in Speciation, by 
Joseph Grinnell, Pp. 179-195, February, 1911 ce. cee lee 
5. The Modesto Song Sparrow, by Joseph Grinnell. Pp. 197-199. Feb- 
TUALLY; TD Ele aos. Suede ee aero acne Roe Nia er ne ant 
6. Two New Species of Marmots from Northwestern America, by H.-S. 
Swarth. Pp; 201-204;. February, 1911-0 ee ee 
7. Mammals of the Alexander Neyada Expedition of 1909, by Walter P: 
Taylor. Pp. 205-307. June, 1911 
8. Description of a New Spotted Towhee from the Great Basin, by J. 
Grinnell.-- Pp) 309-311; = Augsast, 19112 eS 
9, Description of a New Hairy Woodpecker from Southeastern Alaska, by: 
H.-S. Swarth, Pp. 313-318. October, 191%. 2.20202 
10, Field Notes on Amphibians, Reptiles and Birds of Northern Humboldt 
County, Nevada, with a Discussion of Some. of the Faunal Features 
of the Region, by Walter P. Taylor. Pp. 319-436, plates 7-12. 
BODIUAEY; OM nee er a RE eee ee 
Index; pp. 437-446. 


1. The Vertical Distribution of Hucalanus elongatus in the San Diego 
Region during 1909, by. Calvin O. Esterly.. Pp, 1-7. May, 1911 ........ 
2. New and Rare Fishes from Southern California, by Edwin Chapin 
Starks and William M. Mann. Pp. 9-19, 2 text-figures. July, 1911. 
8. Classification and Vertical Distribution of the Chaetognatha of the San 
Diego Region, Including Redescriptions of Some Doubtful Species of 
the Group, by Ellis i. Michael. Pp. 21-186 pls. 1-8. December, 1911. 
4, Dinoflagellata of the San Diego Region, IV. The Genus Gonyaulax, with 
Notes on Its Skeletal Morphology and a Discussion of Its Generic 
and SpecificCharacters, by Charles Atwood Kofoid, Pp, 187-286, 
plates 9-17. ‘ 
5, On the Skeletal Morphology of Gonyaulax catenata (Levander), by 
Charles Atwood Kofoid. Fp. 287-294 plate 18. 
Dinoflagellata of the San Diego Region, V. Cn Spiraulax, a New Genus 
of the Peridinida, by Charles Atwood Kofoid. Pp. 295-300, plate 19, 
Nos..4, 5, and 6 in one cover, September, 1911 2.2.2... coe eeteeeee 
Notes on Some Cephalopods in the Collection of the University of Cali- 
fornia, by 8S. S, Berry. Pp. 301-310, plates 20-21. September, 1911, 
On a Self-Closing Plankton Net for Horizontal Towing, by Charles 
Atwood Kofoid. Pp. 311-348, plates 22-25. 
On an Improved Form of Self-closing Water-bucket for Plankton In- 
vestigations, by Charles Atwood Kofoid.. Pp. 349-352. 
Nos. 8 and 9 in one Cover.. November, 1911 22.2.2. sce cece cece eee 
Index, pp. 353-357. 


The Horned Lizards of California and Nevada of the Genera Phryno- 
soma and Anota, by. Harold C. Bryant. Pp. 1-84, plates 1-9. Decem- 
1,5 ApS IS) AS RS SNRs ce Sees pete EU SAN PERC AM ster AL EUR ily ER eR RE AR ate ME AOS 

On a Lymphoid Structure Lying Over the Myelencephalon of Lepisos- 
teus, by Asa C. Chandler. Pp. 85-104, plates 10-12. December, 1911. 

Studies.on Early Stages of Development in Rats and Mice, No, 3, by 
E. L. Mark and J, A. Long. The Living Eggs of Rats and Mice with 
a Description of Apparatus for Obtaining and Observing Them (Pre- 
liminary paper), by J. A. Long. Pp. 105-136, plates 13-17. February, 
BT Ie ep ee ec tama a ae ate Pas a nee eee GURNEE ap PEERS Ap sn Oi aeas nen Se Sea aT 

4, The Marine Biological Station of San Diego, Its History, Present Con- 

ditions, Achievements, and Aims, by Wm. E. Ritter, Pp. 137-248, 
plates 18-24,-and 2. maps. March, 1912 -o2c os asec Rt wa teeee 

5, Oxygen and Polarity in Tubularia, by Harry Beal Torrey. Pp. 249- 

STS PEERS Us 02: SAN A 2 ora ad os aa A ei UE ee oe ep en aE ant eR re eatin 

6. The Occurrence and Vertical Distribution of the Copepoda of the San 

Diego Region, with particular reference to Nineteen Species, by Cal- 
vin O, Esterly.. Pp. 253-340, 7 text-figures. July, 1912 2.00000... 

7. Observations on the Suckling Period in the Guinea-Pig, by J. Marion 

~ Read. Pp. 341-351. September, 1912 

Haeckel’s Sethocephalus eucecryphalus (Radiolaria), a Marine Ciliate, 
by Charles Atwood Kofoid. Pp, 353-357. September, 1912 -....... 

Index, pp. 359-365. 


bo 


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54 


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1.75 


1.50 
10 


40 


70 
25 


Vol. 10. 


Vol. 11. 


Vol. 12. 


UNIVERSITY OF CALIFORNIA PUBLICATIONS—(Continued) 


(Contributions from the Museum of Vertebrate Zoology.) 

1, Report on a Collection of Birds and Mammals from Vancouver Island, 
by Harry S. Swarth. Pp, 1-124, plates 1-4. February, 1912 2.0.2... 

2,-A New Cony from the Vicinity of Mount Whitney, by Joseph Grinnell. 
E5120 3297s Sanary yc ALD seo Soe Sa era eee ee 


3. The Mole of Southern California, by J. Grinnell and H. S. Swarth, 


Pp, 131-136, 2 text-figures, 
4, Myotis orinomus Elliott, a Bat New to California, by J. Grinnell and 
H. S. Swarth. Pp. 137-142, 2 text-figures. 
Nos. 3 and 4 in one cover. April; 1912 22.20... ectecceeeccene 
5. The Bighorn of the Sierra Nevada, by Joseph Grinnell. Pp, 148-153, 
4 text-fipures: “May, 1902) oc as aie aR hock ceca ee ecleestece 
6. A New Perognathus from. the San Joaquin Valley, California, by 
Walter P. Taylor. .Pp. 155-166, 1 text-figure. 
7. The Beaver of West Central California, by Walter P. Taylor. -Pp. 
167-169, 
Nos. 6 and 7 in one cover. May, 1912 .2..2..0i ich besten 
8. The Two Pocket Gophers of the Region Contiguous to the Lower Colo- 
rado River, in California and Arizona, by Joseph Grinnell. Pp. 171- 
TO SANG TOD eee ENE a REN ek re aes ete Se 
9. The Species of the Mammalian Genus Sorex of West-Central Cali- 


fornia, with a note on the Vertebrate Palustrine Fatnas of the’ 


Region, by Joseph Grinnell. Pp. 179-195, figs. 1-6. March, 1913 ........ 

10. An Account of the Birds and Mammals of the San Jacinto Area of 

Southern California, with Remarks Upon the Behavior of Geographic 

Races on the Margins of Their Habitats, by J. Grinnell and H. S$, 

Swarth. Pp. 197-406, pls. 6-10. October, DOES eS ee ey 
Index, pp. 407-417. . 


1. Birds in Relation to a Grasshopper Outbreak in California, by Harold 
C-Bryant.” Pp. 1-20... November, 1912-3 eo 
2. On the Structure and Relationships of Dinosphaera palustris (Lemm.), 
by Charles Atwood Kofoid and Josephine Rigden Michener. Pp. 21- 
We =-“December- TOV se AME Se hae ace cascthe ene nce een ren 
8. A Study of Epithelioma Contagiosum of the Common Fowl, “by 
Clifford D. Sweet. Pp. 29-51. January, 1913 ....022-2.2 en 
4, The Control of Pigment Formation in Amphibian Larvae, by Myrtle 
E. Johnson... Pp. 53-88, plate 1, March, 1913 20... 0c2. cc 
5. Sagitta californica, n.sp., from the San Diego Region, including 


Remarks on Its Variation and Distribution, by Ellis L. Michael. 


Pp. 89-126; plate Qi Pune AIS ea aon SS he es 
6. Pycnogonida from the Coast of California, with Description of Two 
New Species, by H. V. M. Hall. Pp. 127-142, plates 3-4. August, 1913, 
7. Observations on Isolated Living Pigment Cells from the Larvae - of 
Amphibians by S. J. Holmes. Pp. 148-154, plates 5-6. 
8. Behavior of Ectodermic Epithelium of Tadpoles when Oultivated in 
Plasma, by S. J. Holmes. Pp. 155-172, plates 7-8. 
Nos. 7 and 8 in one cover.. September, 1913 ..-0..0i. cn esceeeteceecenee 


1. A Study of a Collection of Geese of the Branta Canadensis Group from 
the San Joaquin Valley, California, by Harry S, Swarth. Pp. 1-24, 
plates. 1-2, °8-text figs). “November, 1918.20 ee ek 

2. Nocturnal Wanderings. of the California Pocket Gopher, by Harold C. 
Bryant. Pp. 25-29, 1 text fig. November, 1913 2002. 

3. The Reptiles of the San Jacinto Area of Southern California, by Sarah 
Rogers Atsatt. Pp. 31-50. November, 1913 22.52.00 .cl lee 


1,00 . 


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UNIVERSITY OF. CALIFORNIA PUBLICATIONS 
IN 


ZOOLOGY 
Vol. 12, No. 4, pp. 51-294, pls. 3-13, 9 text figs. March 20, 1914 


AN ACCOUNT OF THE MAMMALS AND BIRDS 


OF THE LOWER COLORADO VALLEY 


WITH ESPECIAL REFERENCE TO THE DISTRIBU- 
TIONAL PROBLEMS PRESENTED 


3 


BY 


JOSEPH GRINNELL 


UNIVERSITY OF CALIFORNIA PRESS 
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Vol. 7. - (Contributions from the Museum of Vertebrate Zoology.) 
1. Two New Owls from Arizona, with Description of the Juvenal Plum- — 
age of Strix occidentalis occidentalis: (Xantus), by Harry 8S. Swarth. : 
PD lB HAMA AOL OS Se ee Ae eee eave rar tan dar anes nalle aaogensa bin paysersiiv gene 10 
2. Birds and Mammals of the 1909 Alexander Alaska Expedition, by 
Harry S. Swarth. Pp. 9-172; plates 1-6; 3 text-figures. January, 1911. 1.50 
3. An Apparent Hybrid in the Genus Dendroica, by Walter P, Taylor. 
Pp. 173-177.. February, 1911 
4, The Litnet of the Hawaiian Islands: a Problem in Speciation, by 
Joseph Grinnell... Pp..179-195. February, 1911 -.-22.-......----.ssveceoeenoonnnve Aba VE 
5. The Modesto Song Sparrow, by Joseph Grinnell. Pp. 197-199. Feb- ee 
BT wate KE be Secs esse ian eRe pete Wipro, GWE spore ge eb Orbe Auman a OLE 0555 4 
6. Two New Species of Marmots from Northwestern America, by H. 8. 
Swarth: Pp.-201-204.° February, 19D) nn... esc. cece itcesciventenneseense 2 P05: 
7. Mammals of the Alexander Nevada Expedition of 1909, by Walter P. 
Taylor. “Pp.-205-307. Pune, TOU none coer rare onan era nt cerparees aa 
. Description of a New Spotted Towhee from the Great Basin, by J. 
Grinnell. Pp. 309-311. “August, 1911 —- 2... Bcd ecto Sentinal oat ieceas .05: 
. Description of a New Hairy Woodpecker from Southeastern Alaska, by 
H. S. Swarth. Pp. 313-318. October, 1911 -............. Bre teascep tau wadeoces ae OD 
10. Field Notes on Amphibians, Reptiles and Birds of Northern Humbold ; 
County, Nevada, with a Discussion of Some of the Faunal Features 
of the Region, by Walter P. Taylor. Pp. 319-436, plates 7-12. 
Pre Drita yyy 112s sae e ew Foi sadew cp ncaeeenatnccennncbenet apeutgatiecscbrcyetatnondgseoeneus=uassnte 1.00 
Index, pp. 437-446, 


Vol..8. 1. The Vertical Distribution of Zucalanus elongatus in the San Diego 

Region during 1909, by Calvin O. Esterly. Pp. 1-7... May, 1911 ........ 10 

2. New and Rare Fishes from Southern California, by Edwin Chapin rs 

Starke and William M. Mann. Pp. 9-19, 2 text-figures. July, 1911. .10 . 

. Classification and Vertical Distribution of the Chaetognatha of the San 

Diego Region, Including Redescriptions of Some Doubtful Species of 

the Group, by. Ellis I. Michael. Pp..21-186, pls. 1-8. December, 1911. 

4. Dinoflagellata of the San Diego Region, IV. The Genus Gonyaulaz, with 

Notes on Its Skeletal Morphology and a Discussion of Its Generic 

and Specific Characters, by Charles Atwood Kofoid. . Pp. 187-286, 
plates 9-17. : : 


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+ 


UNIVERSITY OF CALIFORNIA PUBLICATIONS 
IN 


ZOOLOGY 
Vol. 12, No. 4, pp. 51-294, pls. 3-13, 9 text figs. March 20, 1914 


AN ACCOUNT OF THE MAMMALS AND BIRDS 
OF THE LOWER COLORADO VALLEY 


WITH ESPECIAL REFERENCE TO THE DISTRIBUTIONAL 
PROBLEMS PRESENTED 


BY 
JOSEPH GRINNELL 


(Contribution from the Museum of Vertebrate Zoology of the University of California) 


CONTENTS 

PAGE 
USD E508 WY COLE) Se he ec eee Sees ie peceeel oe at a ae 51 
TAM OPAL se cevee os. oeasectaseaesecake a ead ead c eae p as Re ete dee Oo oe ae te ne dEath Unease 53 
IDQSERMOUNO IN. GE TNE) (ONSEN) TANASE ees cere este eco rec cee eo ter occ eeoae 57 
AT alan daenaniM als pOSIULONOL sth ere GV OM ences cere eee eee eee cee sen ere mnee eee eee 62 
Mhevassociamona Wear easy Of at DONre SLO Mase ere ac ee ae eee nte een nee cece a bene eee eee tense 66 
The Colorado River as a highway of dispersal and center of differentiation 

OEE (SYSRVOMOR od eae eee eC ie a ee 97 
The Colorado River as a hindrance to the dispersal of species sce (00) 
The problem of barriers with regard to birds and mammals ~.. Sell 
Gheckalistaotmuhem bind dieters eee ee kw nrc Ree py EN re 110 
General accounts of the birds: local distribution, migration, variation, molt, 

DRUM TOES eee cepae eee ceed eee ae scat nas n fas ee occg cto se vestdgscsuesdescun snes ddestetdeoescccSsoceseecots 113 
Check-listok qian im amma sear cease secs eee cece nce cencer =e 217 
General accounts of the mammals: local distribution, variation, habits .... 218 
VORA IZSY REN DIRS! COAT SO Le Pe RR PR 269 


INTRODUCTION 


The southeastern frontier of California lies in the heart of a vast 
desert region possessing a fauna and flora of notable variety and 
peculiar specialization of forms. Cutting through the center of this 
desert area flows the great Colorado River. Politically, this river 


52 University of California Publications in Zoology — [Vou. 12 


marks the southeastern boundary of California, comprising the interval 
between the Nevada line and the Mexican line, and separates our state 
from Arizona. Biogeographically, the Colorado River completely 
bisects the desert area through which it flows, the resultant two di- 
-visions with their xerophilous fauna and flora being separated, not 
only by the stream itself, but also by the riparian strips of more or 
less width which flank the river immediately on either side. 

Two objects were in mind to justify the selection of the Colorado 
valley for extended field-work in vertebrate zoology: (1) The literature 
pertaining to the birds and mammals of the region was fragmentary, 
relatively little work having been published since the early and incom- 
plete reports of Woodhouse (1853), Cooper (1861, 1868, 1869), and 
Coues (1866). An extended knowledge of the composition of the 
vertebrate fauna of the southeastern frontier of California was a 
desideratum. (2) The effect of the Colorado River, with its riparian 
strips sharply contrasted against the contiguous desert areas, upon the 
distribution of the animals concerned, promised to provide data of 
importance relative to the general problem of barriers. 

Miss Annie M. Alexander, founder of the California Museum of 
Vertebrate Zoology, not only warmly approved of the plan to under- 
take field-work along the lower Colorado River with the above objects 
in view, but generously provided the necessary cost of the expedition 
in addition to her regular appropriation for the support of the museum. 


The plan was carried out in the three months of 1910 from February 


15 to May 15, inclusive. The writer took part in the field work 
in person throughout the entire time, and was assisted by Mr. Frank 
Stephens, Mr. Joseph Dixon, and Mr. L. Hollister Jones. The party 
began work at Needles, and proceeded by boat from place to place 
down the river, the last station being on the California side below 
Yuma and close to the Mexican line. The locations of the various 
collecting stations established are given in the itinerary. 

The three months’ field-work yielded 1,272 specimens of mammals, 
1,374 birds, 443 reptiles and amphibians, 22 sets of birds’ eggs and 
nests, a few fishes, and a collection of the more conspicuous plants. 
All of this material now forms part of the collections of the Cal- 
fornia Museum of Vertebrate Zoology, except the plants, which are 
deposited in the Herbarium of the Department of Botany of the 
University of California, and the few fishes, which are added to the 
ichthyological collection in the Zoological Department of Leland Stan- 
ford Junior University. 


oO 
w 


1914] Grinnell: Mammals and Birds of the Colorado Valley 


The present paper concerns itself with the mammals and birds 
of the region, and is based upon the collections and field-notes obtained 
in 1910, as above enumerated. No attempt has been made as yet to 
work up the reptiles and amphibians, though these promise interesting 
results. The writer’s efforts have been concentrated upon the mam- 
mals and birds, with a view to establishing first of all the systematic 
status of the various included forms; in other words, to determine 
their relationships with similar species or races of the surrounding 
regions. Such determination has been performed in a manner wholly 
satisfactory to the writer in only a part of the critical cases. As must 
always obtain in work of this sort, lack of material in certain direc- 
tions often puts abrupt limitations upon investigation. 

The obvious principle has been followed, that no generalized treat- 
ment of a conerete subject like geographic distribution is justifiable 
upon any but the soundest basis of systematic analysis. 

Because of the close dependence of most mammals and birds upon 
plants, the latter have an important place in any treatment of animal 
distribution. The set of plants secured by the Colorado River Expedi- 
tion has been named by Professor H. M. Hall, of the University of 
California. Names so provided are used throughout the present paper, 
particularly in the discussion of associational areas. 

My sincere acknowledgements are hereby extended to Professor 
Charles H. Gilbert, Department of Zoology, Leland Stanford Junior 
University, and to Professor Charles A. Kofoid, Department of 
Zoology, University of California, for critical suggestions coneern- 
ing general considerations; to Professor H. M. Hall, of the University 
of California, for critical reading of the chapter on associations; 
and to Messrs. Harry S. Swarth and Walter P. Taylor, fellow staff- 
members in the California Museum of Vertebrate Zoology, for help 
from time to time on many points of detail. 


ITINERARY 


Our party assembled at the town of Needles, California, on the 
evening of February 14, 1910, and on the following day our first 
collecting station was established in the river bottom close by. 

A seow was constructed for the transportation of our rather bully 
outfit, while the skiff purchased, being a much readier means of loco- 
motion, enabled us to traverse the river expeditiously where need be. 
Tt was found practicable to divide the party during a portion of the 


54 University of California Publications in Zoology [ Vou. 12 


time, two of the men traveling separately with the skiff and a light 
outfit. All members of the party were working from the same base, 
except where indicated otherwise. The current was depended upon 
almost altogether to carry our boats from station to station. 

The following enumeration shows the location of the various sta- 
tions established from the initial point down the river, the names by 
which they are designated throughout this report, and the time spent 
at each. For their location reference should be made to the map 
(pl. 3). 

1. Needles, California side; camp in river bottom one-fourth mile 
east of the railroad station; February 15 to 18. 

2. Five miles below Needles, California side; camp in river bottom 
in an air line probably slightly less than four miles south-southeast 
of Needles; February 18 to 23. In this and some other cases, even 
where a government map was at hand, the exact location on the flood 
bottom was indeterminable on the map, because of shifting of the river 
channel; for it frequently happens that in a single period of high 
water the topography of the riparian area is profoundly metamor- 
phosed. 

3. Mellen, Arizona side; camp on narrow mud-bar within one-fourth 
mile north of the railroad station locally called Topock (=Mellen, on 
the Needles Special Map, U. S. G. 8.) ; February 23 to March 1. The 
Santa Fe railroad crosses the Colorado River at this point. 

4. Opposite The Needles, California side; camp on high bank about 
one mile northwest of ‘‘B. M. 465’’ of Needles Special Map, U.S. G.S.; 
March 1 to 8. ‘‘The Needles’’ are a group of sharp-peaked hills 
chiefly on the Arizona side of the river, and must be remembered as 
a totally distinct locality from ‘‘Needles,’’ the name of the railroad 
town fifteen miles or more to the northwest. 

5. Foot of The Needles, Arizona side; this station was worked from 
the camp on the opposite side of the river. Certain members of the 
party rowed back and forth morning and evening to tend trap lines; 
March 4 to 7. 

6. Chemehuevis Valley, California side; camp in river bottom at 
lower end of this valley and close to base of hills, probably near 
“BM. 438” of Parker quadrangle, U. 8. G. S.; March 8 to 11. With 
this and the succeeding two stations uncertainty exists as to exact 
location of our base camps. The maps we had at that time were 
inaccurate. The Parker quadrangle, U. S. G. S., was not issued until 
1911. 


1914] Grinnell: Mammals and Birds of the Colorado Valley 55 


7. Lower Chemehuevis Valley, California side; camp at edge of 
whirlpool and in mouth of wash emanating from low hills, probably 
about one mile west-northwest of ‘‘B. M. 418’’ of Parker quadrangle, 
U.S. G. S.; March 11 to 12. 

8. Above Bill Williams River, Arizona side; camp in river bottom, 
of Parker quad- 


” 


probably within two miles above “‘Steamboat Rock 
rangle, U. S. G. S., and hence about eleven miles northwest of the 
mouth of [Bill] Williams River; March 12 to 15. 

9. Parker, Arizona side; camp at the new Santa Fe bridge; this 
was merely an over-night stop, and very little collecting was done; 
Mareh 15 to 16. : 

10. Riverside Mountain, California side; camp at mouth of wash. 
close to “‘B. M. 405”? of Parker quadrangle, U.S. G. S.; March 16 to 22. 

11, 12. Above Blythe, California side. The party separated at 
Riverside Mountain, Stephens and Jones working alone slowly on the 
lookout for beaver sign and making at least two over-night camps on 
the way to Ehrenberg. One of these stops, and the one by Grinnell 
and Dixon, were located some distance apart, in the river bottom 
adjacent to the mesa on the California side, and within six or eight 
miles above Blythe. March 22 to 24. 

13. Ehrenberg, Arizona side; camp in river bottom within one- 
fourth mile below the town; March 24 to 30. 

14, 15, 16. Below Ehrenberg, Arizona side, one station, and near 
Palo Verde, California side, two stations. The party again separated, 
and base camps established at three different points in the river bot- 
tom, estimated to be from ten to twenty-five miles below Ehrenberg. 
The changed course of the river made dependence upon maps, impos- 
sible. March 28 to April 3. 

17, 18. Opposite Cibola, California side, two stations ; main eamp on 
high bank where river swung against mesa; somewhat above, that is, 
to the northwest of, rather than directly opposite, Cibola; Stephens 
and Jones worked from a second station about five miles below the 
main camp, April 5 to 6. March 31 to April 6. 

19. Ten miles below Cibola, Arizona side; camp on narrow terrace 
between river and mesa, about a mile below an adobe ruin; April 6 
to 10. 

20. Twenty miles above Picacho, California side; camp in river 
bottom opposite Lighthouse Rock, and about a mile below the Draper 
ranch. In an air line this station was probably somewhat less than 
twenty miles north of Picacho. April 10 to 17. 


University of California Publications in Zoology (Vou. 12 


Or 
lop) 


21. Hight miles east of Picacho, California side; camp in river 
bottom at lower end of ‘‘Charlie’s Valley, 
below Canebrake Canon. Because of the eastward swing of the river 
in the vicinity of Picacho, eight miles east of Picacho is also eight miles: 
below Picacho. April 17 to 21. 

22. Five miles above (or north of) Laguna, Arizona side; camp in 
river bottom near base of first hills above silted-in area; April 21 to 26. 

23. Four miles above (or north of ) Potholes, California side ; visited 
April 23, from our camp on the opposite side of the river. 

24. Potholes, California side; camp in river bottom just below the 
head-gate; April 26 to 29. 

25. Four miles below Potholes, California side; camp in river bot- 


”? 


which in turn is just 


tom; Grinnell and Dixon; April 29 to May 2. 

26. Five miles northeast of Yuma, California side; camp in river 
bottom; Stephens and Jones; April 30 to May 3. 

27. Five miles above (northeast of ) Yuma, California side ; camp in 
river bottom; Grinnell and Dixon; May 2 to 5. This station was but 
a half-mile or so from the preceding. 

28. Yuma, Arizona side; camp about half a mile up the Gila River 
from its mouth, on the shore towards Yuma and within two miles east 
of that town; Stephens and Jones; May 3 to 7. 

29. Near Pilot Knob, California side; camp on river bank at site 
of the old Hanlon Ranch, marked by a group of date palms. This is 
also the site of the ‘‘ American Girl Pump,’’ of the Yuma quadrangle, 
U.S. G.S.; but only rusting machinery and pipes, and a pile of ashes, 
marked the place at the time of our visit. Grinnell and Dixon; May 
5 to 15; Stephens and Jones, May 7 to 15. The season’s field-work 
was concluded on May 15. 

As will have been noted, our collecting stations were usually estab- 
lished on alternate sides of the river successively. It proved impos- 
sible to propel the heavy scow directly across because of the strong 
current. In two places, however, a limited amount of collecting was 
done on exactly opposite sides of the river, this beimg accomplished 
through the use of the skiff alone. 

From each of the base camps as a center, hunting and trapping 
were carried on as far radially as proved practicable. It was our 
effort to test every sort of ground available, that is, each association 
represented. Three of the four members in the party were constantly 
engaged in trapping for mammals. Trap lines were run in different 
directions, usually from the river’s edge back to the highest parts of 


1914 | Grinnell: Mammals and Birds of the Colorado Valley 57 


the desert within reach, generally not farther than two miles from 
camp. 

It was, of course, possible to carry a single day’s hunt much farther 
back on to the desert. A distance of from four to seven miles from 
the river was occasionally reached in day’s tramps, for instance, at 
Riverside Mountain, twenty miles above Picacho, and at Pilot Knob. 

In the nature of the case, our camps were always pitched on the 
river bank where convenient landings could be made and the boats 
tied up safely. As is to be expected, the writer now regrets that 
extended work was not done at certain points scarcely or not at all 
touched in our 1910 exploration. It would have been a desirable thing 
to work intensively on directly opposite sides of the river, where it 
cut through high ground, at three or more points; for example, at 
The Needles, below Cibola, and at the Laguna Dam. 

It must be remembered, however, that we had practically no 
information to start with, either as to the fauna or local topography 
along most of the route. We had it all to learn by our own efforts; 
and only as experience accumulated could the contrasting sets of facts 
come into relief, thereby directing our enquiries into more productive 
channels. 


DESCRIPTION OF THE COLORADO RIVER 


The portion of the Colorado River explored by the 1910 expedition 
is altogether below the famed Grand Canon. There is no true 
““eanon’’ below the Nevada line, the river flowing through its chan- 
nel in relatively sober fashion. Yuma is very close to 150 miles in 
a direct line due south of Needles. While following a general north- 
to-south course, the many deflections of the river, and especially its 
meandering through the big valleys, make the distance from Needles 
to Yuma by the way of the river approximate 285 miles. The altitude 
of the river at Needles is 460 feet, at Yuma, 125 feet, so that the 
average fall in this portion of the river is only a little more than 
one foot per mile. 

Two features of the Colorado River are extraordinary, when com- 
parison is made with other streams of the North American continent. 
(1) The amount of sediment always carried in suspension is very 
ereat, so that the flowing waters are quite opaque at all seasons of 


58 University of California Publications in Zoology [ Vou. 12 


the year. (2) There is an enormous inerease in the volume of the 
river at the annual period of high water (see Newell, 1904, pp. 123- 
161). 

The sediment load varies from one-tenth to two and one-tenth per 
cent, by weight, according to the rate of the current. The latter varies 
from a maximum of two and one-half miles per hour at low water 
up to even seven miles per hour at flood time in parts of the channel 
where conditions are most favorable. While it is probable that the 
Needles-to-Yuma section of the Colorado River has about reached 
grade, the extraordinary supply of sediment always fed from above 
and the ever fluctuating volume of water, result in a continued pro- 
cess of deposition and erosion, going on simultaneously in different 
parts of the river’s channel. At low water deposition exceeds erosion, 
but at flood time the increased transporting power of the river results 
in removal of large masses of soil from varying parts of the river 
bottom. 

The flow of the river varies from 4,000 to 100,000 eubie feet per 
second. The time of lowest water is in midwinter, that of highest 
flood, in June, at the time of melting snow among the sources of 
the Colorado River, in the Rocky Mountains. The period of high 
water is of short duration, about May 15 to July 1, while throughout 
the year fluctuations of less extent are lable to occur at any time. 

The river’s course is of two markedly different types: where it 
flows between rock walls among hills the channel is of fixed location 
and usually deep and narrow; in the large valleys the river may be 
broad and shallow, but at any rate of constantly shifting channel. At 
The Needles the river passes between high hills and the narrow chan- 
nel thus formed has two sharp bends in it. Such a gorge is locally 
called a ‘‘box canon,’’ and at certain stages of the river is not without 
danger to small boats because of whirlpools and jutting reefs of rock. 
At low water the voyager is astonished at the elevation of the highest 
water marks on the rock walls above the surface of the river at the 
low stage. 

There is good reason for believing that the river is now aggrading 
its channel considerably above the apex of the delta where, of course, 
this process is certainly in progress. At any rate, sedimentation has 
formed broad fiood-bottoms at intervals alone the river’s course. 
These, large or small, are very similar in physical and floral features. 

The most extensive one of these flood-bottoms is sometimes known 
as the ‘‘Great Valley of the Colorado.’’ The settlements of Blythe, 


1914 | Grinnell: Mammals and Birds of the Colorado Valley 59 


Ehrenberg, Palo Verde and Cibola are located in it, and considerable 
activity has of late been directed towards farming the rich bottom 
lands. But the capriciousness of the river has proven a hindrance 
to marked development along this line. 

Shortly below the point where the river emerges from its rock-con- 
fined channel into the broadening valley, meandering begins. The 
detours of the stream increase in extent towards the lower end of the 
valley, the channel swinging from side to side in great curves, marked 
by minor curves along these courses, so that a sort of periodicity of 
meandering is exhibited. The most notable phenomenon in this con- 
nection is the progressive movement of these loops down the valley. 
The result is that in a short period of years, the major portion of 
the river’s flood-bottom is worked over in the path of this irresistible 
and continual shifting of the channel. 

The effect on the flora is obvious. Only in curves of the valley 
sheltered by abutting hills are trees given a chance to reach advanced 
age. The only trees capable of thriving on the unstable portion of 
the flood-bottom are such as grow rapidly, willows and cottonwood. 
As the observer floats along the main channel, winding through the 
ereat valley, especially during rising water, the destructive process 
in question is clearly in evidence. On the outside of each curve the 
river rapidly undercuts the bank, dislodging great masses of the fine 
sedimentary soil, itself laid down but a few years previously. The 
overgrowth of comparatively aged willow and cottonwood topples 
into the stream and is swept away by the swift current. Hundreds 
of acres may be thus appropriated by the river in a few days and 
within a short distance. 

On the other hand, on the inside of each curve, where the current 
is slow, and especially during falling stages of the water, the river is 
rapidly depositing sediment, in other words building up its bank 
towards the general level of the bottom lands. At the annually 
recurring periods of high water, when the entire flood bottom is 
inundated, layers of silt are deposited over the whole, thus tending 
to establish a uniform level. 

The observer, from any appropriate hill-top overlooking the valley, 
ean readily discern the regularly graded heights of tree growth which 
mark the successive ages of the land on which they grow. The year- 
old seedlings but a few inches in height form a crescent-shaped belt 
along the inside of each curve of the river, facing down the valley. 
Paralleling this and next in position back from the river is dense 


60 University of California Publications in Zoology [ Vou, 12 


two-year-old growth, succeeding which is a stand of still older growth. 
Because of the progressive trend of the process it is as a rule the 
oldest growth which becomes subject to the razing action of the river 
as its loops travel down the valley. 

Occasionally cut-offs occur, thus interrupting the symmetry of 
behavior, and the river quickly establishes a new channel, portions of 
the old being left as lagoons, though these are usually short-lived 
because of the rapid sedimentation at recurring times of general over- 
flow. The bottom land immediately adjacent to the channel, where 
the latter has been fixed for some time, is usually higher than the 
lateral tracts. The depressions on one or the other side of the flood- 
bottom are generally drained at the lower end of the valley by a serics 
of sloughs emptying into the main channel just above the constriction 
of the valley where the hills converge. 

At high water these lateral depressions are submerged to a depth 
of as much as twelve feet, as shown by actual measurement of the upper 
limit of the mud marks on the tree trunks. Whatever the water 
touches is discolored by a coat of fine sediment, and one thereby gains 
at any time of the year an accurate estimate of the depth of inundation 
in any part of the flood bottom at the previous period of high water 
(see sectional profiles, figs. A and B). 

In some places the overflow depressions have no drainage outlets. 
After times of flood, the water in them disappears by evaporation, 
which proceeds at a very rapid rate in this excessively arid and hot 
region. Areas of alkaline deposits are left, often stretches of bare, 
baked hard-pan surrounded by concentric belts of halophilous vege- 
tation. 

At rather infrequent intervals along the large valleys the river, 
in swinging, cuts beyond its usual limit and assaults the desert mesa. 
The process tends to add to the area of the regular flood plain and 
conduces to the formation of lateral bluffs. It is noteworthy that 
this aggressive work of the river is much more conspicuous on *iie 
west side of each valley than on the east side. The law of westward 
cutting of north-and-south flowing streams in the northern hemis- 
phere, as brought about by the earth’s rotation, is thus clearly illus- 
trated in the Needles-to-Yuma section of the Colorado River. 

The local use of the terms ‘‘first bottom’’ and ‘‘second bottom’’ 
in the large valleys is worth adopting in general discussions relative 
to ranges of animals. The first bottom is the portion of a valley 
subject to regular overflow, and is clearly marked by the presence of 


1914] Grinnell: Mammals and Birds of the Colorado Valley 61 


willows and cottonwood; the second bottom is of sufficiently higher 
elevation to be chiefly above high-water mark, and is characterized 
by mesquite, salt-bush, and rank clumps of creosote bush. The second 
bottom may be altogether wanting, or it may constitute a broad bench- 
like tract. The surface is often modified by alluvial deposits at the 
mouths of washes leading down from the adjacent desert, and by 
wind-blown sands which heap up about bushes, especially along the 
southeastern borders of the valleys. 

The Laguna Dam has had a pronounced modifying influence on the 
flora and fauna of the vicinity. The dam was built to a height of 
twelve feet above the mean level of the river at that point at the 
time of beginning construction. As soon as it was completed (in 1909) 
the retarded waters above began to deposit silt, and by May, 1910, the 
valley above had been silted in to a depth determined by the top 
of the dam. The water level had been raised conspicuously for at 
least ten miles, and we saw evidences of deepening of the first bottom 
deposits and slowing of current for fully thirty miles, above the dam. 
The cottonwoods of the first bottom within eight miles above the 
dam had all been killed, evidently by the raising of the general surface 
around their trunks; and the mesquites and other vegetation of the 
second bottom had all been drowned out, there thus being no trace of 
second-bottom conditions except for dead stalks. These were replaced 
by vast mud flats growing up to arrowweed. All] this change, of course, 
involved the birds and mammals of the areas affected, in addition 
to the plant life. 

Below the dam reverse changes took place. The water, having 
dropped a considerable portion of its sediment above the dam because 
of the slowing of its current, was able to pick up sediment at a cor- 
respondingly accelerated rate below the dam. This, and the fact of 
a new cut-off having been found by the river in the delta in 1909, thus 
temporarily shortening its channel, resulted in a deepening of the 
channel seven feet below the previous level immediately below the 
dam. Thus the former flood-bottom was, in 1910, far above flood 
level, and in a way to become good second bottom, with appropriate 
metamorphosis in vegetation and fauna. 

Although these changes were local, and due to man’s interference, 
similar ones, due to natural causes, have doubtless occurred from time 
to time in various parts of its course in the river’s history, thus 
repeatedly shifting the riparian strips both in position and total width, 
with corresponding variability in the powers of the river at different 


62 University of California Publications in Zoology (Vou. 12 


points to act as a barrier to the dispersal of some animals and as a 
highway of dispersal for others. 

The effects of the extraordinary and continuous load of sediment 
of the Colorado River, together with the inconstaney of its channel, 
doubtless account directly or indirectly for many of the peculiarities 
in the fauna. As far as known to the writer, in the Needles-to-Yuma 
section of the river valley there are no aquatic molluses or decapod 
crustaceans, or tailed amphibians. Toads and frogs are present but 
not abundant. The fish fauna in the main stream is sparse in both 
species and individuals. 

Our party seined at three different points in the main stream. At 
two of these nothing was caught; in the third, a backwater slough on 
the Arizona side above Mellen, four sorts of fishes were taken, catfish 
(Ameiurus nebulosus), bony-tail (Gila elegans), hump-backed sucker 
(Xyrauchen cypho), and earp (Cyprinus carpio). A huge mmnow 


? 


(Ptychocheilus lucius), ealled locally ‘‘ Colorado salmon,’’ was caught 
with hook and line in back-water on the California side opposite 
Cibola, and was plentiful immediately below the Laguna dam, where 
many were being taken by the Indians living near there. In lateral 
sloughs and overflow depressions carp and eatfish were often observed 
in numbers, and in these relatively clear waters they were far more 
accessible to piscivorous animals than in the opaque water of the 
river itself. It may be said, in conclusion, that the portion of the 
Colorado River under consideration has relatively a very poor aquatic 
fauna and flora, and that this poverty has had its effect in limiting 
the oceurrence of vertebrate animals dependent upon such sources of 
food-supply. 


ZONAL AND FAUNAL POSITION OF THE REGION 


Two schools of faunistie students are represented among American 
zoogeographie writers of the present day. One, of which C. H. Merriam 
(1894, ete.) is the most prominent exponent, sees in temperature the 
chief controlling cause of distribution, and deals with the ranges of 
species in terms of ‘“‘life-zones.’’? The other school, of which C. C. 
Adams (1905, ete.), A. G. Ruthven (1907), and Spencer Trotter (1912) 
are active advocates, assigns to temperature but a minor role, looking 
rather to a composite control, of many factors, resulting in ecologic 
‘“assoeiations,’’ of which plants are essential elements, and which are 


1914] Grinnell: Mammals and Birds of the Colorado Valley 63 


to be further explained on historical grounds. The two sets of areas 
thus defined do not by any means correspond. Yet the reviewer 
cannot fail to note, here and there, places where boundaries coincide, 
and such coincidences are so frequent as to be suggestive of real con- 
cordance in some significant manner. Is it not probable that both 
schools are approximately correct, the difference in mode of treatment 
being due to different weights given the different kinds of evidence, 
or, in other words, to difference in perspective? The opportunity is 
here taken to attempt to bring into accord the systems of the two 
schools. 

The period of field study up to the present time devoted by the 
writer to the animal life of the climatically diversified state of Cali- 
fornia has led him to the recognition of three distinct orders of distri- 
butional behavior as regards terrestrial vertebrates. These are indi- 
cated in the terms: zonal, faunal, and associational. 

Every animal is believed to be limited in distribution zonally by 
greater or less degree of temperature, more particularly by that of 
the reproductive season (see Merriam, 1894). When a number of 
animals (always in company with many plants similarly restricted) 
approximately agree in such limitation, they are said to occupy the 
same life-zone. 

The observation of this category of distributional delimitation 
is particularly easy in an area of great altitudinal diversity like that 
comprised in the southwestern United States. The writer is led to 
wonder if those authors who minimize the importance of temperature 
have ever been privileged to travel, and carry on field studies, outside 
of the relatively uniform eastern half of North America! 

Study of any area which varies widely in altitude and hence 
provides readily appreciable differences in daily temperature from 
place to place brings conviction of the very great effectiveness of 
temperature in delimiting the ranges of nearly all species of animals 
as well as of plants. Particular attention may be ealled to the results 
of a biological survey of Mount Shasta (Merriam, 1899). 

But temperature is not to be considered the only delimiting factor 
of environment, though its possible overemphasis by the Merriam 
school seems to have led some other persons to believe that this view 
is held. In fact, it becomes evident after a consideration of appro- 
priate data that very many species are kept within geographic bounds 
in certain directions only by an increasing or decreasing degree of 
atmospheric humidity (see Grinnell and Swarth, 1913, p. 217). By 


64 University of California Publications in Zoology [| Vou. 12 


the plotting of the ranges of many animals as well as of plants, 
coincidence in this regard is found in so many eases as to warrant the 


29 


recognition of a number of ‘‘faunal areas,’’ on the causative basis of 
relative uniformity in humidity. It is probable that every species is 
affected by both orders of geographie control. 

As to which is the more important, assembled data seem to show 
that more genera and higher groups are delimited by zonal boundaries 
than by faunal boundaries (see Merriam, 1892, p. 49, ete.) The 
arresting power of temperature barriers would therefore seem to be 
relatively the greater. 

In the third category of distributional control there is a con- 
spicuous association of the majority of so-called adaptive structures of 
animals (often of high taxonomic value) with certain mechanical 
or physical features of their environment. An animal may thus 
intimately depend upon certain peculiarities, Inorganic or organic, or 
both, of a given area, and be unable to maintain existence beyond the 
limits of occurrence of those features of the environment. For instance, 
Dipodomys deserti is delimited by soil of certain texture and depth. 
Tracts of relatively uniform environmental condition, including their 
inanimate as well as living elements, are here called associations. 

The geographical distribution of any animal is correctly diagnosed 
in terms of each of the above three groupings. In other words, an 
animal belongs simultaneously to one or more zones, to one or more 
faunas, and to one or more associations. No one of these groupings 
can be stated in terms of the other, any more than a person can com- 
pute liquids by the peck, or weight in miles. The constituent species 
within each of these groupings always belong to the other two. To 
illustrate: the southern white-headed woodpecker inhabits the con- 
iferous forest association of the San Bernardino fauna of the Transi- 
tion zone; the Abert towhee belongs to the mesquite and quail-brush 
associations of the Colorado Desert fauna, of the Lower Sonoran zone ; 
the Pacific shrew belongs to the upland riparian association of the 
northern coast redwood fauna of the Transition and Boreal zones. 

Referring now to the region contiguous to the lower Colorado River, 
we have good reason, both biotic and meteorologic, for assigning it 
all to one zone, namely, the Lower Sonoran, and to one fauna, the 
Colorado Desert; but many associations are represented. In other 
words, the variation in altitude and latitude included is not great 
enough to bring sufficient modification of the characteristically high 
temperature to affect profoundly the distribution of the plant and 


1914] Grinnell: Mammals and Birds of the Colorado Valley ( 


animal life within the region. Nor is there marked variation in atmos- 
pheric humidity, the entire area being swept by air currents of pre- 
vailing dryness. To express the situation in a different way, zonal 
and faunal conditions are remarkably uniform; but associational con- 
ditions are varied, as pointed out in the succeeding chapter. 

The zonal diagnosis, Lower Sonoran, is based in part upon the 
presence of the following determinative genera: 


MAMMALS BREEDING BIRDS 
Ammospermophilus Melopelia 
Sigmodon Miecropallas 
Dipodomys Geocoecyx 
Perognathus Centurus 
Pipistrellus Calypte 
Maecrotus Pyrocephalus 


Guiraca 
Phainopepla 
Toxostoma 
Auriparus 
Polioptila 


The faunal diagnosis, Colorado Desert, is based in part upon the 
presence of the following subspecies and species, selected from the 
entire list as being particularly restricted : 


MAMMALS BREEDING BIRDS 
Odocoileus hemionus eremicus Lophortyx gambeli 
Citellus tereticaudus tereticaudus Otus asio gilmani 
Peromyscus eremicus eremicus Bubo virginianus pallescens 
Reithrodontomys megalotis deserti Dryobates sealaris cactophilus 
Neotoma albigula venusta Centurus uropygialis 
Thomomys albatus Colaptes chrysoides mearnsi 
Dipodomys deserti deserti Agelaius phoeniceus sonoriensis 
Perognathus spinatus spinatus Melospiza melodia saltonis 
Perognathus penicillatus penicillatus Pipilo aberti 
Perognathus intermedius Piranga rubra cooper 
Lepus californicus deserticola Vireo belli arizonae 
Felis oregonensis browni Vermivora luciae 
Canis ochropus estor Dendroica aestiva sonorana 
Vulpes macrotis arsipus Toxostoma crissale 
Mephitis estor Polioptila plumbea 


Procyon pallidus 
Myotis occultus 
Myotis ealifornieus pallidus 


The picked zonal and faunal ‘‘indiecators’’ just named are often 
found in the same association with other elements seemingly less 
sensitive to temperature and humidity. Some of the latter, however, 


66 University of California Publications in Zoology | Vou. 12 


may be present under duress, since their greatest abundance is known 
to be in adjacent zones or faunas. Thus Lynx and Eptesicus are much 
more plentiful in the Upper Sonoran zone or even in the Transition 
zone; and, on the other hand, Sigmodon and Pyrocephalus are genera 
of subtropical abundance. The presence of elements of the latter 
category was probably what led Merriam at one time (1894, p. 233, 
footnote, pl. 14) to refer the lower Colorado River Valley to the 
Tropical zone. All students who employ the life-zone system, now 
unhesitatingly agree in referring the area in question to the Lower 
Sonoran zone. 

The ‘‘western desert tract,’’ of Mearns (1897, pl. 2), is probably 
similar to the ‘‘Colorado Desert fauna’’ of the present paper. The 
former term is not considered apropos in the system adopted by the 
present writer on the ground that an inanimate area is designated 
thereby, rather than an assemblage of living things inhabiting the 


area. 


ASSOCIATIONAL AREAS OF THE REGION 


Since the entire region under consideration all belongs to one zone 
and all to one fauna, according to the definitions of these distributional 
terms given in the preceding chapter, the study of local distribution 
in the Colorado River region pertains chiefly to associations. Perhaps 
nowhere else in America can one find the degree of associational con- 
trast which is presented in the region under consideration. A stream 
of large volume, with paralleling strips of well-watered bottom land, 
maintains its course to the sea through what is considered the hottest 
and most arid desert in the world. 

There is nothing to show that the atmosphere is appreciably more 
humid in the vicinity of the bottom lands or the river itself than 
upon the open desert. The evaporated moisture is quickly dissipated ; 
that is, it becomes diluted to an imperceptible proportion in the desert 
air currents. It appears, therefore, that the great floral differences 
observed between the extreme associations are due primarily to differ- 
ence in amount of soil water available. There are, of course, such 
additional factors as varying alkalinity, and shade (see Spaulding, 
1909). 

It must be understood that the associations here defined are recog- 
nized by the writer primarily because of their service in the treatment 
of animal distribution. Botanists have found it useful to make much 


1914] Grinnell: Mammals and Birds of the Colorado Valley 67 


finer analysis (for example, see Spaulding, 1908). A more or less 
detailed description of the associations observed in our 1910 explora- 
tions is herewith offered, as a necessary prelude to further discussion. 

In the lists of species, grouped according to apparent associational 
preferences, all mammals found by our party are included, and all 
the birds treated in the ‘‘General Accounts’ 
which are of transient occurrence and uncertain forage-ground like 


b) 


except sixteen species, 


swallows, or else so rare as to make even approximate appraisement 
impossible. 

Qualifying terms are appended in each case: whether of exclusive 
(exel.) occurrence in the association under discussion, or of maximum 
(max.) abundance, or of minor (min.) abundance. Obviously a 
species of exclusive occurrence is entered in but one of the associational 
lists, while one marked max. in one list is to be found marked min. in 
one or more of the other lists. It is to be taken for granted that in 
species of easy locomotion, individuals may occur in transit across 
other associations than the one or ones in which it is characteristically 
present en masse. 

There is necessarily more or less uncertainty in many cases, and 
where the doubt is strong, owing to lack of knowledge, a question 
mark is added. With birds, seasonal occurrence is indicated by the 
terms: resident (present throughout the year), winter. (present 
throughout the winter, in certain cases including fall and spring also), 
summer (present during the breeding season, in certain cases including 
spring and fall also), transient (merely passing through during migra- 
tion). 


River Association 
BIRDS 


Gavia immer: exel.; winter 

Sterna forsteri: excl.; transient 

Phalacrocorax auritus albociliatus: 
excl.; winter and transient 

Pelecanus erythrorhynchos: exel.; win- 
ter and transient 

Mergus serrator: excl.; winter 

Anas platyrhynchos: exel.; winter 

Nettion carolinense: exel.; winter 

Querquedula cyanoptera: excl.; tran- 
sient 

Spatula clypeata: excl.; winter 

Dafila acuta: exel.; winter 

Marila affinis: exel.; winter 


Erismatura jamaicensis: exel.; winter 

Chen hyberboreus hyperboreus: exel.; 
winter 

Plegadis guarauna: exel.; transient 

Mycteria americana: exel.; summer 

Ardea herodias treganzai: exel.; res- 
ident 

Butorides virescens anthonyi: exel.; 
transient 

Nycticorax nycticorax naevius: excl.; 
resident 

Grus canadensis: excl.; winter (?) 
and transient 

Fulica americana: min.; transient 


68 University of California Publications in Zoology — [Vow 12 


Pisobia minutilla: exel.; winter Sayornis nigricans: exel.; winter 
Actitis macularius: exel.; winter Corvus corax sinuatus: max.; resident 
Oxyechus vociferus vociferus: min.; Petrochelidon lunifrons lunifrons: 
transient excl.; summer 
Cireus hudsonius: min.; winter Stelgidopteryx serripennis: min.; 
Pandion haliaétus carolinensis: exel.; summer 
transient Anthus rubescens: exel.; winter 


Ceryle aleyon: exel.; transient 


MAMMALS 


Castor canadensis frondator: excel. Procyon pallidus: max. 
Ondatra zibethica pallida: max. 


Remarks upon the River Association—For reasons already 
explained there is relatively little cryptogamic aquatic flora in the 
Colorado River. There is therefore little or no food-supply from this 
source to attract plant-eating ducks. This category of water-birds was, 
in fact, very sparsely represented. 

On the other hand, herons were notably plentiful because of the 
supply of catfish and carp made abundant at intervals by the drying-up 
of overflow ponds. While fishes were not abundant in the main stream, 
they were plentiful in backwater sloughs, where, too, the water was 
more nearly clear because the sediment had a chanee to settle out. 

The ornis of the river appeared to owe its proportionate consistency 
in large measure to the above two circumstances, namely, poverty in 
aquatic plant life, and sporadic abundance of certain fishes in the 
lateral sloughs (see p. 62). 

The single carnivorous mammal (Procyon) belonging chiefly to the 
river association was piscivorous in food habits, foraging along mud 
bars (see pl. 4, fig. 2) and at the margins of overflow ponds and 
sloughs, as do the herons. The two rodents of the river and larger 
paralleling sloughs, which are here included as part of the River 
Association, lived in banks immediately adjacent to the water. The 
beaver fed chiefly upon bark, twigs and foliage of such willows and 
cottonwoods as had fallen over into the water through being under- 
mined by the current. 


1914] Grinnell: Mammals and Birds of the Colorado Valley 69 


WILLow-CoTrronwoop ASsocIATION 


BIRDS 


Melopelia asiatica trudeaui: exel.; 
summer 

Aeccipiter velox: max.; winter 

Aceipiter cooperi: exel.; resident 

Buteo borealis calurus: min.; resident 

Faleo sparverius phalaena: min.; resi- 
dent 

Otus asio gilmani: max.; resident 

Bubo virginianus pallescens: min.; 


resident 

Dryobates sealaris cactophilus: max.; 
resident 

Sphyrapicus varius nuchalis: min.; 
winter 


Centurus uropygialis: min.; resident 

Colaptes cafer collaris: exel.; winter 

Colaptes chrysoides mearnsi: min.; 
resident 

Phalaenoptilus nuttalli nuttalli: max. ; 
winter (as a forager only) 

Phalaenoptilus nuttalli nitidus: max.; 
resident (?) (as a forager only) 

Chordeiles acutipennis texensis: min.; 
summer (as a forager only) 

Archilochus alexandri: max.; summer 

Tyrannus verticalis: exel.; transient 

Myiarchus cinerascens cinerascens: 
min.; transient 

Nuttallornis borealis: exel.;  tran- 
sient 

Myiochanes richardsoni richardsoni: 
max.; transient 

Empidonax difficilis difficilis: exel.; 


transient 

Empidonax trailli trailli: exel.; sum- 
mer 

Empidonax hammondi: exel.; tran- 
sient 


Empidonax wrighti: exel.; transient 

Empidonax griseus: max.; winter 

Pyrocephalus rubinus mexicanus: 
min.; resident 

Molothrus ater obseurus: max.; sum- 
mer 

Xanthocephalus xanthocephalus: 
min.; winter 

Agelaius phoeniceus sonoriensis: max. ; 
resident 


Teterus cuecullatus nelsoni: max.; 
summer 

Teterus bullocki: max.; summer 

Zonotrichia leucophrys gambeli: min. ; 
winter 

Spizella passerina arizonae: min.; 
transient (?) 

Melospiza melodia fallax: min.; win- 
ter 

Melospiza melodia saltonis: min.; 
resident 

Melospiza lineolni lineolni: min.; 
winter 

Pipilo aberti: min.; resident 

Oreospiza chlorura: max.; transient 

Zamelodia melanocephala melanoce- 
phala: max.; transient 

Guiraea caerulea lazula: min.; sum- 
mer 

Passerina amoena: excl.; transient 

Piranga ludoviciana: max.; transient 

Piranga rubra cooperi: excl.; summer 

Vireosylva gilva swainsoni: max.; 
transient 

Lanivireo solitarius cassini: max.; 
transient 

Vireo belli arizonae: max.; summer 

Vermivora ruficapilla gutturalis: 
excl.; transient 

Vermivora celata celata: max.; win- 


ter 

Vermivora celata lutescens: exel.; 
transient 

Dendroica aestiva sonorana: exel.; 
summer 

Dendroica aestiva brewsteri: exel.; 
transient 

Dendroica aestiva rubiginosa: exel.; 
transient 

Dendroica auduboni auduboni: max.; 
winter 


Dendroica nigrescens: max.; transient 
Dendroica townsendi: max.; tran- 


sient 

Dendroica occidentalis: exel.; tran- 
sient 

Geothlypis trichas seirpicola: min.; 
resident 


70 University of California Publications in Zoology — | Vou. 12 


Teteria virens longicauda: exel.; sum- 
mer 

Wilsonia pusilla pileolata: max.; 
transient 

Wilsonia pusilla chryseola: min.; 
transient 

Troglodytes aedon parkmani: exel.; 
winter 


Polioptila caerulea obscura: min.; 


winter 

Hylocichla ustulata ustulata: exel.; 
transient 

Hylocichla guttata guttata: exel.; 
winter 


Planesticus migratorius propinquus: 
max.; winter 


Regulus calendula cineraceus: exel.; 
winter 


MAMMALS 
Odocoileus hemionus eremicus: min. Urocyon cinereoargenteus scotti: max, 
(?) (foraged in all other riparian as- 
Peromyscus maniculatus sonoriensis: sociations) 
max. Mephitis estor: min. 


Sigmodon hispidus eremicus: max. Procyon pallidus: min. 

Reithrodontomys megalotis deserti: Antrozous pallidus pallidus: max. 
min. (?) 

Sylvilagus auduboni arizonae: min. Myotis oecultus: max. (?) 


Felis oregonensis browni: max. Nyctinomus mexicanus: max. (?) 


Remarks upon the Willow-Cottonwood Association —The predom- 
inating plants in this association were: willows of at least two species 
(Salix nigra and Salix fluviatilis), the cottonwood (Populus fremontt), 
and guatemote (Baccharis glutinosa). The latter occurred chiefly as 
an undergrowth where the willows or cottonwoods had reached large 
size. Practically all of the area occupied by this association is subject 
to inundation annually in early summer, of from a few inches to as 
much as twelve feet. Only such plants as can survive this period of 
drowning are able to oceupy the flood-bottom of the river. 

Furthermore, as explained in the general description of the river 
(p. 59), much of the overflow bottom is subject to destructive erosion 
through continual changing of the river’s channel. By this process 
all vegetation in its path is swept away at frequent intervals. Only 
such trees as are of rapid growth are able to maintain a representation 
on the major part of the bottom lands. 

No plants of the willow-cottonwood association (or it may be more 
briefly referred to as simply the willow association) occurred also on 
the upland deserts, and conversely no true desert plant occurred in 
the overflow area. Only one possible exception was observed, the 
serewbean (Prosopis pubescens) which occurred in mixture with wil- 
lows in a few places, where the flood-bottom was old, that is, had not 


1914] Grinnell: Mammals and Birds of the Colorado Valley 71 


been swept by channel-swinging for a long period of years. However, 
in the vicinity of the Colorado River, we failed to observe the serew- 
bean outside of the willow association, although it is known to oceur 
in the mesquite and even the catclaw associations elsewhere; so that 
here the secrewbean may be considered a restrictedly riparian plant. 

One other plant of the willow association requires mention—the 
eane (Phragmites communis), which grows in dense jungles on per- 
manent portions of the river bank from the vicinity of Picacho down. 
The tracts of cane are usually narrow, but, as in Canebrake Canon, 
below Pieacho, may extend continuously close along the river for miles. 
At the higher stages of the river the bases of the stalks are submerged, 
while the drooping ends trail in the water (see pl. 4, fig. 3). A verdant 
sereen on either hand thus intercepts the view of the voyager. 

The exogenous vegetation of the willow association is all of it 
deciduous, so that before the time of leafing-out in early March (see 
pl. 5, fig. 4) a very different appearance is presented from that later 
in the season. The spring growth is luxuriant, and accompanying it 
is a crop of insects which offer prolific food-supply to the abundant 
bird population of that season. Practically all of the birds listed for 
the willow association are either insectivorous or raptorial. Gramin- 
ivorous or spermophilous species are notably absent. 

The greater part of the fifty passerine birds listed for this asso- 
ciation are transients or winter visitants. Only three are permanent 
residents. The most notable characteristic summer visitants are: 
Dendroica aestiva sonorana, Vireo belli arizonae, Piranga rubra coop- 
eri, Archilochus alexandri, and Molothrus ater obscurus. 

Since the willow association ineludes practically the only forest 
elements in the region, we find here a concentration of dendrophilous 
birds, other than foliage-feeders, such as woodpeckers and flyeatchers. 
The latter, in particular, are abundant in both species and individuals, 
and contribute to the great contrast afforded between the life of the 
riverside and that on the desert which in large part lacks them. 

While bird-life is conspicuously more abundant in the willow asso- 
ciation than in any one of the desert associations, just the reverse 
obtains with the mammals. With the exception of the insect-feeding 
bats, which share with the smaller birds the benefits of the insect 
supply, there is but one rodent of wide and plentiful occurrence— 
Peromyscus maniculatus sonoriensis. Three other rodents occur locally, 
notably Sigmodon. Otherwise the only mammals of the willow asso- 
ciation are far-ranging predators. 


72 University of California Publications in Zoology [ Vou. 12 


The paucity of terrestrial mammals in this association is probably 
due to the repressive effect of the annual overflow which cannot fail 
to reduce the food-supply for many days at a time, even if extensive 
mortality does not directly ensue through drowning of individuals. 

As already implied (p. 58), the willow association varies greatly 
in width in different parts of the river’s course. Where the channel 
is constricted by rock walls, as in the box canon at The Needles, all 
trace of it is effaced for many rods. Where the river flows among hills 
patches of willows in ravine-mouths give detached representations to 
one or more elements. On the other hand, the broad valleys are occu- 
pied chiefly by this association which may then be as much as seven 
miles wide and continuous for many miles on one side or the other 
of the meandering channel. Taken by and large, the willow associa- 
tion is the most important one biotically of the entire set of associations 
dependent upon the presence of the river. 


TuLE ASSOCIATION 


BIRDS 

Fulica americana: max.; winter - Melospiza melodia saltonis: min.; 
Oxyechus vociferus vociferus: max.; resident 

winter Geothlypis trichas scirpicola: max.; 
Cireus hudsonius: max.; winter resident 
Xanthocephalus xanthocephalus: max. ; Geothlypis trichas occidentalis: min.; 

winter transient 
Agelaius phoeniceus sonoriensis: Telmatodytes palustris plesius: excl.; 

min.; resident winter 

MAMMALS 

Sigmodon hispidus eremicus: min. Ondatra zibethica pallida: min. 
Reithrodontomys megalotis deserti: Proeyon pallidus: min. 


max. 


Remarks upon the Tule Association.—The river’s habit of overflow 
would be expected to result in rather extensive tracts of palustrine 
flora. As a matter of fact, however, marshes were few and of small 
size. This was probably due to the rapid rate of evaporation of 
overflow water so that favoring conditions did not last long, and also 
to the rapid silting-in of such water basins as ox-bow cut-offs. As 
a result there were either almost lifeless alkali depressions, or lagoons 
practically identical in biotic features with the main river. But in 
a few places there were well-defined palustrine tracts kept wet 


1914 | Grinnell: Mammals and Birds of the Colorado Valley 73 


throughout the year, chiefly by seepage. These were always located 
back from the river near the outer edges of the broader valleys, where 
they were least affected during flood time. They were marked by 
growths of tules, sedge, and salt-grass, sometimes the latter alone, and 
were usually surrounded by the arrowweed or willow association (see 
fig. B). The little open water sometimes attracted a few transient 
ducks and mudhens, but so far as known no water birds outside of 
the Ardeidae remain to breed anywhere along the Colorado River. 

As may be noted from the list, but few animals were found to 
frequent the tule association regularly enough to be considered dis- 
tinetive features of that association. Of these, Reithrodontomys was 
the only mammal finding its center of abundance there. 


ARROWWEED ASSOCIATION 


BIRDS 

Lophortyx gambeli: min.; resident Melospiza melodia saltonis: max.; 
(only as a forager) resident 

Geococcyx californianus: min.; resi- Melospiza lincolni lincolni: max.; 
dent (only as a forager) winter 

Astragalinus psaltria hesperophilus: Pipilo maculatus curtatus: max.; 
min.; resident winter 

Zonotrichia leucophrys leucophrys: Oreospiza chlorura: min.; transient 
min.; winter Vireo belli arizonae: min.; summer 

Zonotrichia leucophrys gambeli: Vermivora celata celata: min.; win- 
min.; winter ter 

Melospiza melodia fallax: max.; win- Dendroica townsendi: min.; transient 
ter Oporornis tolmiei: min.; transient 

MAMMALS 

Peromyscus maniculatus sonoriensis: Sylvilagus auduboni arizonae: min. 
min. Mephitis estor: min. 

Neotoma albigula venusta: min. Spilogale arizonae arizonae: min. (?) 

Perognathus penicillatus penicillatus: 
min. 


Remarks upon the Arrowweed Association—In its purity this asso- 
ciation possesses only one conspicuous plant, the arrowweed (Pluchea 
sericea), which, over extensive areas, grows so densely as to occupy 
the ground to the exclusion of everything else (pl. 5, fig. 5). This 
belt of arrowweed usually oceupies slightly higher ground than does 


74 University of California Publications in Zoology [ Vou. 12 
the willow association, yet not above the high-water mark. It is pres- 
ent quite regularly as a tract along the outer margin of the willow 
association, often only a few feet in width, but sometimes, as for 
example near Pilot Knob, as much as two hundred yards in width. 

The arrowweed grows to a very uniform height, varying in different 
places from three to over eight feet; and the slender straight stems 
stand so close together as effectually to bar rapid progress through 
the more luxuriant tracts. The plant is of perennial growth, but only 
during the season of blossoming, April and May, does it appear to be 
particularly favored by insects. In many places there is mixed willow 
and arrowweed, in fewer places quail-brush and arrowweed, either 
combination being evidently more attractive to birds than the arrow- 
weed alone. 

The only resident bird finding its maximum abundance in the arrow- 
weed association is the song sparrow (Melospiza melodia saltonis). 
Even here the suspicion is aroused that this association is sought more 
for cover because of the peculiar manner of growth of the constituent 
plant, than as a food-producing area. For the song sparrow forages 
freely into both adjacent associations. 

Of mammals, not one ean be said to find its maximum abundance 
in the arrowweed association. The few species regularly trapped there 
were all more prominent constituents of adjacent associations. There 
would be little reason for the recognition of the arrowweed association 
as distinct from the willow association if it were not for its large extent 
and the conspicuous absence from it of a great many of the elements 
abundant in the latter. 


QUAIL-BRUSH ASSOCIATION 


BIRDS 


Lophortyx gambeli: max.; resident 

Zonotrichia leucophrys leucophrys: 
min.; transient 

Zonotrichia leucophrys gambeli: 
max.; winter 


Melospiza melodia fallax: min.; win- 


ter 


Melospiza melodia saltonis: min.; res- 


ident 


Pipilo maculatus curtatus: min.; win- 


ter 


Pipilo aberti: min.; resident 

Oreospiza chlorura: min.; transient 

Guiraea caerulea lazula: max.; sum- 
mer 

Oporornis tolmiei: max.; transient 

Geothlypis trichas occidentalis: max.; 
transient 

Toxostoma ecrissale: min.; resident 

Polioptila plumbea: min.; resident 


| 
1 


1914] Grinnell: Mammals and Birds of the Colorado Valley 


MAMMALS 
Peromyscus maniculatus sonoriensis: Neotoma albigula venusta: min. 
min. Sylvilagus auduboni arizonae: max. 


Remarks upon the Quail-brush Association.—The loeal hunters at 
Needles and other towns along the river apply the term quail-brush 
to Atriplex lentiformis, a plant growing in dense clumps three to eight 
feet in height along the outer margin of the flood-bottom. The belt 
of this plant is so well marked and continuous, and the important 
relations borne to certain animals are so evident, that the writer is 
led to designate it separately by the name quail-brush association (see 
pl. 6, fig. 6). 

Atriplex lentiformis often forms pure growths of a few yards to 
many rods in width on ground at about the upper limit of the 
average annual overflow. It was seen by us only in immediate prox- 
imity to the flood-bottom, and hence was essentially a riparian element, 
different from the other species of Atriplex inhabiting the region. 

The peculiar feature of the quail-brush was its compactly inter- 
lacing network of branches, so dense and resistent that a person 
might throw his whole weight against a thicket only to be hurled back 
by the elastic rebound. The value of the plant to the animals affeet- 
ing it consisted pre-eminently in the protecting cover afforded. Quail 
and cottontail rabbits when frightened took refuge in runways or 
natural spaces beneath its tangled mass; and bush-inhabiting sparrows 
of several species seemed to find ideal shelter in it. 

Although serving thus as a temporary shelter to animals regularly 
foraging in other associations, the quail-brush association also pro- 
vided a safe breeding place for such birds as Pipilo aberti, Gwiraca 
and Melospiza. In places, clumps of quail-brush alternated with 
mesquites, and here Toxostoma crissale found particularly congenial 


ground. 
Mesquitge ASSOCIATION 
BIRDS 
Lophortyx gambeli: min.; resident Centurus uropygialis: min.; resident 
Accipiter velox: min.; winter (only as a forager) 
Dryobates sealaris cactophilus: min.; Myiarchus cinerascens cinerascens: 
resident (only as a forager ?) min.; transient 
Sphyrapieus varius nuchalis: max.; Myiochanes richardsoni richardsoni: 


winter min.; transient 


76 University of California Publications in Zoology 


Empidonax griseus: min.; winter 

Pyrocephalus rubinus mexicanus: 
max.; resident 

Molothrus ater obseurus: min.; sum- 
mer 

Astragalinus psaltria hesperophilus: 
min.; resident 

Astragalinus lawrencei: excl.; winter 

Zonotrichia leucophrys leucophrys: 
min.; transient 

Zonotrichia leucophrys gambeli: min.; 
winter 

Spizella passerina arizonae: min.; 
winter 

Pipilo aberti: max.; resident 

Piranga ludoviciana: min.; transient 

Phainopepla nitens: max.; resident 

Vireosylva gilva swainsoni: min.; 
transient 

Vireo belli arizonae: min.; summer 

Vermivora luciae: excl.; summer 


[ Vou. 12 


Dendroica auduboni auduboni: min.; 
winter 

Dendroica nigrescens: min.; transient 

Wilsonia pusilla pileolata: min.; 
transient 

Wilsonia pusilla chryseola: min.; 
transient 

Oreoscoptes montanus: max.; winter 

Mimus polyglottos leucopterus: max.; 
winter 

Toxostoma crissale: max.; resident 

Heleodytes brunneicapillus couesi: 
min.; resident 

Auriparus flaviceps flaviceps: min.; 
resident 

Polioptila caerulea obscura: min.; 
winter 

Polioptila plumbea: min.; resident 

Planesticus migratorius propinquus: 
min.; winter 

Sialia mexicana occidentalis: exel.; 
winter 


MAMMALS 


Odocoileus hemionus eremicus: min. 

Peromyscus maniculatus sonoriensis: 
min. 

Neotoma albigula venusta: max. 

Perognathus penicillatus penicillatus: 
min. 


Sylvilagus auduboni arizonae: min. 
Felis oregonensis browni: min. (?) 
Lynx eremicus eremicus: max. (?) 
Mephitis estor: max. 

Myotis ecalifornicus pallidus: min. 


Remarks upon the Mesquite Association—This association is given 
the vernacular name of what is perhaps the most widely known plant 
of the region, the mesquite (Prosopis juliflora). Along the Colorado 
River the mesquite is closely restricted to a rather narrow belt along 
the outer edge of the riparian area, mostly above the reach of the 
highest flood water. This belt is of course lacking where hills closely 
abut upon the river, save at the mouths of ravines. Along the great 
valleys it forms a nearly continuous tract consisting of straggling 
clumps or of well-formed, though small, trees, in the latter case some- 
times orchard-like in regularity of spacing. At no point did we see 
mesquites with trunks over eight inches in diameter. 

While evidently requiring abundant water at root, submergence 
of the ground for any length of time kills mesquites, as proven by 


1914] Grinnell: Mammals and Birds of the Colorado Valley 


=I 
~] 


the conditions above the Laguna Dam. Here the whole association 
had been effaced by drowning. On the other hand, the mesquite is 
unable to exist on the desert proper, even in the larger washes. Only 
at the mouths of these did scattering examples extend away from the 
actual river bottom, and then, in the most favored places, for not more 
than a quarter of a mile. 

Man’s occupancy of the region has affected the mesquite associa- 
tion more than any other. The great value of the mesquite trunks 
for fuel has led to its practical disappearance as a tree along much 
of the lower course of the river. The steamboats which once plied 
regularly between Needles and Yuma are said to be chiefly respon- 
sible for this depletion. Several pumping plants contributed to the 
demand for fuel. Mesquite trees are very slow of growth; tracts of 
stumps now mark many areas where luxuriant groves once stood. 

A considerable number of low-growing plants find places as elements 
in the mesquite association, but by far the most important to the 
animal life is the mesquite itself. This shrub, or tree, provides both 
shelter and food, the latter through its fruit and foliage (see pl. 6, 
fig. 7) either directly or by way of insects. The mesquite serves also 
as the host of a parasitic plant (pl. 6, fig. 6), a species of mistletoe 
(Phoradendron californicum), which when in blossom is visited by 
myriads of insects, and which produces an abundant and almost con- 
tinuous crop of berries. Several of the winter and resident birds of 
the mesquite association depend almost wholly on these mistletoe berries 
for their food. Notable among these are: Phainopepla, Mimus, Oreo- 
scoptes, Planesticus and Sialia. 

Four species of breeding birds (Pipilo aberti, Toxostoma crissale, 
Vermivora luciae, and Phainopepla nitens) find in the mesquite asso- 
ciation the center of their abundance. One mammal is characteristic 
of the same association, namely Neotoma albigula venusta. 


SALTBUSH ASSOCIATION 


BIRDS 
Lophortyx gambeli: min.; resident Phalaenoptilus nuttalli nitidus: min.; 
Zenaidura macroura marginella: resident (?) 
max.; winter Chordeiles acutipennis texensis: min.; 
Geococeyx ealifornianus: min.; resi- summer (only as a forager) 
dent Sturnella neglecta: exel.; winter 
Phalaenoptilus nuttalli nuttalli: Poecetes gramineus confinis: excl.; 


min.; winter winter 


7 


Passereulus sandwichensis nevadensis: 


execl.; winter 


Passereulus sandwichensis alaudinus: 


exel.; winter 

Zonotrichia leucophrys gambeli: 
min.; winter 

Spizella breweri: max.; winter 


8 University of California Publications in Zoology 


[ Vou, 12 


Amphispiza bilineata deserticola: 
min.; summer 

Amphispiza nevadensis nevadensis: 
max.; winter 

Lanius ludovicianus exeubitorides: 
min.; resident (?) 

Thryomanes bewicki eremophilus: 
min.; winter 


MAMMALS 


Citellus tereticaudus tereticaudus: 
max. 


Peromyscus eremicus eremicus: max: 


Thomomys albatus: excel. 
Dipodomys deserti deserti: max. 


Dipodomys merriami merriami: min. 


Lepus californicus deserticola: min. 

Sylvilagus auduboni arizonae: min. 

Taxidea taxus berlandieri: excl. (?) 

Myotis ealifornicus pallidus: Max. 
(?) 


Pipistrellus hesperus hesperus: min. 


Perognathus penicillatus penicillatus: 
min. 


Remarks wpon the Saltbush Association—Ranchers in the Colo- 
rado River region distinguish two portions in each of the valleys, 
the ‘‘first bottom’’ and “‘second bottom.’’ These are of course duph- 
eated in reverse position on opposite sides of the river. The first 
bottom is the overflow area, and comprises all the associations treated 
up to this point in the present chapter, from the river to the mesquite 
association, inclusive. These associations together constitute the 
riparian belt proper. The second bottom is in the nature of a terrace 
or bench, and is situated above the reach of high water, extending 
from the mesquite association desertwards to the base of the mesa bluff. 

This second bottom is usually quite level and varies from a mere 
strip, of few yards in width, to a tract as much as a mile wide. It 
is for the most part clothed sparsely with xerophytie or halophytie 
vegetation, a predominating species in which is the saltbush (Atriplex 
polycarpa) ; hence the name adopted for the association represented. 

The soil is almost always of fine sand, often wind-blown. The pre- 
vailing westerly winds have caused a growth of sand-dunes on the 
southeast edges of the second bottom at several points, notably on 
the Arizona side above Mellen. The driving sand is often arrested 
about a bush and as the resulting dune grows, so does the bush com- 
mensurately. A scrubby form of mesquite may thus constitute the 
core of a sand-dune. Several small plants are peculiar to these wind- 
formed hillocks, one of which is the sand verbena (Abronia villosa). 


1914] Grinnell: Mammals and Birds of the Colorado Valley m9 


Favorable conditions for burrowing here attract heteromyid rodents, 
notably the large Dipodomys deserti. A characteristic assemblage 
results which might be appropriately called the aeolian sand associa- 
tion. Its peculiarities are not, however, in the writer’s mind, sufficient 
to warrant giving it more than minor recognition. Future finer 
analysis may possibly justify the separate recognition of the aeolian 
sand association, especially when the reptiles of the desert at large are 
taken into consideration. 

Elsewhere on the second bottom, depressions frequently occur where 
rainwater from adjacent desert slopes leaves by evaporation more or 
less alkali. In cases of excessive deposition, bare white stretches 
result, without any vegetation at all. On somewhat less alkaline 
ground there may be tracts of Spirostachys occidentalis and Suaeda 
suffrutescens, both being shrubby plants popularly included under the 
names pickle-weed and iceplant. 

But the greater portion of the second bottom, as also a strip lead- 
ing back along either side of the desert washes, is marked by the 
saltbush. This Atriplex is quite uniform in appearance wherever it 
grows, forming small but stout-branched bushes seldom more than 
two feet tall. 

The creosote bush (Larrea divaricata), although predominating in 
the next-deseribed association, occurs not infrequently as a minor 
element in the saltbush association. Sometimes individual plants of 


on the desert mesa. Where small alluvial fans from the higher adjacent 


Larrea reach a very large size (pl. 7, fig. 8), much larger even than 


mesa make down on to the second bottom, there is an influx of such 
plants as the coyote melon (Cucurbita palmata), rattle-weed (Eriog- 
onum inflatum), unicorn plant, locally called devil’s-claw (Martynia 
proboscidea), and sandbur (Franseria dumosa), all of which occur 
on sandy parts of the upper mesa as well. Along shallow washes 
through the second bottom there are often extensive thickets of Lyciwm 
andersont. 

In spite of the above indicated variations in floral constitution, 
there is remarkable homogeneity in the animal life of the saltbush 
association. In winter it is the preferred forage ground for a number 
of xerophilous fringillids, as shown in the accompanying lst of birds. 
The generally loose, sandy soil seems to be an attractive feature for 
burrowing rodents of limited fossorial powers such as Dipodomys mer- 
riami merriami. These also find abundant food in the residual seeds 
of numerous small annual plants which for brief periods thrive on 


80 University of California Publications in Zoology 


the open ground between the shrubs. 


[ Vou. 12 


One such plant, gathered exten- 


sively by Dipodomys deserti, is Achyronychia cooperi. 

It will be noted that the food-relations of the birds and mammals 
of the saltbush association and of the willow association are quite the 
reverse of one another; in the latter, mmsectivorous species prevail, in 
the former graminivorous or spermophilous. 


CREOSOTE ASSOCIATION (MgsA) 
BIRDS 


Chordeiles acutipennis texensis: 
max.; Summer 

Sayornis sayus sayus: min.; winter 

Spizella breweri: min.; winter 

Amphispiza bilineata deserticola: 
max.; summer 


Amphispiza nevadensis nevadensis: 
min.; winter 

Piranga ludoviciana: min.; transient 

Salpinectes obsoletus obsoletus: min.; 
winter 


MAMMALS 


Ammospermophilus harrisi harrisi: 
min. (stony) 

Ammospermophilus leucurus leucurus: 
min. (stony) 

Citellus tereticaudus tereticaudus: 
min. (sandy) 

Peromyscus eremicus eremicus: min. 
(sandy) 

Thomomys chrysonotus: exel. 

Dipodomys deserti deserti: min. 
(sandy) 


Dipodomys merriami merriami: max. 


Perognathus bombyeinus: exel. 
(sandy) 
Perognathus formosus: min. (stony) 


Perognathus penicillatus penicillatus: 
min. (sandy) 

Perognathus intermedius: min. 
(stony) 

Perognathus spinatus spinatus: min. 
(stony) 

Lepus californicus deserticola: max. 
Canis ochropus estor: max. (foraged 
at night practically everywhere 

else) 
Vulpes macrotis arsipus: exel. 
Myotis velifer: max. (?) 
Eptesicus fuscus: max. (?) 
Macrotus californicus: max. (?) 


Remarks upon the Creosote Association (Mesa).—The creosote bush 
(Larrea divaricata) was found to be the most widely distributed 
shrubby species of all the desert plants (see pl. 10, fig. 14). It 
occurred in varying abundance from the second bottom and wash- 
sides to the tops of the highest hills. Only the most rocky hill slopes, 
and the periodically eroded wash-bottoms, lacked this plant altogether. 
Yet there were obviously preferred areas of growth, or, still more 
notable in this connection, areas where the creosote bush grew to the 
entire exclusion of all other ligneous vegetation. These areas, where 


1914] Grinnell: Mammals and Birds of the Colorado Valley 81 


Larrea was at least the most conspicuous plant, occurred mostly on 
the more level upland desert mesa. The accompanying distinctive 
assemblage of mammals justifies the recognition of what may therefore 
be appropriately called a creosote (or mesa) association. 

Tongues of typical creosote association often run up from alluvial 
slopes into the rough hill country, following ravines and terraces. On 
the other hand, as already noted, the creosote bush and some of its 
companions in places invade the saltbush association. Along desert 
washes the two associations may be blended in all particulars to such 
an extent that definite diagnosis as one or the other is difficult. 

As will have been observed from the lists, there are only two 
breeding birds of the creosote association proper, Chordeiles acutipennis 
texensis and Amphispiza bilineata deserticola; and neither of these 
are abundantly represented. But graminivorous mammals are plenti- 
ful in species as well as individuals. Not all the species, however, 
are found in exactly the same places. As elsewhere shown, identical 
eround on opposite sides of the river may possess distinct species 
beeause of the action of the river as a barrier. 

Furthermore, segregation of rodent population on the basis of 
ability to find or dig safe retreats is evident. The nature of the 
eround thus bears a controlling relationship. Parts of the desert mesa 
are swept clean of fine sand by the prevalent winds, the resulting sur- 
face consisting of packed gravel, or wind-worn pebbles (pl. 12, fig. 18). 
Other parts of the desert have a sandy soil; and in places accumula- 
tions of sand transported by the wind have grown into sand-dunes, 
having much the character of those described for the saltbush associa- 
tion on a previous page. 

On sandy ground a common small shrub was the sandbur (Fran- 
seria dumosa) ; the rattle-weed (Eriogonum inflatum) was ever sum- 
moning startled attention; and in the vicinity of Pilot Knob a species 
of Ephedra was common. On stony ground often no other plant was 
to be seen than the creosote bush; but everywhere remains of short- 
lived sporadic vegetation gave clue to the source of supply of the seeds 
upon which depended the rodent population. In the vicinity of Pot- 
holes and Pilot Knob the mesa association included scattering clumps 
of round-stemmed cactus, and ocotilla (Mouquieria). 

Each of the two physical types of ground here noted, namely, 
sandy and stony, possesses certain peculiar species of mammals, as 
well as of plants. This segregation is indicated in the list, and might 
again serve as basis for recognizing two separate associations, the 


82 University of California Publications in Zoology [ Vou. 12 
sandy creosote, and the stony, or rocky, creosote; but by giving weight 
to similarities, as well as to differences, such distinction could be of 
but minor rank. Expediency seems to argue against further refine- 
ment in the present stage of study. Then, too, no such precise statis- 
tical gathering of data as regards the animals has yet been done as 
would warrant the exercise of such fine analysis. 


CATCLAW (OR WASH) ASSOCIATION 


BIRDS 


Lophortyx gambeli: min.; resident 

Zenaidura macroura marginella: min. ; 
winter 

Bubo virginianus pallescens: max.; 
resident 

Geococcyx californianus: max.; resi- 
dent 

Dryobates sealaris cactophilus: min.; 
resident 

Phalaenoptilus nuttalli nuttalli: min.; 
winter 

Phalaenoptilus nuttalli nitidus: min.; 
resident 

Archilochus alexandri: min.; summer 

Calypte costae: min.; summer 

Myiarchus cinerascens cinerascens: 
min.; summer 

Icterus cucullatus nelsoni: min.; tran- 
sient 

Ieterus bullocki: min.; transient 

Carpodacus mexicanus frontalis: 
max.; resident 

Astragalinus psaltria hesperophilus: 
max.; resident 


Zonotrichia leucophrys leucophrys: 
max.; transient 

Zonotrichia leucophrys gambeli: min. ; 
winter 

Zamelodia melanocephala melanoce- 
phala: min.; transient 

Phainopepla nitens: min.; resident 

Lanius ludovicianus exeubitorides: 
max.; resident (?) 

Lanivireo solitarius cassini: min.; 
transient 

Oreoscoptes montanus: min.; winter 

Mimus polyglottos leucopterus: min.; 
winter 

Heleodytes brunneicapillus couesi: 
max.; resident 

Thryomanes bewicki eremophilus: 
max.; winter 

Auriparus flaviceps flaviceps: max.; 
resident 

Polioptila caerulea obscura: max.; 
winter 

Polioptila plumbea: max.; resident 


MAMMALS 


Odocoileus hemionus eremicus: max. 

Peromyscus eremicus eremicus: min. 

Dipodomys merriami merriami: min. 

Perognathus formosus: min. 

Perognathus penicillatus penicillatus: 
max. 


Perognathus spinatus spinatus: min. 
Lepus californicus deserticola: min. 
Lynx eremicus eremicus: min. 
Urocyon cinereoargenteus scotti: min. 
Pipistrellus hesperus hesperus: min. 


Remarks wpon the Catclaw (or Wash) Association—Except for 
the Bill Williams and Gila rivers the Needles-to-Yuma section of the 
Colorado River receives no tributary stream, save as an immediate 


1914] Grinnell: Mammals and Birds of the Colorado Valley 83 


result of the very infrequent local cloud-bursts or severe thunder- 
storms. Even the two ‘‘ 
their lower courses following protracted drouth. At frequent inter- 
vals along the valley of the Colorado, well-defined but ordinarily dry 
water-courses leading down from the adjacent uplands record the 


rivers’’ named often go completely dry in 


existence at some time or other of considerable run-off. Among the 
hills, deep, steep-sided ravines show the profound effects of erosion, 
even though, in this era, rains come but rarely. Where the water-course 
drains a large extent of higher country, enough to furnish a volume 
of water sufficient to find its way through the mesa to the river, it 
may occupy a broad valley with low confining banks or bluffs. Such 


““wash’’; and as the biota 


a broad, dry flood plain is called locally a 
of such ‘‘washes’’ is quite distinctive the term ‘‘ wash association’’ has 
been suggested (see pl. 7, fig. 9). Since a prevalent plant in its flora 
is the tenaciously thorny eatclaw (Acacia greggii), the term catclaw 
association may be used as an alternative denomination. Both are 
appropriate terms. 

The vegetation of the catclaw association is the most conspicuous 
of all of the desert associations, for it includes several species which 
reach the stature of trees. The largest of these is the desert 1ronwood 
(Olneya tesota) which grows abundantly in all the larger washes on 
both sides of the river, from the lower Chemehuevis Valley at least 
to the vicinity of Picacho (pl. 8, fig. 10). The branches are leafy but 
thorny, forming ideal refuges for certain small birds and locations 
for their nests. The apparently leafless palo verde (Parkinsonia 
torreyana) is a close companion of the ironwood, and occurs also along 
the smaller ravines into the hills (see pl. 8, fig. 11; pl. 9, fig. 12). Both 
the catclaw and the palo verde were found in practically every wash 
on both sides of the river, from the vicinity of Needles to Pilot Knob. 
The smoke-bush (Dalea spinosa) is a conspicuous element in many of 
the washes from near Riverside Mountain to Pilot Knob. 

As an indication of the size reached by individuals of these truly 
xerophilous trees the following measurements taken by the writer, 
are here presented: An ironwood growing in a wash about one-half 
mile back from the river bottom in lower Chemehuevis Valley, Cali- 
fornia side of the river, was 90 inches in circumference of trunk two 
feet above the ground, and had a height of 31 feet. A palo verde 
nearby was 48 inches in circumference of trunk two feet above the 
ground, and was 28 feet in height. Another palo verde (pl. 9, fig. 12) 
growing in a wash on Californian territory four miles north of Pot- 


84 University of California Publications in Zoology | Vou. 12 


holes, was 60 inches in circumference of trunk two feet above the 
eround, first branch four feet above the ground, and total height 24 
feet. 

It is thus apparent that birds of arboreal habit find only in the 
wash association of the desert a near approach to conditions preferred 
elsewhere, and this doubtless accounts for the conspicuous transient 
arboreal element occurring in this association. But the greatly reduced 
foliage, giving most of the above-named trees the aspect of winter 
leaflessness, results in close resemblance to brush or shrubbery, as 
indieated by the prevalence of the brush-inhabiting category of breed- 
ing birds affecting them. 

The ironwood oceasionally harbors clumps of mistletoe (Phoraden- 
dron californicum) in common with the mesquite of the riparian belt. 
This accounts for the presence of certain berry-eaters. There are also 
berry-produeing shrubs bordering the washes, notably Lycium ander- 
soni and Lycium parishi. The former occurs widely as a character- 
istic member of the ecatclaw association. The latter, a much larger 
thicket-forming shrub, was noted only in small washes in the vicinity 
of Picacho. As already noted, Atriplex polycarpa occurs in an exten- 
sion of the saltbush association leading up along each side of nearly 
all of the larger washes. 

Bird-life is better represented in the catclaw association than in 
any other of the desert associations. Some of the species are closely 
adherent to it, being evidently by structure and habits dependent 
upon the conditions pertaining to thorny brush. But mammals are 
relatively less numerous. Only one rodent finds its maximum abund- 
ance along the washes (Perognathus penicillatus penicillatus, see fig. 
D), and it is possible that even of this species, the metropolis is in 
the adjacent sandy saltbush tract, and that trapping really waylaid 
the individuals foraging at large away from their homing places. 
Times of deluge, even if of rare occurrence, are doubtless accompanied 
by great mortality of ground-dwelling mammals along these washes. 
This factor must be one of no small import in determining the biotic 
constitution on the several levels of the desert surface. 


1914] Grinnell: Mammals and Birds of the Colorado Valley 85 


Saguaro AsSsociaTION 


BIRDS 
Faleo sparverius phalaena: max.; Colaptes chrysoides mearnsi: max.; 
resident resident 

Otus asio gilmani: min.; resident Myiarchus cineraseens cinerascens: 
Micropallas whitneyi: exel.; resident max.; Summer 
Centurus uropygialis: max.; resident Carpodacus mexicanus frontalis: 

min.; resident 

MAMMALS 


(As far as known, same as in creosote association.) 


Remarks upon the Saguaro Association—The conspicuous columns 
of the giant cactus or saguaro (Cereus giganteus) first met our expect- 
ant gaze just below the mouth of Bill Williams River. There, on both 
sides of the Colorado, for a stretch of two miles or more, stood many 
specimens growing on the hill slopes in full view from the river as 
we floated by. A landing was made on the California shore, and sev- 
eral saguaros closely examined for nesting sites of birds. We next 
found these cactuses on the open desert one to three miles east of 
the river on Arizona territory around Ehrenberg. There were so many 
here that they formed a distinct tract, extending across the mesas and 
occupying the interlying washes as well. 

Giant cactuses again came to view some ten miles below Picacho, 
two or three individuals being seen on the California side and many 
on the Arizona side. Finally, on the mesa on both sides of the river, 
three to five miles above the Laguna dam, saguaros were plentiful. 
On the California side from one hundred yards to two miles back from 
the outer edge of the river bottom were about seventy-five individuals 
(see pl. 9, figs. 12, 13). Thirty-seven, big and little, were counted by 
the writer as in sight from one point. 

It was gratifying thus to find this unique plant well represented 
on California ground. For coming with it into the state of California 
were the several birds listed, an assemblage altogether justifying the 
designation of a saguaro association. This association is best devel- 
oped in a large area of southwestern Arizona. The tongue crossing the 
Colorado River above Laguna and Potholes is undoubtedly an exten- 
sion of it. 

As far as we were able to learn, in other plant elements and in 
mammals, the saguaro association was here identical with the creosote 


86 University of California Publications in Zoology — [Vou. 12 


association. It is not improbable, however, that further work would 
disclose the presence of a number of species of both mammals and 
birds, at least on the Arizona side, not now known so far west. 

It is clearly apparent that the critical feature of the saguaro which 
prescribes its avian dependents is nothing else than the favorable 
opportunity offered for the excavation of safe retreats in its trunk. 
And only the two woodpeckers are equipped for making these exca- 
vations. So that, without the woodpeckers to make holes, the other 
birds would be no better off for the presence of the saguaro. As it 
happens, at least one of the species of woodpeckers (Centurus wropy- 
gialis) invariably accompanies the cactus. 

This is a most interesting form of contingent or incidental inter- 
dependence of animal and plant. Since the giant cactus during the 
greater part of the year produces no fruit and harbors no insect life, 
it follows that all the birds using its cavities as roosting or nesting 
places glean their livelihood from the surrounding desert. ‘lhe latter, 
as already stated, presents conditions which seem to be practically 
identical with the creosote association, which is so nearly barren of 
bird life, and of vast extent beyond the limits of the saguaro. One 
form of associational restriction is hereby proven, namely, that by 
dependence upon safe home retreats. 


Enceia (Rocky Hints) Associarion 
BIRDS 


Buteo borealis calurus: max.; resident 

Faleo mexicanus: exel.; resident 

Bubo virginianus pallescens: min.; 
resident 

Aéronautes melanoleucus: excl.; resi- 
dent 

Calypte costae: max.; summer 

Sayornis sayus sayus: max.; resident 

Corvus corax sinuatus: min.; resident 

Carpodacus mexicanus frontalis: 
min.; resident 

Astragalinus psaltria hesperophilus: 
min.; resident (?) (as a forager 
only) 


Spizella passerina arizonae: max.; 
winter 

Spizella breweri: min.; winter 

Spizella atrogularis: excl.; transient 

Amphispiza bilineata deserticola: 
min.; summer 

Stelgidopteryx serripennis: max.; 
summer 

Salpinetes obsoletus obsoletus: max.; 
resident 

Catherpes mexicanus conspersus: 
exel.; resident (?) 


MAMMALS 


Ovis canadensis nelsoni: excl. 
Ammospermophilus harrisi harrisi: 
max. 


Ammospermophilus leucurus leucurus: 
max. 
Peromyscus erinitus stephensi: excl. 


1914] Grinnell: Mammals and Birds of the Colorado Valley 87 

Neotoma intermedia desertorum: Corynorhinus macrotis pallescens: 
excl. excel. (?) 

Perognathus formosus: max. Pipistrellus hesperus hesperus: max. 

Perognathus intermedius: max. (foraging everywhere else) 


Perognathus spinatus spinatus: max. 


Remarks wpon the Encelia (Rocky Hills) Association.—None of the 
hills or ‘‘mountains’’ in the near vicinity of the lower Colorado River 
is of such great altitude as to bring a reduction in temperature to 
an extent sufficient to modify its biotie complexion. The tallest of 
The Needles rises to less than 2,000 feet above the level of the river. 
In other words, as far as observed, the plants and animals of the hills 
show no distributional behavior other than as explained on associational 
grounds. 

But the hill country does exhibit a distinctive association of plants 
and animals, setting them apart sharply from the desert mesa, or the 
riverside. A conspicuous shrub on the rocky steeps, especially at The 
Needles, was the dense Encelia farinosa, with its light gray foliage, 
growing on talus slopes and even in crevices of the cliffs (pl. 10, fig. 
15). Other plants of the same locality were: Atriplex confertifolia, 
Larrea divaricata (sparsely intermixed), Asclepias subulata (visited 
regularly by the Costa hummingbird), Fagonia californica (on the 
hottest slopes of broken rock), Hyptis emoryi (a ‘‘sage-bush”’ five to 
six feet high growing on the sides of ravines and at time of blossoming, 
in March, frequented by hummingbirds), Perityle emoryi (an abund- 
ant composite annual, the seeds of which were much sought after in 
March by fringillids), and Miihlenbergia debilis (a grass growing in 
shaded ravines and providing forage for graminivorous rodents, like 
Perognathus). 

While to distant view the hills seemed more barren of vegetation 
than most of the other associational areas, nearby inspection showed 
abundant remains of inconspicuous annual plants. This in part would 
seem to account for the great numbers of mammals present, as shown 
by our trapping. The additional favorable factor was evidently the 
abundance and availability of natural retreats afforded in the talus 
and fractured outerops. 

With insectivorous and raptorial species, such as the swifts and 
bats, hawks and owls, the rocky hills served merely as home retreats, 
foraging being doubtless carried to the more productive lowlands, 


88 


Vv ‘Sl 
suoyelsossy j/eLuiuey pre /uUe/4 BuUIMOYsS 


University of California Publications in Zoology 


sajpaeN eu, 4? 
Kae sandy pega? Jo ¥yfory jeuoyras /eap] 


MNOZ/YY 


bINYOS/7TrI 


YAY OOYHOTOD 


Encela 


creosole bush 


salt bush 
mesquile 


quail brush 


|arowweed 


| willow 


willow 


arrowweed 


[ quail brush 


mesquile 


salt bush 


creosole bush 


Encelia 


Wedoma deserlorum 


= 


Perognalhus inlermedius 
\Ammospermophilus herrist 


Dipochmys merriami 


Perognalhus penicillalus 
Peromyscus erermicus 
Sy/vilagus arizonae 


Peromyscus sonoriensis 
Neoloma venusta 


Peromyscus SOnOrlensis 
Neolorma venusla 


Peromyscus erermicus 
Sylvilagus arizonaze 


Perognalhus penici/ialus 


Dipodoriys merriam 


Ammospermophilus /eucurus 
Perogsnathus spinalus 
Perognathus formosus 
Neolorma deSerforurl | 
Peromyscus slephens/ 


[ Vou. 12 


1914] Grinnell: Mammals and Birds of the Colorado Valley 


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wed [ie | 
IPS 
= x ® o~ qi i Fiber 
u gh t 
SER [SY 
DWIoO [8 ‘1 
x 1 
> 
urs 2 
~N = 
Ny & ce willow 
g ° y 5 
5 8 = 
v9 
~~ D Neoloma venusla 
S NS arrowweed Peromyscus sonoriens/s 
a 
= ; 
< ke 
z PP wail brush 
° 
g 3 » eS mesguile 
= | 3| a 
72 sf Syiilegus arizonae 
> 
Cro 8 3 saltbush 
a zt Pe 17 Moh. 
= 5 g mesguile erognaltus pericilalus 
oS . 
eae saltbush 
= 9 
Can tr Dpodomys merriami 
& 2 + : 
Ae 9 
To 2 
: < 
= = Immospermophilus leucurus 
a & creosole Lush \\Weolma deserlorum 
8 erosnalhus spinalus 
& » rosnahus formasus 
oy t ‘2 
= & 
. . 


90 University of California Publications in Zoology —- [ Vou. 12 


particularly along the river bottom. Only two insectivorous birds 
were restricted to the Encelia-association, namely, the wrens, Salpinctes 
obsoletus obsoletus and Catherpes mexicanus conspersus. Both are by 
structure adapted to gleaning from crevices of rock surfaces after the 
manner of nuthatches on tree trunks. 


GENERAL DiscussiON OF ASSOCIATIONAL RESTRICTION 


From the preceding description of the conditions in the region 
studied, it is obvious that there are two groupings into which all the 
designated associations can be classed, namely, riparian and desert. 
The riparian set of associations includes those which owe their pres- 
ence to the existence of the river, and is delimited outwardly at the 
mesquite association (see figs. A and B). The Colorado River appar- 
ently exerts no influence beyond the immediate bottom lands, which 
are affected by the underground water supply. All of the desert set 
of associations are represented in varying proportion over the vast 
arid tracts stretching away to the east and west of the river. Often 
they are discontinuous, but recur again and again in the same fauna 
with the same constitution. 

The riparian associations are thus narrow strips of varying width 
closely paralleling the river from north to south and persisting prac- 
tically continuously from the point of emergence of the Colorado River 
from the Grand Cafion to the Gulf of California. In the broad delta 
region the riparian associations spread out so that there are great 
areas of each, doubtless sufficient to be computed by the square mile. 
It is thus possible to trace the elements severally, of each association, 
to places of prevalence over considerable areas, even though those ele- 
ments are, on the upper river, scattered sparsely along a narrow strip. 
Associational diagnosis of species thus often becomes possible when a 
knowledge of local conditions alone would be inconclusive. This prin- 
ciple deserves enlarging upon. 

There were caught in the same trap-line opposite The Needles 
both Peromyscus eremicus eremicus and Peromyscus crinitus stephens. 
It might have been impossible to say from the data gathered at that 
particular point, where the adjacent associations were complexly inter- 
mixed, just what sort of ground each species preferred. But trapping 
previously done in the salt-bush association in the Imperial Valley 


1914] Grinnell: Mammals and Birds of the Colorado Valley oil 


showed the presence there of P. e. eremicus only, while field-work 
among the rocky hills in the vicinity of Victorville, on the Mohave 
Desert, showed P. c. stephensi only. 

As another illustration, along the Colorado River, Melospiza melodia 
saltonis and Pipilo aberti were often found on common ground, 
although evidently averaging differently in associational preference. 
To prove beyond doubt what is the true ecologic niche of each, a 
knowledge of the distribution of each species elsewhere in their respeec- 
tive ranges becomes necessary. In the extensive arrowweed tracts 
around the west end of Salton Sea, Melospiza melodia saltonis is an 
abundant species while the other bird is absent. In the mesquite belt 
not far distant to the west, in the vicinity of Martinez and Torres, 
Pipilo aberti is prevalent, and the song sparrow absent. Hence the 
species of towhee in question may be confidently assigned to the mes- 
quite association, and the subspecies of song sparrow to the arrowweed 
association. 

It is not to be inferred that all species behave in this clearcut 
fashion associationally, any more than that all do so zonally or faunally 
(see Grinnell and Swarth, 1913, p. 220). At the same time the writer 
feels fairly sure of adequate grounds for proposing a general law in 
this regard, namely, that where the faunist happens to meet with a 
heterogeneous assemblage of biotic elements, not subject to clear asso- 
ciational diagnosis in the restricted locality of first observation, assign- 
ment of the species each to a well-defined association becomes possible 
by tracing out their ranges severally into the adjacent areas. 

A concurrent axiom is that if associational analysis is carried far 
enough, no two species of birds or mammals will be found to occupy 
precisely the same ecologic niche, though they may apparently do so 
where their respective associations are represented fragmentarily and 
in intermixture. 

In determining the associational status of mammals we have to deal 
chiefly with elusive animals, of nocturnal habits, which are hidden 
away during the day for the most part beyond reach. Trapping is 
not an altogether certain index to association; for individuals may be 
caught repeatedly in a trap-line which may not happen to intersect 
at all the regular forage ground or breeding home of the species. Indiv- 
iduals forage far and wide beyond the limits of their home territory 
and at the close of the breeding season wander in similar fashion. 
Some species, including nearly all mammals except xerophilous rodents, 
regularly travel far for water. 


oO 
bo 


University of California Publications in Zoology | Vou. 12 


Attention is here called to our records of the capture of pocket mice. 
An unexpectedly large number of species of the genus Perognathus 
yas found to occur in the region traversed. As many as three species 
were taken in one night in traps placed close together. But by testing 
many localities and comparing the results we soon came to know where 
to expect each separate species. The diagrams herewith presented 
(figs. C-F) show in statistical form the associational preferences of 
four species of Perognathus. Providing the same number of trap- 
nights (counting one trap set one night as one ‘‘trap-night’’) was 


= = ne 

2 3 a 

C4 oO = 

Lo} = 2 = = 

3 na 5 s ~— > 

> > ws 

Se 9 Go ee be as 

by = 2 =] 3 =) =o 

o 5 = = 2 a a Om 

> SI o a = a o 8 
om = oS 5 Bq iI 
io) <i) tel m e So A 


Fig. C. Diagram showing frequency of capture of Perognathus formosus in 
the several associations. There were fifty-three individuals trapped, of which 
the associational occurrence was satisfactorily recorded. 


devoted to each association, this method should be fairly accurate. 
There will here occur to the reader ways of securing much greater 
precision in results of field-work in the future. But in fact, as our 
experience grew, far more effort was expended on ground promising 
additional specimens of rare species, than on ground already thor- 
oughly exploited. 

While a mammal or bird may be closely confined to a narrow 
territory characterized by certain conspicuous plants. the critical 
factor or factors of its environment may be quite apart from food 
requirements. As an illustration, the case of Dipodomys deserti 


1914] Grinnell: Mammals and Birds of the Colorado Valley 93 


deserti may be cited. This rodent, taking its entire, irregular and 
discontinuous range into consideration, is closely restricted to those 
portions of the desert affording a deep surface layer of fine sand. 
Areas of aeolian sands constitute its typical home territory (see pl. 11, 
fig. 16). Now, it is quite probably not any peculiarity of food-supply 


s e eS 

a n 

s & = 

3 tf 2 = = 

2 D s = on] 

iS I 2 | o @ ad 

BS is SE Se OSS 
2 =| r=] ¢ = 

A OS eB fe oe oee ee 2 OR 

> = I = = = s ® o~ 
oS 5 Be a = me 
<q (<4 a mn S is {S| 


a 


Fig. D. Diagram showing frequency of capture of Perognathus p. penicil- 
latus in the several associations. Two hundred and seventeen individuals were 
taken, of which associational occurrence was definitely recorded. This species 
is seen to be very widely distributed, yet exhibiting marked preference for 
certain associations. 


attaching to the aeolian sands, which binds the animal to them in this 
case rather than to the mesa at large; but it is the animal’s require- 
ments in the way of retreats for diurnal safety and for breeding. 
These, by nature of its fossorial limitations, Dipodomys d. deserti is 
unable to construct for itself except in ground easy to burrow into to 
a depth commensurate with its own large size and in soil with a 
proper degree of coherence. 


94 University of California Publications in Zoology [ Vou. 12 


In the case of desert quail there is a double need: of daily access 
to water, plus that for refuges constantly within easy reach in event 
of the birds being suddenly threatened by an enemy. The quail-brush 
association affords the ideal shelter. The futile efforts of a coyote 
or fox to dash in pursuit into such an interlacing thorny branch-work 
as is afforded by Atriplex lentiformis can be imagined! 


Willow 
Arrowweed 
Mesquite 
Saltbush 

Wash (Catclaw) 


Fig. E. Diagram showing frequency of capture of Perognathus intermedius 
in the several associations. Eighty-three individuals were trapped, of which 
associational occurrence was definitely recorded. 


In these cases and all others it is self-evident that presence of 
food-supply is the primal associational requisite, whatever other fac- 
tors may be also essential. And kinds of food produced, with regard 
to the structural characters of each animal, determine what kinds of 
associational elements can exist in a locality. An animal having a 
specialized means of procuring food, lke that of a sandpiper, or a 
woodpecker, or a rock wren, is hemmed in by the bounds outlining the 


2 ss 


1914] Grinnell: Mammals and Birds of the Colorado Valley 95 


area in which it is able to get the food necessary for itself by the 
method which its anatomical structure and psychological equipment 
prescribe. 


ote (Mesa) 


Arrowweed 


River 
Willow 
Mesquite 


Fig. F. Diagram showing frequency of eapture of Perognathus s. spinatus 
in the several associations. One hundred and ten individuals were trapped, of 
which associational occurrence was satisfactorily recorded. While the asso- 


ciational preference of this species is almost identical with that of P. inter- 
medius, the ranges of the two species are wholly distinct, being separated 
by the Colorado River. Associational preference is also seen to be similar to 
that of P. formosus, which occupies apparently identical ground. But P. spin- 
atus belongs to a different subgenus (Chaetodipus) from P. formosus (subgenus 
Perognathus), hence the two are not nearly related and probably do not come 
into close ecological competition. 


Not infrequent are the cases where food-supply is located in one 
place, and the refuge or breeding site in another. Animals of such 
habit are necessarily of relatively great powers of locomotion, and 
may live regularly in two or more associations situated remotely from 


96 University of California Publications in Zoology [ Vou. 12 


one another. Incidentally all other associations may be crossed, back 
and forth. Ardea herodias treganzai foraged far and wide in the 
river association, but reared its young in restricted parts of the willow- 
cottonwood association where lofty trees provided safe nesting sites. 
Aéronautes melanoleucus found safe rendezvous in crevices of cliffs, 
but foraged far over the lowlands; so also with certain bats. The 
relations between animals and their environments are manifold and 
complicated; and these interrelations depend both upon the inherent 
peculiarities of the entire organism in the case of the animal and upon 
the physical nature of the environment. 

After a consideration of all the birds and mammals mentioned in 
the present report, as occurring both in the region specifically treated 
and elsewhere as far as the writer’s knowledge goes, associational 
restriction appears to be governed by the following three factors, of 
relative importance in the order named. 

1. Kind of food-supply afforded, with regard to the inherent strue- 
tural powers of each of the animals concerned to make it available. 

2. Presence of safe breeding-places, adapted to the varying needs 
of the animals, in other words, depending upon the respective inherent 
powers of construction, defence and concealment in each species con- 
cerned. 

3. Presence of places of temporary refuge for individuals, during 
day time or night time, or while foraging, when hard pressed by 
predatory enemies, again correlated with the respective inherent powers 
of defence and concealment of each species involved. 


1914] Grinnell: Mammals and Birds of the Colorado Valley OT 


THE COLORADO RIVER AS A HIGHWAY OF DISPERSAL 
AND CENTER OF DIFFERENTIATION OF SPECIES 


According to Gilbert and Seofield (1898, pp. 487, 488) the peculi- 
arities of the fish fauna of the Colorado River bespeak a very long 
period of absolute isolation. A remarkably high percentage of its 
fishes are specifically distinct from those of the other river basins of 
western North America. The same is to be said of the riparian birds 
and mammals. 

So far as known to the present writer, none of the species listed 
with a star in the following table ranges beyond the confines of the 
Colorado River basin, including of course its various tributaries and 
distributaries such as the Gila and New rivers, except sporadically, 
or as accounted for by distal invasion through passes or along sea- 
coastal tracts. In other words, the Colorado River has been in existence 
so long that the conditions imposed by its presence have figured in 
the differentiation of representative species of several families, both 
mammalian and avian. 

The great age of the Colorado River is indicated geologically by 
the vast extent and slow rate of the erosion involved in the formation 
of the Grand Canon. This time-element is justly inferred to have 
been an essential condition in the formation of these species. 


SPECIES WHICH BELONG TO THE RIPARIAN BELT, AND WHICH ARE THEREFORE 
HEMMED IN BY THE PARALLELING DESERT TRACTS. STARRED 
SPECIES ARE PECULIAR TO THE COLORADO SUBFAUNA 


BREEDING BIRDS MAMMALS 
Agelaius phoeniceus sonoriensis* Castor canadensis frondator* 
Melospiza melodia saltonis* Peromyscus maniculatus sonoriensis 
Pipilo aberti* Sigmodon hispidus eremicus* 
Guiraca caerulea lazula Reithrodontomys megalotis deserti 
Piranga rubra cooperi Neotoma albigula venusta* 
Vireo belli arizonae* Ondatra zibethieca pallida* 
Vermivora luciae* Mephitis estor 
Dendroica aestiva sonorana* Proeyon pallidus* 


Toxostoma crissale* 


It may not be amiss to consider these riparian species somewhat with 
regard to origin. By confining our attention to the north-and-south 
valley of the lower Colorado, this becomes, with a knowledge of the 
general status in North America of each group represented, a com- 


98 University of California Publications in Zoology [Vot. 12 


paratively simple matter. The axiom holds, that, because of the ever- 
shifting location geographically of associational, faunal and zonal con- 
ditions, every single element or line of descent, now represented in the 
biota of any one locality must have come either in its present form 
or in some antecedent one from somewhere else. This is certainly 
true of all terrestrial life. Elevation and depression have worked lke 
a seesaw in dislocating faunas. The Colorado valley is a trough, 
hemmed in associationally on either side, and only capable of influx 
of riparian elements at either end. Therefore the riparian species of 
the Colorado fauna can have entered the area under consideration from 
only two directions: from the north and from the south. 

Only two species are clearly seen to have entered the Colorado val- 
ley from the north: Castor canadensis frondator, and Ondatra zibethica 
pallida. The following species or subspecies are believed to have come 
in from the south: Pipilo aberti, Guiraca caerulea lazula, Piranga rubra 
cooperi, Vireo belli arizonae, Vermivora luciae, Toxostoma crissale, 
Sigmodon hispidus eremicus, Neotoma albigula venusta. 

In the remaining seven cases (Agelaius, Melospiza, Dendroica, 
Peromyscus, Reithrodontomys, Mephitis and Procyon) no grounds are 
apparent to the writer for assigning either one over the other direction 
of invasion, and this in spite of whatever may be the marked Austral 
or Boreal distributional affinities of each group concerned. The tide 
of invasion may in fact in these species have tended in one direction 
at one period, in the opposite at another; or, as in Peromyscus mani- 
culatus sonoriensis, the Colorado valley may have acted continuously 
as a narrow bridge where have met and mingled descent-lines from 
both the north and the south. 

The obvious fact that southern representatives prevail over northern 
ones is clearly attributable to the present zonal condition obtaining 
in the region, namely Austral, in its Lower Sonoran division. And 
evidence elsewhere assembled (Grinnell and Swarth, 1913, p. 383) 
points towards an increasing temperature throughout the region. This 
would result in decreasing the favorableness to Boreal forms and 
increasing the availability of the region for immigration of Austral 
types. Consideration of the xerophilous vertebrates as well as of the 
riparian ones leaves little doubt in the writer’s mind but that this has 
been the actual course of events. The northern contingent is on 
the wane, the southern in the ascendency. 

The query presents itself: is the Colorado fauna full? Are all the 
ecological niches, which are available in this area and which have 


1914] Grinnell: Mammals and Birds of the Colorado Valley 99 


occupants in other regions, occupied here? Probably not, for the inter- 
vention of barriers has doubtless prevented the invasion of types 
which, if they could have once gotten there, would have thriven and 
assumed a place as endemic elements in the fauna. Sporadic ineur- 
sion, as of migrants among birds, and strays among both birds and 
mammals, do not appear to the writer to figure in such a process. 
Rather must it be a progressive invasion of the species en masse, 
acquiring, it may be, adaptive modifications as it proceeds. In other 
words, the conquering of the land is the combined result of the 
facilities offered by it plus the relative amenability of each species 
concerned. 

The twelve riparian species and subspecies peculiar to the Colorado 
fauna vary much in degree of difference from their near relatives 
which occupy adjacent differentiation areas. These varying degrees 
of difference might be interpreted as measures of the periods of time 
elapsed since the entrance into the region of each of the types 
involved. That this conclusion is poorly grounded is evident upon 
consideration of the various other elements which must figure in the 
process of species formation. Among these may be suggested: degree 
of isolation, divergence of homologous associational conditions in the 
new region from those in the ancestral, and inherent susceptibility 
to adaptive modification in each of the species concerned. 

In the problem of the origin of the riparian portion of the Colorado 
fauna we seem to have to do with an accentuated kind of isolation. 
For, as already asserted, there is such a thing as more and less isolation. 
In the region here considered, possessing extreme associational con- 
trast, we find the ordinary geographic, or more properly speaking, 
physiographic, isolation coupled with associational isolation. In con- 
sequence of this extra favorable contingency, differentiation of species 
may have progressed with particular celerity, with such distinct forms 
to show for it as Pipilo aberti, Vermivora luciae, Toxostoma crissale, 
and Procyon pallidus. 

The axiom has preseuted itself in this connection that the more 
restricted a species is associationally, that is, the more confined to a 
narrow range of associational conditions, the more subject it is to the 
important factor of isolation; hence the more liable to give rise to 
new incipient strains in different parts of its general range. 

An assertion which seems at first glance opposed to the above is: 
that the less restricted a species is associationally, that is, the more 
widely adaptable to varying conditions, the more numerous the chances 


100 University of California Publications in Zoology | Vou. 12 


for local operation of isolation, because more opportunity for radial 
dispersion to carry the species into distant localities and under extreme 
conditions, and for the ultimate interposition of more or less efficient 
barriers. The factor of distance might here replace the operation of 
associational restriction in segregating descent-line plexuses. 

These are apparently incongruous notions, but the following con- 
ception tends to harmonize them; namely, that, granting the three 
totally different orders of distributionally limiting factors (zonal, 
faunal and associational), it is probable that different species are 
restricted wnevenly with respect to the three; thus a certain wood-rat 
(Neotoma intermedia desertorum) is restricted faunally, but ranges 
widely through zones and associations; a certain wren (7'elmatodytes 
palustris and subspecies) is tightly restricted associationally, but 
ranges widely through faunas and zones; a xerophilous genus of 
rodents (Perognathus) is closely restricted zonally and associationally, 
but ranges rather widely as to fauna. So that both the above asser- 
tions might well be true of a single animal historically and even, in 
different parts of its range, simultaneously! Certainly the first serves 
in explanation of the multiplicity of geographic races or species in 
several widespread groups of birds and mammals. 


THE COLORADO RIVER AS A HINDRANCE TO THE 
DISPERSAL OF SPECIES 


Bats and most birds find in the Colorado River no hindrance what- 
ever to individual travel. Freedom of aerial locomotion gives them 
superiority over any obstruction on the general level of the country 
they inhabit. It is possible that in a few of the resident birds of 
limited flight individuals do not regularly cross the main stream, 
though they readily could do so if such an exigency as that of fire 
sweeping the bottom lands should drive them to it. Geococcysx califor- 
nianus, Pipilo aberti and Toxostoma crissale are birds which probably 
do not often cross the river under normal circumstances. 

Among mammals, carnivores are usually of much wider foraging 
range than rodents. From all the data available it appears that none 
of the carnivores, not even the cats, are averse to swimming the river 
if need be. Among rodents, however, our work showed a number 
of cases in which the Colorado River had effectively checked the 
distribution of species. The following tables show the situation as 
regards all the rodents of the region (see also figs. A, B). 


1914] Grinnell: Mammals and Birds of the Colorado Valley 101 


RODENTS IN WHICH THE COLORADO RIVER ACTS AS AN ABSOLUTE BARRIER 


Association Arizona side California side Association 


Encelia and Ammospermophilus 
Rocky Creosote l harrisi harrisi 


Ammospermophilus ) Encelia and 
leucurus leucurus j Rocky Creosote 


Peromyscus ecrin- 


itus stephensi paces? 


Thomomys chryso- | & 
Sandy Creosote S y y az 
: l notus i 
Ss 
s |Thomomys albatus \ Saltbush 
S J 
S 
O |Perognathus for- )Encelia and 
mosus { Rocky Creosote 
Encelia and Perognathus in- 


Rocky Creosote termedius 


Perognathus spina- )Eneelia and 
atus spinatus { Rocky Creosote 


RODENTS WHICH OccuR ON BoTH SIDES OF THE COLORADO RIVER, BUT WHICH SHOW 
Sticut, ALMOST IMPALPABLE, DIFFERENCES ON THE TWO SIDES 


Association 


Citellus tereticaudus tereticaudus Sandy Creosote and Saltbush 
Dipodomys deserti deserti Sandy Creosote and Saltbush 
Perognathus penicillatus penicillatus Saltbush and Sandy Creosote 


RODENTS WHICH ARE APPARENTLY IDENTICAL ON THE TWO SIDES OF THE COLORADO 


RIVER 

Association 
Castor canadensis frondator River 
Peromyscus maniculatus sonoriensis All Riparian 
Peromyscus eremicus eremicus Saltbush and Sandy Creosote 
Sigmodon hispidus eremicus Willow and Tule 
Reithrodontomys megalotis deserti Tule and Willow 
Neotoma albigula venusta Mesquite and other Riparian 
Neotoma intermedia desertorum Enecelia 
Ondatra zibethica pallida River 
Dipodomys merriami merriami Saltbush and Sandy Creosote 
Perognathus bombycinus Sandy Creosote 
Lepus californicus deserticola Creosote and Saltbush 


Sylvilagus auduboni arizonae Quailbrush and Mesquite 


102 University of California Publications in Zoology [Vou. 12 


The fact is apparent that only members of the strictly desert asso- 
ciations are stopped at the river. And of these the species of the 
most remote associational position are, with one exception, Neotoma 
intermedia desertorum, most effectively delimited. Also degree of 
isolation is in a measure commensurate with amount of difference 
between forms of the same genus. 

It is pertinent to inquire how the Colorado River acts as a barrier 
to those species affected. It appears that in every one of the eleven 
cases the animal in question has no need to visit any water-supply. All 
are species capable of maintaining successful existence without a drop 
of water other than that obtained by chemical elaboration from their 
food. In our three months’ experience we did not once find evidence 
that any individual of any of the eleven species in question had visited 
the river’s edge. 

Furthermore, to the best of our knowledge, all the species are of 
limited foraging range. In the case of the two diurnal chipmunks, 
Ammospermophilus harrist harrisi and Ammospermophilus leucurus 
leucurus, which could be seen, it was seldom that an individual was 
come upon more than fifty yards from its burrow. In the ease of 
Perognathus, which carefully closes the mouths of its burrows for the 
day, after its night’s activity abroad, it was impossible to secure 
definite information on this score except as afforded by trapping; 
but the writer’s impression is that it, too, does not ordinarily venture 
many rods from its retreat. Individuals doubtless travel farther at 
times of rutting, but it is likely that even then the limits of the 
native association would not be far transgressed. 

It is further to be noted that those species finding an insuperable 
check at the river are all closely confined to one general kind of 
associational environment, even though two minor associations, as here 
defined, be oceupied. The river plus intervening associations of an 
unfavorable nature constitutes the total barrier to the rodents in 
question. 

It is true that the element of distance here implied is reduced to 
a negligible quantity where hills closely abut upon the river channel. 
But the major part of the river’s course, probably four-fifths of it 
below the lower end of the Grand Canon, is through valleys of vary- 
ing width, oecupied by riparian associations most adverse in essential 
ecological particulars to the species of the upland deserts adjacent. 

Along the remaining fifth of the river’s course, where the banks 
rise abruptly and are continuous with the adjacent hill slopes, with 


1914] Grinnell: Mammals and Birds of the Colorado Valley 103 


either no trace of riparian tracts or only narrow or interrupted repre- 
sentations, it would seem that chance 7s afforded for such mammals as 
the desert chipmunks and pocket mice to encounter the river itself, 
with all intervening factors removed. 

At our base camp (no. 4) at The Needles, our measurements 
showed the actual width of the stream to be 450 feet at the rather low 
stage of water obtaining at that time (March 4, 1910). At high water 
the river could not have been much more than 150 feet wider (see 
sectional profile, fig. A, and pl. 10, fig. 15). In the box eafion two to 
three miles below, the width of the river appeared to be much less, 
and, because of the precipitous walls, high water would make little 
change in width. 

The two species of Ammospermophilus were seen at points only 
about 850 feet apart in a direct line. All the mammals of the Encelia 
association, as segregated here on the two sides of the river, were 
trapped at this station within one thousand feet of one another. The 
same situation obviously held at several other points along the Colo- 
rado River. The sharp separation of the ranges of nearly related 
vertebrates by a barrier of such narrow width is, to the best of the 
writer’s knowledge, not known elsewhere in North America. 

As to the opportunities for crossing the river by such individuals 
as might get to the water’s edge, only speculation is now possible. 
At times of rising water, riparian mammals are undoubtedly often 
marooned upon islands and finally foreed to swim or to take refuge 
on floating drift. Practically all the riparian species are known to be 
able to swim readily, and are probably in ways just mentioned fre- 
quently carried from side to side of the river. 

The ability of typical desert animals to eare for themselves if east 
into the water is problematical, though a little experimentation would 
go far to proving the point one way or the other. Their powers in 
this line may be inferred to be limited because of the facilities normally 
lacking for putting such powers into practice. For instance, there is 
fair probability that a Reithrodontomys could safely cross a turbulent 
stretch of current, where a Perognathus would perish before any 
chance of reaching the shore. 

Protracted observation along the river brings conviction to the 
observer that no animal of weak swimming powers is likely to sur- 
vive many minutes of exposure to the main current. It is a fas- 
cinating diversion to watch the course of a stick or log adrift in 
the stream. Such an object pursues an exceedingly devious course. 


104 University of California Publications in Zoology — [ Vou. 12 


It may be carried close under the steep outside bank of an ox-bow 
swing, only to be directly thrown back towards the opposite shore. 
Stretches of rough water may be encountered where the object is 
swamped at the crest of every wave. Or, along rocky parts of the 
channel, swirls, large and small, arrest its passage. In the most violent 
of these eddies a twenty-foot log was seen to up-end and sink from 
sight, to reappear after its total submergence, a hundred yards down 
stream. 

At times of falling water a great deal of drift lodges on mudbars 
and projecting reefs of rock. It is imaginable that drift logs might 
be reached by individuals, which freed again with subsequently rising 
water, would carry their passengers until lodged under favoring cir- 
cumstances on the opposite side of the river. In the account of 
Neotoma intermedia desertorum, the only rodent of the Encelia asso- 
ciation not checked by the river, it is suggested that in some such 
way passage was secured from the California to the Arizona side of 
the river. This wood rat now bids fair to occupy much appropriate 
territory in southwestern Arizona not previously possessing an asso- 
ciational homologue, that is a Neotoman representative. Neotoma may 
be looked upon as a more hardy and ecologically less spécialized rodent 
than any of its associational companions. It is certainly much the 
largest, and is notoriously of aggressive disposition as a forager. 

Of the eight species of delimited rodents, not one individual of the 
hundreds trapped was found on the ‘‘wrong’’ side of the river. As 
far as they went, then, our efforts furnished no evidence that even 
an occasional individual does get across. As already shown, there seems 
to be nothing to attract the upland rodents to the water’s edge, so 
that possibility of securing safe transportation on a log or mass of 
drift is doubly remote. Now, supposing that a single individual did 
manage to reach the opposite shore, its species would not necessarily 
be established there. In most cases (not, however, with Peromyscus 
crinitus stephensi and Perognathus formosus) there is already estab- 
lished an associational homologue, with which even a whole family of 
the invaders would have to compete, with the chances at least as much 
against success as favoring it. Hybridization might occur, granted 
that no sexual antipathy arise, but, whatever the immediate results, it 
is the impression of the writer that swamping would eventually be 
likely to wipe out all trace of the invading species. This impression 
is admittedly based upon fragmentary data which has not been sub- 
jected to critical analysis. Whether or not Mendelian behavior in 


1914] Grinnell: Mammals and Birds of the Colorado Valley 105 


inheritance of characters obtains among the rodents here concerned is 
yet to be proven. 

Suffice it to say that all the evidence at hand shows the Colorado 
River to have effectually blocked distribution, in the two directions 
concerned in the eight cases as listed. While this hindrance to distri- 
bution involves the species, it does so through its mechanical action 
upon the frontier individuals of each species. Hypothetically the 
invaders are severally hurled back or else destroyed outright. 

The divergent characters displayed by the upland rodents of the 
two sides of the Colorado River are, in the mind of the writer, to be 
best explained on historical grounds. It is not necessary to believe that 
the specific characters concerned arose in the immediate vicinity of 
the river, though the circumstance of segregation alone is deemed by 
some to suffice as a cause of differentiation. The climatic features 
(zonal and faunal, as well as associational) are identical on the two 
sides of the river. Rather is it reasonable to presuppose separate and 
rather remote centers of differentiation, and convergent dispersal 
through time and space which brought the resulting types to the 
verge of the river, beyond which they were unable to spread. 

It is possible that an arm of the sea continuous with the Gulf of 
California once extended northward into southern Nevada. A sub- 
mergence of only 1,000 feet would divide the present desert areas of 
western Arizona and southern California into two peninsular land 
masses, which might have served as well-isolated centers of differentia- 
tion for various forms which later spread with the elevation of the 
land until their ranges abutted. Unfortunately for this suggestion, 
as I am informed by Professor John C. Merriam, geological evidence 
fails so far to show the existence of such conditions within Pleistocene 
or even Pliocene times. The suggested explanation must therefore be 
discarded in our dealing with the differentiation of present-day species 
and subspecies, especially since even the genera represented and as 
now restricted are not known to have evolved so early as Miocene. 

But another process, recognizable far and wide in dynamic z00- 
geography, may be called into account without assuming any departure 
in the past from topographic and climatic conditions as they are today. 
Comparison of the fauna of the Lower Sonoran plains of south central 
Arizona with that of the Mohave desert plateau in the same zone, 
shows two prevalent character combinations among the nearly related 
component species. The mid-Arizona representatives are usually dark 
colored and large sized; the reverse appearing to hold in the majority 


106 University of California Publications in Zoology — [Vou. 12 


of the Mohave desert forms. There is considerable floral difference in 
the two regions, and minor climatie differences are well known. Dif- 
ferent environments thus impinge upon the animals in these widely 
separated centers, and more or less regular blending of conditions 
oceurs between. Although every factor of environment may be iden- 
tical immediately on the two sides of the Colorado River, the animals 
now there have undoubtedly descended from ancestral lines which have 
invaded the territory from the two opposite directions, bringing with 
them by inheritance the characters developed under the two different 
sets of conditions. 

To express the idea otherwise, from each differentiation area there 
is an outwardly radiating dispersal of descent-lines, involving time as 
well as space. This dispersive process is going on now as it has 
through past time. The eastward-flowing tide of Mohave forms would 
only be arrested by an insuperable barrier, such as the Colorado River. 
The westward invading descent-lines from the Arizona center would 
proceed until stopped by the same barrier. Both sets of forms would 
find themselves along the Colorado Valley under the same associational, 
faunal and zonal conditions; but each set is continually receiving by 
the process of inheritance plus invasion the peculiar characters gen- 
erated on its own side. 

While the Colorado River probably lies in an intermediate position 
between the Mohave and Arizona faunas, the area of intermediate 
conditions of environment is probably relatively narrow. This very 
element of narrowness may be called in to account for the lack of modi- 
fication displayed by the delimited species of the Colorado River fron- 
tiers, for example, in the case of Ammospermophilus. 

Supposing now that the Colorado River does not serve as an 
insuperable barrier, nor ever has done so; invasion would have extended 
from one side to the other as far as associational, faunal or zonal 
barriers permitted. In animals of wide distribution, intergradation 
geographically between the remote extremes would in the end be 
expected to occur. The extremes would not then have differentiated 
so far, at least in quantity of each character developed, because of 
inheritance from the opposite type, again involving time and space, 
concomitantly. Subspecies would have resulted, instead of full species. 
This condition doubtless obtains in some of the birds, as well as in 
some of the rodents listed as being the same on the two sides of the 
river. Take, as an example, Dipodomys merriami merriami of south 
central Arizona, and Dipodomys merriami simiolus of the Mohave 


1914] Grinnell: Mammals and Birds of the Colorado Valley 107 


desert. Our Colorado River series is fairly intermediate between the 
extreme types, though the extremes are not so different as are Ammo- 
spermophilus harrisi harrisi and Ammospermophilus leucurus leu- 
curus. As elsewhere explained, the river is believed to be not so 
much of a barrier to Dipodomys merriami as to the forms of Ammo- 
spermophilus. 

The degree of hindrance, ranging from the condition, as in the 
ease of Castor and Ondatra, where the river offers no bar to perfect 
freedom of crossing, to that where the river is an absolute barrier, 
as in the case of species of Perognathus and Ammospermophilus, 
accords so closely with degree of difference in characters developed on 
the two sides of the river, that adequate ground is afforded for the 
belief that intervention of barriers is a prime factor in the differentia- 
tion of species. And furthermore, it would appear that no two 
species, in birds and mammals, arise except through geographic segre- 
gation. 


THE PROBLEM OF BARRIERS WITH REGARD TO BIRDS 
AND MAMMALS 


The geographical range of any species of animal may be likened to 
a reservoir of water in a canon. The confining walls are of varying 
nature. A concrete dam, absolutely impervious, may retain the water 
at one end. Along either side the basin’s walls differ in consistency 
from place to place. The substratum varies in porosity, at some 
points impervious like the dam, at others permitting seepage of water 
to a greater or less distance from the main volume. The water con- 
tinually presses against its basin walls, as if seeking to enlarge its 
area. And it may succeed in escaping, by seepage through such por- 
tions of its barrier as are pervious or soluble, or by free flow through 
a gap in the walls, if such offers. The area occupied by the water 
will extend itself most rapidly along the lines of least resistance. 

Every species has a center or centers of abundance in which favor- 
ing conditions usually give rise to a rate of reproduction more than 
sufficient to keep the critical area stocked. A tendency to oecupy a 
larger space results because of competition within the species, and 
individuals and descent-lines multiply and travel radially, extending 
those segments of the frontier where least resistance is offered. Such 
radial dispersal takes place slowly in some directions, more rapidly 
in others, according to the degree of passability of the opposing bar- 


108 University of California Publications in Zoology | Vou. 12 


riers. These barriers consist of any sort of conditions less favorable 
to the existence of the species than those in the center of abundance. 

Theoretically, sooner or later and in all directions, every species 
is absolutely stopped. But as a matter of undoubted fact most barriers 
are continually shifting, and the adaptability of the animals them- 
selves may be also undergoing continual modification; so that perfect 
adjustment is beyond the limits of possibility so long as topography 
and climate keep changing. The ranges of species may thus be con- 
stantly shifting. Descent-lines may move about repeatedly over the 
same general region, like sparks in the soot on the back of a brick 
fireplace. 

Yet, in all of our studies, of but a few years’ duration, the time 
element is reduced almost to a negligible quantity, and we may look 
upon the areas occupied by each species as, for the time of our 
observation, fixed. We are thus enabled to compare one with another, 
and because of the large number of the species, we can infer a good 
deal as to the nature of barriers in general as regards birds and mam- 
mals. It is even conceivable that with sufficient refinement in methods 
the inquirer may in time find himself able, from a comparative study 
of the ranges of rodents, for example, to establish the identity of all 
of the external factors which have to do with the persistence of each 
species ; in other words to analyze the ‘‘environmental complex’’ into 
its uttermost elements as regards the existing species of rodents in 
their recent development. 

The most obvious kind of barrier to distribution is that consisting 
of any sort of physical, or mechanical, obstruction. Such obstruction 
affects directly the individuals of a species encountering it, either by 
stopping their advance, or by destroying outright such as attempt 
to cross it. As barriers of this nature are to be cited land to purely 
aquatic mammals, and bodies of water to purely terrestrial, especially 
xerophilous, mammals. In each case width of the barrier has to do 
with degree of impassability. Oceans and continents are most perfect 
and affect a large proportion of the species. The comparatively nar- 
row Colorado River is a barrier of the first rank, but only to a certain 
few desert rodents, as pointed out in a preceding chapter. Mechanical 
barriers, where they exist at all, are clearly recognizable. 

It is to be observed, however, upon considering the birds and mam- 
mals of a whole continent, that by far the greater numbers of species 
are delimited in range irrespective of any direct dependence upon 
actual land and water boundaries; more explicitly, their ranges fall 


1914] Grinnell: Mammals and Birds of the Colorado Valley 109 


far short of coast lines. The barriers here concerned are intangible 
but nevertheless powerful. By their action the spread of species, 
genera, and families is held in check as surely as by any tangible 
obstruction. 

By these invisible barriers the individual may not necessarily be 
stopped at all, as with animals of free locomotion; but the species is 
affected. For example, the mockingbird (Mimus polyglottos leucop- 
terus) in its Californian distribution is closely confined to those parts 
of the state possessing certain definite climatic features; but vagrant 
individuals, especially in autumn, occur beyond the limits of these 
restrictive conditions. Carnivorous mammals are well known to be 
subject to sporadic wanderings on the part of individuals, but the 
species is kept in set bounds by some potent but invisible set of factors. 
The very fact that ¢ndividuals are quite capable of temporarily trans- 
eressing these bounds and yet do not overstep them en masse 
emphasizes all the more the remarkable potency of this category of 
barriers as regards species and higher groups. 

Our geographic studies lead us to designate among these relatively 
intangible barriers: (1) increase or decrease in prevailing temperature 
beyond certain critical limits, according to the species concerned; (2) 
increase or decrease in prevailing atmospheric humidity beyond cer- 
tain limits; (3) modification in food-supply and appropriate breeding 
and foraging ground as regards the inherent structural equipment of 
each animal considered. In these three sorts of barriers will be recog- 
nized the zonal, fawnal and associational delimitation as discussed in 
previous chapters. 


CLASSIFICATION OF BARRIERS TO SPECIES AS REGARDS BIRDS 
AND MAMMALS 


Barriers: 
A. Tangible (mechanical) 
(a) Land to aquatie species 
(b) Bodies or streams of water to terrestrial species 
B. Intangible (non-mechanieal) 
(a) Zonal (by temperature) 
(b) Faunal (by atmospheric humidity) 
(c) Associational 
(1) By food-supply 
(2) By breeding places 
(8) By temporary refuges 
(Each of these three with regard to the inherent struc- 
tural characters of each species concerned). 


110 University of California Publications in Zoology {| Vou. 12 


The above categories are believed to include all the factors com- 
monly involved in the checking of the spread of species of birds and 
mammals. It is possible that inter-specifie competition may sometimes 
occur where associational homologues meet. But even here it becomes 
a matter of relative associational fitness which determines supremacy 
and consequent ultimate limits of invasion of the forms concerned. 

A mountain range is no barrier at all, per se, as frequently alleged. 
Only as it involves zonal or faunal barriers does it affect distribution. 
The same is true of a valley or a desert. 

As far as contemplation of cases has gone, the writer’s experience 
has led him to believe that the outlines of the ranges of all birds 
and mammals may be accounted for by one or more of the factors 
indicated in the above analysis. And as detailed knowledge of the 
facts of geographical distribution accumulates, the delimiting factors 
become more and more readily detectable. By such a study, of com- 
parative distribution, it seems possible that the ranges of birds and 
mammals may become subject to satisfactory explanation. The 
instances included in the list of species discussed in the present paper, 
when considered in connection with many similar ones, point without 
exception towards the existence of the set of factors above specified 
as delimitors. 

When considered in its historical bearing, the problem of barriers 
concerns itself intimately with the origin of species. It is believed by 
the writer that only through the agency of barriers is the multiplica- 
tion of species, in birds and mammals, brought about. 


CHECK-LIST OF THE BIRDS 


Gavia immer (Briinnich) 

Sterna forsteri Nuttall 

Phalacrocorax auritus albociliatus Ridgway 
Pelecanus erythrorhynchos Gmelin 
Mergus serrator Linnaeus 

Anas platyrhynchos Linnaeus 

Nettion carolinense (Gmelin) 
Querquedula eyanoptera (Vieillot) 
Spatula elypeata (Linnaeus) 

10. Dafila acuta (Linnaeus) 

11. Marila affinis (Eyton) 

12. Erismatura jamaicensis (Gmelin) 

13. Chen hyperboreus hyperboreus (Pallas) 
14. Plegadis guarauna (Linnaeus) 

15. Mycteria americana Linnaeus 


Sl ep GA b= SOS i 


ey es) 


1914] Grinnell: Mammals and Birds of the Colorado Valley 


16. 
17. 
18. 


bo tb b& bb bo po 
a 


ww Ww we w 
eS 


eo 
go 


39. 
40. 


42. 


Or or or or 
g fe 


ot 


SU 


yy Se Gor 


— 


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anwn»#n»nn 
PAD 


Ardea herodias treganzai Court 

Herodias egretta (Gmelin) 

Butorides virescens anthonyi (Mearns) 
Nycticorax nyecticorax naevius (Boddaert) 
Grus canadensis (Linnaeus) 

Recurvirostra americana Gmelin 

Fulica americana Gmelin 

Pisobia minutilla (Vieillot) 

Actitis macularius (Linnaeus) 

Oxyechus vociferus voeciferus (Linnaeus) 
Lophortyx gambeli Gambel 

Zenaidura macroura marginella ( Woodhouse ) 
Melopelia asiatica trudeaui (Audubon) 
Cathartes aura septentrionalis Wied 


' Cireus hudsonius (Linnaeus) 


Accipiter velox (Wilson) 

Accipiter cooperi (Bonaparte) 

Buteo borealis calurus Cassin 

Faleo mexicanus Schlegel 

Falco columbarius richardsoni Ridgway 
Faleo sparverius phalaena (Lesson) 
Pandion haliaétus carolinensis (Gmelin) 
Aluco pratineola (Bonaparte) 

Otus asio gilmani Swarth 

Bubo virginianus pallescens Stone 
Micropallas whitneyi (Cooper) 

Geococeyx californianus (Lesson) 

Ceryle aleyon (Linnaeus) 

Dryobates sealaris cactophilus Oberholser 
Sphyrapicus varius nuchalis Baird 
Centurus uropygialis Baird 

Colaptes cafer collaris Vigors 

Colaptes chrysoides mearnsi Ridgway 
Phalaenoptilus nuttalli nuttalli (Audubon) 
Phalaenoptilus nuttalli nitidus Brewster 
Chordeiles acutipennis texensis Lawrence 
Chaetura vauxi (Townsend) 

Aéronautes melanoleucus (Baird) 
Archilochus alexandri (Bourcier & Mulsant) 
Calypte costae (Bourcier) 

Tyrannus verticalis Say 

Myiarchus cinerascens cinerascens (Lawrence) 
Sayornis sayus sayus (Bonaparte) 
Sayornis nigricans (Swainson) 

Nuttallornis borealis (Swainson) 
Myiochanes richardsoni richardsoni (Swainson) 
Empidonax difficilis difficilis Baird 
Empidonax trailli trailli (Audubon) 
Empidonax hammondi (Xantus) 
Empidonax wrighti Baird 

Empidonax griseus Brewster 

Pyrocephalus rubinus mexicanus Selater 


111 


2 


University of California Publications in Zoology [ Vou. 12 


68. Corvus corax sinuatus Wagler 

69. Molothrus ater obseurus (Gmelin) 

70. Xanthocephalus xanthocephalus (Bonaparte) 
71. Agelaius phoeniceus sonoriensis Ridgway 

72. Sturnella neglecta Audubon 

73. Icterus cucullatus nelsoni Ridgway 

74. Teterus bullocki (Swainson) 

75. Euphagus eyanocephalus (Wagler) 

76. Carpodacus mexicanus frontalis (Say) 

77. Astragalinus psaltria hesperophilus Oberholser 
78. Astragalinus lawrencei (Cassin) 

79. Passer domesticus Linnaeus 

80. Poceetes gramineus confinis Baird 

81. Passereulus sandwichensis nevadensis Grinnell 
82. Passereulus sandwichensis alaudinus Bonaparte 
83. Chondestes grammacus strigatus Swainson 

84. Zonotrichia leucophrys leucophrys (Forster) 
85. Zonotrichia leucophrys gambeli (Nuttall) 

86. Spizella passerina arizonae Coues 

87. Spizella breweri Cassin 

88. Spizella atrogularis (Cabanis) 

89. Juneo oreganus thurberi Anthony 


90. Amphispiza bilineata deserticola Ridgway 
91. Amphispiza nevadensis nevadensis (Ridgway) 
92. Melospiza melodia fallax (Baird) 

93. Melospiza melodia saltonis Grinnell 

94. Melospiza lineolni lincolni (Audubon) 

95. Pipilo maculatus curtatus Grinnell 

96. Pipilo aberti Baird 

97. Oreospiza echlorura (Audubon) 

98. Zamelodia melanocephala melanocephala (Swainson) 
99. Guiraeca caerulea lazula (Lesson) 

100. Passerina amoena (Say) 

101. Calamospiza melanocorys Stejneger 

102. Piranga ludoviciana (Wilson) 

103. Piranga rubra cooperi Ridgway 
104, Petrochelidon lunifrons lunifrons (Say) 
105. Hirundo erythrogastra Boddaert 

106. Iridoproene bicolor (Vieillot) 

107. Tachyeineta thalassina lepida Mearns 

108. Stelgidopteryx serripennis (Audubon) 

109. Bombyeilla garrula (Linnaeus) 

110. Phainopepla nitens (Swainson) 

111. Lanius Iudovicianus execubitorides Swainson 
112. Vireosylva gilva swainsoni (Baird) 

113. Lanivireo solitarius cassini (Xantus) 

114. Vireo belli arizonae Ridgway 

115. Vermivora luciae (Cooper) 

116. Vermivora ruficapilla gutturalis (Ridgway) 
117. Vermivora celata celata (Say) 

118. Vermivora celata lutescens (Ridgway) 

119. Dendroica aestiva sonorana Brewster 


1914] Grinnell: Mammals and Birds of the Colorado Valley 113 


120. Dendroica aestiva brewsteri Grinnell 

121. Dendroica aestiva rubiginosa (Pallas) 

122. Dendroica auduboni auduboni (Townsend) 
128. Dendroica nigrescens (Townsend) 

124. Dendroica townsendi (Townsend) 

125. Dendroica occidentalis (Townsend) 

126. Oporornis tolmiei (Townsend) 

127. Geothlypis trichas scirpicola Grinnell 

128. Geothlypis trichas occidentalis Brewster 
129. Ieteria virens longicauda Lawrence 

130. Wilsonia pusilla pileolata (Pallas) 

131. Wilsonia pusilla chryseola Ridgway 

132. Anthus rubescens (Tunstall) 

138. Oreoseoptes montanus (Townsend) 

134. Mimus polyglottos leucopterus (Vigors) 
135. Toxostoma crissale Henry 

136. Heleodytes brunneicapillus couesi (Sharpe) 
137. Salpinctes obsoletus obsoletus (Say) 

138. Catherpes mexicanus conspersus Ridgway 
139. Thryomanes bewicki eremophilus Oberholser 
140. Troglodytes aédon parkmani Audubon 
141. Telmatodytes palustris plesius (Oberholser) 
142. Auriparus flaviceps flaviceps (Sundevall) 
143. Regulus ealendula cineraceus Grinnell 

144. Polioptila caerulea obseura Ridgway 

145. Polioptila plumbea (Baird) 

146. Hylocichla ustulata ustulata (Nuttall) 
147. Hylocichla guttata guttata (Pallas) 

148. Hylocichla guttata nanus (Audubon) 

149. Planesticus migratorius propinquus (Ridgway) 
150. Sialia mexicana occidentalis Townsend 


GENERAL ACCOUNTS OF THE BIRDS: 
RECORD OF SPECIMENS, DISTRIBUTION, VARIATION, 
BIOGRAPHICAL NOTES 


Gavia immer (Briinnich) 
Common Loon 


There is in the Museum an adult-plumaged female specimen (no. 
6403) of this species taken (probably by W. W. Holder) at ‘‘ Mineral 
City in Colorado River’’ (—Ehrenberg) April 4, 1864. 


114 University of California Publications in Zoology — | Vou. 12 


Sterna forsteri Nuttall 


Forster Tern 


Adult male (no. 12616), secured May 4 at the mouth of the Gila 
River, Arizona, near Yuma; clearly a transient. 


Phalacrocorax auritus albociliatus Ridgway 


Farallon Cormorant 


Cormorants were not noted until the vicinity of Laguna Dam was 
reached, and then only in small numbers. April 24 one was seen 
at an overflow pond on the California side above Potholes; and on the 
same date a small flock was observed in the distance flying up the 
river, close over the water. Four individuals were seen below Pot- 
holes, April 29. 

A lone individual (no. 12617) was shot on the river, nearest the 
California shore, five miles northeast of Yuma, May 2. In this speci- 
men the anterior lower parts forward to the chin are lheht drab 
with numerous black feathers scattered uniformly through; naked por- 
tion of gular pouch bright orange yellow; wing 322 mm., tail 164, 
tarsus 65, middle toe with claw 100, culmen 56, depth of bill at base 
17.7. Although immature, this bird is unequivocally referable to 
P. a. albociliatus. 


Pelecanus erythrorhynchos Gmelin 
White Pelican 
Although reported to be extremely numerous at times along the 
river, we saw only stragglers. One was seen March 22 on a mud 


bar in the river above Blythe; April 3 one was noted opposite Cibola 
flying down the river. 


Mergus serrator Linnaeus 
Red-breasted Merganser 
February 18 two small flocks were seen a few miles below Needles; 


March 8 three individuals flew past our boat as we entered Cheme- 
huevis Valley. 


1914] Grinnell: Mammals and Birds of the Colorado Valley 115 


Anas platyrhynchos Linnaeus 
Mallard 
Encountered sparingly: a pair found at a lagoon in Chemehuevis 
Valley, Mareh 10, and female adult (no. 12618) secured; two seen 
at a tule pond near Ehrenberg March 25; two seen in flight along 
the river within two miles above Yuma, May 5. 


Nettion carolinense (Gmelin) 
ireen-winged Teal 


Most numerous of the ducks wintering in the region. One of two 
females (nos. 12619, 12620) taken from a small flock February 26 
near Mellen has scattered through the plumage of the lower surface 
from chin to crissuam many feathers, the exposed terminal portions 
of which are brightly stained with a rusty color. The species was 
further noted in Chemehuevis Valley March 10, near Riverside Moun- 
tain March 17, and opposite Cibola April 4. 


Querquedula cyanoptera (Vieillot) 
Cinnamon Teal 
An adult female (no. 12621) taken on the California side opposite 
Cibola April 3. 
Spatula clypeata (Linnaeus) 
Shoveler 


Four observed on a mud bar opposite Cibola, April 4; one obtained : 
female adult, no. 12622. 


Dafila acuta (Linnaeus) 
Pintail 
One obtained March 10 in Chemehuevis Valley: adult female, no. 
12623. 
Marila affinis (Eyton) 
Lesser Scaup Duek 


Two large flocks seen flying up the river above Ehrenberg March 
24; one noted on a mud bar opposite Cibola April 4. 


116 University of California Publications in Zoology | Vou. 12 


Erismatura jamaicensis (Gmelin) 
Ruddy Dueck 


Four noted on a sand bar in the river four miles above the Laguna 
Dam, April 23; one secured: immature male, no. 12624. 


Chen hyperboreus hyperboreus (Pallas) 


Lesser Snow Goose 
February 16 two small flocks were seen at Needles flying south; 
February 23 a flock was met with on a mud bar on the California 
side between Needles and Mellen. 


Plegadis guarauna (Linnaeus) 
White-faced Glossy Ibis 


A dozen seen May 5 flying down the river along the Arizona shore 
four miles above Yuma. 


Mycteria americana Linnaeus 


Wood Ibis 


A flock of twelve individuals seen April 21 flying up the river, at 
a point about four miles above the Laguna Dam. Said by residents 
of the region to occur abundantly along the river at the present time, 
just as recorded in the ’60’s by Coues (1866, p. 96) and Cooper (1869, 
p. 481). 


Ardea herodias treganzai Court 
Pallid Blue Heron 


Abundant resident along the whole course of the river as far as 
explored. Nesting colonies were observed in trees at many points 
through the large valleys; and one group of nests was noted on a 
pinnacle of rock in the narrow cafion just below The Needles. Ordin- 
arily nests were placed in the tips of the largest cottonwoods in the 
neighborhood. Special predilection was evinced for dead trees stand- 
ing close to the river. This would seem to be because of the clear 
fly-way afforded to and from the nests; and because of the more 
extensive outlook possible. But there were in this region drawbacks 
to these advantages. 

In a number of places the river was rapidly undercutting the 
outside curve of its bank, on which stood oceupied nest trees. Though 


1914] Grinnell: Mammals and Birds of the Colorado Valley 117 


we did not witness such a catastrophe in the few minutes consumed 
in floating by such places, many trees were on the verge of toppling; 
the annual mortality from this local condition alone must be large. 
Evidently the herons have failed to grasp the situation, and cling 
to old habits, even though these entail considerable annual loss. The 
area in which this destruction occurs is so small compared with the 
entire range of the pallid blue heron, and the proportion of the local 
birds which suffer disaster to their nests is also so small, that the 
contingency in question has not effected any change in the habits of 
the birds. 

Ten miles below Ehrenberg, on the California bank of the river, 
was a colony which was visited on March 30. There were approx- 
imately thirty nests, one to three per tree, which were unquestionably 
occupied, besides others in various stages of construction or dilapida- 
tion. The place had evidently been inhabited for at least one year 
previously. One nest, situated forty feet above the ground in a cotton- 
wood, contained three fresh eggs. The time of laying in this region 
is thus indicated. 

Inspection of the nests of a colony on the Arizona side four miles 
above Laguna, April 25, showed young perhaps one-third grown. They 
could be seen from the ground clambering about the nest: platforms. 
Another colony, on the Arizona shore five miles northeast of Yuma, 
‘passed May 2, showed young appearing conspicuously above the nest 
rims, and their hoarse calls were to be heard to a considerable distance. 

Along the whole course of the river, save in the rock-walled box 
canons, blue herons were almost continually in sight. Their chief for- 
aging grounds were the mud bars traversed by shallow diversions of 
the river. The habit of the river of having frequent periods of falling 
water, even when, as in the spring, the aggregate tendency is to rise, 
results in the stranding of many fishes in the shallow overflows as 
the water seeps away or evaporates. This frequently recurring supply 
of fish appears to be the chief source of food of all the species of 
herons occurring in the region. The stomach of one blue heron con- 
tained a semi-liquid mass of fish, identifiable from the large-sized 
scales as carp; another contained a large catfish. One stomach was 
empty save for a single grasshopper leg; this gives a clue as to an 
emergency diet, when the river is rising rapidly. It may be remarked 
that the opacity of the moving water of the main stream is so complete 
as effectually to prevent fishing here by piscivorous birds in the usual 
manner. 


118 University of California Publications in Zoology [| Vou. 12 


Twelve specimens of this heron were secured, two (nos. 12625, 
12626) from the California side near Riverside Mountain, one (no. 
12627) from the Arizona side at Ehrenberg, and nine (nos. 12628— 
12636) from the California side about ten miles below Ehrenberg. 
In the accompanying table of measurements of the fourteen appar- 
ently mature specimens a wide variation will be noted. While the 
largest individuals of the series are males, some females are larger 
than some males. I find nothing to indicate that size increases with 
age, though this might fairly be expected. The average of the series 
accords closely with the measurements given by Court (1908, p. 292) 
for Ardea herodias treganzai (except as resulting from obviously dif- 
ferent methods of measuring, as with middle toe). 

The status of the great blue herons residing in California west of 
the Sierran divide has been recently settled by Oberholser (1912, p. 
550). As compared with these all of the Colorado River skins are 
markedly pale throughout, with reduction in extent of dark areas. 
Bills straw yellow, darkening on culmen, not ‘‘black,’’ as stated by 
Court (1908, p. 291). : 

The weights of two freshly killed males were 514 and 534 pounds, 
of two females 5 pounds each. 


MEASUREMENTS IN MILLIMETERS OF ARDEA HERODIAS TREGANZAI 
FROM THE LOWER COLORADO RIVER AND VICINITY 


Middle Toe Depth of 
Museum No. Sex Wing Tail Tarsus and Claw Culmen Bill 
12625 of 493 186 193 132 157 28.0 
12626 oA 462 162 165 123 142 26.0 
12627" 2 441 164 172 120 142 25.0 
12628 J 488 192 178 128 143 28.0 
12629 ie) 442 168 167 121 143 26.0 
12630 se) 468 172 166 114 140 26.5 
12631 J 466 169 180 129 149 26.8 
12682 2 465 175 173 116 136 26.0 
12633 ref 478 176 180 126 150 28.4 
1263 3 492 184 190 129 154 29.0 
126385 ©) 453 167 162 121 141 25.6 
126386 io) 445 172 158 118 144 25.8 
1070° 2 450 170 169 114 150 25.0 
8041° d 445 174 175 124 141 28.0 
Averages 463 174 173 122.5 145 26.7 


*The plumage of this specimen is not in full breeding condition, and the 
bird may not have been full grown, in respect to flight feathers especially. 

* Pelican Island, Salton Sea, Calif., April 20, 1908. 

“Silsbee, Imperial Co., Calif., April 4, 1909. 


1914] Grinnell: Mammals and Birds of the Colorado Valley 119 


Herodias egretta (Gmelin) 
American Eeret 


Met with only in one place, the recently silted-in area above the 
Laguna Dam. About five miles north of Laguna, on the Arizona side 
and about one half mile back from the river, were extensive shallow 
lagoons sprinkled with numerous water-killed mesquites. Here herons 
found seclusion and feeding grounds. April 22 to 25 we frequently 
saw individuals of the egret, to the number of three at one time; but 
because of the deep mud and water we were unable to stalk them. 
Usually they were seen in flight from one portion of the bog to 
another ; one, however, was seen perched on a dead mesquite, preening. 

We were told that in summer, after the overflow begins subsiding, 
there is a large influx of white herons and ibises from the south. This 
would appear to be accounted for by the abundant food-supply in the 
way of fish left in the drying ponds as the river lowers. This con- 
stitutes a local condition, therefore, serving to modify the seasonal 
movements of these birds. 

There is in the Museum a skin (no. 4492) taken by J. G. Cooper 
at Fort Mohave, January 9, 1861. 


Butorides virescens anthonyi (Mearns) 
Anthony Green Heron 


First seen April 24, five miles above Laguna. Thenceforth com- 
mon all along the river below, and up to the time of our departure, 
May 15. As far as observed up to the latter date the species was still 
in migration. We found no evidences of breeding. Seven specimens 
were taken, nos. 12645-12651, from Potholes, near Yuma, and near 
Pilot Knob, all on the California side. 


Nycticorax nycticorax naevius (Boddaert) 


Black-crowned Night Heron 
Hight specimens secured (nos. 12637-12644). Three, taken March 
25 and April 20, are immatures in the streaked plumage; the rest 
are in more or less perfect adult plumage. 
Since one individual was seen February 18, near The Needles, 
it is probable that the species winters in the region, though sparingly. 


120 University of California Publications in Zoology [ Vou. 12 


March 16 one was seen below Parker. March 25 about fifty night 
herons were found at a tule pond below Ehrenberg. Of these, approx- 
imately forty were in the streaked immature plumage, only the 
relatively small remaining proportion being fully adult-plumaged birds. 
Farther down the river the species was common, being often startled 
from diurnal roosting places in the willows bordering the river. Indi- 
viduals were not seen stalking fish during the day, as did the blue 
herons; but their activity about sloughs and mud bars began with the 
dusk of evening, and, as indicated by their voices, continued all night. 

Among the adult specimens are some, no. 12643 in particular, 
which are extremely pale, almost pure white beneath, with the outer 
surfaces of the closed wings very light lavender. These are noticeably 
paler than the few specimens at hand from elsewhere. We found no 
indication, however, that the species occurs in the region otherwise 
than as a migrant and winter sojourner. 

There is in the Museum an immature specimen (no. 4496) taken 
by J. G. Cooper at Fort Mohave, March 12, 1861. 


Grus canadensis (Linnaeus) 
Little Brown Crane 


Cranes were seen daily, March 1 to 8, in northward flight past 
The Needles. <A large flock spent the night of March 9 on a mud bar 
in the river at the lower end of Chemehuevis Valley. This roosting 
eround was about midway between the high wooded banks of the river, 
and about two hundred yards from either bank. The cranes had thus 
selected a place which could not be approached except in the open, 
and were evidently on their guard all night. They were just opposite 
our camp; every now and then something would disturb them and a 
chorus of sonorous ealls and wing-flappings would ensue for a minute 
or more. 

It is believed that these migrating cranes are more likely to have 
been Grus canadensis than Grus mexicana. I believe the latter to be 
far less common, especially of late years, than generally supposed. 


Recurvirostra americana Gmelin 
Avocet 


There is in the Museum a skin, no. 7069, taken probably by W. W. 
Holder, at ‘‘Mineral City’’ (Ehrenberg), February 12, 1864. 


1914] Grinnell: Mammals and Birds of the Colorado Valley 121 


Fulica americana Gmelin 
Coot 


Noted but sparingly: Six seen on a tule pond below Ehrenberg, 
March 25. Several noted along margins of the main river near Pilot 
Knob, May 9 to 15; here they found concealment among the over- 
hanging masses of cane trailing in the water. Two specimens taken, 
nos. 12652, 12653. 


Pisobia minutilla (Vieillot) 
Least Sandpiper 


A band of eight seen February 24, and another of a dozen Febru- 
ary 27, on the Arizona shore a mile above Mellen; they were at the 
margin of an overflow pond. About a dozen were found similarly on 
the California side eight miles east of Picacho. Two specimens, nos. 
12654, 12655, were secured there April 20. 


Actitis macularius (Linnaeus) 
Spotted Sandpiper 


Frequently observed all along the river, on both sides, practically 
throughout the period of our work. First seen, two individuals, Feb- 
ruary 18, a little below Needles; last seen, six individuals, May 9, near 
Pilot Knob. In spite of the lateness of the last date, there was no 
evidence leading to the belief that the species breeds in the region. 
Two specimens, nos. 12656, 12657, taken March 20 near Riverside 
Mountain and March 31 near Palo Verde, are in winter plumage; a 
third, no. 12658, taken May 1, four miles below Potholes, is in full 
summer plumage. 


Oxyechus vociferus vociferus (Linnaeus) 
Killdeer 


Noted sparingly, not more than a pair at one time, but at a num- 
ber of points and almost throughout the season: Above Mellen, Feb- 
ruary 23; above Bill Williams River, March 14; near Riverside Moun- 
tain, March 17; opposite Cibola, April 1; eight miles east of Picacho, 
April 19; four miles above Potholes, April 23; five miles above Lag- 


122 University of California Publications in Zoology | Vou. 12 


una, April 26; Potholes, April 28; five miles northeast of Yuma, May 
2. In the last four cases the behavior of the birds at the time aroused 
the supposition that they were nesting in the respective vicinities. 
But recalling the tattler-like actions of the killdeer on occasions at 
other seasons, I do not now consider the evidence at all conclusive. 
Two killdeers were preserved, nos. 12659, 12660. 


Lophortyx gambeli Gambel 
Desert Quail 

Desert quail were numerous on both sides of the river almost con- 
tinuously down to the Picacho region. Below this they were notably 
searce, in fact apparently wanting at several of our stations. A few 
were seen at Pilot Knob. Along the whole course of the river the 
quail are close associates of the mesquite and quail-brush (Atriplex 
lentiformis). They also forage widely on the adjacent desert, espec- 
ially up the washes lined with catelaw and ironwood. But in all 
cases the river is their base, as they apparently need to drink both 
morning and evening. Stomachs of birds shot contained masses of 
mistletoe berries, and, at the time the mesquites were just coming into 
leaf, quantities of the tender green foliage of this plant. Nineteen 
specimens of the desert quail were preserved, nos. 12661-12679. The 
combined weight of three males shot February 21, five miles south 
of Needles, was sixteen ounces; of three females fifteen ounces. 

There are in the Museum four skins (nos. 4446-4449) taken by J. G. 
Cooper at Fort Mohave in 1861. 


Zenaidura macroura marginella ( Woodhouse) 
Western Mourning Dove 

First seen February 25, on the Arizona side above Mellen; a flock 
of about seventy-five were scattered out feeding on the ground among 
the creosote and bushes of Atriplex polycarpa. Doves were not again 
noted until March 18, near Riverside Mountain; but thenceforth they 
were seen at nearly every station, and sometimes in considerable num- 
bers. As elsewhere in desert regions, doves visited the water at dusk, 
arriving from a distance over the mesas. Up to May 13 in the vicinity 
of Pilot Knob, the species was still in evidence; it is probably resident 
in the region. 

Four specimens were secured, nos. 12680-12683. 


1914] Grinnell: Mammals and Birds of the Colorado Valley 123 


Melopelia asiatica trudeaui (Audubon) 
White-winged Dove 


First noted on the California side at Potholes, where, on April 29, 
one was heard in a willow thicket and finally secured (no. 12684, ~). 
At our station on the California side, five miles northeast of Yuma, 
the species was common. The hoarse note, so characteristic of this 
dove, was to be heard at almost any hour of the day from the dense 
woods close to the river. As many as three were heard at one time 
from as many directions. Careful stalking usually resulted in diseov- 
ering the performer among the uppermost branches, usually dead ones, 
of the largest cottonwood in the vicinity, surrounded by dense living 
willow and cottonwood timber. The crop of one shot here May 5 
(no. 12685, %) contained thirty-three watermelon seeds and one musk- 
melon seed. In the bottom lands on both sides of the river in the 
vicinity of Pilot Knob white-winged doves were common. One was 
taken on the California side May 12 (no. 12686, 2). This species is 
here a strict adherent to the willow association. 


Cathartes aura septentrionalis Wied 
Turkey Vulture 


First seen, near Mellen, February 25; thenceforth of daily note 
everywhere we went, both in the river bottom and far out over the 
desert. Last observed May 13 at Pilot Knob. We were continually 
bothered in our mammal trapping by these birds. Wherever meat 
bait was used and the steel traps left out during the day unsprung, 
no matter how far back under thick bushes these were placed, and so 
concealing the setting from view, the turkey buzzards were almost 
certain to get caught. Fully two dozen were thus captured, and, being 
seldom severely injured, were usually released. One, no. 12687, was 
saved as a skin, and two more, nos. 12712, 12713, as skeletons. Another 
specimen in the Museum, a skin (no. 5937), was taken probably by 
W. W. Holder at ‘‘ Mineral City’’ (—Ehrenberg), March 10, 1864. 


Circus hudsonius (Linnaeus) 
Marsh Hawk 


February 23 an adult was seen on the Arizona side and an immature 
on the California side, both between Needles and Mellen. April 2 a 
marsh hawk flew north along the California side opposite Cibola. 


124 University of California Publications in Zoology — [Vou. 12 


Accipiter velox (Wilson) 


Sharp-shinned Hawk 


Apparently a common winter visitant to the Colorado River bot- 
tom, adhering closely to the willow belt. Seen in the vicinity of 
Needles, February 16, 20, and 21; at Mellen, February 24; in Cheme- 
huevis Valley March 9; ten miles south of Cibola April 8; twenty miles 
above Picacho, April 14 and 16; and five miles above Laguna April 
24. The latter date probably indicates about the time of departure. 
Two specimens were obtained, nos. 12690, 12691. 

There is also in the Museum a skin (no. 4380) taken by J. G. 
Cooper at Fort Mohave, January 4, 1861. 


Accipiter cooperi (Bonaparte) 
Cooper Hawk 

Seen in February and March; to our surprise this proved to be also 
a breeding species of the region. It was confined almost altogether 
to the timbered bottom lands. Definite points and dates of observation 
were: Mellen, February 26; above Bill Williams River, March 13; 
eight miles east of Picacho, April 19; four miles north of Potholes, 
April 23; five miles north of Laguna, April 24; Potholes, April 29; 
and five miles northeast of Yuma, May 3. 

On April 19 a female bird (no. 12693), with two eggs, was secured 
on the California side eight miles east of Picacho. The nesting site 
was in a tract of tall willows, free from undergrowth, and closely 
paralleled by the river on one hand and an open mud flat on the other. 
The nest tree was relatively slender and leaning, its top about on a 
level with the crowns of the rest of the grove in the vicinity, and 
stood fifty yards from the river’s edge. The nest was about twenty- 
five feet above the ground, lodged among the branches of the willow 
near its top, and was a small, loosely constructed platform composed 
wholly of dry willow twigs. The two eggs it contained were fresh. 
Although the female bird had already begun to sit closely, as we were 
able to determine because the nesting site happened to be only a few 
yards from our camp at this place, the set was incomplete. Dissection 
of the bird showed that two more eggs would have been laid. 

One of the eggs of the set (Mus. no. 765) has sparse speckling 
of liver brown about the large end; both have equivocal nest stains; 
otherwise they are very pale nile blue. They are small, measuring 


1914] Grinnell: Mammals and Birds of the Colorado Valley 125 


in millimeters 45.0 by 37.2 and 45.6 by 36.5. The average for the 
species, as given by Bendire (1892, p. 195), is 49.0 by 38.5. This is 
another instance of diminution in the size of the eggs in the southern 
part of the range of a species. It is not paralleled in this case by 
decrease in size of the birds themselves. The three specimens of 
Cooper hawk obtained from the Colorado Valley (nos. 12692-12694) 
are in no respect smaller than individuals taken in the breeding season 
in the Transition zone, where large eggs are laid. The explanation 
possibly les in the demands of the developing embryo for a larger 
supply of heat-producing food in the colder zone. Even though this 
species incubates closely, the temperature of the egg after deposition 
is doubtless affected to some extent by that of the outside air. 

There is in the Museum an immature female specimen of this hawk 
(no. 4378) taken by J. G. Cooper at Fort Mohave, April 27, 1861. 


Buteo borealis calurus Cassin 


Western Red-tailed Hawk 


This hawk proved to be regularly distributed along the valley of 
the Colorado, both in its timbered portions and where it ent through 
mountains. Yet the species was not nearly as numerously represented 
anywhere along the river as it is ordinarily on the Pacifie slope of 
California. Exact places of observation were: five miles below Needles ; 
both sides of the river at The Needles; above Bill Williams River; 
Ehrenberg; ten miles below Cibola; twenty miles north of Picacho; 
four miles south of Potholes. All adults seen by us closely enough for 
determination in this regard, were in the light phase of plumage 
coloration. There is, however, in the Museum an adult female speci- 
men (no. 4372) taken by J. G. Cooper at Fort Mohave, April 3, 1861, 
which is distinctly in the dark phase. There is also a specimen (no. 
4373), same place and collector, taken March 7, 1861, which is in 
immature plumage, and at least as heavily marked as skins in like 
stage from the California coast region. 


Falco mexicanus Schlegel 
Prairie Falcon 


Two pairs were seen along the steep rock wall of the cafon imme- 
diately below the mouth of Bill Williams River. An evident aerie 
was noted some 200 feet above the water in a hole near the top of the 
face of a cliff. 


126 University of California Publications in Zoology  {Vou. 12 


Falco columbarius richardsoni Ridgway 
Richardson Pigeon Hawk 


There is in the Museum a female specimen (no. 4388) taken by 
J. G. Cooper at Fort Mohave, January 21, 1861. 


Falco sparverius phalaena (Lesson) 
Desert Sparrow Hawk 


First noted March 31, two flying north over the river ten miles 
below Ehrenberg. Next observed April 15, twenty miles above Picacho. 
Otherwise found only in the saguaro belt on both sides of the river 
within five miles above the Laguna Dam. Here the species found 
favorable nesting sites in the deserted burrows in the giant cactuses, 
made by gilded flickers and Gila woodpeckers. A set of five eggs, in 
which ineubation was far advanced, was found eleven feet above the 
eround in an eighteen-foot saguaro on the California side, April 23. 
An elf owl was taken at the same time from a hole in the opposite 
side of the trunk. The excavation in which the hawk’s eggs were laid, 
opened to the south. 

The only specimen secured by the expedition was an adult female 
(no. 12689) shot among the saguaros on the Arizona side of the 
river five miles above Laguna, April 22. This bird is markedly 
different from any one of a series of twenty-seven adult females from 
California, Nevada, Washington, Alaska, and the western United 
States generally, which series I would eall Falco sparverius sparverius. 
Yet it does not accord, particularly in size, with Mearns’s characteriza- 
tion (1892, p. 263) of Falco sparverius deserticola. As compared with 
average examples of sparverius, as represented in the series referred 
to, the specimen in question is decidedly smaller, except bill: wing 178, 
tail 113, tarsus 34, middle toe and claw 31, bill from nostril 11, culmen 
from cere 12, depth of bill 9.4; the browns of the upper and under 
surfaces are more reddish, the brown area on top of head very broad 
and pure cinnamon-rufous without trace of plumbeous shaft-streaks, 
the plumbeous of head reduced to narrow frontal and supercilary 
tracts continuous with one another, shaft-streaks of pectoral region 
narrow, barrings on dorsum very narrow, the widest only 3mm. 


instead of 7. 


1914] Grinnell: Mammals and Birds of the Colorado Valley 127 


It will thus be seen that in all essential respects this specimen meets 
with the deseription of Palco sparverius peninsularis Mearns (1892, 
p. 267), of the southern extremity of Baja California. Mearns 
expressly states in this connection that all of the sparrow hawks 
examined by him from the southern borders of California and Arizona 
are of the large ‘‘deserticolus’’ type. I might dispose of the peculiar 
Colorado River example by referring it to peninsularis without fur- 
ther ado. But I do not consider this a reasonable course, unless later 
acquisition of numerous specimens should show that the sparrow-hawk 
of the saguaro belt here and that of Lower California is normally 
like the one in hand and henee different from that of the surrounding 
regions. This I now doubt, preferring rather to consider the bird in 
question as an extreme of individual variation in the direction of 
peninsularis, but to be identified with the obvious geographic stock- 
form, F. sparverius sparverius (or F. s. phalaena Lesson [= F’. s. de- 
serticola Mearns], if a southwestern desert form be considered different 
from sparverius proper of the north and east). 

There is in the Museum a male specimen (no. 4390) taken by J. G. 
Cooper at Fort Mohave, December 24, 1860. This may or may not have 
been a winter visitant. But it, too, is small as though possibly repre- 
sentative of a resident race. It measures: wing 175, tail 111, tarsus 
31.0, middle toe and claw 31.3, bill from nostril 10.5, eulmen from 
cere 11.1, depth of bill 9.2. In coloration this specimen, also, has the 
black markings of the mantle reduced to mere indications. 


Pandion haliaétus carolinensis (Gmelin) 
Osprey 
One specimen (female, no 12688) taken April 17 on the California 
side of the river seven miles east of Picacho. Another seen flying over 
the river a mile or so above Yuma, May 5. Evidently a migrant 
through the region. 


Aluco pratincola (Bonaparte) 
Barn Owl 


Feathers of a barn owl were seen on the California side at River- 
side Mountain. Its presence was nowhere else detected until we 
reached our last station near Pilot Knob. Here in a low bluff on 


128 University of California Publications in Zoology [| Vou. 12 


the California side, where the river swings northwest against the 
mesa, a nesting site was located as we floated past on May 5. The 
place was revisited on May 10, and the hole in the partly cemented 
gravel face, fifteen feet above the water, was ascertained to contain 
four downy young. Two of these were preserved (nos. 12695, 13979). 


Otus asio gilmani Swarth 
Saguaro Sereech Owl 


The widespread presence of sereech owls was brought to our atten- 
tion by their notes heard almost nightly and at nearly every one of 
our camps from Needles all the way down the river, clear to Pilot 
Knob. The birds appeared to occur both in the willow and cotton- 
wood timber close along the river, and out on the desert. They were 
heard from the precipitous sides of The Needles, the numerous cran- 
nies in the rock walls of which may have afforded the birds daytime 
concealment. At Mellen, Arizona, February 27, a female screech owl 
(no. 12696) was captured in a trap set near the house of a wood-rat 
(Neotoma albigula venusta) under a mesquite and baited with portions 
of a wood-rat found partly eaten in the same trap the night before. 

On the California side, twenty miles north of Picacho, April 16, 
one (no. 12697) was shot at night in a willow when giving the char- 
acteristic deep-toned succession of mellow notes; it proved to be a 
male. 

April 23, on the California side four miles above Potholes, two 
were taken from holes in giant eactuses. A male (no. 12698) was 
taken from an otherwise empty eavity ten feet above the ground. A 
female (no. 12699) oceupied a eavity nine feet above the ground. 
This hole was rather small, evidently originally exeavated by the Gila 
woodpecker; but it held besides the old owl, three newly-hatched 
young (preserved as alcoholics, nos. 13976-13978), and the fresh, 
headless bodies of four mammals: two Perognathus penicillatus penicil- 
latus, one Dipodomys merriami merriami, and one Peromyscus mani- 
culatus sonoriensis. 

The four adults specified above, together with a fifth in the Museum 
(no. 4395) taken by J. G. Cooper at Fort Mohave, February 24, 1861, 
are all typical of the subspecies gilmani as described by Swarth (1910, 
1 de 


1914] Grinnell: Mammals and Birds of the Colorado Valley 129 


Bubo virginianus pallescens Stone 
Western Horned Owl 


The hooting of horned owls was heard almost nightly at every one 
of our camps, from Needles to Pilot Knob. The birds themselves were 
oceasionally seen flying overhead at dusk, and twice, on the California 
side at Riverside Mountain and opposite Cibola, were flushed in the 
daytime from dense ironwood trees in desert washes back from the 
river. Three specimens were shot (nos. 12702-12704). These are 
markedly pale, as compared with Bubo virginianus pacificus of the 
coastal slope of southern California, and in their characters closely 
approximate specimens from central Arizona which in turn have been 
referred to B. v. pallescens (see Oberholser, 1904, p. 182). 


Micropallas whitneyi (Cooper) 
Elf Owl 


The elf owl was detected only in the belt of saguaros which extends 
across the Colorado Valley within six miles above the Laguna Dam. 
The numerous excavations in the trunks of these great cactuses evi- 
dently furnish desirable diurnal retreats for owls of size small enough 
to render them usable. By chopping into many of the perforated 
saguaros we succeeded in capturing two of this species, nos. 12700 and 
12701. The latter was taken on the Arizona side, April 22, from a 
hole twelve feet above the ground. The cactus in which it was located 
was a single column thirty-one feet high and twenty-six inches in 
diameter. In the same trunk, four and one-half feet higher up, was 
an occupied nest of the gilded flicker. The other bird was taken 
April 23 on the California side four miles above Potholes. The cavity 
occupied in this instance was twelve feet above the ground in a three- 
branched saguaro eighteen feet tall and twenty-six inches in diameter. 

In the same trunk one foot lower, but opening on the opposite side, 
was a cavity occupied by a sparrow hawk and its five eggs. Both owls 
were females, and each showed signs of immediate breeding. In one 
ease dissection disclosed five large ova of graded size, the largest being 
an apparently full-sized yolk. 

The locality where our California specimen was secured is some 
two miles west of the river and not far from the old Senator Mine. 
It is in a valley leading down from the hills and locally known as 


130 University of California Publications in Zoology — [ Vou. 12 


the Senator Mine Basin. It was doubtless in the limited tract of 
saguaros here that Brown (1904, p. 46) found the species nesting on 
May 17, 1903, and secured two sets of four eggs each. We noted the 
old sears on the largest cactuses where holes had been chopped into, 
years before our visit. Brown’s was the first authentic record of the 
elf owl for California. But Cooper found the species on the Arizona 
side of the river, near Fort Mohave, whence on April 26, 1861, he 
secured the type, now in the National Museum, though at one time 
the property of the State of California (see Cooper, 1870, p. 443). 


Geococcyx californianus (Lesson) 
Roadrunner 


Safely to be considered a common resident of the whole region 
traversed. Noted at every collecting station on either side of the river, 
from Needles to Pilot Knob. Noted with equal frequeney on the 
upland mesas and hills, and on the first bottom even to the water’s 
edge. The peculiar notes were on several occasions heard from oppo- 
site shores as we floated along, as though birds were answering one 
another from across the water. 

Two examples came into our possession through being caught in 
meat-baited steel traps set for carnivorous mammals. The stomach of 
a roadrunner obtained at Needles contained remains of beetles and 
one half-grown lizard (Cnemidophorus). 

The five specimens secured by us (nos. 12705-12709), and another 
(no. 4369) taken by J. G. Cooper at Fort Mohave, March 26, 1861, 
give a uniform impression at first glance of paleness, as compared with 
roadrunners from the Pacifie slopes of southern and central California. 
This paleness consists in an apparent greater average extent of white 
markings and particularly in the whitening of the buffy lower surface 
and a paling of the buffy margins of the dorsal feathers. The Pacific 
slope series, however, includes specimens taken at all seasons of the 
year, and it becomes apparent at once that the deepest buffy specimens 
in it are those in the freshest (that is, fall) plumage. The variation 
in the Pacifie series in this respect is great, and leads to the conclusion 
that the paleness of the buffy tints of the Colorado desert series, 
which are February to May examples, is not altogether intrinsic, but 


1914] Grinnell: Mammals and Birds of the Colorado Valley 131 


due in part at least to an extreme amount of fading. This does not, 
however, account for the average greater extent of white areas on the 
individual feathers of the desert birds. But this feature is too variable 
in both series to be satisfactorily defined. 

It is of course a remarkable exception if the roadrunner, a ter- 
restrial, permanently resident bird of all the areas it inhabits, should 
not show some geographic color peculiarities. Indeed it is all the more 
strange that it does not show conspicuous differences in color tone 
in the arid and subhumid areas it occupies, when we observe the 
markedly different color tones exhibited by the thrashers, towhees, 
spermophiles, and jack rabbits of the same areas, these being also ter- 
restrial animals. 

The roadrunner’s failure to conform to the rule offers a problem 
for those who would explain animal coloration wholly on the grounds 
of physiological response to meteorological conditions, irrespective of 
adaptive value. 


Ceryle alcyon (Linnaeus) 
Belted Kingfisher 


Our observations would indicate that this species occurred only as 
a migrant through the region. The first individual was noted April 
6 on the California side below Cibola; another was seen on the seventh 
on the Arizona side ten miles below Cibola. In both these cases, and also 
subsequently, the birds were perched on snags intently watching the 
muddy current. But the futility of their scrutiny was apparent in 
that at no time was a kingfisher seen to plunge into the water in the 
characteristic fashion of the species when capturing a fish; the 
opacity of the mud-laden water was most certainly unfavorable to 
this mode of securing fish. Kingfishers, but one at a time, were seen 
April 19, eight miles east of Picacho; May 3, five miles northeast of 
Yuma; and May 5, near Pilot Knob. 

There was not the least doubt in any of these instances that the 
bird seen was the large belted kingfisher and not the little Texas green 
kingfisher. Coues’s oftquoted statement (1866, p. 59) that he observed 
the latter species ‘‘at several points on the Colorado River between 
Forts Mohave and Yuma’’ in the autumn of 1865 remains unconfirmed 


by any later observation. 


132 University of California Publications in Zoology | Vou. 12 


Dryobates scalaris cactophilus Oberholser 
Caetus Woodpecker 

Common resident of the willow and mesquite associations along 
the whole of the explored portion of the river. In but one place was 
the species seen outside of the riparian strip. The exceptional case 
was on the Arizona side above Mellen, where several individuals were 
seen over a mile up the Sacramento Wash, affecting dead or sickly 
palo verde trees. At Needles, February 17 and the few days follow- 
ing, this woodpecker was common in certain portions of the river 
bottom, where, however, its habit of working near the ground in 
young growth, together with its winter quietude, rendered it very 
inconspicuous. Within a month later, those seen along down the river 
had become noisy, and the presence of the species was readily detected, 
even as we floated along from one station to another. 

On April 11, on the California shore twenty miles north of Picacho, 
a nest was found situated twelve feet above the ground in a willow 
stub, the upper part of which was dead and centrally decayed. The 
excavation freshly made in this by the birds opened on the under side 
of this slanting terminal section. There were three eggs on the point 
of hatching, and one newly hatched young, showing the date of egg- 
laying to have been toward the last of March. 

On the California side below Potholes parts of the willow area had 
been burned over as a preliminary to clearing the land. The wood- 
peckers here were much discolored ventrally by contact with charred 
tree trunks, yet they appeared to be thriving quite as well as if arrayed 
purely in their normal colors. 

Fourteen specimens of the Texas woodpecker were secured, nos. 
12714-12727. Besides these there are in the Museum six skins (nos. 
4315-4320) taken by J. G. Cooper at Fort Mohave, December 31, 
1860, and in January and February, 1861; and another taken at 
‘*Mineral City’? (Ehrenberg) March 10, 1864. collector not recorded. 

For the use of the name cactophilus in replacing the older name 
bairdi, formerly in general use for this woodpecker, see Oberholser 
(19105 p. 152). 


Sphyrapicus varius nuchalis Baird 
Red-naped Sapsucker 


Evidently a winter visitant to the Colorado Valley, where we found 
it for a time common among the willow thickets. Localities of capture 


1914] Grinnell: Mammals and Birds of the Colorado Valley 133 
were: the immediate vicinity of Needles, opposite The Needles, and 
Lower Chemehuevis Valley. At the first two places several were seen 
that were not also taken; at the latter point but the one, taken March 
11, was noted. This probably indicates the approximate date of 
departure from the region, as we saw none thereafter. Willows were 
the trees attacked by this woodpecker; but in one ease a single large 
mesquite, and only this one out of many in the vicinity, had been 
selected for bleeding, and its main trunk and larger branches were 
plentifully bored. I visited this tree many times during the space of 
three days, March 2 to 4, opposite The Needles, and invariably found 
a sapsucker working about the borings. I shot two of the birds at this 
mesquite, and there was still one there the last time I visited the tree, 
although I had never seen but one at a time there. 

The five specimens secured (nos. 12728-12732), all from the Cali- 
fornia side, are characteristic of the subspecies. 

There is a skin in the Museum, no. 4312, taken by J. G. Cooper 
at Fort Mohave, February 20, 1861. 


Centurus uropygialis Baird 
Gila Woodpecker 


A common and characterstie resident the whole length of the region, 
from Needles to the vicinity of Yuma. Found at every station on both 
sides of the river. While regularly present in the willows and cotton- 
woods of the river bottom, the species occurred also up the desert 
washes a mile or more from the edge of the riparian strip. This was 
the ease on the Arizona side above Mellen, and on the California side 
opposite Cibola, and on the Arizona side ten miles below Cibola. At 
the latter two points Gila wooapeckers were feeding on mistletoe 
berries in the ironwoods. At the last named place, an excavation 
fifty-four inches above the ground in a dead broken-off branch of a 
palo verde contained a single fresh egg, April 7. The nest cavity 
appeared to be an old one, perhaps dug out the year previous. On 
the California side twenty miles above Picacho a nest was found 
twenty-five feet above the ground in a broken-topped living willow, 
the core of which was rotten. There were three small young, April 15. 

The Gila woodpecker was one of the birds conspicuously associated 
with the giant cactus. Where a scattering growth of this plant reached 
nearly to the river on the Arizona side east of Ehrenberg, several of 
these birds were encountered far out on the mesa and many of the 


134 University of California Publications in Zoology [| Vou. 12 


larger cactuses contained from one to four holes each. At the date 
they were examined, March 27, egg-laying had not yet commenced. 

Where the better defined belt of saguaros meets the Colorado above 
the Laguna Dam, with well represented tracts on both sides of the 
river, the Gila woodpecker abounded. The availability of satisfactory 
sites would therefore seem to be the prime factor in governing the 
local distribution of this species. For the cactus itself certainly 
would not seem to afford food, except during the fruiting season, or 
other attractions. 

In this vicinity nests were found in saguaros as follows: On the 
Arizona side, five miles above Laguna, April 24, a set of three fresh 
egos, twelve feet above the ground; on the California side, four miles 
above Potholes, April 23, nest (not investigated) twenty-four feet 
above the ground; another nest, same place and date, fourteen feet 
above the ground, with four small young. 

In the same vicinity Gila woodpeckers were nesting also in dead 
cottonwoods in the river bottom. The birds appeared to find favorable 
foraging grounds both in the riparian tracts and out on the desert. 

A series of twenty-three adult specimens was taken, nos. 12733- 
12755: 

There are five other skins in the Museum, nos. 4330-4334, taken 
at Fort Mohave by J. G. Cooper, December 24 and 25, 1860, and 
February 4 and March 12, 1861. It is to be remarked that not an 
iota of difference, save as is accountable for by extraneous causes, is 
observable between the Cooper birds obtained fifty years ago and the 
freshly obtained material. Considering the rather intricate color 
pattern and the opportunity thus afforded for the detection of deter- 
minate or any other sort of variation, this is indicative of constancy 
of characters under natural conditions through a long series of genera- 
tions. 


Colaptes cafer collaris Vigors 
Red-shafted Flicker 


Evidently a common winter visitant. Noted daily the third week 
of February in the willow timber of the river bottom below Needles, 
and early in March in the narrower willow association on both sides 
of the river opposite The Needles; likewise in Chemehueyis Valley. 
The last individual of the species observed, was shot in the willows on 


1914] Grinnell: Mammals and Birds of the Colorado Valley 135 


the California side near Riverside Mountain, March 16. Five speci- 
mens in all were secured, as listed in the table presented under the 
discussion of Colaptes chrysoides mearnsi. 

There is in the Museum another skin (no. 4325) taken by J. G. 
Cooper at Fort Mohave March 12, 1861. 


Colaptes chrysoides mearnsi Ridgway 


Mearns Gilded Flicker 


Detected by our party only in a restricted area within six miles 
north of Laguna Dam. This area of occurrence coincided with the 
westward extension across the Colorado Valley at this point of a belt 
of the giant cactus, or saguaro. Here this woodpecker was fairly com- 
mon, probably as a permanent resident. While closely restricted on 
the desert to the saguaro belt, at least two pairs were nesting in dead 
cottonwood stumps in the drowned-out area of the river bottom. A 
nesting hole located here was eighteen feet above the ground, in a 
large stub. It is probable that the species occurs, or has oceurred in 
the past, along the bottom timber up and down the river, though not 
seen by us. The year 1910 may have been one of a series of unfavor- 
able years when the range of the species was retracting through dying- 
off of the frontier individuals. We saw near Pilot Knob and at 
several places above Picacho excavations in willows and cottonwoods, 
which were too large for the Gila woodpecker, and in all probability 
were made some years before by gilded flickers. Moreover, the Museum 
contains two of Cooper’s specimens, taken at Fort Mohave, consider- 
ably above Needles, February 23 and April 2, 1861 (nos. 4328, 4329). 
Cooper (1870, p. 411) found ‘‘two pairs’’ there, in cottonwoods. 

Nests of the gilded flicker were found by us, in the saguaro belt 
above referred to, as follows: On the Arizona side, April 22, exeava- 
tion sixteen and one-half feet above the ground in cactus thirty-one 
feet high, contained two fresh eggs; April 24, excavation twenty feet 
above the ground, not investigated. On the California side, April 23, 
excavation ten and one-third feet above the ground, in cactus twenty- 
eight feet high, contained one infertile egg and two small young. The 
two parent birds taken with the latter had their gullets distended with 
a mass of small black ants and ant larvae. It was in this same place 
that Brown (1904, p. 46) found a nest of this bird in 1903. 


136 University of California Publications in Zoology {| Vou. 12 


Of the five specimens secured by us only one has the shafts and 
under surfaces of the wings and tail yellow as in the two Cooper 
specimens from Fort Mohave. Reference to literature shows that this 
yellowness is generally supposed to be a constant character of chrysoides. 
In fact I find but four recorded exceptions: Brewster (1883, p. 25) 
mentions a specimen taken at Tucson, Arizona, as “‘having the yellow 
of the wings and tail replaced by orange, while the shafts of many of 
the feathers show an even stronger reddish cast, those of the rectrices 


» Brewster remarks fur- 


at their bases being especially deep in color. 
ther that ‘‘this departure from the normal coloring undeniably nar- 
rows the gap which separates chrysoides from mexicanus [=collaris], 
but it may be merely a chance reversion, or, what is perhaps still 
more likely, the specimen in question may be a hybrid.’’ Coues (1903, 


p. 602) observes that ‘‘gradation between this form [chrysoides| and 


C. mexicanus [=collaris| has not yet been observed.”’ 

Swarth (1905, p. 27) discusses at length an aberrant specimen from 
the Papago Indian Reservation, Arizona, evidently in the same eate- 
gory as Brewster’s. Swarth considers this a hybrid between chry- 
soides and collaris, although he goes on to say that he found these two 
species in that region ‘‘breeding almost side by side, practically 


” 


without mixing,’’ and this seems strange ‘‘when we consider the 
extensive hybridization that takes place in the northwest, where 
collaris and luteus come together.’’ 

Breninger (1898, p. 13) reports two supposed hybrids between the 
‘“oilded’’ and ‘‘red-shafted’’ flickers. But these are probably of the 
same nature as Brewster’s and Swarth’s. 

The abnormal appearance of four out of five of the flickers col- 
lected by us on the lower Colorado, and the circumstance that this 
abnormality consists in a redness approximating the color of corres- 
ponding areas in collaris, led us to conclude at the time that we had 
there found chrysoides and collaris hybridizing. The chief characters 
of the specimens in question are shown in an accompanying table; also 
the corresponding characters of those examples of collaris taken by 
us in the Colorado Valley. The latter, it should be emphasized, are 
believed to have been winter visitants to the region, and not repre- 
sentative of the breeding species. 

No. 12761, male of typical yellow chrysoides, and no. 12762, female 
with under surfaces of wines and tail coral red, were a mated pair, 
the parents of the young found in a saguaro on the California side. 
These young, preserved as alcoholics, are in too early a stage for the 


1914] Grinnell: Mammals and Birds of the Colorado Valley 137 


colors in the rudiments of their feathers to be ascertained. Nos. 
12763 and 12764, both with reddish coloration, were a mated pair taken 
in the overflow bottom on the Arizona side. No. 12765, of reddish 
type, was taken on the desert on the Arizona side. 

The salient fact shown by this comparative examination is that 
in all other characters the specimens aberrant in color of wings and 
tail, are perfectly typical of chrysoides (that is, of its subspecies 
mearnsi). None of the phenomena consequent upon hybridization is 
evinced in other particulars, such as general size, proportional dimen- 
sions, extent of dorsal barring, colors of body and head. In all these 
characters there is no nearer approach of the red-shafted chrysoides 
to collaris, than of the yellow-shafted chrysoides. 

My conclusion is that the strain of chrysoides occurring at the 
present time in the lower Colorado Valley shows proneness to replace- 
ment of yellow by red, without there having been any interbreeding 
with another species. This may be accounted for chemico-physiologic- 
ally, as in the case of the linnet of the Hawaiian Islands, where, how- 
ever, the change has been from red to yellow. (See Grinnell, 1911, 
p. 191.) According to this idea there has been some cause, germinal 
or somatic, affecting the oxidation of the basic chromagen, so that in 
the critical instance the process is going beyond the stage of yellow- 
production and reaches the red-production stage. The phenomenon 
at the present time may be purely sporadic; on the other hand it may 
be an incipiency of a character which, beginning in this center of 
differentiation, may become fixed and heritable over a larger area, 
thus constituting the arising of a new species. The flickers collected 
were only a few out of the large population which probably occupies 
the extensive tract of saguaros widening out to the eastward of the 
river in Arizona. Observation in the same region at successive times 
in the future will afford data indicative of the real significance of 
the phenomenon. 

It is quite evident that the aberrant examples described by Brew- 
ster and by Swarth from central Arizona, as referred to above, are of 
the same nature as the Colorado Valley specimens. The chances are 
that they were not hybrids. So far as shown by the literature at hand, 
no unquestioned hybrids have been found between chrysoides (or any 
of its subspecies) and collaris or cafer. 


138 University of California Publications in Zoology — [Vou. 12 


MEASUREMENTS IN MILLIMETERS AND COLOR FEATURES 0F COLAPTES 
CHRYSOIDES MEARNSI FROM THE COLORADO VALLEY 


Color of 
lower surface Whole top Throat, 
Dorsal of wings and of head to fore-neck and 
No. Sex Wing Tail Culmen barring tail level of eyes sides of head 
12761 3b 152 100 40 barely chrome* bright clear gray, 
indicated yellow cinnamon no. 9 
12762 2 145 87 35 distinet but pale coral bright clear gray, 
reduced red cinnamon no. 9 
12763 3 147 91 35 slightly pale saturn bright clear gray, 
indicated red cinnamon no. 9 


12764 2 149 88 34 well-defined pale coral bright clear gray, 


though narrow red cinnamon no. 9 
12765 ©) 142 93 36 well-defined pale coral bright clear gray, 
though narrow red cinnamon no. 9 
4328" iS) 154 94 38 narrow but chrome bright elear gray, 
distinet yellow cinnamon no. 9 
43297 io) 147 91 37 well-defined chrome bright clear gray, 
though narrow yellow cinnamon no. 9 


1 Cooper's Fort Mohave specimens. 
? Color names from Ridgway's Nomenclature of Colors, 1886. 


MEASUREMENTS IN MILLIMETERS AND COLOR FEATURES 0F COLAPTES 
CAFER COLLARIS FROM THE COLORADO VALLEY 


Color of Top of 
lower surface head between Throat, 
N i$ Dorsal of wings and lighter super- fore-neck and 
No. Sex Wing Tail Culmen barring tail ciliary areas sides of head 
12756 9) 161 110 37 broad and deep coral broceoli clear gray, 
conspicuous red brown no. 6 
12757 Jd 165 108 39 broad and deep coral broceoli clear gray, 
conspicuous red brown no. 6 
12758 Jo 161 109 — broad and deep coral broccoli _ clear gray, 
conspicuous red brown no. 6 
12759 3 165 111 41 broad and deep coral broeecoli clear gray, 
conspicuous red brown no. 6 
12760 J 170 113 38 broad and deep coral broceoli clear gray, 


conspicuous red brown no. 6 


1914] Grinnell: Mammals and Birds of the Colorado Valley 139 


Phalaenoptilus nuttalli nuttalli (Audubon) 
Nuttall Poor-will 


Phalaenoptilus nuttalli nitidus Brewster 
Frosted Poor-will 


Poor-wills were first noted the evening of February 28 at Mellen; 
then March 3, at The Needles; thenceforth at every one of our stations 
all the way down the river to Pilot Knob, May 12. The mellow eall 
was heard practically every night when the wind was not blowing. 
As with the nighthawks, the poor-wills appeared to spend the day out 
on the desert. One individual was flushed near Ehrenberg from the 
stony surface of a wash in the shade of a palo verde. In the evening 
the birds appeared at dusk in the river bottom, alighting in charae- 
teristic fashion on spaces of bare ground or in roads, not infrequently 
on mud bars in the river, or skimming low over the water itself. 
Their proneness to alight on the wet mud at the edge of the water 
was evidenced by the packed balls of mud firmly dried on the toes of 
several of the birds shot. In the evening of April 19, eight miles east 
of Picacho, fully a dozen poor-wills were observed. There was a lot 
of broken-down fencing close to the river near our camp at this point; 
the birds congregated here as a basis for foraging and consociating, 
resting on the posts as well as on the prostrate poles, evidently in 
preference to alighting on the sticky mud of an overflow depression 
on the one hand or the river margin on the other. 

The series of eighteen specimens of this bird secured at once aroused 
inquiry because of the large range displayed in size and pattern of 
coloration. Even in the field, as the specimens were collected, con- 
spicuous differences were noted and the suspicion aroused that really 
two subspecies were represented, one being the resident breeding form, 
the other a winter visitant. The first clue to discrimination came 
through observing the state of activity of the reproductive organs. 
Of two birds shot the same evening at Riverside Mountain, one, no. 
12780, with testes small, was relatively large, dark, and coarsely 
marked ; the other, no. 12781, with testes very large, was small in gen- 
eral size, narrow-barred, and notably pale. In five other birds, field 
examination showed the same relative state of affairs (see acecompany- 
ing tables). It is of course regrettable that state of reproductive 
activity was not recorded of all the poor-wills taken. 

The seven birds, dissected in the field by the writer, would seem 
to provide adequate basis for the conclusion that two races were repre- 


140 University of California Publications in Zoology | Vou. 12 


sented, in spite of the fact that of the remainder of the series it is 
difficult to allocate certain examples with one or the other category. 
This has finally been done, however, as shown in the tables. The first 
character employed for separation, and the only one holding through- 
out the present series, is the style of black marking on the medial 
scapulars. In the resident form, for which I am using the name 
nitidus, these markings are narrow and more V-shaped or hastate; 
in the presumably migratory form, which I am ealling nuttalli, these 
markings are broad, sagittate to rhomboid. I am inclined, however, 
to believe that this character, like the others pointed out beyond, will 
be found to fail in some eases; for a skin at hand from the Huachuca 
Mountains, Arizona (no. 10324), is otherwise good nitidus as here 
understood, but the scapular markings are broadly rhomboid. In the 
Colorado series, believed to represent two forms, any of the remaining 
characters are not separately diagnostic in every case. But the series 
is found divisible with fair precision as indicated in the tables, by 
an aggregate of characters. 

Besides the scapular marks above specified, the black barring across 
the primaries averages narrower in nitidus than in nuttalli, and their 
transverse trend is zigzag in the former, more squarely defined in 
the latter. The transverse dark barring on the posterior lower sur- 
face averages narrower again in mitidus, coarser in nuttalli. The 
ground-color, whitish or hoary, is not only more extended in nitidus 
but of a clearer white tone. In general size nitidus is distinctly the 
smaller, and it appears to have a more rounded wing, indicative of 
less extended migratory flight. 

While in each of these characters variation leads from one extreme 
to the other, there are clearly two modes. The theory suggests itself 
as accounting for this, that there are two unstable phases in the 
poor-will, irrespective of locality, sex or age, a condition which has 
been verified in the case of certain owls. But the facts as here pre- 
sented militate against either that idea or the contention that the 
paleness develops with individual senility. All the above characters 
are paralleled in such birds as are represented by races occupying the 
deserts on the one hand and the more northern and less arid regions 
on the other. Some of the nuttalli examples are precisely like poor-wills 
in the Museum collected from Humboldt County, Nevada. No skins 
examined from anywhere north of the Colorado Valley show char- 
acters of nitidus. 


1914] Grinnell: Mammals and Birds of the Colorado Valley 141 


The inference is that the nuttalli of the Colorado Valley are winter 
visitants from the Great Basin region. That these, however, are 
identical with typical nuttalli, described from South Dakota (aceord- 
ing to A. O. U. Check-List, 1910, p. 197), is problematical. I have 
had no opportunity for comparison of relevant material. Neither is 
there assurance that the Colorado Valley nitidus is typical of the 
form bearing that name and deseribed from the Nueces River, 
southern Texas (Brewster, 1887, p. 147). In the original deseription 
of the latter, restriction of dark markings and general paleness are 
emphasized characters, which are in accord with the Colorado Valley 
nitidus. Ridgway (1887, p. 588) adds the character of small size. 
The difference in facial mottling, mentioned by both of the authors 
just cited, is not clearly in evidence in the Colorado Valley material. 

It is to be remarked that Phalaenoptilus nuttalli californicus, repre- 
sented by specimens from central and southern California west of the 
desert divide, is well differentiated from both of the forms here pointed 
out. Three races of the poor-will thus occur within the state of Cali- 
fornia, and as there are good grounds for believing that P. n. nuttalli 
summers east of the Sierran divide in northeastern California, all 
three probably breed within the limits of the state. 

In this connection sixty-seven adult-plumaged poor-wills have been 
examined. 


LIST AND DIMENSIONS IN MILLIMETERS OF PHALAENOPTILUS NUTTALLI 
NUTTALLI FROM THE COLORADO VALLEY 


12780 
12785 
12791 
12777 
12788 
12783 


* Dissection showed reproductive organs to be inactive. 


9th primary 


exceeds Wing 

Sex Locality Date Wing Tail 10th by formula 
ois Riverside Mt., Calif. side Mar. 18 145.5 89.0 5.5 9-8-1 0-7 
3 Opposite Cibola, Calif. side Apr. 92 140.0 88.2 too worn 
fe 8 mi. E. Pieacho, Calif. side Apr. 18 136.3 84.0 too worn 
Jb Chemehuevis Val., Calif. side Mar. 10 133.8 80.3 4.7 
Jb 10 mi. 8. Cibola, Ariz. side Apr al 142.4 82.0 too worn 
J Ehrenberg, Ariz. side Mar. 27 144.3 82.0 7.4 

Average of the males, 140.4 84.2 5.9 
9 8 mi. E. Picacho, Calif. side Apr. 18 138.1 7.5 9-8-7-10 
2 20 mi. N. Picacho, Calif. side Apr. 10 144.2 7.1 9-8-—7-10 
2 Opposite Cibola, Calif. side Mar. 31 142.7 8.7 9-8-7-10 
2 Riverside Mt., Calif. side Mar. 18 139.0 4.7 9-8-7-10 
2 Chemehuevis Val., Calif. side Mar. 10 138.0 by) 9-8-10-7 
2 5 mi. N. Laguna, Ariz. side Apr. 22 141.0 5.5 9-8-7-10 

Average of the females 140.5 6.4 

Average of the 12 specimens 140.4 6.2 


pate ey Sy 4 


tb po b& po 


142 University of California Publications in Zoology [Vou. 12 


LIST AND DIMENSIONS IN MILLIMETERS OF PHALAENOPTILUS NUTTALLI 
NITIDUS FROM THE COLORADO VALLEY 


9th primary 
exceeds Wing 
0. Sex Locality Date Wing Tail 10th by formula 


81 ' Riverside Mt., Calif. side Mar. 18 132.8 79.8 7.9 9-8-7-10 
86 <' Opposite Cibola, Calif. side Apr. 2 135.4 82.8 6.0 8-9-7-10 
87 Opposite Cibola, Calif. side Apr. 3 133.1 83.7 5.7 9-8-7-10 
78 6 Above Bill Williams River, 
Ariz. side Mar. 14 140.3 82.1 10.0 9-8-7-10 
Average of the males, 135.4 82.1 7.4 
82 9 Above Blythe, Calif. side Mar. 23 130.7 77.3 5.2 9-8-7-10 
92 9 8 mi. E. Picacho, Calif. side Apr. 19 131.9 77.0 6.5 8-9-7-10 
Average of the females, 131.3 77.1 5.8 
Average of the 6 specimens, 134.0 80.3 6.9 


1 Dissection showed reproductive organs to be much enlarged. 


Chordeiles acutipennis texensis Lawrence 
Texas Nighthawk 


First seen on March 9, on the California side in Chemehuevis 
Valley—one individual at dusk flying north. Next, March 27, on the 
Arizona side at Ehrenberg. Thereafter common at all stations, appear- 
ing each evening at dusk over the river and its flood plain. Not 
observed anywhere in the riparian strips during the day, but often 
flushed from the desert floor, usually from the scanty shade of a 
creosote bush. 

A specimen shot on the Arizona side five miles above Laguna, April 
24, had a spine-cushion of a cactus firmly stuck into the right carpal 
joint. This impediment, evidently picked up from the ground some 
time previously, had proven so serious a handicap as to result in the 
bird’s emaciated bodily condition. It would probably have resulted 
finally in the death of the bird, as the spines were deeply imbedded 
in the tissues. Accidents of this sort may not be infrequent and thus 
introduce an additional factor into the economy of desert bird-life. 

Near Pilot Knob, May 8, a Texas nighthawk was flushed from its 
complement of two fresh eggs. These lay on bare gravel between 
pebbles in the shade of a creosote bush on the hot glaring mesa. They 
are of the size and type of coloration usual with this species. 

The eight skins preserved, nos. 12766-12773, show no peculiarities 
as compared with specimens from the San Diego and San Joaquin 


1914] Grinnell: Mammals and Birds of the Colorado Valley 143 


districts of California. It is quite possible that the species is a com- 
paratively recent invader of the latter regions, hailing originally from 
the more arid deserts, with which its pale coloration would appear 
to associate it most closely. 

There are in the Museum two skins (nos. 4185, 4186) taken by 
J. G. Cooper at Fort Mohave, April 23 and May 23, 1861. 


Chaetura vauxi (Townsend) 
Vaux Swift 

Appeared as a migrant. First seen at Potholes, April 29, two 
individuals; next, one, on the following day, four miles south of 
Potholes. Very many were observed May 4 along the California side 
five miles northeast of Yuma; these were flying in a general northerly 
direction across the woods and river, irrespective of the local trend 
of the latter. May 5 a flock was seen at the mouth of the Gila, near 
Yuma. May 9 the last was seen, a single individual among cliff 
swallows near Pilot Knob. Two specimens secured, nos. 12774, 12775. 


Aéronautes melanoleucus (Baird) 
White-throated Swift 


Observed as follows: March 1 and 5, on the Arizona side among 
The Needles, seen to enter a slanting crack in the face of the cliff; 
April 4, on a precipitous mountain peak about five miles west of the 
river opposite Cibola, seen to enter and leave a crevice in the rock 
wall; April 19, eight miles east of Picacho, flying overhead; May 2, 
four miles south of Potholes, overhead. Not more than two individuals 
were noted at any of the above localities. 


Archilochus alexandri (Bourcier and Mulsant) 
Black-chinned Hummingbird 


First noted March 3, one of each sex, on the California side oppo- 
site The Needles; next March 7, several on the Arizona side at the foot 
of The Needles. Here they were feeding about the flowering bushes 
of Lyciwm anderson. Noted on the California side in the lower Chem- 
ehuevis Valley, March 9, one specimen; several on the Arizona 
side above Bill Williams River, March 13 and 14; several at River- 


144 University of California Publications in Zoology — [ Vou. 12 


side Mountain, March 17 to 21. At our station above Blythe, March 
22 and 23, this hummer was for the first time numerous, both males 
and females occurring along the desert edge of the riparian bottom, 
feeding among bushes of Lyciwm andersoni. At Ehrenberg the last 
week of March and opposite Cibola the first week in April, the species 
was abundant in the desert washes, feeding about the profusely blos- 
soming palo verdes. Noted at all stations below the latter point, 
though in lesser numbers. 

The males were more seldom seen, and the females became closely 
restricted to the willow strip along the river, in which association we 
were convinced that this was the only species of hummingbird breed- 
ing. The males were not seen in the willows, but only in the mesquite 
association and up the desert washes. The females foraged everywhere 
except on the desert mesa, but nested exclusively in the willows. 
Nest-building was in progress April 12, on the California side twenty 
miles north of Picacho. Nests, each with two small young, were found 
April 18 and 20, on the same side eight miles east of Picacho. Many 
full-grown young-of-the-year were feeding about the lavender flowers 


Fig. G. Vertical sectional diagram of nest (no. 769) of Archilochus alex- 
andri. Note position as regards supporting stalk, and construction of rim (see 


text, p. 145). X %. 


1914] Grinnell: Mammals and Birds of the Colorado Valley 145 


of ironwoods in washes near Pilot Knob, May 13. The breeding season 
of this hummingbird in this region is thus definitely indicated and 
shows itself to be just about one month earlier than on the Pacifie slope 
of southern California. 

Of the nests above referred to, one was placed eight feet above 
the ground on a dead willow twig, in a close-set grove of tall willows. 
The other was four feet above the ground on a slanting dead stalk of 
arrowweed beneath a large spreading willow. In structure and com- 
position the nests are of the compact type peculiar to this species as 
compared with those of the Costa. The body of the nest in one (no. 
769) is closely felted cottonwood and willow down. The whole outer 
surface is closely beset with small yellow and green leaves gathered 
where dried, on the lower portions of arrowweed stalks, and attached 
with web. A conspicuous feature is the extreme constriction of the 
rim, as shown in the accompanying sectional diagram (fig. G). Even if 
the supporting stalk were bent down to a horizontal position, there 
would be no likelihood of eggs or young falling out, because of the 
depth of the cavity and narrowing of the neck at the top. This max- 
imum in complexity of construction, accompanying, as it does, situation 
upon a tall, slender, and easily swayed stalk, shows marked adaptation 
in the habits of the bird to meet the special conditions imposed by its 
nesting site. 

The four newly hatched young (numbered 13985, 13986) are of 
interest in one particular in that they show a double row of down 
filaments along the dorsal tract, thus indicating the presence of a 
partial natal plumage, which plumage has been supposed to be unrepre- 
sented in the Trochilidae. 

Eighteen adult examples of the black-chinned hummingbird were 
preserved, nos. 12794-12811. 

There is also in the Museum a skin (no. 5882) taken by W. W. 
Holder at ‘‘Mineral City’’ (Ehrenberg) March 20, 1864. 


Calypte costae (Bourcier) 
Costa Hummingbird 
First seen February 26, a male flying northward near Mellen, 
Arizona. Next noted March 1, opposite The Needles, and for the 
andersoni which were at this time profusely laden with flowers. <A 
succeeding week very common along ravines on both sides of the river 
at The Needles. They were feeding about the spiny bushes of Lycium 
andersoni which were at this time profusely laden with flowers. <A 


146 University of Califorma Publications in Zoology — [Vou. 12 


tall-stalked milkweed (Asclepias subulata) growing high among the 
precipitous peaks was also an attraction; so, too, a sage (Hyptis 
emoryi). The latter was about the only shrub growing on the walls 
of steep-sided gulches at Riverside Mountain, and here two old nests 
of the Costa hummingbird were found:on horizontally projecting or 
drooping branches of the plant. Another nest found March 16 was 
five feet above the bed of a wash and contained two eggs; on the 19th 
the eggs were hatched. 

This hummingbird was pre-eminently a desert species; it was only 
casually noted even as a forager within the riparian set of associa- 
tions. It was thus complementary in its local distribution to Archil- 
ochus alexandri. It was more abundant in the early part of the season 
than later. This may have been due to local variation in favoring con- 
ditions as we traversed the region; or it is quite likely that the males 
and, as soon as able, the young and females largely departed from the 
desert floor to seek feeding grounds on the higher desert ranges. 

Besides the localities mentioned above, the Costa hummingbird 
was noted in Chemehuevis Valley, above Bill Williams River, opposite 
Cibola, twenty miles above Picacho, and at Pilot Knob. At the latter 
point, May 13, full grown juvenals were feeding about flowering 
ironwoods. 

Seventeen adult specimens of the species were obtained, nos. 12812— 
12828. 


Tyrannus verticalis Say 
Western Kingbird 


First seen, two individuals, March 24, at Ehrenberg; next, three 
individuals, March 29, at the same place; thereafter noted frequently 
at various points down the river. On the Arizona side five miles above 
Laguna, where there were many dead cottonwoods, many kingbirds 
were to be seen in pairs May 25; and their behavior led me to suspect 
them to be about to nest. The same condition held in the river bottom 
on the California side four miles below Potholes. No actual nests 
were seen, however. Many individuals believed to be in migration were 
seen at Pilot Knob, May 12. Seven specimens of this bird were taken, 
nos. 12829-12835. 

There is also in the Museum a skin (no. 4295) taken by J. G. 
Cooper at Fort Mohave April 25, 1861. 


1914] = Grinnell: Mammals and Birds of the Colorado Valley 147 


Myiarchus cinerascens cinerascens (Lawrence) 


Ash-throated Flycatcher 


Our capture of an example of this flyeatcher as early as February 
19 in a willow tract on the California side five miles below Needles, 
would appear to indicate that the species winters in the region. This 
idea is further borne out by Cooper’s observation (1870, p. 317) that 
he encountered an individual at Fort Mohave January 15. On March 
8 in the willow groves in Chemehuevis Valley we saw three ash-throated 
flyeatchers; on March 9, two. Thenceforth the species was noted 
regularly. It was common above Bill Williams River, affecting a 
tract of mesquites as well as the willows; also at Riverside Mountain 
and above Blythe. At Ehrenberg, besides scattered individuals, sev- 
eral pairs were encountered March 27 in the saguaro belt back on the 
mesa; in each case the birds had obvious claims on some old wood- 
pecker hole in a huge eactus trunk. Opposite Cibola and ten miles 
below Cibola individuals were seen to enter rotted-out cavities in iron- 
woods. 

The species was less common at our stations twenty miles north 
of and eight miles east of Picacho. But in the saguaro belt on both 
sides of the river and within five miles above the, Laguna Dam the 
ash-throated flycatcher was one of the characteristic breeding birds. 
The woodpecker holes in the giant cactuses are pre-eminently suited 
to these birds as nesting sites. A nest on the Arizona side was 
chopped out April 21 and found to contain one fresh egg. The nest 
cavity was five feet above the ground, and was bedded with a mass 
of burro hair and duck feathers. Another nest, found on the Cali- 
fornia side, April 23, was twelve feet above the ground in a cavity 
fourteen inches deep; composed of a closely packed mass of rabbit 
fur; there were four eggs in which incubation was well along. 

Ash-throated flycatchers were further observed, though more spar- 
ingly, at our stations at Potholes, four miles south of Potholes, five 
miles northeast of Yuma, and at Pilot Knob. 

A specimen (no. 4293) is in the Museum taken by J. G. Cooper 
at Fort Mohave April 13, 1861. 

Our series of twenty-five specimens (nos. 12836-12860) was secured 
as a result of an unsuccessful attempt to detect the presence of some 
one of the Mexican forms which come regularly into southern Arizona. 
While it is probable that this series includes transient individuals, it 
is not possible to pick them out, and the majority are, for certain, 


148 


University of California Publications in Zoology 


[Vot. 12 


breeding birds. I cannot discern in the series as a whole any departure 


from the characters of M. c. cinerascens of the coast slope of Cali- 


, fornia. The accompanying table shows the size and range of variation 
in certain respects of the Colorado Valley birds. 


MEASUREMENTS! IN MILLIMETERS OF A SERIES OF MYIARCHUS CINERASCENS 
CINERASCENS TAKEN IN THE COLORADO VALLEY FROM NEEDLES TO 


No. Wing 
12837 103.0 
12838 100.0 
12839 99.3 
12840 101.3 
12841 102.5 
12842 98.4 
12843 99.5 
12844 101.6 
12845 102.3 
12848 98.4 
12850 101.6 
12851 98.0 
12854 98.5 
12858 102.0 
Average 100.5 


Maximum 108.0 
Minimum 98.0 


Mid-point 


a 100.5 


of Range 
Variation 


either side 


of mid-point 


215% 


YUMA, FEBRUARY 19 TO MAY 9, 1910 


Males 


Tail 
95.0 
91.5 
92.5 
92.0 
93.0 
89.0 
90.0 
95.0 
92.3 
89.2 
91.9 
92.0 
92.2 
93.1 
92.0 
95.0 
89.0 


92.0 


3% % 


Culmen 
17.8 
17.2 
17.4 
17.8 
16.9 
18.2 
18.0 
18.7 
18.0 
17.5 
18.0 
Wf 
18.2 
19.7 
17.9 
18.7 
16.9 


17.8 


5% 


Width of 
bill at 
nostrils 


mt fa | 
10D 


ty 


UCR Gepost SS) Si =] 
NPwWH DD 


“Io 
co mw 


“I 


594% 


Females 
No Wing Tail 
12836 93.0 87.0 
12846 92.5 85.5 
12847 93.5 85.0 
12849 97.7 89.3 
12852 92.2 82.7 
12853 94.0 84.7 
12855 91.7 83.8 
12856 92.3 85.3 
12857 96.0 88.4 
12859 94.0 88.5 
12860 95.5 87.2 
Average 93.8 86.1 
Maximum O77, 89.3 
Minimum 91.7 82.7 
PRONE Nery) ai) 
of Range 
Variation 


either side (314% 
of mid-point 


1See text under Molothrus ater obscurus, p. 157. 


3% % 


Width of 

bill at 

Culmen nostrils 
16.4 8.0 
17.0 tet 
16.7 Wel 
17.2 Ten 
17.5 7.4 
16.9 8.0 
17.6 7.2 
17.8 et 
18.4 7.8 
17.8 7.3 
18.0 7.2 
17.4 7.6 
18.4 8.0 
16.4 {eal 
17.4 7.0 
534% 6% 


The table shows also the decided sexual difference in size in this 
species, and further that in both sexes there is greater variation in 


the size and relative narrowness of the bill, than in wing and tail 


leneth. Incidentally it would appear that females are subject to 
greater variation than males. 
are presented are comparable as regards stage of plumage wear and 


age; they are, at youngest, birds of the previous year. 


All of the birds whose measurements 


1914] Grinnell: Mammals and Birds of the Colorado Valley 149 


Sayornis sayus sayus (Bonaparte) 
Say Phoebe 


In February and early March this species was met with in small 
numbers out on the desert mesas as well as on the lower slopes of the 
hills. But it was not found in the wooded bottom lands of the broader 
valleys nor along the river banks except where the rocky hills abutted 
close upon the water. The bird was evidently wintering in the region, 
as the several individuals seen in the town of Needles upon our arrival 
there, February 15, were much smoke-begrimed, showing that their 
plumage had been exposed to the local conditions for some consider- 
able time. 

The Say phoebe was further noted at Mellen, among The Needles, 
and above Bill Williams River, on the Arizona side; and, on the Cali- 
fornia side, opposite The Needles. As the season advanced this species 
was met with only sparingly and in the vicinity of nesting sites. A 
pair was noted at Ehrenberg March 28, flying about some adobe ruins. 
On the California side opposite Cibola, April 3, a pair was found in 
a steep-sided ravine two miles back from the river. A last year’s nest, 
of usual contruction and containing one dried egg, was found on a 
narrow shelf of rock itself a part of an overhanging conglomerate 
wall. <A single bird was noted ten miles south of Cibola near some 
adobe ruins. Another was taken April 12 among the hills twenty 
miles north of Picacho. The species was last seen at Potholes, where 
one was observed on a telephone wire near the head works of the canal, 
April 29. Seven specimens were preserved, nos. 12861-12867. 


Sayornis nigricans (Swainson) 
Black Phoebe 


Met with only during the early part of the season, and only close 
along the river, where it was fairly common. Favorite perches were 
tips of snags on mud bars, or twigs of trees fallen over into the river 
through undereutting of the bank. The species was noted at Needles, 
five miles below Needles, at Mellen, on both sides of the river in the 
vicinity of The Needles, in Chemehuevis Valley, at Riverside Mountain, 
and at Ehrenberg. One, the last noted, was seen on the California side 
opposite Cibola, April 5. As far as it goes, the evidence would seem 
to show that the black phoebe is only a winter visitant in the Colorado 
Valley. Five specimens were preserved, nos. 12868-12872. 


150 University of California Publications in Zoology — [Vou. 12 


Nuttallornis borealis (Swainson) 
Olive-sided Flycatcher 


One seen May 6 perched at the top of a willow stub in bottom 
land on the California side near Pilot Knob. Two seen near the same 
place, May 10. Undoubtedly migrants. 


Myiochanes richardsoni richardsoni (Swainson) 
Western Wood Pewee 


First observed April 20, when a male was secured on the California 
side, eight miles east of Picacho. Next noted at Potholes, April 29; 
thenceforth in small numbers nearly every day, at the stations four 
miles south of Potholes, five miles northeast of Yuma, and at Pilot 
Knob. Most numerous at the latter point, May 6 to 12, where as 
elsewhere they were closely confined to the willow bottoms. Evidently 
only a migrant through the region. Eight specimens taken, nos. 
12918-12925. 

There are also in the Museum three skins (nos. 4305-4307) taken 
by J. G. Cooper at Fort Mohave, May 21 and 22, 1861. 


Empidonax difficilis difficilis Baird 
Western Flycatcher 


First noted April 3, on the California side opposite Cibola, observed 
almost daily thereafter throughout the trip, the last being seen the day 
we left, May 15. Still, there is no evidence to indicate that the species 
remains to breed anywhere in the region. The migration appeared not 
to have reached its height until the second week of May when, in the 
willows of the bottom lands on both sides near Pilot Knob, western 
flycatchers were continually in evidence, through their unmistakable 
notes. 

The fourteen skins preserved (nos. 12896-12909) are representa- 
tive of the following localities: Arizona side: ten miles below Cibola, 
five miles north of Laguna; California side: opposite Cibola, eight 
miles east of Picacho, four miles south of Potholes, Pilot Knob. One 
specimen (no. 12909) is aberrant in that the lower surface has a 
peculiar blanched appearance, the sides of the belly and crissum being 


1914] Grinnell: Mammals and Birds of the Colorado Valley 151 


distinctly whitish. This results in a casual similarity to #. trailli; all 
of the other characters are normal for difficilis, however. 

There is also in the Museum a skin (no. 43802) taken by J. G. 
Cooper at Fort Mohave, May 20, 1861. 


Empidonax trailli trailli (Audubon) 


Traill Flycatcher 


First detected April 28 and 29, when several were discovered in 
a tract of willows on the California side within a mile below Potholes. 
There were several of the birds, keeping in the upper foliage of the 
trees on ground overflowed from the adjacent canal to a depth of two 
feet or less. 

On the same side of the river five miles northeast of Yuma several 
Traill flyeatchers were observed May 4 and 5 close along a rapidly 
filling slough lined with willows. On both sides of the river in the 
vicinity of Pilot Knob the species was frequently observed up to the 
date of our departure, May 15. The birds were never detected away 
from dense willow growths close to the water, just such a type of 
loeality as is chosen elsewhere for nesting ground. This fact and the 
behavior of the birds led me to believe that they would nest in the 
vicinity. If so, this is the only Empidonar to breed in the lower Colo- 
rado Valley. Two specimens taken, nos. 12910, 12911, both males, are 
quite like more northern and western breeding birds. 


Empidonax hammondi (Xantus) 


Hammond Flycatcher 


First observed April 3, on the California side opposite Cibola; 
next April 6, at the same place; thereafter almost daily all along down 
the river, usually in the mid-branches of large willows. Thirteen 
specimens taken (nos. 12883-12895) from: Arizona side: ten miles 
below Cibola; California side: opposite Cibola, twenty miles above 
Picacho, eight miles below Picacho, Potholes, Pilot Knob. At the 
latter point the species was noted up to and including May 11. It 
appears that the species is only a transient through the region. 


E. 


Bae. 

E S38 gah 

=) ° 2S 

El ne = see 

Sng 8 noes. pag 

Exposed = sa BS = = a 

No. Sex Wing Tail Tarsus culmen fon] Es As Bs EEE 

E. hammondi 12894 fof 69.3 56.1 15.0 8.8 7 5.4 3.3 A 4.2 
wrighti 12873 fot Taz0 61.8 18.4 10.1 8.1 6.0 3.9 8. 3.0 

E. griseus 12879 dé 72.0 59.2 18.8 12:9 10.0 6.5 4.6 8. 0.9 


152 University of California Publications in Zoology [ Vou. 12 


Empidonax wrighti Baird 
Wright Flycatcher 


In only one instance was the presence of this species ascertained, 
and the specimen secured happened to be the first Empidonax taken: 
no. 12873, male, shot from the upper branches of a willow in the then 
leafless first-bottom timber, on the California side, five miles south 
of Needles, February 19. 

In spite of the largely increased extent of material illustrating this 
genus, the relative characters of Empidonax griseus, Empidonax 
wright, and Empidonax hammondi remain somewhat subtle. It would 
seem that wrighti, as I now recognize it, is much less common in the 
southwest even as a transient than formerly supposed, since out of 
the thirty-nine examples of the genus taken by our Colorado expedi- 
tion but one is referable to wrighti. 

To show upon what characters I base the discrimination of the 
wrighti example in the present case, a comparison with a picked 
average specimen of hammondi and of griseus is here given. All are 
males of apparently equal age and stage of plumage wear, this being 
very slight. The color differences are minute: hammondi is slatiest, 
griseus ashiest, wrighti intermediate; wrighti is greenest dorsally and 
pectorally; the outer web of outer tail-feather is distinctly white 
nearly to its tip in griseus, grayish white in wrighti, and but shghtly 
paler than rest of feather in hammondi. The lower mandible is 
entirely blackish brown externally, in hammondi, dull or leghter 
brownish in wrighti, while in griseus it is blackish brown at tip and 
abruptly straw yellow for its basal two-thirds, brightest along the 


rami. 


COMPARATIVE MEASUREMENTS IN MILLIMETERS OF THREE CLOSELY 


RELATED SPECIES OF EMPIDONAX OCCURRING TOGETHER 
IN SPRING IN THE COLORADO VALLEY 


trices in closed tail 


1914] Grinnell: Mammals and Birds of the Colorado Valley 153 


The above are the measurements and features of coloration which 
I find of diagnostic value, in discriminating these three flycatchers. 
They apply satisfactorily to the transient flyeatchers of Arizona and 
California. But the breeding ranges of these forms west of the Rockies 
are still imperfectly made out and there is undoubtedly extensive con- 
fusion in the literature. 


Empidonax griseus Brewster 
Gray Flycatcher 

With very little doubt this flycatcher is a winter visitant; but it 
was nowhere common. The first one noted was shot February 22, on 
the California side five miles below Needles. The next was secured 
March 10, also on the California side, in Chemehuevis Valley. Three 
were taken on the California side near Riverside Mountain; two on 
the same side opposite Cibola; and two on the Arizona side ten miles 
below Cibola. The last two were taken April 6 and 7. Nothing more 
was seen of the species after the latter date. Of the nine examples 
secured by us (nos. 12874-12882) all are males. This may be indic- 
ative of separate areas of wintering for the two sexes, or at least that 
those individuals wintering farthest north are males. 

Two specimens (nos. 4300, 4301) taken by J. G. Cooper at Fort 
Mohave, April 11 and 27, 1861, are also males. 


Pyrocephalus rubinus mexicanus Sclater 
Vermilion Flycatcher, 


An adult male seen at Needles, California, February 14. <A few, 
not more than six in all, noted March 27 to 29 in the vicinity of 
Ehrenberg, Arizona, where they remained in the sparse mesquite growth 
at the margin of the bottom farthest from the river. Fairly common 
April 1 to 5 on the California side opposite Cibola, where closely con- 
fined to a narrow strip of mesquite close to the river and along a 
lagoon. Here a nest was found, April 2, containing three eggs in 
which ineubation was far advanced. It was fifty-four inches above the 
ground, saddled on the bare forking branch of a dead mesquite stand- 
ing in an open area thirty-five yards from the river bank. 

The nest (no. 767) is shght in bulk, but firmly constructed, mainly 


of straight short dry twigs held together and to the large supporting 


154 University of California Publications in Zoology [ Von. 12 


branches with much cobweb; the rather shallow cavity (48 mm. wide 
by 17 deep) is lined with plant fibers, a few horse and cow hairs, and, 
most liberally, with down feathers from the desert quail. In spite of 
the absence of foliage on the tree, the nest was difficult to see even 
from a distance of fifteen feet, because of its small height above the 
branches on which it rested, and because its outer surface of gray 
twigs blended admirably with the striately shredded bark of the dead 
branches. One large branch extended above the nest, and gave partial 
protection from the sun, and certain other small branches may have 
helped some; but the sitting bird must have been compelled to endure 
the direct and intense sunshine a good part of the day. The nest was 
discovered only by flushing the bird, which sat very closely, and 
returned promptly when the intruder left the immediate vicinity. 
Only the female was seen on or close to the nest. The far more bril- 
liantly colored male remained at a distance of twenty to fifty yards 
from the nest, selecting conspicuous perches on dead mesquites, from 
which it sallied forth after passing insects in ordinary flyeatcher 
fashion. It sang musically early in the morning before sunrise, the 
song resembling somewhat that of the black phoebe, though of greater 
length and not so loud. 

The nest and pair of birds were finally collected. The abdominal 
surface area of the female was found to be conspicuously modified for 
ineubatory function, the skin being greatly thickened owing to the sub- 
cutaneous vascular development. The male had no such feature. This 
must have been an exceptionally early laying for the species in this 
vicinity, as other females taken showed no signs of breeding. 

The vermilion flycatcher was next noted on the Arizona side five 
miles above Laguna, where it was fairly common on overflow land 
among drowned and living cottonwoods and dead mesquites. On the 
California side, four miles below Potholes, the species proved to be one 
of the most numerous of the birds present. Here the valley widens out; 
much land is being reclaimed for farming, and intersecting irrigation 
ditches supplied from the Laguna Dam conduct water throughout the 
region. There is much waste land as yet, and burning has killed many 
of the cottonwoods. In this sort of locality the vermilion flyeatchers 
were found to be far more numerous than anywhere else; it would 
appear that here is a native bird of the Colorado Valley which will 
ereatly augment in numbers with the settling of the region. 

A peculiarity of the bird which impressed me was that by apparent 
preference it avoided green foliage, almost invariably perching on 


1914] Grinnell: Mammals and Birds of the Colorado Valley 155 


dead branches or exposed tree-tips, where the brilliant colors of the 
male rendered each individual most conspicuous. Yet this same sort 
of perch, because of its being a good vantage point from which to eet 
the widest survey for passing insects, was also preferred by the other 
flyeatchers, such as wood pewees and western kingbirds. No instance 
came under my observation of the conspicuousness of the male ver- 
milion flycatcher causing it to be molested save by human hunters. 

A half-grown juvenal was taken May 2 at this last station. The 
species was seen nowhere else than as above stated. Eighteen specimens 
were secured (nos. 12938-12955). 


Corvus corax sinuatus Wagler 
Western Raven 


Ravens were to be seen at practically every point of observation 
along our route from Needles to Pilot Knob. They were always noted 
singly or paired and were usually quiet. No young of the season were 
observed up to the time of our departure from the region. Ravens came 
to notice most frequently along the river as we floated down the swift 
current. Our boat was sometimes quietly carried into close proximity 
to an unwitting raven as it foraged for fish on some sand bar. 

At a heronry on the California side, ten miles below Ehrenberg, 
a pair of ravens was observed circling close over some nests which had 
just been deserted by the startled herons. Nearby, a dead cottonwood 
stub had under it a number of broken heron egg shells which doubtless 
betokened a source of the ravens’ food supply. The stomach of one 
of the ravens shot contained only some mammal hair and one spider. 

In each of the three ravens secured the ‘‘concealed’’ grayish white 
of the hind neck is much whiter and more extended than in ravens from 
California west of the desert divide. In the latter it is ordinarily dis- 
tinetly hight gray; in the Colorado River birds it is so much lighter as 
to give a first impression of pure white contrasted with the black 
surface-plumage when the feathers are parted. In fact this impression 
was so vivid that for a time in the field we thought we had secured 
examples of Corvus cryptoleucus. But comparison of skins shows the 
concealed white of cryptoleucus to be actually much more snowy. And 
there is, of course, no difficulty in distinguishing sinwatus from cryp- 
toleucus when measurements are compared, the former being very much 


the larger (see accompanying table). 


156 University of California Publications in Zoology [ Vou. 12 


The Colorado River ravens, as shown by the accompanying table, 
are of the usual sinwatus dimensions. The whitening of the basal or 
down portions of the nuchal feathers in desert representatives of the 
corax group would seem to be a modification in the direction of the 
condition as it finds its extreme in cryptoleucus, and probably because 
of coming under the same conditions. But no use of the feature is 
apparent. 


MEASUREMENTS IN MILLIMETERS OF CORVUS CORAX SINUATUS 
FROM THE COLORADO VALLEY 


Sty os ane s 

: a 33 & fg 2 a 

5 ce! Fa sy ao =a =z 2 

No. Sex Locality Date fe & Sie & mg b&b A 
12914. Riverside Mt., Calif. Mar. 16 440 242 70.0 60.0 74.0 52.5 36.3 25 
12913 g 10m.S. Ehrenberg, Calif. Mar. 30 414 222 66.8 59.5 69.2 50.1 34.3 25 
12912 @ 10m.S8. Cibola, Ariz. Apr. 7 381 224 64.8 61.7 67.4 48.0 33.0 26. 
43677 2 Fort Mohave, Ariz. Jan. 6 410 212 64.7 58.2 65.6 45.3 31.9 24 
4368" 2 Fort Mohave, Ariz. Jan. 6 423 221 65.8 59.0 68.9 49.0 32.9 25. 


* Collected by J. G. Cooper in 1861. 


Molothrus ater obscurus (Gmelin) 
Dwarf Cowbird 

Cowbirds were first seen when we reached Ehrenberg, March 25. 
Here, on the Arizona side, flocks aggregating approximately one hun- 
dred individuals were staying close about the town. They were either 
roosting quietly in mesquites among the adobe ruins, or flying about in 
compact, unanimously acting flocks, or feeding in company with other 
blackbirds in eattle pens. In one case a bunch of fully forty were 
feeding close together on the ground in a ealf corral, all with uptilted 
tails and quivering bodies, a mannerism peculiar to the species. They 
were eating millet seeds, and the crops of those taken were crammed 
with this material. One flock was seen flying north along the California 
side opposite Ehrenberg. On the same side, opposite Cibola, during 
the first week of April a number were seen, now in pairs haunting the 
bottom lands close to the river. 

The species was next seen April 17 on the California side twenty 
miles north of Picacho at the Draper ranch, four individuals; then 
eight miles east of Picacho, April 20, a small flock. On the Arizona 
side again, five miles north of Laguna, April 24 to 27, several pairs 


bill at base 


tw to tH & 


1914] Grinnell: Mammals and Birds of the Colorado Valley 157 


were seen among cottonwoods. On the California side, from the vie- 
inity of Potholes south all along the river to the Mexican line, the 
species was numerous throughout the bottom lands. Here up to May 
15 cowbirds were to be seen usually in pairs frequenting the densest 
willow tracts, as well as foraging on the mud bars at the water’s edge. 
When prospecting in thickets near the ground they were usually quiet 
and gave the impression of sneaking, as if intentionally planning to 
avoid observation. But when affecting the taller tree tops and open 
places, they kept flying about actively, often intently pursuing one 
another, and giving frequent voice to an attenuated squeal. The species 
confines itself strictly to the river bottom: none were seen out on the 
desert, even in flight overhead. 

Fifty-three skins of the dwarf cowbird were obtained, nos. 12956— 
13008. 

In addition to these there are in the Museum two skins, one (no. 
4347) taken by J. G. Cooper at Fort Mohave, April 4, 1861, and 
another (no. 5963), of unknown collector, from ‘‘Mineral City”’ 
(=—Ehrenberg), April 20, 1864. 

The acquisition of so large a series of Molothrus ater obscurus (pre- 
viously sparingly represented in collections) from so limited an area, 
warrants a detailed examination of its characters. The fifty-three 
specimens are all comparable, as far as age is determinable, and are in 
comparatively unabraded plumage. The variations in measurements 
are therefore intrinsic, except, of course, for an indefinite but small 
amount of error which seems unavoidable. Although but four dimen- 
sions are presented, it is believed that these are a close index to any 
other correlated features that could be selected for measurement. And 
the ones adopted would appear to be capable of more exact determina- 
tion than, for example, length of body and wing expanse. 

The wing is measured, according to prevalent custom, from the bend 
of the carpus in closed wing to tip of longest primary; but even here 
a shght error may result from the mode in which the wing has been 
“*set’’? in drying, that is, according to the sharpness of the curve alony 
its outer edge. The tail is measured from the point between the inser- 
tions of the median rectrices into the uropygium to the tip of the 
tail; and here too the proximal point from 


” 


longest rectrix in ‘‘closec 
which to measure is not quite satisfactory. The culmen is rather more 
accurate of measurement in the cowbirds because it is a well-defined 
ridge coming to an end rather abruptly on the forehead. The depth of 
bill is most unsatisfactory because of the lability to slight separation 


158 University of California Publications in Zoology [| Vou. 12 


or abnormal closeness of the mandibles incurred in drying, and because 
of the conical shape of the beak. The method of taking the measure- 
ment here presented is to secure with calipers the diameter of the bill 
from the highest point on the culmen along a line through the nostril. 
This does not, therefore, take into account the extreme base of the lower 
mandible, where it is difficult to find any constant poimt from which 
to measure. 

It will be noted from the tables that, in both sexes, the average is 
practically identical with the mid-point of range; that the wing length 
is least variable; that males are more variable than females; that the 
bill of the female is relatively shorter and thicker than that of the 
male, and its shape is much more constant; that in the male the varia- 
tion in the shape of the bill involves length much more than depth, 
so that a wide range in degree of slenderness results, rather than in 
actual volume; that in matter of general size of body (using the wing 
and tail length as an index) females are nine per cent smaller than 
males. 

Previously (1909, p. 281) the writer has given a table of measure- 
ments of the then available specimens of Molothrus ater obscurus. 
There were eleven males, from Arizona and southeastern California. 
Comparison with the dimensions of thirty-one other males as here 
presented shows remarkably close accord in all particulars save one, 
due to depth of bill being taken to extreme base of lower mandible 
instead of through nostril. It is of interest to note that among the 
entire series of fifty-three cowbirds from the lower Colorado River 
there is not one showing any closer approach to the Nevada form, 
Molothrus ater artemisiae, than in the smaller series above referred to 
and used originally for comparison. The apparent absence of M. a. 
artemisiae from the Colorado Valley as a migrant is contrary to pre- 
vious ideas as to its route of migration to and from the Great Basin 
region. But what other route may be followed is still a matter of 
conjecture. 

As regards color the Colorado series is remarkably uniform, except 
in one respect: the steely reflections of the dark body plumage in the 
male vary from deep violet to bronzy. This variation is exhibited 
similarly in M. a. artemisiae and in M. a. ater. In obscurus, however, 
the brownness of the head averages perceptibly paler than in the 
Atlantie race, ater. It is in the female that obscurus shows marked 
color characters as compared with ater; and this has apparently been 
overlooked in current descriptive works (as, for instance, by Ridg- 


1914] Grinnell: Mammals and Birds of the Colorado Valley 159 


way, 1902, p. 210). The entire plumage in obscurus is very much 
paler than in ater. The former is drab throughout, slightly paler ven- 
trally and very much paler, more clay color, on the throat; the narrow 
dusky shaft-streaks on the dorsum, chest and sides are clearly indicated. 
The female of the eastern form is dark slaty hair brown with less 
abruptly contrasted grayish throat. 


MEASUREMENTS! IN MILLIMETERS OF THIRTY-ONE MALES OF 
MOLOTHRUS ATER OBSCURUS FROM THE LOWER 
COLORADO VALLEY 


Ratio % of 
Depth of bill depth of bill 


No. Wing Tail Culmen at nostril to culmen 
12960 96.8 65.3 15.5 8.5 55 
12961 97.8 64.7 15.2 8.7 57 
12962 96.5 63.8 16.0 8.3 55 
12963 102.3 68.3 15.3 8.6 56 
12964 100.7 69.6 14.8 8.2 55 
12965 100.4 66.4 16.2 8.4 52 
12966 101.0 67.5 16.0 8.5 53 
12967 103.3 66.7 16.0 8.8 55 
12968 98.7 64.5 15.5 8.4 54 
12969 97.3 65.0 15.7 8.0 51 
12970 100.9 66.4 16.6 8.4 51 
12971 101.4 66.0 16.5 8.7 53 
12972 103.0 69.6 15.8 8.5 54 
12976 100.0 67.4 16.0 8.4 52 
12979 100.8 65.5 15.6 8.3 53 
12981 98.5 69.3 16.1 8.6 53 
12986 102.6 69.9 16.5 8.5 52 
12987 103.7 68.5 15.9 8.4 53 
12988 98.9 64.7 15.6 8.0 51 
12989 96.7 66.2 16.1 8.2 51 
12990 99.8 67.0 16.9 8.3 49 
12992 103.1 70.5 17.0 8.4 49 
12993 97.4 66.2 ali fei 8.2 48 
12995 101.3 70.0 17.0 8.2 48 
12998 103.2 69.4 16.0 8.0 50 
138000 99.4 64.6 Iie} 8.6 50 
13001 101.1 67.0 16.8 8.1 48 
138005 102.5 70.0 15.7 8.4 53 
13006 102.0 69.7 17.2 8.1 47 
13007 99.1 63.6 16.4 8.3 51 
13008 97.0 67.0 16.0 8.1 51 
Average 100.2 67.1 16.1 8.4 52 
Maximum 103.7 70.5 17.3 8.8 57 
Minimum 96.5 63.6 14.8 8.0 47 
Mid-point of Range 100.1 67.0 16.1 8.4 52 


Variation on either | 3.6% 5% 7.4% 4.8% 10% 
side of mid-point {f 


1See text, pp. 157-158. 


160 University of California Publications in Zoology [ Vou. 12 


MEASUREMENTS IN MILLIMETERS OF TWENTY-TWO FEMALES OF 
MOLOTHRUS ATER OBSCURUS FROM THE LOWER 
COLORADO VALLEY 


Ratio % of 
Depth of bill depth of bill 
No. Wing Tail Culmen at nostril to culmen 
12956 90.5 59.6 14.7 8.0 54 
12957 91.7 60.8 15.0 8.4 56 
12958 89.8 59.9 14.0 8.0 57 
12959 91.9 61.3 15.1 8.3 55 
12973 88.8 60.3 14.5 7.8 54 
12974 89.2 58.0 14.4 Ue 53 
12975 89.7 59.7 14.5 8.0 55 
12977 92.3 58.6 14.6 7.8 53 
12978 89.0 59.7 14.8 7.9 53 
12980 91.8 62.0 14.6 8.1 55 
12982 89.0 59.5 15.2 8.0 53 
129838 91.1 60.3 14.4 Ceti 53 
12984 91.4 61.6 15.0 8.0 53 
12985 92.9 6337/0) Wee : 7.5 ats 
12991 94.6 65.0 14.9 8.2 55 
12994 90.4 G2'9.-" | ee: mee a 
12996 : 91.6 59.3 14.8 el 52 
12997 92.7 62.0 15.4 7.8 51 
12999 93.4 63.3 15.0 8.1 54 
13002 89.0 59.0 14.7 8.0 54 
13003 93.0 59.5 15.0 8.3 55 
13004 93.3 59.8 14.9 8.2 55 | 
Average 91.2 60.7 14.8 8.0 54 
Maximum 94.6 63.7 15.4 8.4 57 
Minimum 88.8 58.0 14.0 7.5 51 
Mid-point of Range 91.7 60.8 14.7 8.0 54 
WUE CGN ERE \ 3.2% 4.6% 4.8% 5.6% 5.5% 
side of mid-point 


* Defective. 


Xanthocephalus xanthocephalus (Bonaparte) 


Yellow-headed Blackbird 


The single specimen secured, an adult male (no. 12915), was shot 
from a flock of cowbirds at Ehrenberg, Arizona, March 28. No others ; 
of the species were seen at that point. But on April 24, on the Arizona 
side five miles north of Laguna, a male and two females were seen over- | 
head flying south. And on May 5, on the California side four miles | 
south of Potholes, a small flock of yellow-headed blackbirds was seen 
flying north along the river bank. 


1914] Grinnell: Mammals and Birds of the Colorado Valley 161 


There is in the Museum an adult male (no. 4346) taken by J. G. 
Cooper at Fort Mohave March 31, 1861; and another skin (no. 5959) 
taken by W. W. Holder at ‘‘Mineral City’? (=Ehrenberg) March 8, 
1864. The species is probably a regular winter visitant to the Colo- 
rado Valley. 


Agelaius phoeniceus sonoriensis Ridgway 
Sonora Red-winged Blackbird 


First seen March 6, when a large flock was encountered flying north 
along the Arizona shore at the foot of The Needles. Next noted at 
Ehrenberg, Arizona, during the week beginning March 24. Here not 
more than fifteen individuals all told were staying close about the town, 
associated with the cowbirds and Brewer blackbirds which foraged in 
the cattle corrals. On the same side of the river, ten miles below Cibola, 
April 7, four individuals were seen in flight up the river. Several 
small flocks came to notice at our station on the California side eight 
miles east of Picacho, April 18 to 20. On both sides of the river 
above Laguna and at Potholes, April 22 to 29, many were seen, mostly 
in pairs or small flocks pursuing their way up the river. It was 
observed in nearly every instance that these presumably migrating 
birds, instead of taking a more direct route northward up the valley, 
closely followed the river bank even when through the meandering of 
the stream this course was sinuous in the extreme. 

At Potholes and below, evidence was unmistakable that red-wings 
were preparing to nest, though no nests had been built as yet up to 
the time of our examination. At one place in particular, on the 
California side of the river, about five miles northeast of Yuma, a 
large colony had evidently settled for breeding purposes. Here in 
the late afternoon of May 2, on the inside of a large bend of the river, 
we were afforded opportunity of securing a good series of specimens 
and of observing the behavior of the birds. The males were by them- 
selves, spaced out, and perched at the tips of willow saplings in an 
extensive dense tract of these on ground which at a little higher stage 
of the rapidly rising river would have been wholly inundated. The 
male birds were singing continually, and displaying themselves con- 
spicuously in short cireling flights, but each one behaving independ- 
ently of any other. The females on the other hand were quietly for- 
aging on the ground in close flocks in a much younger growth of 


162 University of California Publications in Zoology | Vou. 12 


willow seedlings some three hundred yards off. The inference was 
that the males had selected the appropriate site for the location of 
nests, but that pairing off had not yet occurred. Here, as with many 
other birds of the Colorado Valley, it was evident that nesting time 
is deferred until the period of highest water, which custom doubtless 
obviates danger of destruction of nests through flooding. Elsewhere 
in the Austral zone red-winged blackbirds are earing for broods of 
young at so late a date as May 2. 

The species was common near Pilot Knob flying back and forth 
overhead, evidently between breeding and feeding grounds somewhere 
in the vicinity. Nowhere was it seen outside of the confines of the 
river flood plain, so that the desert mesa forms the practical barrier 
to its farther spread laterally. 

Thirty-three specimens were preserved, nos. 13009-13041. 

The red-winged blackbirds of the Colorado Valley show themselves 
to be markedly different from those of the Pacific slope of southern 
California. The latter are Agelaius phoeniceus neutralis Ridgway ; the 
former are here referred to Agelaius phoeniceus sonoriensis, following 
Ridgway’s last diagnosis (1902, p. 337). Although the type locality 
of sonoriensis is stated by Ridgway (1902, p. 338) to be Mazatlan, 
Arizona had previously been fixed as type locality by Nelson (1900, 
p. 126), and the A. O. U. Check-List (1910, p. 233) further specifies 
Camp Grant, Arizona. I have no Mexican material for comparison. 

The Colorado Valley form, as compared with neutralis, has the bill 
in both sexes very much longer and slenderer. In some of the males 
this feature is extraordinarily pronounced: the culmen, gonys and 
lateral outlines are all concave, resulting in an acicular sharpness of 
the bill. An opposite variation, however, leads to a moderately pointed 
bill in six out of twenty-six males, which can be matched by picked 
examples of neutralis. These neutralis-like males are all but one 
obviously immature. It is possible, of course, that some of these may 
actually be transient or casual visitants of newtralis. But it appears 
to me now as more likely that they are individual extremes of the 
locally native form, which in the material at hand varies towards 
them. 

The six females collected are all conspicuously different in colora- 
tion, as well as in shape of bill, from neutralis. They have the colora- 
tion ascribed to sonoriensis, the black streaking below being much nar- 
rower and more distinctly contrasted against the extended white 
ground; also dorsally and on the sides of the head the pale edgings 


1914] Grinnell: Mammals and Birds of the Colorado Valley 163 


of the feathers are much lighter and more extensive. In the males 
no color peculiarities are apparent. 

There is in the Museum a skin (no. 4350) taken by J. G. Cooper 
at Fort Mohave, January 13, 1861. It is a male, obviously immature. 
It has a markedly thick bill and much dark rusty on the back. While 
probably not referable to sonoriensis, I am not able to place it satis- 
factorily. It possibly represents some northern race which winters 
in the region. 


Sturnella neglecta Audubon 
Western Meadowlark 

Numerous February 15 to 18 on the first bottom close to the river 
at Needles, California. Here there had been some attempt to induce 
the growth of a coarse pasture grass by irrigation, and the resulting 
cleared field scantily clothed with grass formed the chief local attrac- 
tion for the meadowlarks. A few were seen February 22 five miles 
below Needles on a section of mesa by the river. Three individuals 
were seen on some grass land on the Arizona side, above the mouth 
of Bill Williams River March 13 and 14; and a lone individual was 
shot April 22 on the Arizona side five miles north of Laguna. Twelve 
specimens were secured (nos. 12926-12937), all but one near Needles. 

There is in the Museum a skin (no. 4359) taken by J. G. Cooper 
at Fort Mohave January 1, 1861; and another, without date, from 
““Mineral City’? (Ehrenberg) taken by W. W. Holder. 


Icterus cucullatus nelsoni Ridgway 
Arizona Hooded Oriole 

Common and breeding about the site of the old Hanlon ranch on 
the California side near Pilot Knob. Here these birds affected prim- 
arily the large date palms which afforded characteristic nesting sites. 
One nest was noted suspended in a mistletoe clump in the cottonwood 
above the ruins of the American Girl pump. One hooded oriole was 
discovered far up a wash on the mesa, sipping nectar in company with 
hummingbirds in a profusely blossoming ironwood. Six of these orioles 
were secured May 6 to 14, nos. 13055-13060. Elsewhere the species 
was observed twice on the Arizona side, one seen April 26 five miles 
north of Laguna, and one heard April 9, ten miles below Cibola; and 
once on the California side, one heard April 11, twenty miles north 
of Picacho. 


164 University of California Publications in Zoology — [Vou. 12 


Icterus bullocki (Swainson) 
Bullock Oriole 


Nests of this species were conspicuous in the taller willows early 
in the season while the trees were yet leafless, all along the river from 
Needles down. The first birds to appear were noted on the California 
side opposite Cibola, April 5. There were five of them, all males, feed- 
ing about the blossoms of palo verde trees. From this date on, almost 
daily, and at every station, the species was noted in increasing num- 
bers, until at Potholes the last of April, and elsewhere down the river 
to Pilot Knob, this was one of the numerous and characteristic birds 
of the willow-cottonwood association. Fourteen examples were pre- 
served, nos. 13041-13054. Another specimen in the Museum (no. 
4342), an adult male, was taken by J. G. Cooper at Fort Mohave April 
3, 1861. 

Although there are doubtless breeding individuals among these, as 
well as probable transients, I am unable to find any differences by 
which the Colorado Valley birds may be distinguished from those of 
western California or northern Nevada. 


Euphagus cyanocephalus (Wagler) 
Brewer Blackbird 
Common about the town of Needles, California, February 15, where 
they were foraging indifferently along the railroad tracks and in the 
streets. Very many were seen similarly in the town of Parker, Arizona, 
March 16. Small flocks, aggregating about forty individuals, were 
noted at Ehrenberg, Arizona, March 24 to 29, and two secured (nos. 
12916, 12917). Here the birds were usually mingled with the flocks 
of cowbirds feeding in chicken yards and cow pens. 
There are four skins in the Museum (nos. 4355-4358) taken by 
J. G. Cooper at Fort Mohave January 4, 1861. Evidently a winter 
visitant in the Colorado Valley. 


Carpodacus mexicanus frontalis (Say) 
California Linnet 


Locally common all along our route, from Needles to the vicinity 
of Yuma. Large flocks were foraging over the Sacramento Wash on 
the Arizona side near Mellen the last week in February. Elsewhere 


1914] Grinnell: Mammals and Birds of the Colorado Valley 165 


only small companies were noted, either on the river bottom proper 
or out on the desert. As the season advanced linnets were more fre- 
quently noted in pairs, often along the deep, narrow ravines and 
precipitous hillsides where the mountains reached the neighborhood of 
the river. Nests with eggs were found ensconced among the tangled, 
sharp-pointed twigs of old smoke-bushes on desert washes both at 
Riverside Mountain, March 18, and Ehrenberg, March 26. Four miles 
below Potholes the birds were common on the bottom lands, affecting 
especially those cottonwood trees laden with mistletoe clumps. Five 
miles above Laguna, a linnet’s nest was found in a cavity in the side 
of the main shaft of a saguaro. Twenty miles north of Picacho a 
nest with small young was found April 14 on the thorny upper branch 
of an ironwood. 

The series of twenty-eight skins secured (nos. 13061-13088) shows 
the desert linnets to be in no perceptible way different from those of 
the coast region of California. 

There is in the Museum a skin (no. 4169) taken by J. G. Cooper 
at Fort Mohave, January 26, 1861. 


Astragalinus psaltria hesperophilus Oberholser 
Green-backed Goldfinch 


More or less common all along the river; evidently resident. First 
noted on the California side opposite The Needles, several, March 2. 
On the Arizona side at the foot of The Needles, March 4 to 6, large 
flocks were congregated in the central portions of extensive dense 
mesquite thickets where, perched from three to four feet above the 
ground, they were certainly safe from marauders; here they sang 
volubly in chorus until dusk settled. During the day they were seat- 
tered out over the hills feeding on the tlower-heads of Perityle emory?. 
California side, at Chemehuevis Valley, common, both in the palo 
verde washes back on the mesa, and in the mesquites and willows near 
the river; noted similarly, still in flocks, at Riverside Mountain, 
March 16 to 20, and above Blythe, March 23. On the Arizona side in 
the vicinity of Ehrenberg there were but few noted, and these only in 
pairs along the mesa margin of the river bottom; on March 25 two 
partially constructed nests were found each four feet above the ground 
in arrowweeds. 


166 University of California Publications in Zoology — [ Vou. 12 


Goldfinches of this species were henceforth noted in small numbers 
as follows: opposite Cibola, ten miles below Cibola, twenty miles north 
of Picacho, eight miles east of Picacho (full grown young taken), four 
miles below Potholes and near Pilot Knob (family of young). Twenty- 
five specimens were taken, nos. 13091-13115. These are identical with 
the form inhabiting the Pacifie slope of California. Of the sixteen 
males in the series, not one is of the arizonae or mexicanus type of 
coloration, that is, with unusual extensions of black in the plumage. 


Astragalinus lawrencei (Cassin) 
Lawrence Goldfinch 

Observed only on the California side at Riverside Mountain, March 
17: a flock of eight was discovered in the upper branches of a dead 
mesquite on the verge of the river bank, singing and ealling in char- 
acteristic fashion. Two were procured, nos. 13089-13090. 

A single specimen (no. 4164) is in the Museum, taken by J. G. 
Cooper at Fort Mohave January 28, 1861. The species is probably 
a winter visitant to the Colorado Valley, though not in abundance. 


Passer domesticus Linnaeus 
English Sparrow 
Several dozen were apparently well established in the station park 
and about the railroad yards of Needles. They had their retreats in 
the palms and eucalyptus by the station. One lone individual was 
seen in some willows on the Arizona side half a mile above the railroad 
station at Mellen, February 24. 


Pocecetes gramineus confinis Baird 
Western Vesper Sparrow 

Met with in but two places: on the California side, in Chemehuevis 
Valley, where a specimen, no. 13124, was secured March 10; and on the 
Arizona side, above Bill Wilhams River, March 12 and 13, where 
three specimens were taken, nos. 13125-13127. A few others were 
seen in the latter locality, where they were all found in a growth of 
tall coarse grass paralleling the mesquite belt. 

There is in the Museum a skin (no. 4155) taken by J. G. Cooper 
at Fort Mohave, March 6, 1861. This sparrow probably oceurs in 
the region throughout the winter. 


1914] Grinnell: Mammals and Birds of the Colorado Valley 167 


Passerculus sandwichensis nevadensis Grinnell 
Nevada Savannah Sparrow 
Four Savannah sparrows referable to this form were taken: no. 
13116, California side, opposite The Needles, Mareh 3 (the only one 
of the genus seen here) ; nos. 13117, 13118, Arizona side, above mouth 
of Bill Williams River, March 13; and no. 13119, Arizona side, five 
miles north of Laguna, April 22. 


Passerculus sandwichensis alaudinus Bonaparte 
Western Savannah Sparrow 

Five specimens secured, all on the Arizona side: nos. 13120, 13121, 
above Bill Williams River, March 13, and nos. 13122, 13123, 13128, 
five miles north of Laguna, April 22. At both places small areas of 
grassy land on the river bottom back near the mesa afforded the attrac- 
tion to the species. The last-mentioned specimen is so nearly an inter- 
mediate in all characters between P. s. alaudinus and P. s. nevadensis 
as to be allocated uncertainly with one or the other. At least six Sav- 
annah sparrows were observed in a tract of low herbage lying near the 
river, five miles northeast of Yuma, May 2; these were the last to 
be noted. 


Chondestes grammacus strigatus Swainson 
Western Lark Sparrow 
One specimen taken (no. 13129) at edge of bottom near mesa on 
California side at base of Riverside Mountain, March 17. Not other- 
wise seen by our party. 


Zonotrichia leucophrys leucophrys (Forster) 
White-crowned Sparrow 

Appeared in the region only as a transient. First noted in Cheme- 
huevis Valley, California side, March 11, when an adult female (no. 
13130) in worn winter plumage was shot from a flock of Z. 1. gambeli. 
This must have been an exceptionally early arrival, or possibly a 
stray sojourner through the winter. For no more were identified with 
certainty until April 11, when a male (no. 13131) was taken on the 
California side, twenty miles north of Pieacho. This bird is in the 
midst of the first prenuptial molt, as is also another (male, no. 13133) 


168 University of California Publications in Zoology | Vou. 12 


taken April 18, eight miles east of Picacho. Another (female, no. 
13132) taken on the same date and at the same place, and one (male, 
no. 13134) secured April 21, on the Arizona side five miles north of 
Laguna, have nearly or quite completed the prenuptial molt (whether 
first or later, I am not able to judge). 

From April 16 to 26 many crown sparrows of this form were 
observed closely enough to make identity certain. They sometimes 
formed scattering companies in brush along desert washes; or, in 
company with Zonotrichia leucophrys gambeli, they invaded the mes- 
quite and arrowweed associations nearer the river. None were seen 
after April 26. It would appear that molting and migration occur 
simultaneously. Still there is nothing inconsistent with utility in this. 
For the prenuptial (or spring) molt in this sparrow, as with gambeli, 
is only partial; the flight feathers (primaries, secondaries and rec- 
trices) are not involved, save that, in case any have been lost by 
accident, new ones grow in at this time. 


Zonotrichia leucophrys gambeli (Nuttall) 
Intermediate Sparrow 


A winter visitant to the region in large numbers, occurring in 
seattered flocks with greatest frequency along the river, in the mesquite, 
quail-brush and salt-bush associations. Thence they were to be found 
up the brushy margins of desert washes at least as far as two miles 
from the river. Although often found foraging over open ground, 
these sparrows appeared never to venture so far from dense bushes 
but that they could on alarm find quick shelter. 

In the vicinity of the railroad station at Needles many intermediate 
sparrows were to be observed at all times of the day foraging over 
the tracks, often beneath trains, where they appeared to find plenty 
to eat in the grain and refuse sifting from the cars. These birds had 
evidently been in the neighborhood all winter, for those shot February 
15 and 16 had their plumage begrimed by coal dust to such an extent 
in some cases that the birds at a distance appeared to be of a uniform 
deep plumbeous color. No birds of such extraneous peculiarity were 
observed farther than two miles from the station, perhaps showing 
habitual close adherence to a local feeding ground selected upon arrival 
in the fall. 


1914] Grinnell: Mammals and Birds of the Colorado Valley 169 


To enumerate localities of occurrence would be to list every place 
explored along down the river to the collecting station on the Arizona 
side five miles north of Laguna. By the time we reached this point, 
April 21, intermediate sparrows had become notably fewer in numbers 
than immediately before. Still there were several to be seen each day 
in the arrowweed and willow thickets until the 26th of April, when the 
last for the season were seen at the same place, and one, a female, 
taken. 

There is in the Museum a skin (no. 4177) taken by J. G. Cooper 
at Fort Mohave, January 26, 1861. 

Of thirty individual specimens shot before the completion of the 
prenuptial molt, April 15, fifteen are in the brown-and-gray-crowned 
first winter plumage. As there was, as far as I know, no intention 
of selecting adults in preference to immatures, these figures probably 
give an approximate proportion of birds-of-the-year at that season 
(February 15 to April 15) that is, fifty per cent. 

The whole series taken, thirty-four specimens (nos. 13136-13169), 
serves well to indicate the period occupied in the prenuptial molt. The 
earliest individual showing molt, adult female, March 27 (no. 13160), 
shows many new feathers unsheathing in the mid-dorsal and pectoral 
tracts. An immature of the same date (male, no. 13161) shows a few 
unsheathing feathers in the mid-dorsal region only. An adult male, 
April 6 (no. 13162), is well along toward completion of the molt, 
though many old feathers persist in the capital region. It is of course 
to be remembered that the prenuptial molt does not involve any of the 
flight feathers, or, in this species, the wing and tail coverts. 

That the molt program is not exactly simultaneous in all indiy- 
iduals is shown by an adult female (no. 13163) taken April 7, in which 
no sign of molt is apparent. An immature (male, no. 13165), April 
10, is in the midst of the molt all over the body. In another specimen 
(female, no. 13166), April 18, of age not with certainty recognized, 
the molt is approaching completion. In the remaining three specimens, 
females, April 20, 24, and 26, feather growth in all the tracts involved 
is evidently complete, though in each individual a few old abraded 
feathers still remain in the post-auricular and pectoral regions. 

It would thus appear that the spring molt occupies a period of not 
to exceed twenty-five days. Because of lack of uniformity in date of 
inception in different individuals or rate of the process, this is prob- 
ably somewhat more than the time occupied in any one individual. 
The nuptial dress differs in color from the adult winter plumage in 


170 University of California Publications in Zoology — [Vou. 12 


the clearer ash of the anterior lower parts, in the darker clay color 
of the flanks, the clearer white of the belly, and the decidedly darker 
tone of the dorsal streaking, sepia rather than bay. 


Spizella passerina arizonae Coues 
Western Chipping Sparrow 

A single example noted February 20, on the California side, five 
miles below Needles. On the Arizona side, among the precipitous hills 
constituting The Needles, a large scattering flock was seen March 5. 
The birds were foraging among the sparse creosote and Encelia bushes 
on the steep slopes. Thereafter observed on the California side in 
Chemehuevis Valley, at Riverside Mountain, Blythe, opposite Cibola 
and eight miles east of Picacho (last April 18), and on the Arizona 
side above Bill Williams River, at Ehrenberg, and ten miles below 
Cibola. At all of these points chipping sparrows were frequently met 
with, either in the bushy ravines among desert hills, or in the brush 
of the different associations paralleling the river. 

I had concluded after seeing none for many days after April 18, 
that the species had left the region about that time. But as a matter 
of surprise the species was found once again on April 30 and May 1, 
on the California side four miles south of Potholes. Here the birds 
were notably common in the newly settled bottom land characterized 
by open fields, groves of mistletoe-laden cottonwoods and intersected 
by irrigation canals. They were in full song, and not in flocks as 
were the last previously seen, but in pairs, on or near the ground; or 
the males were singing alone from the bare upper branches of fire- 
killed trees. The two males shot had their testes greatly enlarged. 
Thus all the evidence obtained indicates that the species was breeding. 
Yet no nest was actually found, and the previously unknown occur- 
rence of the species in so low a life-zone brings an element of prob- 
ability that these individuals were merely tarrying late in the region 
before leaving for breeding grounds in a higher zone. Moreover the 
specimens secured differ in neither coloration, general size, nor pro- 
portions from breeding birds from the Sierras and northward. The 
breeding of the chipping sparrow on the lower Colorado is problem- 
atical; if a fact, it is a remarkable instance of extreme range of a 
species through zones from low Lower Sonoran to Canadian, inclusive. 

Thirty specimens were obtained by us (nos. 13170-13199) and 
these show the time of the partial prenuptial molt to oceupy some- 


a} 


1914] Grinnell: Mammals and Birds of the Colorado Valley ila 


what less than one month (in different individuals, March 9 to April 
3). The prenuptial molt in Spizella passerina arizonae involves in 
both sexes and apparently all ages, only the capital tracts, and, per- 
haps in only a few individuals, part of the pectoral region. This is 
apparently the same condition of affairs found in Spizella passerina 
passerina, of eastern North America, as described by Dwight (1900, 
peplg9)) 

Since there are in the Museum two skins (nos. 4160, 4161) taken 
by J. G. Cooper at Fort Mohave, December 24, 1860, it is evident that 
the western chipping sparrow passes the winter in the Colorado Valley. 


Spizella breweri Cassin 
Brewer Sparrow 

Since this sparrow was noted as early as February 19, when it 
was common a few miles south of Needles, a fair inference is that it 
is a winter sojourner in the region. At any rate, from the above date 
on until the middle of April, scattering flocks were of frequent note 
at all our collecting stations, on either side of the river, down to a 
point on the California side four miles north of Potholes. Here several 
individuals were seen April 23, the last for the season. As with the 
chipping sparrow, with which the Brewer sparrow was not infrequently 
associated, the latter occurred both on the desert uplands and along 
the river bottom lands. The preferred feeding grounds were probably 
on the desert, the river being visited only for water. 

The series of specimens of Spizella breweri secured (nos. 13201— 
13226) shows that there is a partial prenuptial molt, beginning (as 
shown by no. 13219, a male) as early as March 11, and continuing at 
least until March 27. This molt, however, is very limited in extent, 
involving only the chin and throat in addition to replacement of a 
part of the feathers of the crown, particularly anteriorly, and the chest. 
There is a scarcely discernible change in color, resulting in slightly 
whiter throat and clearer ash and clay color on the crown, certainly 
not enough to warrant the molt from the standpoint of color signifi- 
cance. Neither does the very small amount of wear which the speci- 
mens in hand up to the first of March appear to have incurred, seem 
sufficient to make a spring molt imperative. Yet the latter would 
appear the best explanation, restricted area of replacement pointing 
towards relative importance of the process. A spring molt may have 


2, University of California Publications in Zoology | Vou. 12 


been more essential as a renewal of worn plumage formerly than at 
the present time. 

Cooper, in at least three places (1861, p. 122; 1869, p. 475; 1870a, 
p. 75) made record of Spizella pallida from Fort Mohave. He states 
that two specimens were secured. In the second citation Cooper even 
emphasized that it was ‘“‘not S. Brewerii’’ he had. A little later 
(1870b, p. 209) he for the first time gives Spizella breweri from Fort 
Mohave, then without mention of S. pallida. It always seemed prob- 
able, therefore, that Cooper’s ‘‘ 
other authors, was really a misidentification of brewert. There are 
no Cooper specimens of Spizella in the Museum of Vertebrate Zoology 
bearing either name. At my request Dr. C. W. Richmond kindly 
looked into the matter at the United States National Museum. He 
writes me that there is a specimen of Spizella breweri, so identified 
by Ridgway and now no. 208619, labeled in Cooper’s handwriting 
“Spizella pallida,’’ and taken by him at Fort Mohave, April 2, 1861. 
This is evidently one of the skins upon which Cooper’s record of 
pallida was originally based, so that there is now no doubt but that 
brewert was the species really involved. Needless to say, Spizella 
pallida is not known from any station in this latitude so far west. 


pallida,’’ although quoted freely by 


Spizella atrogularis (Cabanis) 
Black-chinned Sparrow 


California side opposite The Needles, in a ravine among the hills, 
March 3, male taken (no. 13200); the only one of the species seen 
on our whole trip. 


Junco oreganus thurberi Anthony 


Sierra Junco 


In willow bottom, on California side at base of Riverside Mountain 
March 17, female taken (no. 13135); the only junco of any sort seen 
by us anywhere in the region. 

There is a well-preserved skin in the Museum (no. 4143) taken by 
J. G. Cooper at Fort Mohave, December 24, 1860, and as stated by 
Cooper (1870, p. 200) a few probably winter regularly in the Colorado 
Valley. 


1914] Grinnell: Mammals and Birds of the Colorado Valley 173 


Amphispiza bilineata deserticola Ridgway 
Desert Sparrow 

This is a bird of the upland deserts; not one was seen in the 
riparian belt. Around the base of Riverside Mountain, March 21, 
several were noted. On the desert back of Ehrenberg, Arizona, March 
27, a few were seen; on the California side, opposite Cibola, April 3, a 
pair was located in a ravine, where they appeared to have established 
a nesting site; and on the Arizona side ten miles south of Cibola, April 
9, two birds were noted in full song. Three skins were obtained, nos. 
13324-13326. 

There are also in the Museum two skins (nos. 4135, 4136) taken 
by J. G. Cooper at Fort Mohave, March 18 and April 4, 1861. 


Amphispiza nevadensis nevadensis (Ridgway) 
Nevada Sage Sparrow 


Common early in the season on the desert, especially in the salt- 
bush and sandy creosote associations. On the California side five 
miles south of Needles, and on the Arizona side above Mellen, Febru- 
ary 20 to 28, many were observed. Their habit of skulking along the 
eround among the close-set bushes doubtless often resulted in their 
eluding observation. 

Although intermediate sparrows occupied the same sort of ground 
and were present in much larger numbers, the two species kept in 
separate companies. The sage sparrows were prone to space themselves 
out much farther apart than Zonotrichia, and flushed singly rather 
than nearly simultaneously. 

Many sage sparrows were observed on both sides of the river at 
The Needles, March 1 to 5. Nine specimens preserved, nos. 13227- 
13235. 

There is also in the Museum a skin (no. 4137) taken by J. G. 
Cooper at Fort Mohave, January 26, 1861. That the species winters 
in the region is therefore practically established. 


Melospiza melodia fallax (Baird) 

Rocky Mountain Song Sparrow 
Common as a winter visitant to the riparian associations. Five 
miles below Needles on the California side, on both sides of the river 
at The Needles, on the Arizona side above Bill Williams River, and 


174 University of California Publications in Zoology | Vou. 12 


at Riverside Mountain, on the California side, the presence of this 
song sparrow was established by the taking of specimens. The last 
capture for the season was made at the last-named station, March 17. 
The ten specimens secured (nos. 13236-13245) are quite uniform in 
characters and are so exactly like birds from northern Nevada as 
to leave no doubt as to the source of those individuals of fallax visit- 
ing the Colorado Valley in winter. 

Song sparrows were very often heard when it proved impossible to 
rout them out from the dense arrowweed thickets in which they took 
refuge. It thus happened that the identity of most of the song 
sparrow population known to occur in a locality could not be ascer- 
tained. Judging from specimens shot, up to the middle of March, 
there were about as many fallax as saltonis. After this time fallax 
had departed from the region for its summer home, and the resident 
form, saltonis, was the only song sparrow remaining. No difference 
in habits or notes was detected between the two forms, and they 
appeared to possess identical preferences in regard to feeding grounds 
and cover. Both often ventured a few yards into open grassy or weedy 
places oceasionally found adjoining the quail-brush belt; and both 
forms occurred through the willows to the water’s edge, where, 
especially at early morning or late evening, they foraged among drift 
or through root tangles. 

The name montana of Henshaw (1884, p. 224) is here considered 
synonymous with the older fallax, of Baird (1854). The latter has 
been shown to be incorrectly employed for the “‘desert’’ song sparrow 
resident along the Gila and Colorado rivers (see Grinnell, 1909, p. 
269), and I now fail to see good grounds for recognizing two song 
sparrows from the Rocky Mountain region. 


Melospiza melodia saltonis Grinnell 


Salton Sink Song Sparrow 


This was the form of song sparrow resident along the Colorado 
River. Song sparrows were noted in the vicinity of every one of our 
collecting stations from Needles and Mellen to Pilot Knob. Through 
the big valleys, as we floated along near shore especially in April and 
May, we were often almost continuously within hearing of them. The 
presence of fallax early in the season renders somewhat uncertain the 
oceurrence of saltonis at the two points (Needles and Mellen) where 


1914] Grinnell: Mammals and Birds of the Colorado Valley 175 


no specimens of it were actually secured. But elsewhere abundant 
material testifies to its status. 

The occurrence of this sparrow appeared to be correlated closely 
with the extent of the arrowweed association, or of the almost as 
attractive young willow growth. The lower ends of the large valleys 
thus appeared to offer the best conditions. <A series of lagoons below 
Ehrenberg margined with tules and surrounded by arrowweed was 


“ce 


notable for its song sparrow population, as was also the ‘“‘drowned- 
out’’ area just above the Laguna dam. Below Potholes, in a growth 
of mixed cane and young willow, song sparrows were notably numer- 
ous; also wherever there was a pure growth of cane, as along ‘‘ Cane- 
brake Canon’’ and on the California bank near Pilot Knob. 

All the above areas are yearly subjected to overflow, and this may 
be held to account for the fact that even up to May 15 not a single 
new nest or young bird of this species was found. In the vicinity of 
Salton Sink young have been taken as early as March 30, and are 
common by the middle of April. It would appear that the birds have 
in the former region habitually come to defer their nesting time until 
after the high-water period, the last of May and early June. It was 
further observed that most of the old nests (which were considered 
with fair certainty to have belonged to song sparrows) were in arrow- 
weeds or willows above the highest mud mark. The coating of mud on 
vegetation up to a regular height was indicative of the previous sum- 
mer’s flood level. So that, again, the song sparrows have here accom- 
modated themselves to the extraordinary conditions by building their 
nests much higher above the ground than is usual elsewhere. 

The fifty-one specimens (nos. 13246-13296) of Melospiza melodia 
saltonis obtained by our party, together with three other skins (nos. 
4178-4180) taken by J. G. Cooper at Fort Mohave in January, 1861, 
are in no perceptible degree different from the topotype series from 
Salton Sink. (See Grinnell, 1909, p. 268.) The larger series now 
available increases the range of variation in all characters, but not so 
much that any single specimen, even when considered by itself, would 
raise a question as to its identity. The Colorado Valley birds, as far 
as I am able to judge, are not in the least divergent towards fallax, 
it being understood that the latter name applies to the Rocky Moun- 
tain song sparrow. It would appear that saltonis is very sharply 
defined both geographically and specifically. It is of further note that 
Cooper’s specimens, taken fifty years ago, are identical with the 
average saltonis as it is today. 


176 University of California Publications in Zoology | Vou. 12 


Melospiza lincolni lincolni (Audubon) 
Lincoln Sparrow 


First observed February 26 at Mellen; this is not indicative, how- 
ever, that the species had not been present all winter in suitable 
places. Indeed, it seems quite probable that at least a portion of 
those individuals encountered later had wintered in the Colorado 
Valley. Very common the first week in March on both sides of the 
river in’ the vicinity of The Needles; also in lower Chemehueyis Valley 
and above Bill Williams River up to March 15; then, in reduced num- 
bers, at Riverside Mountain and Ehrenberg, at the latter point up to 
March 29. Finally one was taken opposite Cibola April 5, and the 
last for the season April 7, ten miles below Cibola. At all these points 
the Lincoln sparrows were confined to the riparian belt, where they 
affected brush and nearby grass patches, especially at the outer margin 
of the arrowweed association. They were usually found in close com- 
pany with song sparrows, though not in the least gregarious; in fact, 
many were shot by mistake as flushed singly from low herbage, during 
our pursuit of song sparrows. 

Among the twenty-seven specimens (nos. 13297-13323) consti- 
tuting the series of Lincoln sparrows preserved, much variation is 
displayed. Most of the specimens are quite typical of Melospiza lin- 
colni lincolni as breeding in the Sierra Nevada and Rocky Mountains; 
but several (six in particular, nos. 13301, 13304, 13311, 13313, 13320, 
13322) vary strongly in the direetion of Melospiza lincolni gracilis 
(—=M. 1. striata of the A. O. U. Check-List, 1910, p. 276). The peculi- 
arities of these lie in the slightly smaller bill, shorter wing and broader 
black streaking, especially on the dorsum. None, however, are quite 
like extreme gracilis, and this fact, together with the presence in the 
series of various intermediates, would seem to indicate that the Colo- 
rado specimens did not any of them hail from breeding stations in the 
coast belt proper (Sitkan district) but perhaps from an interlying 
area towards the interior. This is the more probable, too, in consid- 
eration of the fact that in no case was any strictly humid coast belt 
subspecies found wintering in the Colorado Valley. Possibly the Lin- 
coln sparrows breeding in the northern Rocky Mountain region are 
subject to greater variation especially towards darkening, and so 
the dark (and incidentally smaller) Colorado Valley examples may 
not be essentially gracilis. Material is lacking to demonstrate their 
status satisfactorily. 


1914] Grinnell: Mammals and Birds of the Colorado Valley 1 


-] 


“a 


Pipilo maculatus curtatus Grinnell 
Nevada Towhee 


Oceurred in the river bottom only, and even there not in large 
numbers. One was seen on the California side near Needles, February 
17, and another was taken on the same side five miles below Needles, 
February 20. One was secured and others heard on the California 
side of the river in the lower Chemehuevis Valley, March 9. One 
was heard on the Arizona side at Parker, March 15. And individuals 
were heard, and one finally secured, March 17, on the California side 
near Riverside Mountain. In nearly all cases the birds kept close to 
the arrowweed and quail-brush associations, from within which it was 
difficult to see them or drive them out, though they were readily 
heard. 

As none was seen after March 29, when two were routed out of 
an arrowweed thicket near Ehrenberg, this bird is doubtless to be 
considered only a winter visitant to the region. Careful examination 
of the three specimens obtained (nos. 13337-13339) showed them to 
be unlike the P. m. montanus resident in the mountains to the eastward 
in Arizona, and yet quite different from the P. m. megalonyx resident 
in the southern Sierran and San Diegan districts of California. Cast- 
ing about for comparable specimens disclosed the fact that Colorado 
River birds were practically identical with examples representing a 
previously unnamed form, inhabiting in summer portions of the Great 
Basin region at least of northern Nevada, eastern Oregon and north- 
eastern California. The Colorado Valley birds were thus evidently 
winter visitants from this region and not from any nearer locality, as 
far as known (see Grinnell, 1911, p. 309). 


Pipilo aberti Baird 
Abert Towhee 


Abundant resident of the riparian strip along both sides of the 
river with scarcely a break the whole way from Needles to the Mexican 
line. Of the four recognized riparian associations—willow, arrow- 
weed, quail-brush and mesquite—the latter two were the ones for 
which preference was most often shown. Although foraging a few 
yards outwardly from the mesquite belt, in no instance were these 
birds found to have followed up the desert washes away from the 


178 University of California Publications in Zoology — { Vou. 12 


river, although the lines of catclaw or palo verde would seem to offer 
congenial cover. The Abert towhee seems to be closely bound to the 
vicinity of water, in spite of its evident adaptation in color and 
structure to a region of desert conditions. 

Although we were camped almost continually within the riparian 
strip, we saw no evidences of the nesting of this species until April 
27, on which date near Potholes a half-grown juvenal was found. 
This, however, appeared to be exceptional, as many adults in pairs 
were to be seen. A juvenal was taken near Pilot Knob, May 14, 
where, again, there were no further evidences of nesting. In 1908, 
Museum collectors found the Abert towhee around the northwest end 
of Salton Sea, where nests were found plentifully in April; and in 
the same locality in 1911, Van Rossem (1911, p. 136) found eggs 
March 20 and 21. It would appear that the Abert towhee, as in 
the case of the song sparrow, has along the Colorado River deferred 
its regular time of nesting until relatively late in the season, so as to 
avoid the period of rising floods. 

A series of forty-six specimens (nos. 13340-13385) obtained by 
the present expedition shows striking uniformity in external char- 
acters; that is, individual variation both in respect to color and size 
lies within very narrow Imits. Taking a single feature: twenty-two 
adult males show a wing length of 89.9 to 95.1 millimeters. The 
average is 92.3, the mid-point of the range is 92.5, and the variation 
each side of the mid-point is but 2.8 per cent. In twenty-one adult 
females the average wing length is 86.8 millimeters; the extremes are 
83.0 and 89.4, the mid-point of the range is 86.2, and the variation 
on either side of the mid-point is 3.7 per cent. 

Three specimens are in the Museum (nos. 4115-4117) taken by 
J. G. Cooper at Fort Mohave, February 19 and April 4 and 24, 1861. 


Oreospiza chlorura (Audubon) 
Green-tailed Towhee 


First seen March 14, on the Arizona side above Bill Williams River, 
one or two individuals; next April 7, on the same side ten miles 
below Cibola, two seen; then April 15, on the California side twenty 
miles north of Picacho, several; two were seen April 26 on the 
Arizona side, five miles north of Laguna; and one, on the California 
side, May 4, five miles northeast of Yuma, and May 8 and 10, one 


Oh 


1914] Grinnell: Mammals and Birds of the Colorado Valley 179 


each day, near Pilot Knob. In all these cases the birds were in the 
riparian tract, in willows, arrowweed or quail-brush. Although not 
numerous, the species is clearly a well-established transient through 
the region. 

Five specimens secured, nos. 13332-13336, April 7 to 24, show no 
sign of even a partial prenuptial molt; if such is actually lacking, 
the species in this respect resembles Pipilo rather than Zonotrichia, 
again showing the affinities of Oreospiza to be nearer to the former 
genus than to the latter. 

Two specimens are in the Museum (nos. 4123, 4124) collected by 
J. G. Cooper at Fort Mohave, February 13 and March 11, 1861. 


Zamelodia melanocephala melanocephala (Swainson) 
Black-headed Grosbeak 


First observed April 12 on the California side twenty miles north 
of Picacho, one male; next April 25, eight miles east of Picacho, two 
males; then April 25, five miles north of Laguna, two males. May 4, 
on the California side five miles northeast of Yuma, many were 
observed, apparently as a result of arrested migration; for the pre- 
vious afternoon and all night, as well as on the day of observation, 
a strong west wind, laden with dust, was blowing across the desert. 
The black-headed grosbeaks in company with several other transient 
species were restlessly foraging through the sheltering willow timber 
of the river bottom. During the succeeding week, practically until 
we left the region, the species was numerous. On May 13, especially, 
another windy day, around the east and north bases of Pilot Knob 
many black-headed grosbeaks were observed towards evening roving 
out over the desert in companies, as if preparing for through flight. 

The above observations, in addition to the fact that dissection of 
specimens taken showed no signs of immediate breeding, leads to the 
conclusion that the species occurs in the region only as a transient. 
A peculiar thing was the late arrival here, as compared with that on 
the Pacific slope of southern California; in fact, the greater part of 
the migration appeared to take place here in early May, while the 
black-headed grosbeaks of the other region had arrived a full month 
previously and in the majority of cases already had nests and eggs. 
It would appear likely that the grosbeaks passing through the Colo- 
rado Valley are not bound for the Pacific Coast region, but rather to 


180 University of California Publications in Zoology [| Vou. 12 


the Great Basin and northern Rocky Mountain region, where the 
spring of the Transition zone dawns much later in the season, forcing 
a much later nesting season. 

This idea is further borne out by a comparison of specimens. 
Breeding birds from the Pacific Coast, selected as being fully adult, 
are smaller throughout, with particularly less tumid bill than the Colo- 
rado Valley birds of obvious maturity. The latter, however, in these 
particulars, as well as in certain points of coloration, as extent of 
black on head and extra amount of white on wings and tail, are 
almost exact counterparts of breeding birds in the Museum collection 
from the Pine Forest Mountains of northern Nevada. The May birds 
of the Colorado Valley were thus evidently bound for the elevated 
interior to the northward. The differences above referred to, as 
exhibited between Pacific Coast black-headed grosbeaks and those of 
the Rocky Mountain plateau, would appear to warrant recognition in 
nomenclature, if for no other reason than to serve in designating their 
separate course of migration. Ridgway (1901, p. 619) presents data 
showing the differences in size and proportions, and also (J. c., p. 617) 
refers to one of the color differences. But he does not consider the 
races so distinguished worthy of naming. His synonymy (l.c., p. 
620) would appear to show that the Rocky Mountain race should prob- 
ably be called Zamelodia melanocephala melanocephala (Swainson) 
while the Pacific Coast race should bear the name Zamelodia melan- 
ocephala capitalis (Baird). 

Eleven examples of the black-headed grosbeak were secured on the 
Colorado expedition, nos. 13433-13443. Among these is an aberrancy : 
No. 13439 is a male of otherwise full plumage, except for the tail, in 
which there is one old full-length, sticky and bedraggled feather, and 
ten short feathers, newly unsheathing; this unusual condition at this 
season was probably induced by loss of tail-feathers through some 
accident. 


Guiraca caerulea lazula (Lesson) 
Arizona Blue Grosbeak 
First seen May 1, four miles below Potholes, two males; the next 
day one male. Common at our station on the California side five miles 
northeast of Yuma, May 3 and 4, as also on both sides of the river 
in the vicinity of Pilot Knob. Observed here up to the time of our 
departure, May 15. There was reason to believe that this Lower 


1914] Grinnell: Mammals and Birds of the Colorado Valley 181 


Sonoran species was preparing to breed in the river bottom: none were 
seen outside of the willow and arrowweed associations, and here, 
especially in clumpy tracts of willow, the males were present and in 
full song, spaced out over the appropriate territory as if settled for 
the nesting season. Females were doubtless present also, but only one 
was seen, probably because of their extremely retiring mood during 
the season of courtship. 

The eight specimens secured (nos. 13444-13451) all on the Cali- 
fornia side of the river at the three stations above named, together 
with one more (no. 4132) taken by J. G. Cooper at Fort Mohave, May 
6, 1861, are all of the large-billed race mhabiting southern Arizona. 
They differ markedly from the race summering in central and southern 
California west of the Sierran divide, in much larger and more tumid 
bill. (See Grinnell, 1911, p. 163). 


Passerina amoena (Say) 
Lazuli Bunting 

First seen, one of each sex, April 11, twenty miles above Picacho; 
next, a few examples April 24 and 25, five miles above Laguna; then 
April 29 at Potholes, and almost daily thereafter at all points to the 
vicinity of Pilot Knob where last seen May 8. In all eases the birds 
were observed in the riparian bottom, usually in willows close to the 
river. There was no evidence of breeding, and the probabilities are 
that the lazuli bunting occurs in the region merely as a transient. 
Four examples were preserved, nos. 13390-13393. 

There is a specimen in the Museum (no. 4157) taken May 20, 
1861, by J. G. Cooper at Fort Mohave. Two of the five specimens 
have larger bills than any breeding bird at hand from the Pacifie 
slope of California. This might be interpreted as being a parallel 
to the case with the black-headed grosbeaks, the Colorado River bunt- 
ings being migratory to the Great Basin region. The material at 
hand, however, is insufficient to elevate the supposition to even a 
probability. And apparently there is, besides, a wide range of varia- 
tion of the characters in question. 


Calamospiza melanocorys Stejneger 
Lark Bunting 


On March 8, on the California side opposite The Needles, a flock 
of a dozen lark buntings alighted in the tops of some tall willows close 


182 University of California Publications in Zoology  [Vor. 12 


to the river. Four were shot, nos. 13327-13330, all proving to be males 
in winter plumage. In these the lower parts are irregularly blotched 
with black. Two more were seen the same day in a willow on the 
California side a few miles below, in Blankinship Valley. A solitary 
female (no. 13331) was secured ten miles below Cibola, on the Arizona 
side, April 8. This bird was perched upon the wall of an adobe ruin 
on a spur of the mesa abutting upon the river. 


Piranga ludoviciana (Wilson) 
Western Tanager 


Common as a migrant. First seen, a male. on the Arizona side 
five miles above Laguna, April 25; another male noted there the next 
day. Several were seen at Potholes, April 29, and four miles south 
of Potholes, May 1. 

Five miles northeast of Yuma, on the California side, May 4, a 
very conspicuous arrested migration was in evidence among many 
species of transient birds. A heavy westerly wind blew all day, and 
the thick willow and cottonwood growth close to the river furnished 
both shelter and forage. Western tanagers were present in large num- 
bers. I counted twenty in sight at one moment, fiying serially along 
the edge of the woods. In the vicinity of Pilot Knob, up to May 15, 
this species was numerous, both on the bottom lands, and out on the 
desert. On the 7th and 13th, many western tanagers, singly and in 
pairs. were seen flying to the northwest low over the desert mesa north 
of Pilot Knob, frequently alighting for a moment in creosote bushes 
or ocotillos before resuming their transit. Eight specimens of this 
tanager were obtained, nos. 13415-13422. 


Piranga rubra cooperi Ridgway 
Cooper Tanager 


First seen April 20 on the California side of the river eight miles 
east of Picacho: a single full-plumaged male which was seeured. It 
attracted attention through its typical tanager call note, “‘pritit’’; but 
the bird was difficult to discern in spite of its brilliant red attire. amid 
the vivid green of the new foliage of the willow thicket in which it 
ensconced itself. 


1914] Grinnell: Mammals and Birds of the Colorado Valley 183 


The species was next observed April 30 and May 1 in the bottom 
lands four miles below Potholes (in California), and then, on the 
same side of the river, five miles northeast of Yuma, May 3 and 4. At 
each of these places a dozen or more individuals were seen. On both 
sides of the river in the vicinity of Pilot Knob, May 5 to 15, this 
tanager was frequently observed. While the call note was closely 
similar to that of the western tanager, present as a migrant in numbers 
at the same time, the song was quite different, being a clear, full-toned 
warble more nearly like that of the black-headed grosbeak, yet with 
an individuality of its own. 

The Cooper tanager, as far as our observations went, is strictly 
confined to the willow association. Not one bird was seen even so far 
from this association as the mesquite belt. The species is evidently 
a regular summer visitant. 

The ten specimens secured, nos. 13423-13432, are all males. One 
of these was thought to be a female until dissected, as it is in a greenish- 
yellow phase of coloration. However, there are scattering pale reddish 
feathers in nearly every feather tract, though there is no indication 
that any molt is in actual progress. This is probably a case of 
arrested molt. 

The notable absence of female specimens in our series resulted in 
spite of our greater effort to secure that sex. It was not that the 
males were more conspicuous, though this may have been a factor 
against that sex, but that the females were at this season extremely 
wary. As a pair was approached the female was always first to take 
alarm. I was repeatedly able to distinguish females at a distance as 
soon as males, in some cases sooner, after having located the pair by 
their notes. The red of the male appeared to be neutralized by the 
shimmering green of the surrounding foliage, so that the male was no 
more quickly discerned than the female. The male, however, was far 
more conspicuous in flight across open spaces. 

There are in the Museum two skins (nos. 4202, 4203) taken by 
J. G. Cooper at Fort Mohave, April 29, 1861. 


Petrochelidon lunifrons lunifrons (Say) 
Cliff Swallow 
This swallow is evidently a common breeding species alone the 
Colorado River wherever appropriate nesting sites are afforded. But 
we found the birds remarkably late in arriving and in nest-building. 


184 University of California Publications in Zoology [ Von. 12 


Many cliffs, from the vicinity of The Needles on down the river, were 
seen with remains of nests adhering to their faces in rows or patches. 
The first birds noted were a very few flyimg high overhead near 
Riverside Mountain, March 17. Next seen near Ehrenberg, March 26. 
Several were seen April 12 coursing over the river and bottom lands 
twenty miles north of Picacho. Not noted again until May 5, when 
at Yuma a large colony was seen constructing nests on the steep rocks 
on the Arizona side near the prison. The birds were gathering mud 
at the edge of the river on the California side, thronging back and 
forth over the unusually narrow channel at this point. 

On the California side in the vicinity of Pilot Knob two large 
nesting colonies were in evidence. One was on a sandstone bluff of 
some twenty-five feet height past which the main current of the 
river swung with exceptional speed. Here the nests were attached, 
to the number of ninety-seven, mostly as yet incomplete, to the irregu- 
larly eroded surface of the rock from ten to twenty feet above the 
surface of the river at its present stage—evidently above the reach 
of the highest level at flood time. Four of the finished nests were 
investigated on May 9 and found to contain in two eases three fresh 
eggs, in two cases one fresh ege. 

On the same day, sixteen of the birds were shot for specimens. 
The skiff was repeatedly rowed out from shore a little below the 
colony, and as it was swept down stream by the current was pulled 
back again by means of a long rope manipulated by a man on shore. 
Those swallows flying nearest the boat upstream in front of the colony 
were shot at, so that the dead birds would float past within reach. 
Of the sixteen secured, fifteen were males. There is no way that 
I know of to determine sex until the birds are in hand. This would 
seem to indicate that the females at this time in the breeding season 
are most retiring, and that the males are the ones to make demonstra- 
tions of alarm when the colony is invaded by an enemy. 

The other colony located had taken possession of the concrete walls 
in the head-gates of the Imperial Canal at Andrade. The nests were 
being constructed in crowded rows in the lateral upper corners beneath 
the woodwork bridging the piers. The birds were gathering mud at 
the margin of the canal nearby, alight with uplifted wings. Four 
shot were all males. 

The superintendent of the canal, stationed at Andrade, complained 
that the swallows proved a nuisance by dirtying up the painted wood- 


1914] Grinnell: Mammals and Birds of the Colorado Valley 185 


work. He had tried to drive them away repeatedly in previous years 
by destroying their nests. But the birds persisted in returning. Last 
year he had procured a quantity of poisoned wheat and spread it out 
on nearby ground, without, however, bringing any perceptible results ! 

Our nineteen specimens, nos. 13396-13414, show slight peculiarities 
as compared with examples from central California and Illinois. They 
are smaller: wing 100.0 to 108.0, averaging 103.2 mm. (see Ridgway, 
1904, p. 48); the frontal ight patch is cinnamon instead of whitish; 
the feathers of the dorsum are more conspicuously edged with clear 
white; the belly is whiter. These differences are indicative of the 
probable fact that the cliff swallows of the lower Colorado Valley are 
near in derivation to the forms in Mexico, particularly Petrochelidon 
lunifrons tachina Oberholser, which is recorded north on the east side 
of the Mexican plateau to southern Texas. 


Hirundo erythrogastra Boddaert 
Barn Swallow 


Observed but once: one, evidently a migrant, flying overhead at 
Potholes, April 29. 


Iridoprocne bicolor (Vieillot) 
Tree Swallow 


Recognized but once: March 23 a small number were noted flying 
back and forth over the second bottom on the California side at our 
collecting station above Blythe. One specimen taken, no. 13394. Evi- 
dently only a transient through the region. 


Tachycineta thalassina lepida Mearns 
Northern Violet-green Swallow 


First noted March 4 opposite The Needles. No birds were actually 
seen; but a tumult of the characteristic twitterings high overhead 
belonged unmistakably to this swallow. The sky was dazzlingly bright 
so that small objects at but a moderate distance were invisible. Next 
seen with certainty, ten or so, on the Arizona side, March 7; then 
above Bill Williams, Mareh 13. <A few violet-green swallows were 
seen above Blythe March 23, in company with other swallows: simi- 


186 University of California Publications in Zoology — [Vou. 12 


larly at Ehrenberg, March 26 and 28; two seen opposite Cibola, April 
5: and a few ten miles south of Cibola, April 8. One specimen taken, 
no. 13395. The species occurs in the region as a transient only. 


Stelgidopteryx serripennis (Audubon) 
Rough-winged Swallow 


First noted February 20, three individuals, five miles below Needles 
on the California side; next noted at Mellen, Arizona, February 23, 
thenceforth observed daily and at every station all the way down 
the river to Pilot Knob. This swallow forages alike over the river 
and flood-plain, and out over the desert where I saw it as far as I 
went (opposite Cibola fully six miles from the river). At Ehrenberg, 
March 24 to 29, many rough-winged swallows were to be seen close 
about the old adobe houses. They roosted by the half-dozen on wires 
stretched overhead for drying meat. Rafter holes in the walls, up 
under the thatches of those houses still possessing roofs, would appear 
to offer attractive nesting sites for these swallows. Individuals were 
seen flying up to such openings, though actual nest-building had not 
yet commenced. 

Back in the hills, two to five miles west of the river and twenty 
miles north of Picacho, one or two pairs were noted in nearly every 
ravine. April 16 a nesting site was located in a naturally eroded hole 
in the face of a conglomerate wall. The cavity was too deep and 
too small to investigate. But the actions of the birds plainly indicated 
that it held either eggs or young. 

At a number of places below Picacho, and down to Pilot Knob, 
nesting holes of rough-winged swallows were observed in banks, but 
always above flood level. In no ease did we see birds attempting to 
nest in banks subject to undereutting or overflow, although these would 
appear to be in some eases otherwise quite suitable, suggesting extra- 


’ 


ordinary powers of anticipatory ‘‘diserimination’’ on the part of the 
birds. Near Pilot Knob, May 14, a nest burrow was found in the 
wall of a wash meeting the river at right angles, and only some fifty 
feet from the brink of the main bank. This burrow was about eight 
feet above the floor of the wash; the terminal cavity was found to 
contain one fresh egg. Twenty-six specimens of this swallow were 


secured, nos. 13452-13477. 


1914] Grinnell: Mammals and Birds of the Colorado Valley 187 


Bombycilla garrula (Linnaeus) 
Bohemian Waxwing 


Known from the region only from the female specimen, now no. 
4207 in this Museum, taken by J. G. Cooper at Fort Mohave, January 
10, 1861. 


Phainopepla nitens (Swainson) 
Phainopepla 


Everything indicated that this bird was common as a permanent 
resident of the region. It was, however, closely restricted to two 
narrow belts paralleling the river, one on each side; namely as con- 
stituting the mesquite association. The close coincidence of the range 
of the bird with the plant association in question was here clearly 
due solely to the preferred food afforded in constant and abundant 
quantity by the berries of the mistletoe parasitic upon the mesquite. 
Judging from experience elsewhere, there is reason to believe that 
the phainopepla would have availed itself of edible berries in what- 
ever part of the region these might have been produced. Yet the 
fact remained that in the Colorado Valley the bird’s presence and 
distribution was remarkably controlled by those of the mesquite ; 
where there were no mesquites, as at our station near Pilot Knob, not 
a phainopepla was seen; where the mesquite had amassed itself into an 
extensive belt, the birds abounded. 

In certain places, as on the Arizona side above Mellen, and on 
the California side opposite Cibola, this bird was, within the riparian 
strip, the most abundant single species. In such localities the birds 
overflowed in small numbers a little way up contiguous desert washes, 
foraging about palo verdes or ironwoods, which latter plant occasionally 
bore mistletoe clumps. 

The phainopepla nested earlier than most of the resident birds of 
the region. Bob-tailed young out of the nest were seen April 12 on 
the California side, twenty miles above Picacho. April 5, on the same 
side opposite Cibola, two nests were found, each containing two eggs 
nearly hatched. In one of these cases the nest was eight feet above 
the ground on a branch of an ironwood; in the other, sixty-two inches 
above the ground on a mesquite limb. A nest with three fresh eggs 
found April 11 at the first-named locality, was seven feet above the 


188 University of California Publications in Zoology | Vou. 12 


ground in a mesquite. These situations are representative of the 
average predilections of the species in other regions. 

Thirty-two specimens of the phainopepla were preserved, nos. 
13478-13509. There are in the Museum five specimens (nos. 4196— 
4200) taken by J. G. Cooper at Fort Mohave, December 24, 1860, 
and January 17 to April 13, 1861. 


Lanius ludovicianus excubitorides Swainson 
White-rumped Shrike 


Shrikes proved unexpectedly scarce in the region explored. In 
spite of our special exertions, only four specimens were secured, nos. 
13386-13389. Besides these, there are two more in the Museum (nos. 
4205, 4206) taken at Fort Mohave by J. G. Cooper, April 18, 1861, 
and December 26, 1860. The six specimens show much variation 
among themselves, and not enough of uniformity in any one character 
to suffice for recognition, as I had anticipated, of a Lower Sonoran 
form of the excubitorides type different from an Upper Sonoran or 
Transition one. It is quite possible that some of the variation 
exhibited, as in degree of paleness and size and outline of bill, may 
be due to the fact that in the series some individuals were resident 
birds and others winter visitants from the Great Basin region. At 
any rate, the small series is inadequate as a basis for any satisfactory 
study of subspecifie status. I am therefore including all under the 
name exccubitorides, with the suggestion that it will probably be 
found after more extensive collections are worked over that further 
subspecific separation will have to be made. 

Our first specimen was taken February 15 at Needles, and lke 
several other birds of that neighborhood, the plumage was seriously 
discolored by coal smoke. Another was taken five miles below Needles, 
February 22, and one additional pair seen there. A pair was seen 
on the Arizona side near Mellen, February 24 and 27. Next noted 
at Ehrenberg, where a lone individual was secured March 28; next on 
the same side five miles above Laguna, where one was taken April 24. 
Last seen May 15 near Hanlon Junction, just north of Pilot Knob. 
In all these cases, the shrikes were in desert washes from a half-mile 
to two miles from the margin of the riparian belt. As usual they 
chose the most open ground productive of an adequate food-supply. 


1914] Grinnell: Mammals and Birds of the Colorado Valley 189 


Vireosylva gilva swainsoni (Baird) 
Western Warbling Vireo 


First noted April 1, opposite Cibola. Thenceforth of almost daily 
observation at all our stations from there to Pilot Knob, where still 
common May 14. At times numerous in the willows; a few noted in 
the mesquite belt. Evidently a plentiful migrant through the region. 
Fourteen specimens taken, nos. 13537-13550. 


Lanivireo solitarius cassini (Xantus) 
Cassin Vireo 

Occurred only in April and as a transient. First seen on the 7th 
of that month on the Arizona side ten miles below Cibola. One was 
found on the 9th singing volubly from an ironwood in a desert wash; all 
the rest seen were in the willow belt. On the California side, twenty 
miles above Picacho, several were noted April 10 and 11. Thereafter 
none were seen anywhere. Four specimens, nos. 13533-13536. 

There is also in the Museum a skin (no. 4247) taken by J. G. 
Cooper at Fort Mohave, May 14, 1861. 


Vireo belli arizonae Ridgway 
Arizona Least Vireo 


First detected early in the morning of March 8, on the California 
shore opposite The Needles. Heard repeatedly the same day from 
both banks as we floated down to lower Chemehuevis Valley. Nests, 
a year or more old, were seen both in the vicinity of The Needles and 
above, and the inference was that the species arrived throughout the 
region in full force on the above date. Thenceforth the species was 
met with at all stations all the way down the river, being one of the 
most characteristic avifaunal elements in the riparian strip. The bird 
foraged in all of the component associations, but was perhaps best 
represented in the willow association, especially where there was an 
undergrowth of guatemote (Baccharis glutinosa). 

On the Arizona side above Bill Williams River, March 14, I was 
able to make some observations on local distribution. Here the willow 
association was narrow but well defined, and the vireos were closely 
confined to it. A singing male “oceupied each segment of about 200 
yards in this belt, just about the same spacing as the Lucey warbler 


190 University of California Publications in Zoology | Vou. 12 


in the adjacent mesquite belt. Each pair of vireos was closely delimited 
in its forage beat by that of its neighbor. Each pair in its own area 
actively resented encroachment by others of its own species. The 
vireos worked a rather low zone of foliage, from the ground up to a 
height of six or eight feet, just about the same, again, as in the case 
of the Luey warbler. It is of interest to note that the Sonora yellow 
warblers, which arrived much later, after the willows had leaved out, 
were spaced much closer, but foraged through a greater depth of 
verdure, from the summits of the willows to within five or six feet 
of the ground. The vireo’s domain was but slightly impinged upon. 

At Ehrenberg, Arizona, a newly started nest was found, March 29. 
It was at this date about half completed, and was attached to the 
forking stalk of a guatemote five feet above the ground. It would 
appear that many nests meet with disaster from their being built, 
as they so often are, in openings between thickets. These openings 
serve as passage ways for browsing cattle, which as they crowd through, 
force the supporting branches aside and demolish the nests. Evidence 
of a number of instances of this type of catastrophe came to notice. 
A partly completed nest found on the California side near Pilot 
Knob, May 12, the owners being seen in the vicinity, would, together 
with the previous instance, indicate a leneth of breeding season of 
at least six weeks. Repeated trials, following such common accidents 
as the above, may account for this. 

A nest found April 24 on the Arizona side, five miles above Laguna, 
was loeated three and one-half feet above the ground on a horizontal 
willow branch, beneath and darkly shaded by several small willow 
trees growing close together at the margin of an overflow slough. The 
nest preserved (no. 759), is of normal construction for vireos of this 
genus. The chief constituents are weathered mesquite and willow bark 
strips and spiderweb and cocoons; the inner lining is of fine round 
grass stems and shreds of dry grass blades. The eggs, four in number, 
were advanced about one-third in the process of incubation. The 
ege-shells are pure white, dotted very sparsely about the large ends 
with bay and hazel. 

The series of twenty-three adult specimens taken (nos. 13510- 
13532) establishes clearly the identity of the Colorado Valley birds 
with Vireo belli arizonae of south central Arizona. The range of this 
geographie race is thus carried over into the confines of California 
along the southeastern frontier of the state. 


1914] Grinnell: Mammals and Birds of the Colorado Valley 191 


Vermivora luciae (Cooper) 
Lucey Warbler 


First seen March 10, on the California side in lower Chemehuevis 
Valley ; there were three individuals foraging quietly in company with 
Audubon warblers and ruby-crowned kinglets in the sunlit summits 
of willows. The one shot was a male. The Lucy warbler is undoubt- 
edly absent from the region in winter, and the above observation 
indicates approximately the date of its arrival. 

Next noted March 14 on the Arizona side above Bill Williams 
River. Here, at least four males were located in mesquites, and as 
they were in full song and spaced apart, had doubtless settled upon 
breeding locations. The song is unmistakable, as far as all other birds 
of southern California and Arizona are concerned. It resembles the 
song of the Sonora yellow warbler in length and frequency of utter- 
ance and somewhat in quality, but with a distinct hurried and lisping 
effect reminding one of the song of the Lazuli bunting. 

On the California side, both at Riverside Mountain and above 
Blythe, Lucey warblers were numerous, and very closely confined to the 
narrow belt of mesquite. The singing males, each representing the 
forage area and nesting site of a pair, were spaced out very uniformly, 
so that an estimated strip of about 200 yards in length belonged to 
each. The birds foraged out to a limited extent from the mesquites 
towards the river into the arrowweed and willows, and away from 
the river at the mouths of washes into the ironwoods and palo verdes. 
But the metropolis was always most emphatically the mesquites. At 
this time, March 18 to 23, the mesquites were just coming into leaf, 
and the new yellow-green foliage was prolific of insect life and formed 
both a productive food-source and an excellent cover for a low-foliage 
feeder, such as the Luey warbler pre-eminently is. 

At Ehrenberg, Arizona, the last week of March, the species was 
common, as also on the California side opposite Cibola the first week 
of April, and on the Arizona side, again, ten miles below Cibola. At 
the latter point a nest, nearly completed, was found April 8. It rested 
upon a loosened skein of bark on the under side of a slanting mesquite 
trunk, and was five feet above the ground. 

On the California side the Luey warbler was fairly common in 
the vicinity of our stations twenty miles above and eight miles east 
of Picacho, at both points being closely adherent to the mesquite strip. 


192 University of California Publications in Zoology — [Vou. 12 


Not one was observed anywhere below the last-named station, and this 
fact we ascribed to the lack of mesquite at the remainder of the points 
visited, this wood having been cut out for fuel. 

It was a particularly gratifying circumstance that a nest of the 
Luey warbler was found by us on California territory, thus adding 
a species to the state list of breeding birds. As above implied, indiy- 
iduals were encountered in fair numbers at our station twenty miles 
above Picacho. Here, as usual, they affected the mesquite belt, but 
strangely enough the nest found was situated in a nearly dead iron- 
wood, at the base of a hill rising abruptly from the river bottom. This 
tree (see pl. 12, fig. 19) evidently owed its failing condition to the 
rising water level in the ground in which it grew, a circumstance as 
fatal to an ironwood, as it is, up to a certain degree, propitious to a 
mesquite. There were young mesquites in the immediate vicinity. 

The nest was built thirty-five inches above the ground in the crotch 
where a steeply inclined branch sprang from the main trunk, which 
was here about one foot in diameter. The nest was sheltered from 
above by a two-inch strip of loosened bark and an outstanding twig. 
The nest was thus practically within a closed cavity save for an 
approach in one direction, as shown in the photograph (pl. 13, fig. 20). 
This falls within the known predilection of the species elsewhere, as in 
the upper Gila Valley of south central Arizona (Gilman, 1909, p. 168), 
where out of ‘‘twenty-three nests observed, twelve were in natural cay- 
ities, four under loose bark, four in woodpecker holes, and three in 
verdin’s nests.”’ 

The nest in question was evidently a relining of a previous year’s 
structure, the latter being distinguishable by its compactness and 
admixture of dried mud, as if it had been sifted full of wind-blown 
dust and then drenched by heavy rain. There was no evidence that 
the tree had been submerged to this depth. The new portions of 
the nest (no. 754) are loosely formed of various feathers (mostly 
down-feathers of desert quail), mingled with weathered shreds of 
grass. There is a scanty lining of fine hairs. The cavity of the 
nest is 45 mm. across, by 22 mm. deep. 

The three eggs, taken April 12, were considerably incubated, and 
thus constituted a full complement. The shells are pure white, with 
a fine and abundant speckling of vinaceous, vinaceous-cinnamon and 
cinnamon-rufous, chiefly in an agglomerated ring about the large 
ends. Some of the markings nearest the pole at the large end are 


1914] Grinnell: Mammals and Birds of the Colorado Valley 193 


splotehy rather than punctulate. The eggs are rounded-ovate and 
measure in millimeters 14.0 by 11.7, 14.6 by 11.8, and 14.0 by 11.6. 
Only the female parent was observed in the vicinity of the nest and, 
as noted by Gilman, there was a notable lack of expressed anxiety. 
The bird merely remained among distant mesquites, uttering an ocea- 
sional faint, one-syHabled alarm note. 

Twenty-five specimens of this warbler were obtained, nos. 13579- 
13603. 

The Lucey warbler was originally discovered at Fort Mohave, which 
is on the Arizona side of the river. Its deseriber, J. G. Cooper (1861, 
p. 120), remarks as follows: ‘‘This bird was common at Fort Mojave, 
near Lat. 35°, in the Colorado Valley, where it arrived about March 
25th, and remained until I left there, the twenty-eighth of May. I saw 
none along the Mojave river, on the route westward. I collected five 
male specimens and one female.’’ 

Two of Cooper’s specimens are in the Museum collection: female, 
no. 4266, April 5, 1861; male, no. 4267, April 11, 1861. These may 
with propriety be considexed co-types, at least. Although no single 
specimen had been designated as type, Ridgway (1902, p. 474) indicates 
that a specimen which he considers the type is in the United States 
National Museum; this is probably one of the five males referred to 
by Cooper. Both our specimens have the original label, on faded blue 
note-paper, giving full data, entirely in Cooper’s own handwriting. 
The female has ‘‘Iuciae, n.s. Cooper’’ in one corner in the same faded 
ink as on the rest of the label, except that both specimens have 
“Tueiae, J. G. C.’’ interlined in blacker ink, just beneath the original 
““Helminthophaga,’’ evidently inscribed by Cooper at some later time. 


Vermivora ruficapilla gutturalis (Ridgway) 
Calaveras Warbler 


First noted April 7 to 9, on the Arizona side of the river, ten 
miles below Cibola. Here they were not uncommon in the upper foliage 
of blossoming willows. Next observed on the opposite side, twenty 
miles north of Picacho, April 11, with similar mode of occurrence ; 
then five miles above Laguna, April 25, and at Potholes, April 29. 
Here, and four miles south of Potholes, up to May 2, this was one of 
the commonest warblers. One was seen five miles northeast of Yuma, 
May 3; and in the vicinity of Pilot Knob, May 6 to 15, a few were 


194 University of California Publications in Zoology — [Vou. 12 


seen daily. The species was thus well represented as a migrant 
through the region. 

Twelve specimens were taken, nos. 13551-13562. No. 13522, an 
adult male, shows conspicuously a character not mentioned in text 
descriptions, namely, a mixture of long, fine, black hairs (filoplumes) 
in plumage of sides and especially the flanks. Examination of appro- 
priate series of specimens shows this character to be possessed in 
varying degrees, even in female first-winter plumage, by both Vermi- 
vora ruficapilla gutturalis and V. r. ruficapilla, but not by other 
members of the genus. 


Vermivora celata celata (Say) 
Orange-crowned Warbler 


Apparently the only member of the genus present in the Colorado 
Valley through the winter. That it does winter here is quite apparent 
from its being observed by us almost daily during the latter part of 
February, and early March, whenever we were at work in favorable 
localities. Later its identity was obscured by the arrival of its close 
relative, Vermivora celata lutescens, which, as a migrant, outnumbered 
V. c. celata. 

The orange-crowned warbler was closely confined to the riparian 
strip, where it foraged singly at a low level in the tangle of under- 
brush in the willow association, or, less often, in the arrowweed 
thickets. Its presence was betrayed as a rule through the sharp, 
single call note. Though usually repeated at not infrequent intervals, 
this clue was not always alone sufficient for the discernment of the 
bird, because of the impenetrability of its cover. 

The first bird was seen in the river bottom near Needles, February 
17. On the 20th, five miles below Needles, a specimen (no. 13574) was 
secured. Although observed frequently, no more were obtained until 
April 8, when one (no. 13575) was taken on the Arizona shore, ten 
miles below Cibola. On the California side, eight miles east of 
Picacho, another (no. 13576) was taken April 20. And two examples 
were secured on the same side near Pilot Knob, May 9 and 14 (nos. 
13577, 13578). 

These five specimens are unequivocally celata, save that one shows 
a slight but notable aberraney. This is no. 13577, a male, in which 
the grayness characteristic of celata pervades the plumage as usual, 


1914] Grinnell: Mammals and Birds of the Colorado Valley 195 


except facially, where it is replaced on the chin, lores, superciliary 
line, lower eyelid and fore part of auriculars with the clear greenish 
yellow peculiar to V. c. lutescens. This abrupt replacement is bilater- 
ally uniform and so conspicuously in evidence as to suggest the style 
of coloration in the yellow-faced gray-bodied verdin. It is probable 
that the warbler in question is a hybrid between V. c. celata and V. ec. 
lutescens, having been bred in a region where the ranges of these 
two forms meet. Such a locality is Prince Wilham Sound, Alaska 
(see Grinnell, 1910, p. 409). 


Vermivora celata lutescens (Ridgway) 
Lutescent Warbler 


Common as a migrant, chiefly in April and May. Two specimens 
taken February 23 and 28 at Mellen, Arizona, may have been winter- 
ing in the region; no more were encountered until March 29, at Ehren- 
berg, when two in full song came to notice; thereafter the species was 
observed almost daily, though not in numbers. The willow strips 
formed the main forage-ground. The last specimen was taken May 7, 
on the California side near Pilot Knob. 

The eleven specimens preserved (nos. 13563-13573) are varyingly 
grayer than breeding birds (May and June) from the Pacifie Coast. 
But it is clear that a slight ashy obscuration in some degree accounts 
for the duller yellow of the early spring birds. The underlying yellow 
of the male appears just as intense in February and March examples 
as in June birds. Wear removes most or all of the ashy feather tips. 


Dendroica aestiva sonorana Brewster 
Sonora Yellow Warbler 


First encountered April 8, a single adult male, on the Arizona side 
ten miles below Cibola; next noted April 10, two in song on the 
California side, twenty miles north of Picacho; many on the morning 
of the 17th near the same place. When we reached our station on 
the California side eight miles east of Picacho, April 17, Sonora yellow 
warblers were abundant. As this was a particularly favorable locality, 
more so than the last, it is fair to infer that the species had arrived 
in force some days previously. Even so, the first appearance of yellow 
warblers along the Colorado River was later than the usual date of 


196 University of California Publications in Zoology [ Vou. 12 


arrival of the closely related Dendroica aestiva brewsteri at the same 
latitude in the San Diegan district of southern California; and this 
in spite of the lower zone of the Colorado Valley. 

The Sonora yellow warblers were abundant at all suitable places 
along down the river from Picacho, the stations being indicated in the 
accompanying table of measurements. They adhered closely to the 
willow association, the mode of their forage zone lying somewhere in 
the crown foliage of the willows and cottonwoods, the height above the 
ground of course varying with the stature of the trees. The females 
were hard to locate, but the incessant song of the male birds rendered 
the latter easy to detect. Observation on April 19 in a very extensive 
willow tract in ‘‘Charlie’s Valley,’’ eight miles east of Picacho, showed 
them to be regularly spaced out through the top foliage at an estimated 
frequency of four per acre. Thus, at this point, a quarter of an acre, 
with a depth of perhaps twelve feet of foliage, was the forage allow- 
ance of each nesting pair of warblers. This was a greater congestion 
of warbler population than in most places, because of the evident 
unusual favorableness of the environs. 

At another place, five miles above Yuma, May 3, there was only 
one singing male to two acres of willows. With a minimum of 640 
yellow warblers to the square mile, their aggregate numbers in the 
Colorado Valley must reach an enormous total, considering the vast 
area of first bottom. Since there is in the Museum collection a female 
specimen (no. 4265) of Dendroica aestiva sonorana taken by Cooper 
May 4, 1861, at Fort Mohave, it is reasonable to suppose that this 
form breeds north in uniform abundance along the valley to the Nevada 
line. In spite of the numbers of the birds, we failed to run across 
nests. This may, however, have been due to nesting being deferred 
until later than the middle of May. 

The series obtained of this warbler (forty-seven specimens, nos. 
13604-13650) provides so much material of a bird not heretofore 
adequately represented in our collections, that a rather detailed exam- 
ination of its characters seems warranted. Furthermore, so many 
examples of one species of migratory bird from so limited an area, 
and taken practically within a month’s time at one season, would 
seem to be a basis from which to obtain a fair idea of normal variation. 

Referring first to the accompanying table of measurements, it will 
be noticed in the males that the average and the mid-point of the 
range practically coincide; that the variation in the six respects varies 


1914] = Grinnell: Mammals and Birds of the Colorado Valley 197 


from six to nine per cent on either side of the mid-point; that the 
wing length is most constant; that in average dimensions females are 
smaller than males except that the bills in the former are slightly the 
larger. 

Although little more than three weeks apart in time of capture, 
the May birds are distinctly more worn than those of April 18. This 
undoubtedly accounts for the slight lessening in wing length, and 
also modification in wing formula, as exhibited bottomwards in the 
appropriate columns. In obtaining the wing formula as given, the 
primaries are numbered consecutively from the carpal joint distally. 
The numbers are arranged in order of relative length of the primaries, 
the first given being the longest; thus 8-7-6-9-5-4-3-2-1 means that the 
eighth primary is longest, the seventh next in length, and so on. 
Since in this bird the primaries proximally from number 5 in all 
individuals decrease in length with regularity, only the variable 
(outer) portion of the formula is given. The variation in wing for- 
mula as shown in the table appears to be greater than it really is, for 
in most cases 6, 7, and 8 are very nearly of a length, 9 being propor- 
tionally shorter but in no case as short as 5. Hight and 7 are most 
frequently of greatest length. 

In coloration the females are paler, being more ashy and less yellow 
than in either D. aestiva aestiva or D. aestiva brewsteri. The males, on 
the other hand, have the yellow more intense and extensive, involving 
the whole head and back, and represented on the wings and tail by 
much broader edgings. The green mantle of brewsteri is represented 
in sonorana by a more restricted area of dull yellow, the whole head 
and rump being brightly yellow. The yellow of the crown is often 
tinged strongly with chestnut, and the dorsum is streaked narrowly 
with a darker tone of the same color. The underparts are even more 
narrowly chestnut-streaked than in brewsteri. Variation in the latter 
respect is considerable, from a case where the ventral streaking is 
searcely perceptible to the opposite extreme, where the streaking is 
about as well defined as in average brewsterv. 

Sonorana, as shown by the present series, is an easily recognized 
form, in spite of its great range in individual variation. For, among 
so many characters, where one fails, others are left to hold to. 


[ Vou. 12 


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200 University of California Publications in Zoology | Vou. 12 


Dendroica aestiva brewsteri Grinnell 
California Yellow Warbler 
Appeared as a migrant through the willow bottom, individuals at 
large being distinguished from the breeding D. a. sonorana by their 
silence. Five specimens taken, nos. 13653-13657, bear locality and 
date as follows: Male, California side, twenty miles north of Picacho, 
April 12; male and female, same side, five miles northeast of Yuma, 
May 4; male and female, same side, near Pilot Knob, May 9 and 6, 


respectively. 


Dendroica aestiva rubiginosa (Pallas) 
Alaska Yellow Warbler 


A late migrant along the willows, secured only on the California 
side near Pilot Knob; two specimens: male, no. 13651, May 9; female, 
no. 13652, May 14. 


Dendroica auduboni auduboni (Townsend) 
Audubon Warbler 


Varyingly common as a winter visitant or transient on either side 
of the river from the first day of our work, February 15, and from 
our first station, Needles, until the last day of our work at our last 
station, May 15, at Pilot Knob. Full-plumaged males were still noted 
on the latter date, which would indicate unexpectedly late migration. 
The species was seldom noted outside of the willow association, and 
then only a short distance away, in the mesquite or a little beyond. 
Twenty-five specimens were taken, nos. 18658-13682. 

There is also in the Museum a skin (no. 4257) taken by J. G. 
Cooper at Fort Mohave, February 24, 1861. 


Dendroica nigrescens (Townsend) 
Black-throated Gray Warbler 
First seen April 2, on the California side opposite Cibola, a solitary 
female. Next seen on the Arizona side ten miles below Cibola April 9, 
a male; then more frequently, sometimes in small droves with other 
warblers, twenty miles north of Picacho, eight miles east of Picacho, 
five miles north of Laguna, four miles south of Potholes and at Pilot 
Knob. The last one was noted at the latter place May 9. The species 


1914] Grinnell: Mammals and Birds of the Colorado Valley 201 


was noted all along only in willows or mesquites. Seven specimens 
taken, nos. 13683-13689. Evidently occurs through the region as a 
migrant only. 


Dendroica townsendi (Townsend) 
Townsend Warbler 


First noted April 26, on the Arizona side five miles north of Laguna, 
two seen and another secured. At Potholes, April 28 and 29, fully 
twenty-five were noted; four miles south of Potholes May 2 one was 
taken, and another five miles northeast of Yuma, May 4. In the vic- 
inity of Pilot Knob several were seen May 13 and 14. At every other 
place the birds were exclusively in the willows, but here they were in 
arrowweed as well. Evidently a transient. Nine specimens taken, 
nos. 13690-13698. 


Dendroica occidentalis (Townsend) 
Hermit Warbler 


First observed, a male, April 20, on the California side, eight miles 
east of Picacho; next, a male, on the Arizona side five miles north of 
Laguna, April 25; then at Potholes on the 28th and 29th, several; 
five miles northeast of Yuma, May 3, a male; and near Pilot Knob, 
May 9, a female. A through migrant. Six specimens taken, nos. 
13699-13704. 


Oporornis tolmiei (Townsend) 
Tolmie Warbler 


First noted, a male, April 12, on the California side of the river, 
twenty miles north of Picacho. Next observed, in numbers, both sexes, 
April 25 and 26, on the Arizona side five miles above Laguna. Then, 
on the California side, almost daily in varying numbers, at Potholes, 
four miles below Potholes, five miles northeast of Yuma, and in the 
vicinity of Pilot Knob. The species was still common in the latter 
locality, May 14. 

In all the above places, the species adhered closely to the arrow- 
weed and quailbrush associations, frequently visiting the water’s edge 
through the willows, especially during midday. The brush associa- 


202 University of California Publications in Zoology — [| Vou. 12 


tions are preferred doubtless because nearest like their summer asso- 
ciational habitat. Ten specimens taken, nos. 13705-13714. 

There is also in the Museum a skin (no. 4255) taken by J. G. 
Cooper at Fort Mohave, April 24, 1861. 


Geothlypis trichas scirpicola Grinnell 
Tule Yellowthroat 


A cursory glance at the sixteen specimens of yellowthroats obtained 
shows clearly that as regards size and color two diverse types are repre- 
sented. Observation in the field showed that there were two categories 
as regards behavior. And these two assortments, as far as data justi- 
fies, appear to coincide; namely, the smaller, grayer and duller birds 
were quiet, and doubtless migrating, while the larger, more brightly 
colored birds were in full song, evidently the breeding subspecies. 
Both forms were found in the densest available cover, the resident 
birds, however, being always met with in the immediate vicinity of 
water, while some of the migratory individuals were in quail-brush on 
the second bottom many rods from the river or nearest slough. It was, 
of course, impossible to distinguish the forms by appearance in the 
field: in fact, some of the specimens themselves are only with difficulty 
and some uncertainty placed in one or the other category. 

After an independent study of the Museum’s series of yellowthroats 
from the western United States, British Columbia and Alaska, I have 
arrived at the same conclusions as expressed by Swarth (1912, p. 72). 
These conclusions are that there is no recognizable Pacifie coast race 
(arizela), the name occidentalis applying rightly to all breeding yellow- 
throats of North America west of the great plains, except those of the 
San Francisco Bay region (G. t. sinuosa) and those of southern Cali- 
fornia and Arizona (G. t. scirpicola). 

The breeding males of the Colorado Valley agree with those of 
southern Arizona and the San Diego district in southern California, 
and differ from occidentalis in large size throughout, and in brighter 
coloration. The latter feature consists in a deeper toned yellow below 
and an extension of this yellow posteriorly to include the abdominal 
area; in brighter yellow of the crissum; in the flanks being washed with 
a darker tone of clay color; in the dorsum being pervaded with golden 
yellow instead of being of a grayish east, as in occidentalis; and in 
the grayish area on the head bordering the black mask posteriorly 


1914] Grinnell: Mammals and Birds of the Colorado Valley 203 


being very broad, tending to cover in some cases the whole pileum, 
though becoming concealed beneath an olive wash posteriorly, and 
with a mixture of yellow. 

Extreme examples of scirpicola are conspicuously different from 
the nearest approach to it in appearance among many specimens of 
occidentalis from Nevada, northern California, British Columbia and 
southeastern Alaska. But among the considerable number of speci- 
mens from the range of scirpicola, there are some which have so far 
defied every effort to distinguish them satisfactorily from occidentalis. 
The problem is complicated by the fact that there are yellowthroats 
moving through the range of scirpicola at the same season that the 
resident birds are nesting, that is, until the last week in April at least. 

The twelve Colorado Valley examples referred to scirpicola (char- 
acterized by Grinnell, 1901, p. 65) are listed in an accompanying table 
to show locality and date of capture, and measurements. One speci- 
men, no. 13723, deserves special comment because of an extraordinary 
aberraney in coloration: the whole throat is clear, intense cadmium 
yellow, in marked contrast with the normal canary yellow of the chest 
and remaining lower parts; otherwise the bird is lke the average. 
This extra intensification towards orange, though over a restricted area, 
is probably a color change of significance along the same line as dis- 
cussed in this paper under Colaptes chrysoides. 

The tule yellowthroat was closely restricted to dense vegetation 
erowing close beside or over water. Tules bordering sloughs formed 


MEASUREMENTS IN MILLIMETERS OF GEOTHLYPIS TRICHAS SCIRPICOLA 
FROM THE COLORADO VALLEY 


Mus. No. Sex Locality Date Wing Tail Tarsus Culmen 
13720 g Riverside Mt., Calif. side Mar. 18 57.7 54.5 20.0 11.1 
18721 og Riverside Mt., Calif. side Mar. 20 56.5 52.8 20.3 11.0 
13723 og Ehrenberg, Ariz. side Mar. 29 55.1 53.3 19.7 11.8 
13728 g 5mi.N. Laguna, Ariz. side Apr. 25 538.9 50.5 20.9 11.5 
138729 g 4mi.S Potholes, Calif. side May 2 57.0 52.1 20.0 11.2 
13731 g 5 mi. N.E. Yuma, Calif. side May 4 BY EO, aly PAU eee 
13732 g 5 mi. N.E. Yuma, Calif. side May 3 58.6 56.7 20.2 11.6 
13733 g 5 mi. N.E. Yuma, Calif. side May 4 57.3 53.6 20. 12.1 
13734 g Near Pilot Knob, Calif. side May 6 8.9) 00:6) sees IES 
13735  g Near Pilot Knob, Calif. side May 14 57.5 55.0 20.8 12.0 

Average of the males, 56.6 53.6 20.3 11.5 
13725  Q Opposite Cibola, Calif. side Apr. 1 538.5 49.5 19:8 11.9 


13730 2 4 mi.S. Potholes, Calif. side May 2 54.8 49.3 20.2 11.7 


204 University of California Publications in Zoology — [Vou. 12 


the usual habitat. Occasionally dense growths of very young willows, 
being then similar in habit to tules, offered the proper conditions. 
Along the lower course of the river, thickets of cane which clothed the 
abrupt banks were the chief resort. Where tules failed, as near Pilot 
Knob, this was the only accepted cover. On April 17, as we floated 
through Canebrake Cafion, three to seven miles below Picacho, the 
songs of yellowthroats were heard almost continually. Here they were 
inhabiting the jungles of cane which grew down into the river along 
both shores. Unfortunately we did not have the time to devote to 
search for nests, and information in regard to breeding habits is 
lacking. 


Geothlypis trichas occidentalis Brewster 
Western Yellowthroat 


Occurred as a migrant along the valley, as noted under the pre- 
ceding heading. The four specimens taken and referred to this sub- 
species are all males, and were obtained as follows: No. 13722, Blythe, 
California side, March 23; 13724, ten miles below Cibola, Arizona side, 
April 9; 13726, 13727, twenty miles north of Picacho, California side, 
April 11 and 12. These migrants were quite likely bound for the 
Great Basin to the northward, being indistinguishable from specimens 
from northern Nevada. 


Icteria virens longicauda Lawrence 
Long-tailed Chat 


First seen April 25, above Laguna, one individual; three in the 
same place the next day. On the 27th, at Potholes, several were seen ; 
and thenceforth, at Potholes and four miles below, five miles northeast 
of Yuma, and on both sides of the river in the vicinity of Pilot Knob, 
chats were abundant. Twenty-four specimens were taken, nos. 13769-— 
13792. 

There is in the Museum a skin (no. 6423) taken by J. G. Cooper at 
Fort Mohave, April 25, 1861. 

This bird was everywhere closely confined to the willow associa- 
tion, at any rate never seen beyond the arrowweed. In point of actual 
numbers, they were probably not so numerous as yellow warblers or 
song sparrows; but in volume of noise they exceeded all other birds 
combined. These chats of the Colorado possess an amazingly large 
vocabulary in imitation of other sounds. On May 3 a few minutes of 


1914] Grinnell: Mammals and Birds of the Colorado Valley 205 


attention to a single individual was enough for unmistakable recog- 
nition of notes or songs of Abert towhee, flicker, kingfisher, Bullock 
oriole, and tanager. The chat is here a far better mocker than the 
mockingbird itself. 


Wilsonia pusilla pileolata (Pallas) 
Alaska Pileolated Warbler 


Appeared commonly as a migrant through the riparian strip. Six- 
teen specimens (nos. 13736-13751) were taken, from April 19 to May 
12, inclusive, representing the following localities: California side, 
eight miles east of Picacho, April 19 and 20; Arizona side, five miles 
north of Laguna, April 21, 24 and 25; California side at Potholes, 
April 27; same side, four miles south of Potholes, May 1 and 2; same 
side, five miles northeast of Yuma, May 4; same side near Pilot Knob, 
May 8 and 12. 


Wilsonia pusilla chryseola Ridgway 
Golden Pileolated Warbler 


Common as a migrant, through the riparian strip. First seen 
March 9 and 10, one each day, on the California side in Chemehueyis 
Valley. Next noted March 20, one taken, near Riverside Mountain; 
thenceforth of almost daily observation in suitable places at nearly all 
stations until May 2, when the last one with certainty identified was 
taken on the California side four miles below Potholes. Intermediate 
points and dates of capture were: California side above Blythe, March 
23; Arizona side below Ehrenberg, March 26; ten miles below Cibola, 
Arizona side, April 7, 8 and 9; California side twenty miles north of 
Pieacho, April 11 and 12; same side, eight miles east of Picacho, 
April 18 and 20; Arizona side, five miles above Laguna, April 25 and 
26; four miles below Potholes, April 30, as well as May 2, as noted 
above. The series of specimens taken consists of seventeen examples 
(nos. 18752-13768). 

At times pileolated warblers were numerous in tracts of willow, 
but since it was impossible to distinguish between the race pileolata 
and the race chryseola without shooting, the proportion present of 
these two forms between April 19 and May 2 could not be judged. All 
individuals shot were, however, preserved. Between these dates, thir- 
teen pileolata were taken and seven chryseola. This proportion, nearly 


206 University of California Publications in Zoology | Vou. 12 


two to one, for this period may therefore be near the truth. But it 
is further observable from the data presented that pileolata was a 
later migrant through this region than chryseola. 


Anthus rubescens (Tunstall) 
Pipit 
At the time of our arrival at Needles, February 15, pipits were 
numerous close along the river, both on the grassy areas near the 
Indian camp and on the mud bars at the water’s edge. They were 
thenceforth observed through March at many points along the river on 
both sides, especially when we were floating down from station to 
station. They were still common the first week of April, opposite 
Cibola. But none were seen later than April 8, when a single bird 
was noted flying up river, ten miles below Cibola. 
Five specimens taken, nos. 13715-13719. 


Oreoscoptes montanus (Townsend) 
Sage Thrasher 


Observed only on Mareh 23 and April 2 and 3, so probably a 
transient purely. On the first specified date three were noted on the 
California side, above Blythe. On the latter two days fully a dozen 
were encountered on the same side opposite Cibola. In each ease the 
birds adhered closely to mistletoe-bearing mesquite and ironwood, 
the latter extending up the desert washes. Four specimens taken, nos. 
13793-13796. 


Mimus polyglottos leucopterus (Vigors) 
Western Mockingbird 


Common early in the season all along the river on both sides. 
Almost exclusively confined to the mesquite belt and to the ironwoods 
a short distance up washes, especially where these trees were laden 
with mistletoe. The berries of this parasite appeared to be the chief 
or only food of the mockingbird. 

Mockingbirds were especially common the last week in February 
in company with robins, bluebirds, and phainopeplas in the mesquite 
belt near Mellen, Arizona. The day (March 15) we floated down 
from our last station above the mouth of Bill Williams River to 
Parker, we heard the singing of mockingbirds from either side of the 


1914] Grinnell: Mammals and Birds of the Colorado Valley 207 


Colorado River at frequent intervals all along. At Riverside Moun- 
tain and above Blythe, on the California side, they were still com- 
mon. <A few were noted the first week in April on the California side 
opposite Cibola. Three individuals were seen on the Arizona side 
ten miles below Cibola, April 9, and the last for the season were noted 
April 19, eight miles east of Picacho. Nine specimens of the mocking- 
bird were secured (nos. 13797-13805), one on this latest day. 

There were no indications, either through dissection or judging 
from the behavior of the many birds observed, that the species nests 
in the region. It appears most probable that the species is only a 
winter visitant from the higher Upper Sonoran Zone to the northward 
in eastern California and southern Nevada. 


Toxostoma crissale Henry 
Crissal Thrasher 

A very characteristic element of the fauna of the Colorado bottom 
along the whole portion of its course explored by us. All evidence 
shows that it was a permanent resident, and closely restricted at all 
times to the outermost riparian association, namely the mesquite belt. 
Wherever this belt was strongly represented, the presence of crissal 
thrashers was to be recognized through their song or call note. But 
it was a difficult matter to shoot specimens, as the birds were ever 
alert, and kept close to or upon the ground beneath dense cover. 
Locally the thrashers were found foraging among catelaws and iron- 
woods up desert washes within a mile of the river bottom, and in some 
places they had temporarily forsaken the mesquites and invaded the 
willow tracts. The total absence of mesquite in any section of the 
river valley, however, was a sure indication of the absence of thrashers. 

The species was noted at the following localities: California side, 
five miles below Needles; Arizona side, above Mellen (more numerous 
than at any other point); both sides of the river in the vicinity of 
The Needles; Arizona side above Bill Williams River; California side 
at Riverside Mountain, above Blythe, opposite Cibola, twenty miles 
above Pieacho, and eight miles east of Picacho; Arizona side, five 
miles above Laguna; and California side in the vicinity of Pilot Knob. 
Mellen was the only station where definite indications of breeding 
were found. A female taken February 24 showed conclusively that 
incubation was in process; and another contained large ova. Still no 
young-of-the-year were encountered during the succeeding two and 


one-half months. 


208 University of California Publications in Zoology — | Vou. 12 


Twelve specimens of the crissal thrasher were preserved, nos. 
13806-13817. 

There is also a specimen in the Museum (no. 4226) taken by J. G. 
Cooper at Fort Mohave, January 2, 1861; and another (no. 6112) 
taken by W. W. Holder at ‘‘ Mineral City’? (Ehrenberg) March 20, 
1864. 


/ 
Heleodytes brunneicapillus couesi (Sharpe) 


Cactus Wren 

Met with in small numbers at most of the collecting stations. 
Where desert washes led down to the river bottom, cactus wrens 
occurred locally in the mesquite belt; but otherwise the species was 
restricted to the neighborhood of tracts of ironwood, catclaw, and 
cactus back from the river. In other words, it was a desert species, 
not properly a member of the riparian assemblage. 

This wren was evidently resident wherever found, as nests were 
always to be seen in the vicinity. Specific points of occurrence were: 
in the Sacramento Wash near Mellen; both sides of the river in the 
vicinity of The Needles; Arizona side above Bill Williams River; Cali- 
fornia side at Riverside Mountain; and same side above Blythe, oppo- 
site Cibola, twenty miles above Picacho, eight miles east of Picacho 
and four miles north of Potholes. At the latter place in a wash in 
the giant eactus belt, about two miles back from the river, a nest was 
found April 23 containing four eggs in which incubation was far 
advanced. The nest was of usual construction, and was situated five 
feet above the ground in a dense cholla cactus. Other nests were 
seen at different points, in ironwood, palo verde and mesquite. 

Nine specimens of the cactus wren were taken, nos. 13818-13826. 


Salpinctes obsoletus obsoletus (Say) 
Rock Wren 

Up to the middle of March common widely on the desert mesas 
as well as among hills. Noted at every station from the vicinity of 
Needles and Mellen south to Riverside Mountain. Thenceforth 
observed only in pairs in restricted localities affording appropriate 
nesting sites. On the California side opposite Cibola, April 3, a pair 
of rock wrens were found in a ravine about two miles from the river. 
Their nest was located in a hole in the roek wall of a gully, only about 
five feet from the bed. The floor of the opening, and also of some 


1914] Grinnell: Mammals and Birds of the Colorado Valley 209 


adjacent cavities, was paved with flat pieces of stone about an inch 
in diameter. The nest itself was out of reach. Several rock wrens 
were observed among the hills twenty miles north of Picacho. On 
April 16 a bird was observed to fly into a eranny with a bill full of 
insects, thus indicating young at this date. A bird taken on the 
hillside near Potholes, April 27, showed evidences that the breeding 
season was past. 

Five specimens were secured (nos. 13841-13845). 

There are also in the Museum two skins (nos. 4277, 4278) taken 
by J. G. Cooper at Fort Mohave, January 31 and February 10, 1861. 


Catherpes mexicanus conspersus Ridgeway 
Canon Wren 


Observed only where precipitous rock walls furnished the ideal 
environs for the species, and only sparingly then. At least three 
pairs were located on the river side of the pinnacles constituting The 
Needles, and one specimen obtained March 5. Another pair was 
observed on the California side opposite. The song was heard repeat- 
edly as we passed through the whirlpool canon just below The Needles. 
Again met with in deep, narrow ravines in some red conglomerate 
hills above Bill Williams River, where one example was taken March 15 
and another seen. From the canon walls on either side of the river 
in its course immediately below the mouth of Bill Willams River, the 
song of this wren came often to our ears as we drifted along. 

March 17 and 18 canon wrens were heard in ravines well up on 
Riverside Mountain. As none were seen after the latter date, although 
other favorable localities were visited, it is possible that the species is 
only a winter visitant to the region from the higher desert ranges in 
the Death Valley region. Still it is known to breed in similar localities 
in the same life-zone elsewhere. 

The two examples secured (nos. 13848, 13849) are doubtfully 
referred to conspersus. Lack of material prevents a satisfactory study 
of the ease. 


Thryomanes bewicki eremophilus Oberholser 


Desert Bewieck Wren 


Common as a winter visitant to the region. Observed chiefly in 
the sparse brush margining the washes leading down from the desert 
interior. The ecatelaw and larger creosote bushes appeared to afford 


210 University of California Publications in Zoology [ Vou. 12 


both productive foraging grounds and safe retreats. It was rarely 
that this wren was seen near the river, and then only as far as the 
salt-bush belt. The range of the western house wren in the willow 
association appeared to be not at all impinged upon by that of the 
desert Bewick wren. This again shows the local dissociation of birds 
of the same or nearly the same habits, even in their winter habitats. 
It is to be inferred that there are inherent preferences of the two 
species for cover of the two different sorts. 

The desert Bewick wren was observed on the California side five 
miles below Needles, opposite The Needles, and in the lower Cheme- 
huevis Valley; on the Arizona side in the vicinity of Mellen, at the 
foot of The Needles, and above Bill Williams River. At least one 
specimen was secured at each of the above points. The last for the 
season were observed March 21 at Riverside Mountain. The series of 
nine specimens (nos. 13827-13835) are uniform in their exhibition 
of the characters assigned to this race by its original describer (Ober- 
holser, 1898, p. 427). The great length of tail alone serves to dis- 
tinguish Thryomanes bewicki eremophilus from any of the other seven 
forms of bewicki occurring within the state of California (Grinnell, 
1910, p. 309). The Colorado Valley birds are with a high degree 
of probability visitants from a summer habitat lying on the higher 
desert mountains in the vicinity of Death Valley, California and 
Nevada, where this form has been recorded as breeding (Oberholser, 
1898, p. 429). 

There are in the Museum three skins (nos. 4280-4282) of this wren 
taken by J. G. Cooper at Fort Mohave, January 1, March 6 and 21, 
1861. 


Troglodytes aédon parkmani Audubon 
Western House Wren 


Common as a winter visitant to the river bottom only, and even 
here confined almost exclusively to the willow association. Not one 
individual was seen anywhere on the desert proper. The species was 
noted in the willow thickets near Needles, February 15, and thence- 
forth at all stations, on both sides of the river, until the last of March. 
At Ehrenberg but one was noted (and obtained) March 29. The last 
for the season was seen April 9, a single individual, in a pile of drift 
on the river bank on the Arizona side ten miles below Cibola. 

Five specimens preserved (nos 13836-13840). 


1914] Grinnell: Mammals and Birds of the Colorado Valley 211 


There is also a skin in the Museum (no. 4287) taken by J. G. 
Cooper at Fort Mohave, January 22, 1861. 

As there are no known characters by which to recognize house 
wrens breeding in the Great Basin region from those on the Pacific 
coastal slope, there is no way of deciding where those individuals win- 
tering in the Colorado Valley come from. But it is most likely that 
they hail from the interior somewhere, as do practically all of those 
birds where this point can be definitely determined. 


Telmatodytes palustris plesius (Oberholser) 
Western Marsh Wren 


Observed only in dense but low vegetation such as tules, along 
lagoons in the river bottom. A single example was seen and secured 
(no. 13846) on the California side at the lower end of Chemehuevis 
Valley, March 11. Near Riverside Mountain March 17 several were 
seen, and one secured (no. 13847). On the Arizona side a mile or 
so below Ehrenberg, March 25 to 29, there were many around a series 
of ponds. Nothing was seen of the species after the latter date. Since 
no old nests or other indications of breeding were detected, it is not 
to be presumed that marsh wrens are more than winter visitants to 
the Colorado Valley. 


Auriparus flaviceps flaviceps (Sundevall) 
Verdin 

The most numerous and widespread resident species of bird in the 
whole region. The only essential condition for the presence of this 
species appeared to be stiff-twigged thorny bushes or trees of some 
sort. This requisite was met with in a variety of situations, as in 
the screwbeans of the first bottom, mesquites of the second bottom, 
and eatelaw, ironwood, palo verde and daleas of the desert washes. 
The birds appeared to have already paired off by the latter half of 
February; each pair had a particular beat or forage area, focussing 
at one or more nests. Nests were occupied, at least as roosting places, 
throughout the season, and as nests were not constructed in other 
than the thorny bushes above named, the local range of the species 
was predetermined. While the birds were often seen in willows, 
arrowweed, and even low shrubs of Atriplex and sandburr, these were 


212 University of California Publications in Zoology | Vou. 12 


always within a limited radius of nests. As far as observation went, 
these birds do not need to visit water; some were met with as much 
as three miles away from the river up desert washes. 

Nests containing eggs or young were found as follows: April 5, 
California side opposite Cibola, eggs four, incubation advanced; April 
7, Arizona side ten miles below Cibola, eggs four, incubation far 
advaneed; same date and place, three eggs incubated and one newly 
hatched young; same date and place, eggs five, incubation nearly 
complete; April 11, California side, twenty miles above Picacho, eggs 
four, incubation advaneed; April 12, same place, three small young 
and one egg about to hatch; April 22, on the Arizona side five miles 
above Laguna, a brood of two-thirds grown young was encountered. 
The breeding period thus would appear to be of remarkable uniformity 
among all individuals of the species. 

The above six nests varied from 38 to 96 inches above the ground, 
averaging 69 inches. These, of the usual firm-walled, globular type 
and constructed of stiff, thorny twigs, with the laterally placed opening 
seareely larger than the diameter of the bird, would thus appear 
admirably adapted for the exclusion of the parasitic cowbird, of much 
ereater size, as well as a defence against depredators of various sorts. 
I can see no other reason for so specialized a structure. Yet the 
plumbeous gnateatecher, occupying almost an identical range, and with 
open nests, lays the same number of eggs, as though it were no more 
subject to fatalities than the verdin. This is true also of the Sonora 
yellow warbler (see Brown, 1903, p. 47). 

A series of twenty-four verdins was taken, nos. 13922-13945. Two 
of our specimens, taken at Needles February 15 and 16, are much 
discolored with coal soot. 

Besides these, there are in the Museum collection five specimens 
(nos. 4238-4242) taken in 1861 by J. G. Cooper at Fort Mohave. The 
dates of capture are: February 19, March 9, 15 and 30, and April 5. 
These old skins are perceptibly paler than the freshly obtained speci- 
mens, most probably due to their having been exposed to strong light 
in a show ease at some time or other. 


Regulus calendula cineraceus Grinnell 
Ashy Kinglet 
Evidently a characteristic winter visitant, as it was one of the com- 


monest birds of the willow bottom in the vicinity of Needles at the 
time of our arrival in the region, February 15. It was thenceforth 


1914] Grinnell: Mammals and Birds of the Colorado Valley 213 


noted regularly down along the river until the first week in April, 
when it became scarce; the last one noted was secured April 18, on 
the California side eight miles east of Picacho, this being the only 
individual seen at this point. The other localities of capture, lying 
between the two above-named, are as follows: on the California side: 
opposite The Needles, Chemehuevis Valley, Riverside Mountain, Blythe, 
opposite Cibola, twenty miles north of Picacho; on the Arizona side: 
Mellen, foot of The Needles, Ehrenberg, ten miles south of Cibola. 
Tn all these places the kinglets were seldom seen outside of the willow 
association. 

The series of twenty-four specimens obtained, nos. 13946-13969, 
is uniform in its exhibition of the characters assigned to Regulus calen- 
dula cineraceus (see Grinnell, 1904, p. 25). As compared with 
examples from eastern North America, they are distinctly more ashy- 
hued anteriorly both above and below. Especially is this peculiarity 
marked over the top and sides of the head. There is a slightly.greater 
general size in the case of the southwestern race. It is therefore prob- 
able that the Colorado River birds are visitants from the mountains 
of the southwest, whence kinglets of the same characters have been 
secured in summer, rather than from the forests of the far north. 

There is also in the Museum a skin (no. 4272) taken by J. G. 
Cooper at Fort Mohave, January 18, 1861. 


Polioptila caerulea obscura Ridgway 
Western Gnateatcher 


First seen at Needles, February 16; thenceforth noted regularly 
at all stations along down the river until April 6, when the last were 
noted ten miles below Cibola. As the specimens taken the first week 
in April, when Polioptila plumbea was nesting, showed no signs of 
immediate breeding, it seems probable that the western gnatcatcher is 
only a winter sojourner in the Colorado Valley. It occurred chiefly 
in the bushes lining the desert washes leading back from the river; 
a few individuals were met with in the mesquite and willow associa- 
tions. 

The series of thirteen skins preserved (nos. 13874-13886) repre- 
sent the following localities : California side, five miles south of Needles ; 
both sides of the river at The Needles; California side at Riverside 
Mountain and opposite Cibola; and Arizona side, ten miles below 
Cibola. 


214 University of California Publications in Zoology [Vou. 12 


There is also in the Museum collection a skin (no. 4216) taken by 
J. G. Cooper at Fort Mohave, March 26, 1861. 

The spring molt (which involves only the body plumage) is well 
along in the bird (a male) of the latter date and in birds of April 4, 
5 and 6 appears to be nearly or quite complete. In the females of 
these dates there is, however, a want of clearness and continuity in 
the gray of the dorsum and pileum, and upon close examination this 
is found to be due to an interrupted and irregular replacement of 
feathers. This results in a mixture of new clear gray feathers and 
worn ashy ones. 

Of the April birds there are six females and one male. The latter 
(no. 13882), in comparison with series of Polioptila caerulea obscura 
from western California, has an extraordinary amount of black on 
the forehead, there being a frontal band of an approximate width of 
five millimeters, with a conspicuous extension posteriorly over each 
eye. Moreover, two of the females (nos. 13880, 13881) show a distinct 
supra-loral black line on each side, the two lines converging and 
meeting over the base of the culmen. This tendency to blackness on 
the crown may characterize a race wintering in the Colorado Valley 
and breeding in the Upper Sonoran zone of desert mountains to the 
northwestward. Material is not at hand to permit following up the 
matter. 


Polioptila plumbea (Baird) 
Plumbeous Gnateatcher 


A common resident along the whole line of exploration from Needles 
to the Mexican line. The series of thirty-five specimens secured (nos. 
13887-13921) represents the following localities: California side, at 
Needles and five miles below Needles; Arizona side, above Mellen and 
at the foot of The Needles; California side in Chemehuevis Valley, 
at Riverside Mountain and above Blythe; Arizona side, at Ehren- 
bere; California side, opposite Cibola, twenty miles above Picacho, 
eight miles east of Picacho, and near Pilot Knob. 

There are also in the Museum two skins (nos. 4219, 4220) taken by 
J. G. Cooper at Fort Mohave, February 19 and March 19, 1861. 

The desert wash association, with its catclaw, palo verde, ironwood 
and smaller woody and stiff-twigged plants, was the preferred habitat 
of this bird, though it occurred also in the Atriplex and mesquite belts 
along the river, even straying occasionally into the arrowweeds and 


1914] Grinnell: Mammals and Birds of the Colorado Valley 215 


willows. It did not, however, seem to visit water, and so its presence 
did not at all depend upon the proximity of the river. 

April 19, on the California side, eight miles east of Picacho, a nest 
was found in a smoke-bush (Dalea). It was four feet above the ground, 
of usual structure, and contained five small young. April 23, on the 
same side, four miles above Potholes, a brood of nearly grown young 
was flitting about at large. The breeding time in this region is thus 
indicated. 


Hylocichla ustulata ustulata (Nuttall) 
Russet-baecked Thrush 


As elsewhere in the west this bird proved to be a late arrival. The 
first individuals were noted May 4 and 5, five miles northeast of 
Yuma, one each day. On both sides of the river in the vicinity of 
Pilot Knob, May 9 to 15, the species was of frequent observation, 
chiefly at daybreak in the willows close along the river. One was 
seen, however, in a rocky ravine on Pilot Knob itself, towards late 
evening of the 14th. The species is most certainly only a transient 
through the region. The three specimens taken (nos. 13850-13852) 
are unequivocally referable to H. wu. ustulata. 


Hylocichla guttata guttata (Pallas) 
Alaska Hermit Thrush 


Hermit thrushes proved not to be characteristic winter visitants, 
as expected, and they were rare even as migrants. Unmistakable notes 
of one were heard at daybreak from a willow thicket in Chemehuevis 
Valley, March 9. Heard again similarly, ten miles below Ehrenberg, 
March 31. <A single individual secured (no. 13853) on the California 
side twenty miles north of Picacho, April 10. This bird, the last of 
the species noted, was in a dense willow growth close to the river. 
The specimen is identical, in size and coloration, with examples of this 
race from the Prince William Sound region of Alaska. 


Hylocichla guttata nanus (Audubon) 
Dwarf Hermit Thrush 
There is in the Museum a skin (no. 6432) taken by J. G. Cooper at 
Fort Mohave, January 25, 1861. Although obviously faded on its 


ventral surface, by size and dorsal color this specimen is distinctly 
referable to nanus. 


216 University of California Publications in Zoology [ Vou. 12 


Planesticus migratorius propinquus (Ridgway) 
Western Robin 


Fairly common as a winter visitant to the bottom lands. Often 
met with in the dense willow woods, on the ground foraging for insects 
in the layers of dead leaves. Also one of the regular feeders on 
mistletoe berries. At Needles, California, February 17, several indiv- 
iduals were seen hopping about on the grass in the railway parking. 
On the Arizona side above Mellen a number were found among the 
mistletoes of the mesquites, February 23 to 31. A few were encoun- 
tered on each side of the river at The Needles, March 1 to 7. One was 
seen March 14, on the Arizona side above Bill Willams River; a 
few on the California side at Riverside Mountain, March 17 to 21; 
several on the same side opposite Cibola, April 1 to 5; and one, the 
last noted, on the Arizona side, ten miles below Cibola, April 7. 

Seven specimens secured, nos. 13854-13860. 

There is also in the Museum a skin (no. 4224) taken by J. G. 
Cooper at Fort Mohave, March 7, 1861. 


Sialia mexicana occidentalis Townsend 
Western Bluebird 


Common, as a winter visitant, feeding in flocks on mistletoe berries 
wherever these were obtainable, usually in the mesquite belt. Numerous 
at Needles, California, and immediately below, February 14 to 22; at 
and above Mellen, Arizona, February 23 and 31; and on both sides 
of the river at The Needles, March 1 to 7. The species was last seen, 
a very few, March 13, on the Arizona side above Bill Williams River. 

There is a skin (no. 4212) in the Museum taken by J. G. Cooper 
at Fort Mohave, February 22, 1861. 

The thirteen specimens secured by us (nos. 13861-13873) are not 
exactly typical of S. m. occidentalis, as occurring in the northwest 
coast district of the United States. Yet I do not find grounds for 
allocating them with S. m. bairdi, of the Rocky Mountain area, nor 
with S. m. anabelae of northern Lower California. They are very 
much like birds from the Sierra Nevada, and I have little doubt are 
winter visitants to the Colorado Valley from there. Sierra Nevada 
birds are customarily referred to S. m. occidentalis, and at present 
I cannot do anything better than employ this name for the specimens 
in hand. Coloration furnishes characters difficult to weigh properly, 


1914] Grinnell: Mammals and Birds of the Colorado Valley 


because of much seasonal variation in tone of browns and blues through 
fading and abrasion, and because of wide individual variation in extent 
of chestnut areas. General size, too, is an inconstant feature (see Grin- 


nell and Swarth, 1913, p. 318). 


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CHECK-LIST OF MAMMALS 


Ovis canadensis nelsoni Merriam 
Odocoileus hemionus eremicus (Mearns) 


Ammospermophilus harrisi harrisi (Audubon and Bachman) 


Ammospermophilus leucurus leueurus (Merriam) 
Citellus tereticaudus tereticaudus (Baird) 
Castor canadensis frondator Mearns 
Peromyscus maniculatus sonoriensis (Le Conte) 
Peromyscus crinitus stephensi Mearns 
Peromyscus eremicus eremicus (Baird) 
Sigmodon hispidus eremicus Mearns 
Reithrodontomys megalotis deserti J. A. Allen 
Neotoma albigula venusta True 

Neotoma intermedia desertorum Merriam 
Ondatra zibethica pallida (Mearns) 
Thomomys chrysonotus Grinnell 

Thomomys albatus Grinnell 

Dipodomys deserti deserti Stephens 
Dipodomys merriami merriami Mearns 
Perognathus bombyeinus Osgood 

Perognathus formosus Merriam 

Perognathus penicillatus penicillatus Woodhouse 
Perognathus intermedius Merriam 
Perognathus spinatus spinatus Merriam 
Lepus ecalifornicus deserticola Mearns 
Sylvilagus auduboni arizonae (J. A. Allen) 
Felis oregonensis browni Merriam 

Lynx eremicus eremicus Mearns 

Canis ochropus estor Merriam 

Vulpes macrotis arsipus Ellot 

Urocyon cinereoargenteus scotti Mearns 
Mephitis estor Merriam 

Spilogale arizonae arizonae Mearns 

Taxidea taxus berlandieri Baird 

Procyon pallidus Merriam 

Corynorhinus macrotis pallescens Miller 
Antrozous pallidus pallidus (Le Conte) 

Myotis oceultus Hollister 

Myotis californicus pallidus Stephens 

Myotis velifer (J. A. Allen) 

Pipistrellus hesperus hesperus (H. Allen) 
Eptesicus fuscus (Beauvois) 

Nyctinomus mexicanus Saussure 

Macrotus ealifornicus Baird 


218 University of California Publications in Zoology — [Vou. 12 


GENERAL ACCOUNTS OF THE MAMMALS: LOCAL 
DISTRIBUTION, VARIATION, HABITS 


Ovis canadensis nelsoni Merriam 
Desert Bighorn 


Mountain sheep still occur in parts of the country on both sides 
of the river. Among The Needles, on the Arizona side, March 5, fresh 
feces and footprints showed plainly where a band of at least five had 
been grazing. The point of this occurrence was not more than half 
a mile from the river and 300 feet in elevation above it. 

We were assured of the existence of a flock of fully thirty-five 
sheep on the Arizona side east of Cibola. These sheep were reported 
to visit regularly certain springs in the Chocolate Mountains. 

On the California side, as we were informed, sheep used to range 
over Riverside Mountain, but they have not been seen there of late 
years, probably because of the mining activity in that vicinity. Pros- 
pectors told us that they had always taken every opportunity to 
obtain ‘‘fresh meat’’ wherever they might be in the desert hills; 
and as the camps of the prospectors are usually made at the water 
holes which the sheep depend upon in the long, dry seasons, it is not 
difficult to account for the disappearance of sheep in a mineralized 
region ! 

West of the river, 16 to 22 miles above Picacho, we found sheep 
to be quite common. At least five had been killed within the previous 
year by residents along the river. Two of these were shot from a 
rancher’s house and within a few hundred yards of the river, which 
here flows between steep hills, with relatively little bottomland inter- 
vening. This, by the way, is in the near vicinity of Lighthouse Rock, 
and in the sketch of the river including this landmark, given in Ives’ 
Report upon the Colorado River of the West (1861, page 52, figure 7), 
two unmistakable bighorns are portrayed at the water’s edge. 

We learned nothing to indicate that the sheep in this neighborhood 
ever visit the river for water at the present time. There are springs 
back in the hills to which the sheep regularly resort. We saw five 
sheep, and secured two, a ewe (no. 10588) and a male lamb (no. 
10589). These were found about three miles west of the river among 


rough hills. 


1914] Grinnell: Mammals and Birds of the Colorado Valley 219 


There is a possibility that the bighorns on the Arizona side of 
the Colorado differ slightly from those on the California side. The 
latter unquestionably belong to the race nelsoni. Mearns (1907, p. 240) 
describes a race (Ovis canadensis gaillardi) from the Gila Mountains, 
Yuma County, Arizona. This locality is less than seventy-five miles 
from the locality near Cibola where sheep are known to occur, and if 
gaillardi is a really distinct form the sheep of Cibola are perhaps 
referable to it rather than to nelsoni. 


Odocoileus hemionus eremicus (Mearns) 
Burro Deer 


No trace of deer was found by us anywhere, nor had anyone we 
talked with seen deer along the Colorado within four years. We were 
told of their occurrence in numbers many years before, when they 
were to be found both in the river bottom and back through certain 
desert ranges, where there are springs which the deer could visit 
regularly for water. 

In August, 1902, Mr. Frank Stephens saw two deer on the Cali- 
fornia side of the river west of Cibola. The animals were jumped 
from a wash among ironwood trees and made off across a wash. Sign 
was fairly plentiful, and deer were said to be common at that time 
on both sides of the river. 

Members of our party in 1910 looked over the same ground closely 
without finding sign. The rapid settlement of the river bottom around 
Palo Verde probably accounts for the disappearance of deer throughout 
that section. At the Draper ranch twenty miles above Picacho a single 
old buck had been seen ‘‘about’’ five years previously. 


Ammospermophilus harrisi harrisi (Audubon and Bachman) 
Harris Ground Squirrel 


Twenty-six specimens secured, from localities all on the Arizona 
side of the river, as follows: Mellen, 18; foot of The Needles, 1; above 
Bill Williams River, 2; Ehrenberg, 3; 10 miles below Cibola, 2. None 
was seen at any locality where not also taken, so that the species 
appears not to occur near the river along its course below Cibola. Our 
work shows, further, an extreme associational restriction, namely, to 
rupestrine conditions as furnished in the rough hills and on those 
parts of the desert mesa where the wind keeps the weathered detritus 


220 University of California Publications in Zoology [Vou 12 


removed, leaving a surface of firmly packed gravel or pebbles. The 
burrows of the Harris ground squirrel are most often to be found 
opening out beneath a creosote bush (see pl. 12, fig. 18), which plant 
occurs throughout the range of the squirrel. Up to the last date of 
capture, April 9, no young had been observed, although on March 5, 
at Mellen, a female taken contained six embryos. 

In no instance was this rodent found to have strayed on to any 
part of the river bottom, even on to the edge of the sandy second 
bottom. But where the river passed among the steep-sided hills, as at 


LIST AND MEASUREMENTS IN MILLIMETERS OF AMMOSPERMOPHILUS 


HARRISI HARRISI FROM THE ARIZONA SIDE OF THE 
COLORADO RIVER 


3 | 

2 a 3 

s rh as 22 oa ne 

$2 43% gi Ess 383 & 
Mus. No. Sex Locality Ax i Bim Eo! is Cie) (Sis 

105381 9 £Near Mellen Feb. 25 225 70 389 5 388 22.0 | 
105382 og Near Mellen Feb. 26 246 85 40 5 404 23.7 

10533 og Near Mellen Feb. 27 226 84 40 8 39.38 22.57 | 

105384. 2 Near Mellen Feb. 27 217 #76 39 8 385 22:00 a 

10585 og Near Mellen Feb. 27 232 70 40 . 39.4 2216 | 

10586 @Q Near Mellen Feb. 27 215 70 38 . 38:4 227 @ 
10537 g Near Mellen Feb. 27 222 66 37 . 38.5 22:2 
105388 9 Near Mellen Feb. 27 225 70 40 39.7 22.5 
105389 og Near Mellen Feb. 27 224 68 39 40.8 238.5 
10540 g Near Mellen Feb. 27 243 80 40 40.1 22.5 
10541 Q Near Mellen Feb. 28 205* 50* 39 38.7 22.5 
10542 Q Near Mellen Feb. 28 230 70 38 39.1 23.0 
10548 g Near Mellen Feb. 28 220) 168) ) 3360) ees 22.1 
10544 ¢ Near Mellen Feb. 28 220 67 38 38.5 238.2 
10545 og Near Mellen Mar. 1 230 70 38 40.1 28.4 
10546 og Near Mellen Feb. 28 220 77 38 7 383 22.4 
10547 Q Near Mellen Feb. 28 219 78 39 7 38:6 “2216 
10548 2 Foot of The Needles Mar. 5 230 86 38 8 385 22.4 
10549 @Q Above Bill Williams River Mar. 13 23.0) O03 8) se 22.5 
10550 @Q Above Bill Williams River Mar. 15 235) 73. (38), = ee 22.8 
10551 Q Ehrenberg Mar. 25 220 70 39 39.2 23.0 
10552 gf = Ehrenberg Mar. 26 223 75 40 39.5 22.8 
10553 @Q Ehrenberg Mar. 29 240 75 41 39.8 22.8 
10554 @Q 10 miles below Cibola Apr. 8 225 72 38 39.8 23.4 
10555 ¢ 10 miles below Cibola Apr. 9 225 72 38 40.2 22.5 
Average of the 24 adults 227 «= 73.4 38.7 6.9 39.38 22.7 


* Not averaged, tail defective 


1914] Grinnell: Mammals and Birds of the Colorado Valley 221 


The Needles, harrisi was observed as close as within one hundred yards 
of the water’s edge. The animals did not, however, even here, show 
the least disposition to visit the water for any purpose whatever; so 
that the chances of being transported across to the opposite shore are 
practically nil. That in this part of the range of harrisi the Colorado 
River is the limiting barrier is a fact beyond question. There is a per- 
sistent statement in literature (for example, Elliot, 1905, p. 97) to the 
effect that harrisi oceurs in California; but I am not cognizant of any 
basis for this assertion and I doubt its correctness. 

Two races of Ammospermophilus harrisi have been distinguished 
within the state of Arizona. Mearns (1896, p. 444, and 1907, pp. 303— 
309) has characterized a form A. h. saxicola from extreme south- 
western Arizona, thus restricting harrisi proper to central Arizona. I 
fail to find grounds, either in tone of coloration or relative length of 
tail, to warrant separate recognition of the Colorado River series 
from the animal of the vicinity of Tucson. Even Mearns’ own tables 
of measurements do not bear out the size characters of his saxicola, 
which name might be expected to be usable for our specimens. The 
measurements of the Colorado Valley series are given herewith, as is 
also a table showing comparison with the near-related form leucurus, 
of the opposite side of the river (see pp. 220, 223). 


Ammospermophilus leucurus leucurus (Merriam) 
Antelope Ground Squirrel 

Seventeen specimens obtained, all on the California side: 5 miles 
below Needles, 1; opposite The Needles, 5; Chemehueyis Valley, 2; 
Riverside Mountain, 1; opposite Cibola, 2; twenty miles above Picacho, 
4; eight miles east of Picacho, 2. This rodent was not notably numer- 
ous anywhere, and was closely restricted to rocky or gravelly ground, 
chiefly among hills. It occupied the same ecologic niche on the Cali- 
fornia side that harrisi did on the Arizona side; and where the hills 
closely abutted upon the river, as at The Needles, the two species 
existed in places not more than three hundred yards apart, but with 
the river between. In no ease was there any evidence of crossing; 
all individuals captured or seen were unquestionably one form or the 
other, and always appropriately segregated as regards the river. 

No young of lewcwrus were taken; but a female taken five miles 
south of Needles, February 20, contained eight embryos, and another 


222 University of California Publications in Zoology 


[Vot. 12 


taken at Chemehuevis Valley, March 11, contained five embryos. It 
is notable that the weather on the first date was very cold for the 
region. It seems to be a rule among the Sciuridae of the Colorado 
desert area that the young-of-the-year are well grown long before the 


period of intense summer heat. 


Ammospermophilus l. leucurus and A. h. harrisi are perfectly 


distinct but closely related species. 


The following comparative table 


serves to show the differences between them, which differences must 
not be overemphasized as contrasted with the close general resemb- 


lanees. 


AMMOSPERMOPHILUS L, LEUCURUS 


General size slightly less; body 
149.4 mm. 

Tail relatively shorter, 44 per cent of 
body. 

Audital bullae slightly smaller. 


External ear slightly larger, 7.1 mm 

General coloration paler above, less 
brown more gray. 

White side-stripes more conspicuous. 


Tail beneath pure white. 

Tail carried appressed over the rump, 
displaying under surface conspicu- 
ously as if there were a white 
rump patch. 

Tail often rapidly vibrated from side 
to side, causing the white to 
flicker 


AMMOSPERMOPHILUS H. HARRISI 


General size greater; body 153.6 mm. 


Tail relatively long, 48 per cent of 
body. 

Audital bullae slightly higher, more 
inflated. 

External ear slightly smaller, 6.9 mm. 

General coloration darker, distinetly 
brownish dorsally. 

White side-stripes narrower and not 
so long. 

Tail beneath grizzled gray. 

Tail carried vertically or inclined 
backward or forward, but snow- 
ing no conspicuous white. 


Tail often waved or jerked in fore- 
and-aft are. 


In the matter of audital bullae, harrisi parallels Thomomys chrys- 
onotus, in the inereased size of bullae on the Arizona side of the river. 
A curious anomaly appears in the reverse proportion in the size of 
the external ear in the two squirrels, lewcwrws of the California side 
having larger ears than harrisi. There are similar reverse ratios 
between ears and bullae in certain southern California Perognathus. 

The most striking difference between Ammospermophilus harrisi 
and A. leucurus is, as noted in the table, in the color of the under 
surface of the tail and in the manner of carrying this appendage. 
It may be inferred that the musculature connected with the tail is 
quite differently developed in the two species, together, of course, 
with correlated nervous and circulatory differences. 


1914] = Grinnell: Mammals and Birds of the Colorado Valley 223 


The range of harrisi is practically restricted to Arizona and parts 
of Sonora. The range of leucwrus and its geographic races almost 
completely surrounds that of harrisi. But, as far as I know, the 
ranges of the two nowhere overlap, nor has there been found any 
ease of hybridization between the two. It is certain that only the 
existence of the Colorado River keeps the ranges of the two separate 
in this segment of their respective peripheries. What would have 
happened if the river had not served as an absolute barrier is, of 
course, wholly conjectural. But there is no doubt in my mind that in 
such event both the ranges and the characters of the two forms 
would be far different from what they now are. 


LIST AND MEASUREMENTS IN MILLIMETERS OF 4MMOSPERMOPHILUS 
LEUCURUS LEUCURUS, FROM THE CALIFORNIA SIDE OF 
THE COLORADO RIVER 


Mus. no. Sex Locality 4 PSI rs & Zé 5S é 5 S 
10556 2 5 miles below Needles Feb. 20 177 40* «385.50 7) 336.8 = 2.0.8 
10557 § Opposite The Needles Mar. 2 220 60 35 37.3 22.7 
10558 2 Opposite The Needles Mar. 2 222 65 38 39.1 21.3 
10559 + g Opposite The Needles Mar. 2 218 65 38 38.2 22.6 
10560 @G Opposite The Needles Mar. 3 220 60 38 37.8 22.0 
10561 @ Opposite The Needles Mar, 7 215 60 40 = of.d) 214 
10562 ¢ Chemehuevis Valley Mar. 11 221 76 38 7 385 22.0 
10563 @Q Riverside Mountain Mar. 20 225 67 39 se fess 23.2 
10564 2 Opposite Cibola Apr. 4 235 75 38 Sy ee oe 
10565 2 Opposite Cibola Apr. 5 190 43" 40 9 39:0! 22.3 
10566 2 20 miles above Picacho Apr. 12 220 76 37 (ie aN.9) 82222 
10567 2 20 miles above Picacho Apr. 13 220 67 37 J) BOOT 
10568 g 20 miles above Picacho Apr. 13 215 60 38 - 40.0 23.1 
10569 2 20 miles above Picacho Apr. 15 194 61 34 6 364 21.1 
10570 @ Chemehuevis Valley Mar. 11 200 2 37 7 37.4 22.0 
10571 @ 8 miles east of Picacho Apr. 20 Pale “GB a eee 22.2 
10572 = g 8 miles east of Picacho Apr. 21 220 70 39 38.0 22.2 

Average of the 17 adults 215 65.6 37.5 7.1 38.0 22.1 


* Not averaged; tail defective. 


224 University of California Publications in Zoology — [Vou. 12 


Citellus tereticaudus tereticaudus (Baird) 
Round-tailed Ground Squirrel 


Fairly common at intervals from Needles to the Mexican line. Six- 
teen specimens were collected (nos. 10637-10651, 10722) on both sides 
of the river, as follows: California side: five miles below Needles 1, 
opposite Cibola 2, near Pilot Knob 4; Arizona side: at Mellen 5, five 
miles northeast of Laguna 3. The species was seen at no other places, 
and the inference is to be drawn that it is of quite interrupted distri- 
bution. This is explained when its associational position is taken 
account of. 

Citellus tereticaudus is a sand-dwelling mammal, resembling in 
its preferences Dipodomys deserti, Peromyscus eremicus and Perog- 
nathus penicillatus. The squirrel, however, is even more restricted 
than either of these, for a rather large extent of arenaceous territory 
seems to be necessary to support a representation. Where the second 
bottom was broad, with accumulations of wind-drifted sand about the 
seattering Atriplex and creosote bushes, tereticawdus was best repre- 
sented. Such a place was afforded on the Arizona side above Mellen. 
On the desert mesa similar conditions were rare, but where they did 
oceur to any extent, as near Pilot Knob, the round-tailed ground 
squirrels were usually in evidence. The burrows open up among the 
stems of partly buried bushes, the animals foraging out over the bare 
intervals where seeds from annual plants are found sifted into the 
surface sand. 

No young were seen by our party; but two females taken opposite 
Cibola, April 3 and 4, contained six and four embryos respectively. 
There are in the Museum two half-grown young (no. 7767, 7768) 
taken by Charles L. Camp at Needles, July 15 and 19, 1909. 

The series of Citellus tereticaudus, when segregated into two groups 
according to the side of the river from which the individuals were 
obtained, shows appreciable diversity in external features. The speci- 
mens from the Arizona side are distinetly darker, more brownish than 
those from the opposite (California) side, even where obtained within 
but a very few miles of one another. This diversity in color is most 
marked at Needles, less so lower down the river. It would appear 
that the case is quite comparable to that in Perognathus penicillatus, 
where the darkest individuals hail from the east side of the river north 
of The Needles. 


1914] Grinnell: Mammals and Birds of the Colorado Valley 225 


In measurements of the two groups, the Arizona-side examples of 
tereticaudus are shorter-tailed, as shown in the following comparison. 
Average of eight adults from the Arizona side as compared with aver- 
age of seven adults from the California side (in parenthesis) is: total 
length, 234.1 (240.6) ; tail vertebrae, 76 (88.3) ; hind foot, 35.1 (35.5) ; 
ratio, tail to body, 48 per cent (58 per cent). These measurements 
of the California side examples are close to those of many others of 
the Colorado Desert west of the river. The shortness of tail is fairly 
well borne out in the measurements of Citellus tereticaudus from east 
of the river along the Mexican line as given by Mearns (1907, p. 338). 
The present series, however, is inadequate to settle the point satis- 
factorily. 

It will be observed that in relative darkness of the Arizona side 
examples, we have here a parallel to the case in Ammospermophilus, 
while in ratio of tail to body the reverse relationships are presented. 
The degree of difference exhibited in Citellus, even if actual, is very 
slight as compared with the differences between Ammospermophilus 
h. harrisi and A. l. lewcurus. 


Castor canadensis frondator Mearns 
Sonora Beaver 


Signs of beavers were seen at many places along the river through 
the big valleys, from near Riverside Mountain to Pilot Knob. We 
heard of their occurrence also above Needles. At the present time, 
however, beavers are scarce along the Colorado River, this being said 
to be due to the unrelenting pursuit of them by professional trappers. 
Various evidences of the late presence of trappers were seen by us 
below Ehrenberg. 

Very little of the beaver sign seen during our cruise of the river was 
fresh. The first noted was on the California side near the base of 
Riverside Mountain, Mareh 16. Many young cottonwoods had been 
felled at the edge or within twelve feet of the main river channel, 
where its bank sloped steeply into deep water. The trees had been 
eut off about one foot above the ground and all felled down-stream, 
or, rather, diagonally, down-stream and towards the river. The largest 
cottonwood was fourteen inches in diameter where cut, the next two 
seven inches in diameter. Other cuttings were noted down to willow 
shoots one-half inch in diameter. 


226 University of California Publications in Zoology [ Vou. 12 


s) 


A stick ‘‘house’’ was located at the edge of the river a short dis- 
tance below these cuttings. It resembled a mass of drift caught among 
the snags resulting from the undercutting of the timbered bank. This 
house was three and one-half feet in height, by fourteen and sixteen 
feet in two diameters on the ground. It consisted of small branches 
and broken saplings from the driftwood of the river and some brush 
dragged in from the land side. A little earth and dead leaves had 
been worked into the interstices. Less than a dozen beaver-cut sticks 
were in evidence in the whole structure. There were no fresh signs 
was found on 


2? 


in or around this house. But a recently used ‘‘slide 
the bank a few yards off. 

Opposite Riverside Mountain, and below Ehrenberg in at least two 
places, beaver cuttings were found on the Arizona side. And on the 
California side from near Ehrenberg to below Palo Verde considerable 
work was noted. In one place bark had been stripped within a day 
or two from a cottonwood sapling lying in the water, where it had 
recently fallen as a result of undermining by the river. 

It was thought that the continual undercutting of its timbered 
banks by the river itself, thus precipitating many green trees into the 
water, provided an immediate food-supply for the beavers, so that even 
if present in numbers they would gain an easy living without them- _ 
selves having to fell trees. It is possible, too, that the habitually 
unsettled behavior of the river accounts for the loss of the dam-building 
propensity, or at least for its futility on the part of the beavers of 
the Colorado Valley. We saw no signs anywhere of dams such as are 
reported by Mearns (1907, pp. 354-358) as occurring on the Verde 
River, Arizona, a tributary of the Gila, which in turn flows into the 
Colorado. 

A used ‘‘house’’ located at the margin of a slough near Palo Verde 
was cut into by Stephens and Jones on April 3. It proved to be three 
feet in height and twelve feet across at the base. It consisted of 
branches and small saplings cut by the beavers when in full leaf, and 
laid compactly. The mass was eight inches thick over the nest cavity. 
The interior space was two and one-half feet high by four to five feet 
across. At one side the bank had been dug away to make the floor 
comparatively level, as the house was built on a sloping bank near 
its top. There was an under-water entrance at one side, and an open- 
ing through the wall above ground. A fresh willow sapling had been 
hauled through this opening butt first. The beds were merely hollows 
in the earth floor. 


+ 


1914] Grinnell: Mammals and Birds of the Colorado Valley 227 


On both sides of the river in the vicinity of Pilot Knob there was 
considerable beaver sign, consisting of well-beaten trails up the banks, 
foot and tail prints, and cut willow saplings. At all points where 
prospects seemed at all favorable, traps were set, but our nearest 
approach to success was the securing of a front foot (no. 10731) in 
a trap on the Arizona shore opposite Pilot Knob. The shore sloped 
so gently at this place that the trap could not be set in water deep 
enough to insure drowning of the victim and yet occupy a position 
anywhere near the foot of the used slide. The animal had evidently 
pulled loose. 

Further troubles in our efforts to trap beavers resulted from the 
continual rising or falling of the river and from the heavy deposit 
of silt. The latter so rapidly filled in the steel traps (Newhouse nos. 
3 and 4), even when set upon broad strips of bark and staked out two 
feet or so from the actual bank, that beaver or any other animals could 
walk upon them without setting them off. 

At the last trapping place, the lowering of the water repeatedly 
exposed both the traps and stakes, but tracks showed that during the 
night a beaver had gone past, nevertheless, indicating no particular 
shyness or suspicion. 

From the reports given us by different people along the river, it is 
probable that beavers were abundant in suitable parts of its course up 
to a few years ago. As many as 250 skins are said to have been taken 
in a single year by one trapper. With observance of the protective 
laws now in foree in both Arizona and California there is a chance 
for beavers to become sufficiently plentiful again to warrant a brief 
open season each year when the fur is prime. I see no reason why 
the Colorado River should not produce a regular output of skins, 
provided the number taken annually be adjusted to the rate of increase. 


Peromyscus maniculatus sonoriensis (Le Conte) 
Sonora White-footed Mouse 


An abundant inhabitant of the bottomlands everywhere, this in 
spite of the annual overflow which might be expected to drown out 
a mammal of this non-aquatie genus. 

The 65 specimens secured (nos. 10100-10164) represent the follow- 
ing localities: California side: five miles below Needles, opposite The 
Needles, Chemehuevis Valley, at Riverside Mountain, above Ehren- 
berg, near Palo Verde, eight miles below Picacho, four miles south of 


228 University of California Publications in Zoology (Vou. 12 


Potholes, five miles northeast of Yuma, and near Pilot Knob; Arizona 
side: Mellen, Ehrenberg, twenty-five miles below Ehrenberg, five miles 
northeast of Laguna, near Yuma. 

The species thus proved to be regularly distributed all along the 
Colorado Valley. Yet we failed to trap it anywhere on the nearby 
desert outside of the mesquite belt. In other words, this Peromyscus 
has appropriated the river bottom, which, in turn, is tabooed by the 
two desert species of the region, eremicus and stephensi. It is to be 
inferred that sonoricnsis used the Colorado Valley as a highway of 
immigration through the region, and found it suitable for permanent 
occupancy. The maniculatus division of the genus Peromyscus is 
notorious for its success as an invader. Single subspecies representa- 
tive of this group range through the habitats of dozens of other species 
of small rodents, from the hottest, Lower Sonoran, Colorado desert to 
the Boreal zone on the San Bernardino Mountains. It is thus a 
remarkably hardy animal; yet, as far as known, it nowhere thrives 
to the total exclusion of other small rodents, save in the case of slightly 
different subspecies on certain islands. 

The aggressiveness of Peromyscus m. sonoriensis was illustrated 
by our experience along the Colorado. It was the only wild mammal 
which found its way on to our boats; individuals were twice routed 
out of the cargo of supplies on the scow. This boat was most of the 
time moored alongside the shore, either touching the bank at one corner, 


or with a gangplank out, so that any venturesome animal could easily . 


go aboard. 

At a point five miles northeast of Yuma, Sonora white-footed mice 
were trapped April 30 on a river bar covered with a very young growth 
of willow. This was at a time when the rising water had already cut 
the place off from connection with the main shore. Footprints of mice 
were plentiful on the dry, fine silt on the higher part of the island 
thus formed. In several other places this rodent was trapped on 
portions of the flood-bottom elevated in such a degree that they would 
have been first cut off from the shore as the water rose and then set 
adrift. In some of these cases safety could have been secured by 
swimming short distances through relatively quiet water, or by taking 
refuge on floating drift. It is highly probable that in the rush of 
flood waters a large mortality must annually occur. 

It is probable also that the paucity of snakes and other enemies 
in the flood-bottom gives this mouse relative immunity from those 
dangers which beset the small mammals out on the desert; so that 


1914] Grinnell: Mammals and Birds of the Colorado Valley 229 


sonoriensis can stand the fatalities due to the river’s irregular habits 
and still maintain a large population, without any modifieation in 
birth rate. 

Mice of the Peromyscus maniculatus group are known to be much 
more prolific than Perognathus or Dipodomys, not so much beeause of 
the larger litters but because there are several litters each year in the 
first-named rodent, and only one as a rule in the last two. On the 
Colorado the results of our collecting give but little information as 
to the breeding of P. m. sonoriensis. A female taken March 25 con- 
tained five embryos; one taken May 5 contained three embryos. 


Peromyscus crinitus stephensi Mearns 
Stephens Canon Mouse 


As judged from the results of our three months’ trapping, this 
was the least numerous of the smaller rodents of the region. Only 
thirteen specimens were procured, apportioned by locality as follows: 
Opposite The Needles, five (nos. 9978-9982); above Ehrenberg, one 
(no. 9983) ; twenty miles above Picacho, two (nos. 9984, 9985) ; Pot- 
holes, two (nos. 9986, 9987) ; Pilot Knob, three (nos. 9988-9990). 

All of these localities are on the California side of the river. It 
would thus appear that in its lower course the river forms an absolute 
barrier to the eastward spread of this mouse. Yet it has been taken 
on the Arizona side of the Colorado, whence, considerably above Fort 
Mohave, Osgood (1909, p. 233) has recorded three specimens. 

The associational preferences of this mouse are most pronounced. 
All of the specimens were captured near or among rocks in ravines or 
on steep hillsides. It is thus a rupicoline species with much the same 
associational exclusiveness as Perognathus spinatus. As in the case 
of the latter, each mountain mass probably possesses its more or less 
completely isolated colony of these mice, separated from its neighboring 
colony by the uninhabited interval of level mesa or valley. 


Peromyscus eremicus eremicus (Baird) 
Desert White-footed Mouse 


The series of 109 specimens of this mouse (nos. 9991-10099) are 
representative of the following localities: Arizona side: Mellen, foot of 
The Needles, above Bill Williams River, Ehrenberg, twenty-five miles 
below Ehrenberg, ten miles below Cibola, five miles north of Laguna, 
and Yuma; California side: opposite The Needles, Chemehuevis Valley, 


230 University of California Publications in Zoology [ Vou. 12 


Riverside Mountain, Palo Verde, opposite Cibola, twenty miles above 
Picacho, eight miles below Picacho, Potholes, five miles above Yuma, 
and Pilot Knob. 

The wide occurrence of Peromyscus eremicus through the region is 
thus indicated. Yet our trapping showed distinet associational pref- 
erences. The overflow bottom is evidently rarely invaded, there being 
but slight overlapping of the habitat of Peromyscus maniculatus 
sonoriensis; and the rocky hills and mesas are eschewed, so that in 
this direction the domain of Peromyscus crinitus stephensi is seldom 
encroached upon. P. eremicus was found in greatest numbers on 
arenaceous alluvium not included in the overflow area of the river 
bottom. It was present on the second bottom outside the mesquite 
belt, and along desert washes leading down from the interior; also 
on the desert mesa where wind-blown accumulations of fine sand 
afforded the same sort of ground, that is, ground of a fine but loose 
consistency, easy to burrow into. 

The breeding season of the desert white-footed mouse, as indicated 
by the following dates, covers at least three months. Nearly or quite 
full-grown, blue-pelaged young were taken as early as March 2, and 
from then on at increasingly frequent intervals until the last of April. 
Females were taken March 5 with three embryos, March 8 with three 
embryos, April 13 with four embryos and April 19 with four embryos. 


Sigmodon hispidus eremicus Mearns 
Western Desert Cotton Rat 

This rodent proved to be strictly riparian in its associational prefer- 
ences, and, furthermore, was found only along the lower course of 
the river, below Ehrenberg. It belongs to a southern zone, tropical 
or semi-tropical, and is one of the lowest zonal elements entering 
the region; moreover, it has not pressed its way far up the river. 

I should judge its habits and food requirements to be closely 
similar to those of Microtus, of cooler regions. But as far as known, 
the ranges of Sigmodon and Microtus do not quite meet anywhere. 
Certainly there are no Microtus in the Colorado River bottom as low 
down as Needles. Sigmodon thus has a clear right of way, from a 
competitive standpoint. 

The three localities of capture are all on the California side; yet 
there is every reason to suppose that cotton rats occur on the Arizona 
side as well, being surely as readily able to swim as harvest mice. 


1914] Grinnell: Mammals and Birds of the Colorado Valley 231 


In the bottomlands a few miles below Palo Verde, cotton rats were 
found in apparent abundance March 31 to April 3. They were here 
inhabiting a tule patch at the edge of a slough and a dense patch of 
seedling willows adjacent. There were no runways in evidence, but 
a dozen or more little piles of cut willow twigs were found lying on 
the ground one to four feet from the edge of the water. 

The twigs were from four to eight inches in length with teeth-cuts 
at each end, and there were from half a dozen to twenty of these 
twigs ina pile. A few bits of tule stems were found in one place, and 
in another a rather thick green tule stem was nearly eaten through. 
The stomach of one of the rats taken contained a finely masticated 
mass of material judged to be tule stem. 

Five miles northeast of Yuma, Sigmodon was found April 30 to 
May 3 inhabiting a tract of wire-grass bordering a back-water slough 
near the river, and flanked by young willow growth. Near Pilot Knob, 
May 10, specimens were caught in thickets of cane surrounded by 
dense arrowweed. 

A half-grown young one was caught on the last named date. <A 
eravid female was caught on April 30. 

The weight of one of the largest males taken (no. 10626) was 
eight ounces; that of a smaller male (no. 10632), though quite adult, 
414 ounces. A very great range in size is also shown from the accom- 
panying table of measurements, which lists all of the specimens taken 
by the expedition. So great is the discrepancy between nos. 10626 
and 10627, and other apparently fully adult males in our series, that 
the existence of two distinct species was at first suspected. The two 
large examples referred to present measurements far above any of the 
specimens listed by Mearns (1907, p. 453), and some twenty per cent 
above the typical size of the race eremicus as given by Bailey (1902, 
p. 107). There are also apparent other differences concurrent. with 
the great size, namely, rather coarser pelage, greater hairiness of tail, 
and a faintly pinkish cast to the coloration on the rump and sides. 

I sent specimens illustrating this state of affairs to Mr. Vernon 
Bailey, who gives it as his opinion that the differences are due to 
age; in this genus ‘‘old males often outgrow the bounds of specific 
characters.’’ In support of this contention, that the extremes are con- 
specific, is the consideration of geographical probabilities: to-wit, that 
experience teaches that it is not to be expected that two closely related 
species in a genus exist together in exactly the same locality. 


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1914] Grinnell: Mammals and Birds of the Colorado Valley 233 


On the other hand, these aberrant examples might be recognized 
as mutants, that is, variants of phylogenetic significance though not 
geographically isolated. After going over the ground, I lean per- 
sonally towards the notion of the existence of an exceptionally wide 
range in ordinary fluctuational variation, as accounting for the 
extremes in question. This is coupled with the inadequacy of the 
material available, so that the distinctions are emphasized. With large 
series more nearly normal frequency would be expected. 

In this connection it may be pointed out that an alleged race, 
Sigmodon hispidus arizonae Mearns (see Bailey, 1902, p. 108), has 
been described from Fort Verde, Yavapai County, Arizona, with char- 
acters close to those appertaining to the Colorado River giants in 
question. In size these two specimens from the Colorado equal or 
exceed that of the type of arizonae, so that the existence of a separate 
race based on size is questionable. 


Reithrodontomys megalotis deserti J. A. Allen 
Desert Harvest Mouse 


This rodent was confined strictly to the riparian strips. Although 
not found by us above Ehrenberg, it probably does occur in suitable 
environs as far up the Colorado as these are afforded, that is, up to 
the steep-walled canons above Fort Mohave. Specimens were trapped 
close to the water’s edge, usually in grassy places adjacent to tracts 
of willows. In some eases these mice were on islands cut off by chan- 
nels from connection with the shore. In practically all cases the 
ground they occupied would at high water be entirely submerged, so 
that the mice would either have to swim to higher ground or take 
refuge in the willows and drift piles. Doubtless they swim freely and 
are also swept back and forth across the river channels at flood time. 

Twenty-one specimens were preserved (nos. 10165-10185). The 
localities of capture were: Arizona side: Ehrenberg, five miles north 
of Laguna, and Yuma; California side: near Palo Verde, and five 
miles northeast of Yuma. Young over half grown were taken May 2, 
and on the same day a female containing five embryos. j 


Neotoma albigula venusta True 
Colorado Valley Wood Rat 


This species of wood-rat was in distribution the exact associational 
complement of Neotoma intermedia desertorum: it was restricted to 


234 University of California Publications in Zoology — [ Vou. 12 


the riparian strips, and no evidence was forthcoming of its occurrence 
out on the desert mesa, in the hills, or even up the desert washes. As 
compared with rodents of the desert proper this species might for 
the sake of emphasis be termed semi-aquatic, for it was trapped within 
a few feet of the main river, and its tracks were noted on fresh mud 
at the water’s edge. Individuals were caught on willow-grown islands, 
which were subject to submergence with rising water, so that the 
animals would be forced to live in trees or take to water. No signs 
of nests were observed in trees anywhere, and it seems reasonable to 
suppose that these wood rats voluntarily swim narrow channels, 
especially when pressed by hunger. 

Although foraging widely over the bottomlands, the permanent 
abodes of these rats appeared to be chiefly located at the lateral rims 
of the riparian belts, just at the upper reach of high water. This, too, 
marked the belt of mesquites, so that the mesquite association can 
be confidently assigned as the distinctive habitat of Neotoma a. venusta. 

At no point did we find the ‘‘enormous nests’’ of this species 
described by Mearns (1907, p. 474) as found by him below Yuma. 
His name, Neotoma cumulator, for this species was selected on this 
account. A few small stick houses were found by us in the mesquite 
strip above Mellen, and nests of similar scant proportions, at other 
places down the river. In all cases the rats appeared to have ready 
access to subterranean burrows. At the mouth of the Gila River, near 
Yuma, many burrows were found in a tract of guatemote without 
trace of stick nests, and a number of the animals were caught. It is 
quite possible that recurring unusually high flooding discourages the 
rats in the portion of their range explored by us from amassing much 
material, since it is subject to being floated off. 

We found small young as early as February 24 and 25 at Mellen, 
and from that date on to May 5, near Yuma. It is probable that the 
breeding season is just about closed by the time the annual overflow 
begins, so that young as well as adults are able to seek safety for the 
brief period of exile by assuming arboreal habits or swimming to 
higher ground. 

The series of 68 specimens of Neotoma albigula venusta obtained 
(nos. 10463-10530) represent localities of capture as follows: Cali- 
fornia side: near Riverside Mountain, 2; Palo Verde, 2; twenty miles 
above Picacho, 8; eight miles below Picacho, 2; five miles northeast 
of Yuma, 8; Pilot Knob, 3. Arizona side: Mellen, 12; Ehrenberg, 3; 
twenty-five miles below Ehrenberg, 1; ten miles below Cibola, 7; five 


1914] Grinnell: Mammals and Birds of the Colorado Valley 235 


miles northeast Laguna, 6; near Yuma, 14. Two adult males from 
twenty miles above Picacho and near Yuma weighed nine ounces each; 
an adult female from the former station weighed seven ounces. 

No differences are observable between the specimens from the two 
sides of the river. The river is evidently in searcely any degree ¢ 
barrier to distribution in this species. Rather has the Colorado bottom 
served as a highway of invasion for the species from its center of distri- 
bution, which is manifestly to the southward, the desert on either side 
acting as confining walls. An examination of the series of specimens 
shows an apparent slight decrease in size up the river, that is, away 
from the assumed center of dispersal. But since there is also con- 
tinued, though diminished, growth of individuals with age, and because 
of the few examples from any one of the uppermost stations, this 
direction in variation up the river is not to be considered as established. 
Much more material is needed. 


Neotoma intermedia desertorum Merriam 
Desert Wood Rat 


The 41 specimens taken (nos. 10424-10462, 10716, 10717) indicate 
stations of occurrence as follows: California side: opposite The 
Needles, 3 specimens; Riverside Mountain, 2; opposite Cibola, 1; 
twenty miles above Picacho, 13; eight miles below Picacho, 3; Potholes, 
2; Pilot Knob, 11. Arizona side: foot of The Needles, 3; above Bill 
Williams River, 1; ten miles below Cibola, 2. 

None of our specimens was trapped within the riparian strips; 
all were taken on the desert proper, though at The Needles, where 
because of the abrupt rock walls the riparian element is in places 
reduced to a mere nearly vertical band, desert wood-rats or signs of 
them were found within a few yards of the river on the opposite sides. 

Here scanty accumulations of sticks were observed in crevices 
among loose boulders on hillsides or in clefts of the walls of ravines. 
At Riverside Mountain, some two miles back from the bottomlands, 
nests of large size were noted among boulders; this was true, too, at 
Pilot Knob, where also sign was seen and the rats themselves caught 
around clumps of desert tea far out on the mesa. 

Fifteen of the specimens taken at the different stations are young 
of varying sizes. Two very small young of 172 and 167 mm. length, 
respectively, were trapped March 6 and 7 at The Needles, indicating 
early breeding. Others nearly as small were taken at Pilot Knob, May 


236 University of California Publications in Zoology | Vou. 12 


10; so that there is not such constancy in breeding time in this species 
as In certain other rodents of the region. 

A careful comparison of the six specimens secured from the Arizona 
side with the much larger series from the California side, shows both 
eranially and as regards external features, no tangible differences 
which might be expected to occur (and do occur in other cases) in a 
mammal of the desert proper and whose range is divided by the 
Colorado River. , 

It appears that this species of wood-rat has not previously been 
found to the east of the Colorado River (see Goldman, 1910, pp. 76, 
77). In fact, in his revision of the genus Neotoma, Goldman, in dis- 
cussion of Neotoma lepida and its subspecies stephensi (which are 
forms not dissimilar to desertorum), states (1910, p. 80) that ‘‘the 
ranges of the two |lepida and desertorum] are completely separated by 
the effective barrier of the Colorado River.’’ While the results of our 
work detract from this statement as to fact, the implication remains 
the same, namely, that the Colorado River may have been at one time 
of prime service in effecting the isolation of the lepida stock, particu- 
larly in the upper (northeastern) course of the river (see Goldman’s 
map, p. 77). 

I sent my Arizona-side examples of desertorum to Mr. Goldman 
with the request that he examine them closely and give his opinion 
as to their possible approach in characters to NV. l. stephensi. The 
latter form is, by the way, the nearest wood-rat of the same group 
recorded from east of the river, its nearest station being (Goldman, 
1910, p. 80) the Hualpai Mountains, Arizona, only about fifty miles 
east of The Needles. Mr. Goldman rephed that my specimens were 
‘“typieal desertorum,’’ and hence not bearing any significant resemb- 
lance to stephensi. 

It would appear, therefore, that the desertorwm, as found by us 
on the Arizona side of the Colorado at the three stations named, is of 
probably direct and relatively recent descent from the stock on the 
California side. While this wood-rat does not inhabit the river bottom- 
lands, as does Neotoma a. venusta, it is quite possible that individuals 
forage down to the water’s edge where the riparian strip is reduced 
in width or practically wanting, as at The Needles. In event of rats 
becomigg marooned on drift-rafts at periods of rising water such 
animals would be in a position to be ferried across the river; for it is 
not unlikely that such rafts would be carried by the swerving currents 
to opposite banks. No matter where such waifs should be landed they 


1914] Grinnell: Mammals and Birds of the Colorado Valley 237 


would, ultimately, barring accidents, tend to reach the same sort of 
environs they were used to, namely, the rocky hill slopes. 

While such a series of propitious events is in but a remote degree 
possible, it appears to me the most logical way of explaining the 
extension of the range of desertorwm across the Colorado to the Arizona 
side. As above intimated, this occurrence, if a fact, does not militate 
against Goldman’s hypothesis that lepida and desertorwm may have 
been held apart by the Colorado River, and are still so held apart 
in most of its course. 


Ondatra zibethica pallida (Mearns) 
Pallid Muskrat 


Evidence, hearsay and direct, indicates the presence of muskrats all 
along the Colorado River, from above Needles to below the Mexican 
line. They occur both in the main stream and in the various diverging 
sloughs of the big valleys. Our nine specimens (nos. 10652-10660) 
were all secured on the California side of the main stream, three near 
Palo Verde and six near Pilot Knob. 

No signs of houses were seen anywhere, the muskrats appearing to 
resort entirely to holes in banks where the current was sluggish. Near 
Palo Verde, April 1, a system of burrows was dug out by Stephens 
and Jones. This system of holes was in the bank of a tule-bordered 
slough about a half mile above its confluence with the main river. 
The entrances all opened considerably beneath the surface of the 
water at its level at the time. From these the burrows sloped upwards 
as they extended back from the slough, until those parts farthest 
from the water were two to three feet above its level at that stage. 

Some green tule stems were found in the passage-ways. One of 
the blind leads was filled in with packed tule stems, mostly dead ones, 
but moist and crushed. The regular nests consisted of dry tule stems, 
some of them shredded, and laid two to three inches deep. Some of 
the short blind leads showed fresh claw scratches in the earth at their 
ends. The passage-ways were ordinarily five inches in diameter, in 
places more, and usually kept near the surface of the ground, follow- 
ing its irregularities pretty closely. In places there was not more 
than three inches of earth above the burrow, and there was seldom as 
much as a foot. The system was in fact discovered by one of the 
party’s stepping upon a thin place and breaking through. 

The impression gained from a survey of the uncovered burrows 
was that the excavator had at the beginning burrowed up from the 


238 University of California Publications in Zoology [ Von. 12 


slough five to eight feet, then gone back a third of the way and 
started a branch to the water for another entrance. After finishing 
that, it had come half way back on the branch and dug another branch 
from it inland, then a branch from that to the water again. Roughly 
speaking, a series of connected Y’s thus resulted, with ends alter- 
nately in the water and in the bank landwards. It would appear as 
though such a plan of runways were well adapted to eluding enemies, 
both terrestrial and aquatic. An adult male and two two-thirds-grown 
young were trapped in the entrance burrows. 

In the vicinity of Pilot Knob the muskrats were living along the 
steep northern bank of the main river, where the water was overhung 
by a dense growth of cane. A number of willow branches sagging 
into the water and drift logs caught in the tangle of cane (pl. 4, fig. 3) 
showed themselves to be the nightly rendezvous of muskrats. On 
these, at the farthest projecting portions, and usually not over four 
inches above the surface of the water, capsule-shaped pellets of exere- 
ment to the number of three to a dozen or more marked the perching 
places of the rats. This excrement was usually fresh, as the logs would 
go awash with the frequent cross-river winds. Number 0 steel traps 
set on these logs without attempt at concealment, either with or with- 
out bait (apple or potato), caught five adult muskrats, May 10 to 14. 
Four out of the five were drowned, the traps being provided with 
chains which in turn were nailed to the logs. A half-grown youngster 
was shot on the latter date as it swam among some trailing cane stalks. 

Muskrats have recently invaded the Imperial Valley along the 
irrigation canals leading from the Colorado River. I visited this 
valley in February, 1912, and was told there that the California Devel- 
opment Company found it necessary to hire men to shoot and trap 
muskrats because of the damage done in burrowing through the 
levees. The rats even follow the smaller ditches out into the farms 
at Calexico, Heber, and El Centro. At Calexico, February 8, I 
obtained a fresh specimen shot by a boy on a farm elose by, on the 
California side of the Mexican boundary. 

The entire series of ten specimens of the pallid muskrat in the 
Museum displays the now well-known features peculiar to the race 
represented, namely, small size, pale color and relatively scanty pelage 
(see Mearns, 1907, p. 495). The weights of the five full-grown male 
animals were: 31, 24, 21, 26, and 27 ounces, respectively; average 
26 ounces. 


1914] Grinnell: Mammals and Birds of the Colorado Valley 239 


Thomomys chrysonotus Grinnell 
Ehrenberg Pocket Gopher 


The only place on the Arizona side of the river where gopher sign 
was seen was on the mesa within two miles back from Ehrenberg. Here 
two sets of mounds were found on low sandy ridges and a single gopher 
caught from a burrow, March 27. This specimen is a male, young adult 
(no. 10617) and proved so different from any previously described 
race of gopher as to warrant making it the basis of a new specific 
name (see Grinnell, 1912, p. 174). This is evidently an upland species, 
that is, not riparian, and its nearest relationships are suggested to 
be with forms to the east and north rather than with 7. perpallidus 
or 7. albatus of the desert region west of the river. 


Thomomys albatus Grinnell 
Imperial Valley Pocket Gopher 

At only one station on the California side of the river did we 
find any sign of gophers. This was at our last camp, east of Pilot 
Knob, where on the site of the old Hanlon Ranch several sets of work- 
ings were located, and eight specimens taken (nos. 10618-10625), May 
7 to 10. Four of the animals were young, not more than half grown. 

These workings were all on the first bottom just at the outer mar- 
gin of the arrowweed association, in ground barely reached at the 
highest level of overflow. The absence of gophers in the greater por- 
tion of the Colorado bottom is reasonably explained by the occurrence 
of the yearly overflow which would drown them out. The colonies 
invading the river bottom do so only at points which they reach from 
some more favorable center of distribution back from the river. 

As already shown (Grinnell, 1912, p. 172), our gophers from the 
Pilot Knob station belong to the species occupying the alluvial delta 
region including Imperial Valley, which species is distinet from both 
the one represented on the Arizona side at Ehrenberg (Thomomys 
chrysonotus) and the one of the western end of the Colorado desert, 
at Palm Springs (Thomomys perpallidus). 

The characters distinguishing albatus from chrysonotus cannot 
be reasonably considered as due alone to the action of the river as a 
barrier, for the former species belongs to a different association 
(saltbush) from the latter which, as far as known, adheres to the 


240 University of California Publications in Zoology — [ Vou. 12 


creosote association. Thus the two species might exist on the same side 
of the river, with complementary ranges. Still, as far as known, 
neither species occurs on the opposite side of the river, the river thus 
serving as the factor of absolute delimitation in the respective cases. 
It is probable that the two species had latterly a separate course of 
origin, having been derived from distantly located stocks. 


Dipodomys deserti deserti Stephens 
Big Desert Kangaroo Rat 


The thirty-nine specimens secured were preserved as follows: nos. 
10352-10381, skins with skulls; nos. 10708, 10709, complete skeletons ; 
nos. 10724-10730, alcoholics. The following localities are represented : 
Arizona side: Mellen, 23; Ehrenberg, 2; twenty-five miles below Ehren- 
berg, 1; five miles northeast of Laguna, 3. California side: Riverside 
Mountain, 1; near Pilot Knob, 9. 

This rat shows strong preferences for the aeolian sand association. 
It was found present practically wherever such accumulations reached 
an extent of an acre or more of sufficient depth to contain the burrows. 
This essential depth was seen to be at least a foot. Because of the 
limited powers of digging, as shown both by weak incisors and small 
front feet and claws, only such loose and fine-grained substratum could 
be oceupied by this rodent. The packed floor of the mesa or the hard- 
pan of the hillsides was uninhabitable. 

The presence of this Dipodomys could always readily be recognized 
by means of conspicuous burrows, the mouths of which were usually 
at all times open. The looseness of the ground in which the burrows 
are dug results in their frequent caving in when men or horses walk 
over them. When the wind had not acted to efface them, the tracks 
of the rats could be plainly seen on the surrounding sand surface 
(pl. 11, figs. 16, 17). These tracks showed imprints of the hind feet 
and tails, indicating the tripedal mode of ambulation characterizing 
the Heteromyidae. 

The associational restriction of this large kangaroo rat gives the 
impression that it possesses a colonial habit. But it was observed that 
where proper conditions were continuous, as at the north base of 
Pilot Knob, the local distribution over the suitable area was practically 
uninterrupted. The burrows occur in small groups, each group prob- 
ably representing but a single home center and occupied ‘at times by 
a single adult, at others by an adult pair, and at others by adult and 


1914] Grinnell: Mammals and Birds of the Colorado Valley 241 


young. Tracks were to be seen far and wide between the scattering 
eroups of burrows. 

At Mellen, February 25, specimens of Dipodomys deserti trapped 
had their cheek-pouches filled with the minute blossoms and stem ends 
of Achyronychia coopert, an inconspicuous plant growing prostrate 
upon the surface of the broad, sandy intervals between the creosote 
and Atriplex polycarpa bushes (pl. 11, fig. 16). 

The series of specimens from Mellen, on the Arizona side at the 
north, is faintly darker in color dorsally on an average than the series 
from near Pilot Knob on the California side at the south. Otherwise 
I can see no differences between specimens from the two sides of the 
river. 

Like Peromyscus eremicus and Perognathus penicillatus, Dipo- 
domys deserti inhabits the second bottom in places along the river, 
though much less continuously than these smaller rodents. Doubtless, 
as with them, individuals occasionally forage on to the first bottom, 
and thus run the chance of being transferred from one side of the river 
to the other by means suggested in other parts of this paper. 


Dipodomys merriami merriami Mearns 
Merriam Kangaroo Rat 

A series of 168 specimens of this small kangaroo rat were pre- 
served, and besides these many were discarded. The Museum numbers 
are: 10186-10351 (skins with skulls), 10710-10712 (skeletons), 10733— 
10741 (alcoholics). The following localities are represented: Arizona 
side: Mellen, 18; foot of The Needles, 6; above Bill Williams River, 6; 
Ehrenberg, 10; twenty-five miles below Ehrenberg, 3; ten miles below 
Cibola, 8; five miles northeast of Laguna, 12; Yuma, 2. California 
side: five miles below Needles, 3; opposite The Needles, 4; Cheme- 
huevis Valley, 10; near Riverside Mountain, 6; above Blythe, 3; oppo- 
site Cibola, 19; twenty miles north of Picacho, 21; eight miles below 
Picacho, 12; Potholes, 5; Pilot Knob, 20. 

As indieated above, this was a widespread and abundant desert 
rodent, being taken in nearly every line of traps outside of the over- 
flow bottom. Its greatest abundance oceurred on sandy ground 
adjacent to desert washes, on tracts of aeolian sands on the desert 
mesa, and on the second bottom paralleling the river on either side 
through the broad valleys, but wholly above the reach of the highest 
floods. In a number of cases specimens were trapped on the packed 


242 University of California Publications in Zoology — [Vou. 12 


eround of the upper mesa, and occasionally on rocky hillsides. These 
instanees probably show great extent in foraging radius, the home 
burrows being in the softer ground of neighboring ravine bottoms or 
in depressions where sand could lodge. For, as far as observed, the 
burrows are always dug by the animals themselves, and their burrow- 
ing powers are weak. 

Their small size, however, does not require anywhere near the depth 
of workable soil that the huge D. deserti needs. Probably the far 
greater restriction in distribution of the latter species compared with 
that of D. merriami is determined by this difference in depth of soil 
required. 

The burrows of D. merriami are not easy to locate, as the entrances 
are left smoothly closed during the day. But where the fine wind-laid 
sand composed the surface of the ground, the hind foot and tail tracks 
showed clearly the routes taken by the rats to and from their burrows. 

The breeding season is at its height in April, though the following 
dates show considerable latitude. Females with embryos were taken 
on February 19, March 5 and 6, April 1 to 4, 19 and 21, and May 8. 
On each of the above dates but one animal is concerned, except during 
the period from April 1 to 4, when observations were made upon at 
least ten, and April 19, upon two. In sixteen cases there were two 
foetuses, in two cases three. 

The above data indicate a relatively slow rate of reproduction: 
but one litter per year is raised and there are seldom more than two 
young to a litter. This indicates a much safer existence, individually, 
for this kangaroo rat than for the ground squirrels and white-footed 
mice of the same habitat. In the case of these latter rodents litters 
consist of four to eight young, and in some species, at least, two or 
more litters are born each year. Kangaroo rats must be relatively 
very successful in escaping the numerous enemies that assail the rodent 
population of the desert. 

The writer finds it impossible to refer the Colorado River series of 
Dipodomys merriami, as a whole, to the subspecies D. m. simiolus (type 
locality, Palm Springs, on the western arm of the Colorado Desert), 
as might have been expected. The material shows great range in 
variation; some of the specimens, particularly in the series from the 
lower course of the river, are, it is true, very close to simiolus. On 
the other hand many, especially from the Arizona side of the river 
above Ehrenberg, are not with certainty distinguishable from near- 
topotypes of D. m. merriami, from south-central Arizona. It proves, 


1914] Grinnell: Mammals and Birds of the Colorado Valley 243 


however, impossible to assort the series into two subspecies on any 
satisfactory basis, and, since the average appears nearest D. m. mer- 
riami, all are placed under this name. 


Perognathus bombycinus Osgood 
Yuma Pocket Mouse 

Twenty-five specimens were procured (nos. 9956-9977, 10742— 
10744), three being preserved as alcoholics, and the rest as skins- 
with-skulls. Only two localities are represented, near Ehrenberg on 
the Arizona side of the river, where eighteen specimens were caught 
March 25 to 30, and near Pilot Knob on the California side of the 
river, where seven were taken May 7 to 15. Those at Ehrenberg were 
caught on areas of wind-blown sand, especially where heaped about 
the bases of creosote bushes, on the mesa a mile or more back from the 
river. At Pilot Knob the species was found on aeolian sand accumula- 
tions and in shallow sandy washes on the desert mesa. 

In all cases Perognathus bombycinus was found on common ground 
with Dipodomys deserti, Dipodomys merriami, Peromyscus eremicus, 
and Perognathus penicillatus. The five rodents named thus have very 
similar associational preferences. While Perognathus bombycinus was 
not found by us as near the river as the second bottom (as were all the 
other rodents named) it is fair to assume that it may so occur where 
conditions favor. And like the others of the same association, transfer 
of individuals from side to side is likely to have taken place at inter- 
vals in the past. This might be advanced as a reason for the close 
similarity of the representations of bombycinus on the two sides of 
the river. 

The original description of Perognathus bombycinus (Osgood, 1907, 
pp. 19, 20) was based on a single specimen from Yuma, Arizona, and 
two from just over the Mexican line in Sonora. Our present series 
bears out to the dot the cranial characters assigned by Osgood, namely, 
as compared with Perognathus panamintinus bangsi, enormously 
inflated audital bullae and mastoids, the latter conspicuously pro- 
tuberant posteriorly, and narrow interparietal. Externally our speci- 
mens of bombycinus are distinguishable from bangs? in their very pale 
coloration, which consists in lighter ground color (dilute pinkish buff) 
and almost obsolete black tippings to the hairs dorsally. Although, 
as compared with bangsi, bombycinus has greatly enlarged audital 
bullae, the external ear is not of appreciably greater size. 


244 University of California Publications in Zoology — [ Vou. 12 


The measurements of twenty adult males from our series are, aver- 
ages and extremes: total length 139 mm. (130-149), tail vertebrae 79 
(70-85), hind foot 18.5 (17-19.5). 

I am unable to find any differences between the specimens from 
Ehrenberg and those from Pilot Knob, the species being thus identical 
on the two sides of the river. We found no traces of pocket mice of 
this group anywhere else along the river. Osgood (1907, p. 20), how- 
ever, records typical bangsi from Needles, California. It is quite pos- 
sible that there is a hiatus between the ranges of bangsi and bomby- 
cinus. At any rate, among the large series of bangsi in the Museum 
from the Mohave desert and from the west side of the Colorado desert 
from San Gorgonio Pass to the Mexican line, there is no individual 
showing close enough approach in characters to bombycinus to war- 
rant classing it as an intermediate. Although the affinities of bomby- 
cinus are clearly with the panamintinus group, there is good evidence 
for carrying the former as a full species. 

Curiously, only one out of the entire twenty-five bombycinus taken 
was afemale. This, and the fact that no young-of-the-year were found, 
would seem to show that up to May 15 the breeding season had not 
yet begun. For experience with other rodents in different places 
indicates that as soon as the young are born the females forage abroad 
actively, and are then caught in at least equal proportions to the males. 


Perognathus formosus Merriam 
Long-tailed Pocket Mouse 


A series of 44 specimens was obtained, 40 skins-with-skulls (nos. 
9652-9691) and 4 aleoholies (nos. 10794-10797). These were taken 
exclusively on the California side of the river and represent localities 
as follows: five miles below Needles, 6; opposite The Needles, 12; 
Chemehuevis Valley, 8; Riverside Mountain, 5; near Blythe, 1; Pot- 
holes, 5; Pilot Knob, 7. 

The known range of this species is thus carried southeast along 
the west side of the Colorado River to the Mexican line (see Osgood, 
1900, pp. 40, 41). The distribution is not, however, continuous, for 
marked restriction is shown to hilly country and rough mesas. The 
immediate valley of the Colorado is avoided, as also the broad, flat and 
low desert depressions between the mountain ranges. 

Yet within the rough country trapping showed that Perognathus 
formosus occurred most frequently upon the narrow strips of loose, 


1914] Grinnell: Mammals and Birds of the Colorado Valley 245 


sandy soil along ravine bottoms or along washes cutting the elevated 
mesas. This closer associational phase of occurrence is significant on 
comparison with the local distribution of Perognathus spinatus. For- 
mosus and spinatus both live west of the river, and both occupy the 
same general associations (see figs. C, F). But locally the two are 
often found on separate ground, spinatus adhering most closely to 
the vicinity of boulder-strewn hillsides and fractured outcrops, while 
formosus prefers smoother ground, as noted above. 

At Riverside Mountain formosus was caught on the mesa, but not 
next to the abruptly rising base of the mountain where spinatus was 
found. But opposite Cibola, where pocket mice were phenomenally 
abundant, examples of Perognathus spinatus, P. formosus and P. peni- 
cillatus were all caught within a radius of fifty feet. The first two 
species were found in a single trap on successive mornings. 

As further considerations with regard to the occurrence of for- 
mosus and spinatus on common ground, it is to be remembered that 
the two species belong to separate subgenera (Perognathus and Chae- 
todipus, respectively), and thus have structures so different that impor- 
tant differences in food or other requirements are suggested ; so that 
the forms are probably complementary instead of violently competitive. 

The breeding time of Perognathus formosus is indicated by the 
capture of blue-pelaged young at Potholes and Pilot Knob on April 
28 and 29, and May 10 to 14. Preceding the first-named date no 
signs of breeding were in evidence, so that relative lateness in this 
annual function is indicated. 


Perognathus penicillatus penicillatus Woodhouse 
Colorado Desert Pocket Mouse 


A series of 196 specimens preserved, 184 as skins-with-skulls (nos. 
9692-9875), two as skeletons (nos. 10720, 10721), and ten as alco- 
holics (nos. 10798-10807). The following localities are represented: 
California side: five miles below Needles, 8; opposite The Needles, 10; 
Chemehuevis Valley, 17; near Riverside Mountain, 16; near Blythe, 
2; above Ehrenberg, 1; opposite Cibola, 35; twenty miles above Pic- 
acho, 10; eight miles below Picacho, 8; Potholes, 1; four miles below 
Potholes, 1; five miles northeast of Yuma, 5; near Pilot Knob, 8. 
Arizona side: Mellen, 12; above Bill Williams River, 2; Ehrenberg, 


16; twenty-five miles below Ehrenberg, 3; ten miles below Cibola, 27; 
five miles northeast of Laguna, 8; near Yuma, 6. 


246 University of California Publications in Zoology | Vou. 12 


In addition, the Museum contains four skins (nos. 5636, 5637, 5825, 
5826) taken in 1861 by Dr. J. G. Cooper at Fort Mohave, on the 
Arizona side twelve miles above Needles. 

It is at once patent that of the five distinct species of Perognathus 
found along the lower Colorado, P. penicillatus is both most abundant 
and most widespread. Very many more individuals were trapped day 
by day than were preserved. 

It may be asked why explicit record is not here presented, not 
only of specimens saved, but of all the individuals trapped. The reason 
is that so similar in external appearance are the various species of 
Perognathus, that especially with our previous unfamiliarity with 
them, we were liable to many mistakes in identification in the field. 
After museum study was made of the suites of specimens brought home, 
with skulls cleaned and measurements tabulated, it was found that 
errors had actually oceurred in a number of cases thus checked up. 
Perognathus penicillatus and P. intermedius had been confused on 
the Arizona side, and P. penicillatus and P. formosus on the California 
side, especially in the cases of immatures. Field determinations were 
therefore totally disregarded. 

It may scarcely be necessary to remark that penicillatus (plus 
angustirostris, if the form designated by the latter name be consid- 
ered tenable), intermedius, spinatus, and of course formosus, are all 
distinguishable with absolute certaity (for characters see Osgood, 
1900). No difficulty was experienced in allocating all specimens as 
soon as cleaned skulls were available. 

Although obtained at all collecting stations and from all associa- 
tions, reference to the graph for frequeney of occurrence (see fig. D) 
clearly shows that Perognathus penicillatus has marked preferences for 
one particular environment. This preferred habitat is characterized 
by ground of fine-grained sand. The second bottom along the river 
and the broad washes of the flatter parts of the desert furnish the 
ideal conditions; and in the most typical of these penicillatus is either 
the exclusive or the prevailing member of the genus present. 

Penicillatus is the only species invading alkali depressions at the 
edges of the broad valleys back from the river (where Suaeda grows), 
and is the only species occurring regularly on parts of the overflow 
area of the river bottom. Referring again to the frequency record 
(fig. D), it is shown that penicillatus was found in every member of 
the riparian group of associations, even into the willows. Individuals 
were always, however, as far as record and memory serves to estab- 


1914] Grinnell: Mammals and Birds of the Colorado Valley 247 


lish the point, trapped on dry surfaces, indicating a preference for 
dryness. On thoroughly dried-out, rather high sand banks along the 
river where the bottom strip was narrow, sign was noted to within a 
few feet of the edge of the swift current. Individuals were trapped 
on sand bars, at the time high and dry, but so slightly elevated that 
rising water would at first form islands of them and then engulf them 
completely. 

No doubt, as with Peromyscus, aggressive individuals of Perog- 
nathus penicillatus are often thus caught and set adrift, thus giving 
a chance for transfer from one side of the stream to the other. As 
discussed in a general way elsewhere, there seems to be significant cor- 
relation between the fact of this free invasion of the river bottom 
and existence and similarity of the representatives of the species on 
both sides of the river. 

The breeding season of penicillatus is of greater duration than that 
of the other pocket mice of the region, as shown by the following data. 
A female taken April 19 contained five embryos; another taken May 4 
contained four embryos. Gray-coated young-of-the-vear were taken as 
early as March 5 and at intervals from then on through April and 
into May. 

After study of the Colorado River series in connection with the 
120 additional specimens in the Museum from various loealities in 
the Salton Sink region of the Colorado desert, I have come to the 
conclusion that the recognition of the name angustirostris as apply- 
ing to any of the Colorado River specimens is impracticable. Osgood 
(1900, pp. 45-47) deseribed a race angustirostris from the Colorado 
desert, type locality Carriso Creek, west side of the desert. The 
Museum has a good topotype series, and also a series from Mecea, in 
the bed of the Salton Sink at the northwest end of Salton Sea. 


““angustirostris’’ were, as compared 


The characters assigned to 
with penicillatus, small size, less massive skull, and longer and more 
slender rostrum. Osgood referred his few specimens from the Colorado 
River, from Ehrenberg and above, to penicillatus, while those from 
Fort Yuma and all localities to the west he listed under angustirostris. 

Close scrutiny of the extensive material now at hand shows great 
instability in all the alleged characters. There is as notable fluctuation 
in size from place to place up and down the river, as across the Colorado 
desert. The topotypes of angustirostris include some skulls as massive 


as many from Mellen, Arizona. 


248 University of California Publications in Zoology _{ Vou. 12 


One thing is apparent, that the Mecca series includes more small 
specimens, with narrow-snouted skulls, than series of like extent from 
elsewhere. There is thus a tendency towards the existence of a race, 
of the characters assigned by Osgood to his angustirostris, in Salton 
Sink. But if the upper Colorado Valley specimens are penicillatus, so 
also are those from all the way down to Yuma and Pilot Knob and 
thence across to the east flank of the Coast Range in eastern San Diego 
and Riverside counties. 

In view of the locally fluetuating variations through the region 
under consideration, the inclusion of all under the one name penicil- 
latus seems now the wisest course. Should the penicillatus of Wood- 
house prove to represent a truly distinct race, occupying the elevated 
north central deserts of Arizona, as hinted by Osgood, then angustiros- 
tris would be the name to be used for the subspecies of penicillatus 
oceurring all along the Colorado Valley below the Grand Canon, and 
through the Colorado desert. 


Perognathus intermedius Merriam 
Intermediate Pocket Mouse 


A series of 82 specimens taken, nos. 9876-9955, saved as skins- 
with-skulls, and nos. 10785, 10786, as alcoholics. The localities of cap- 
ture were all on the Arizona side of the river, as follows: Mellen, 19; 
foot of The Needles, 10; above Bill Williams River, 2: Ehrenberg, 13; 
twenty-five miles below Ehrenberg, 1; ten miles below Cibola, 27; five 
miles northeast of Laguna, 10. 

The preferments of this pocket mouse are to all appearances iden- 
tical with those of the Harris ground squirrel. Both rodents are 
restricted to rocky hills and the hard-surfaced, coarse-graveled mesa. 
In both situations scattering creosote bushes form the prevailing vege- 
tation, though on the hills Hncelia farinosa is an additional conspicuous 
plant. Perognathus intermedius was in no case found so near the 
river as the second bottom (see diagram, fig. E). In fact, only three 
out of the 82 individuals caught were found in sandy desert washes, 
and these could have readily reached the points of capture in foraging 
down from home centers on adjacent hills or mesa. 

Our series of record stations, as above named, carry the known 
range (see Osgood, 1900, pp. 52, 53) of P. intermedius to the west, 
and mark its limits in that direction as being at the east side of the 


1914] Grinnell: Mammals and Birds of the Colorado Valley 249 


Colorado River bottom. The presence of the species was ascertained 
at every station where appropriate ground was trapped, from Mellen 
nearly to the Laguna dam. 

Among all the specimens taken, only two were juvenals. These 
were taken above Laguna on April 24 and 25. None of the numerous 
females taken earlier in the season contained embryos, so that the 
breeding season would appear to be deferred in the case of this 
species until the advent of hot weather. 


Perognathus spinatus spinatus Merriam 
Spiny Pocket Mouse 


This pocket mouse was found only on the California side of the 
river. A series of 125 specimens was taken, nos. 9536-9651 (skins- 
with-skulls), 10713, 10714 (skeletons), 10787-10793 (alcoholies). 
Localities are represented as follows: opposite The Needles (practically 
topotypes of the species), 14; Chemehuevis Valley, 1; Riverside Moun- 
tain, 26; opposite Cibola, 36; twenty miles above Picacho, 32; eight 
miles east of Picacho, 6; Potholes, 4; Pilot Knob, 6. 

The spiny pocket mouse proved to have much the same associational 
preferments as the antelope ground squirrel, namely, the hills and 
rough-surfaced mesas of the desert. It was found close to the river 
only where the riparian bottomland associations were pinched in to 
merest traces of their elements by the abutment of the hills. Places 
of this nature, where spinatus was caught within as near as a hundred 
yards of the water’s edge, and yet on ground perfectly appropriate 
to the species, were opposite The Needles and twenty miles above 
Picacho. 

At the latter locality the only exception to the above statements 
came to notice. An adult female was caught among the willows on 
first bottom and within seventy-five feet of the water, hence where 
proper conditions of topography and rising water might have resulted 
in transporting the animal to the opposite side of the river. But the 
excessive rarity of such a combination of circumstances probably 
accounts for the fact that Perognathus spinatus has never been found 
to the east of the Colorado River. The strip of bottomland at the 
point where this wanderer was captured was only about three hundred 
yards wide, and the species was found to occur commonly on the 
hillsides down to the outer edge. The individual probably occurred 


merely as a forager. 


250 University of California Publications in Zoology (Vou. 12 


Opposite Cibola, as already noted under P. formosus, the spiny 
pocket mouse occurred on a broad desert wash, strewn with boulders. 
Specimens were trapped there, not only among boulders but around 
seragely stumps of ironwood. 

The breeding season apparently does not begin till April. On the 
13th and 19th of that month females taken contained four embryos 
each. On April 28 and May 9 half-grown young were trapped. 


Lepus californicus deserticola Mearns 
Colorado Desert Jack Rabbit 


The eight specimens secured (nos. 10682-10689) came from the 
following localities: California side: five miles below Needles, twenty 
miles above Picacho, and near Pilot Knob; Arizona side: Ehrenberg, 
and five miles north of Laguna. Specimens from the two sides of the 
river are identical as far as I can see; and this is to be expected. For 
although the jack rabbit belongs primarily to the desert mesas, it 
forages also all over the flood-bottoms. I was told that this species 
of rabbit swims strongly when forced to take to the water. 

Jack rabbits occurred through the region in 1910 in very small 
numbers. At some stations none at all were seen, though old sign was 
usually plentiful. The greatest numbers were observed on the mesa 
back of Ehrenberg, but not more than half a dozen could be seen 
during a three hours’ circuit. 

Two females shot near Ehrenberg, March 28 and 29, contained two 
and three embryos respectively. 

The weight of an adult male (no. 10682) was four pounds, four- 
teen ounces; of an old female (no. 10683) six pounds. 


Sylvilagus auduboni arizonae (J. A. Allen) 
Arizona Cottontail 


The year of our exploration of the Colorado Valley was stated by 
the residents of the region to be an off-year for both jack rabbits and 
cottontails. Whatever the usual conditions, both kinds of rabbits were 
_unexpectedly searce. 

The cottontails were chiefly riparian in their loeal distribution. In 
fact, the only individuals seen desertwards beyond the mesquite belt 
were in the salt-bush association closely adjacent, and (opposite The 
Needles) up a catelaw wash not to exceed one-half mile from the river. 


1914] Grinnell: Mammals and Birds of the Colorado Valley 251 


The quail-brush association was the most preferred local habitat, the 
rabbits finding ideal refuges there beneath the thickets of Atriplex 
lentiformis. They ranged all over the bottomlands, and since individ- 
uals were seen on islands already cut off by rising water from main- 
land connection, it is probable that many rabbits are washed adrift each 
year and are carried from side to side of the river. We were assured 
that cottontails had been seen swimming in the river during flood time. 

We procured twenty-one specimens of the cottontail (nos. 10661- 
10681), representing the following localities: Arizona side: Mellen, 
foot of The Needles, five miles northeast of Laguna; California side: 
opposite The Needles, Riverside Mountain, Blythe, opposite Cibola, 
eight miles east of Picacho, five miles northeast of Yuma, near Pilot 
Knob. A close examination of the comparable specimens from the 
two sides of the river shows no differences. This is explainable on 
the ground that to the distribution of this rodent the river forms 
no permanent hindrance. 

The weight of an adult male (no. 10678) was 24 ounces, of an adult 
female (no. 10679), 26 ounces; both shot near Mellen. 

Partly grown young were caught March 20 and April 3. A female 
taken May 6 contained two large embryos. 


Felis oregonensis browni Merriam 
Yuma Cougar 


We were told of the occurrence of cougars at several points along 
the river from Riverside Mountain south. They appear to range chiefly 
through the densely wooded bottomlands. One, however, was reported 
as having been seen in the rough hills constituting Riverside Mountain. 
In the vicinity of Ehrenberg, Cibola, and Potholes I was given more 
or less definite accounts of them. At a point on the California side 
four miles below Potholes, we saw fresh footprints of one within fifty 
yards of the main river channel. 

From a rancher, J. C. Draper, who lives on the California side, 
eighteen miles north of Picacho, I purchased two skins with skulls, 
with the following history. Mountain hons had not been previously 
seen in his neighborhood for at least ten years. In the autumn of 
1909 his hogs began to disappear, until eight were gone. Finally the 
tracks of a lion were found, and then the fresh trail where a 125-pound 
hog had been dragged a quarter of a mile through the brush. The 
beast was discovered nearby, treed by a dog, and shot. This was on 


252 University of California Publications in Zoology [ Vou. 12 


the California side of the river one and one-half miles below Draper’s 
house, about November 15, 1909. 

Meanwhile the hogs had become thoroughly frightened and had 
taken to swimming the river twice daily, to forage for mesquite beans 
on the Arizona side, where they appeared to feel safer. But in Decem- 
ber traces of a lion were discovered on that side. After a pig and 
a coyote had been killed by the lion, the latter was trailed and treed 
by a dog, and shot by Draper. This was directly opposite the Draper 
house, on December 29, 1909. Draper stated his belief that the lions 
swam back and forth across the river at will. 

Both lions were males, the first (no. 10586) measuring before skin- 
ning, 6 feet 6 inches in length, the second (no. 10587) 6 feet 4 inches. 
The skull of the first is imperfect, the whole base having been shot 
away; but that of the second is entire. The skins are flat, in fairly 
good condition, and measure, as now tanned, no. 10586: total length 
2120 mm., tail 680; no. 10587: total length 2085, tail 710. 

They show the following characters. Pelage: short, fine-haired and 
smooth; longest hairs on middle of back 17mm. in no. 10586, and 
24 mm. in no. 10587 (in oregonensis from western California the pelage 
is much heavier and coarser, 28 mm. long) ; tail slender due to shorter 
clothing of hair (much more bushy in Pacific slope Felis). Coloration 
pale: clay color on upper parts deepening towards hazel down middle 
of back, and becoming whitish on mid-lower surface and on insides 
of legs and thighs. Close examination discloses an admixture mid- 
dorsally of black-tipped tawny hairs and fewer pure white hairs; but 
uniform pale hazel hairs predominate. Outer sides of legs and tops 
of feet, pale clay color; hair of soles of feet between pads, prout 
brown. Tip of tail dorsally (in both specimens), dark burnt umber for 
about 50 mm. from tip; tail otherwise dusky tawny dorsally and pale 
clay color ventrally. Back of ears blackish, dulled because of admix- 
ture of white hairs, especially towards tip and inner margin. Ears 
relatively lightly clothed. Face pale, of color of back but duller; 
black patches at bases of whiskers not conspicuous, much mixed with 
whitish. Whiskers wholly white, except three or four dorsalmost ones 
which are blackish, light tipped. 

While the skulls are small (see accompanying table of measure- 
ments) as compared with oregonensis of western California, they are 
larger than the type of ‘‘Felis aztecus browni’’ (Merriam, 1903, p. 
73) as shown by the describer’s measurements in the two particulars 
civen by him. They agree much more closely with the measurements 


1914] Grinnell: Mammals and Birds of the Colorado Valley 253 


given of the type of Felis hippolestes aztecus (Merriam, 1901, p. 593). 
Since the sagittal crest of our no. 10587 is very highly developed, 
it may be an older animal. Merriam’s transverse diameter of bulla 
in the type of brown is 16mm., while in our smallest specimen it is 
19; his upper carnassial length in brown is 20.5; in our least it is 
21.5, which, however, belongs to the otherwise largest skull. The type 
of browni came from the Colorado bottom, Arizona, twelve miles 
below Yuma. 

There thus appears to be considerable variation, and it is probable 
that there is really less to distinguish browni from aztecus than the 
original description of the former indicates. However, our material 
proves that there is a well-marked desert form of the cougar, char- 
acterized, as compared with oregonensis, by smaller size, paler colora- 
tion, and shorter pelage. 


meter of 


Greatest ¢ 


a 


CRANIAL MEASUREMENTS IN MILLIMETERS OF FELIS OREGONENSIS 
FROM THE COLORADO VALLEY 
s 
= st 
ao oe AES 
= £3 ai = Sea & £ rt 
¢ ay el Se Ep a) 
wb = : ie 2a & & a Va B s ee wo 38 
2 sae =e sc Ni Tes @ e928 F2° 2 & 
: oy =o S 23 se f S fi Be Soh BS 38 
5 3 a8 E E5 a 2 = =a BES as Gok 
cI Se as Nic a o2 O2 Ake pea Sos! 
PEIASUMMEEC WEN fscsec | Gececiciy 4 ocent A Se OMS an (2m ens sip ete STS G27 
10587 ¢ 198 161 127.5 360 41 180 62 19 1384 22.5 
1 Measured from plane of inferior surface of palatines vertically to highest point on 


surface of frontals. 


Lynx eremicus eremicus Mearns 
Desert Wildeat 


Two bobeats were trapped by our party: a female (no. 10604) in 
a patch of screwbean trees in the river bottom on the California side 
five miles below Needles, February 21; and a male (no. 10605) in a 
wash lined with palo verde and eatclaw on the Arizona side north of 
Mellen, February 27. In addition to these specimens, both saved as 
skins and complete skeletons, there is in the Museum a skin with the 
skull inside (no. 5620) taken by W. W. Holder at Mineral City 
(Ehrenberg), Arizona, in 1864. 


BROWNI 


Ine 


left upper 


bo 09 
NID 


254 University of California Publications in Zoology — [ Vou. 12 


The three skins agree in relative pale tone of coloration as com- 
pared with appropriately selected material from the Pacifie slope of 
southern California (Lynx eremicus californicus). The paleness 
consists in more extended white-tipping to the hairs. The Mellen 
specimen is slightly the darkest; but there is as much or more varia- 
tion in skins from a single locality elsewhere. 

The measurements, in-the-flesh, of the two taken in 1910 are: 


Total Tail Hind Height Tuft of 
Mus. no. Sex length vertebrae foot of ear hair on ear 
10604 2 815 150 175 63! 207 
10605 Jb 780 150 170 75 26 


* Measured on dry skin. 


There are in the Museum three skins of wildeats from Victorville 
on the Mohave Desert, which are also somewhat pale in comparison 
with Pacific Slope skins. Aside from this character of paleness, I am 
unable to find any other diagnostic features in wildeats of the desert 
region, even after carefully following Mearns’s description (1897, p. 
457). Eremicus and californicus are certainly very close, though 
present material appears to me to warrant separate recognition (see 
Stephens, 1896, pp. 210, 211). 

The weight of the male wildeat caught near Mellen was 18 pounds. 
The stomach of this animal contained the fragments of at least two 
wood-rats. These had not been chewed finely, but had been chopped 
up into chunks about an inch long; the feet were still entire. 


Canis ochropus estor Merriam 
Desert Coyote 


Coyotes were only fairly common along the Colorado Valley. They 
were occasionally seen on the desert mesas by daylight, but were 
rarely heard. It was evident from tracks on the mud bars that they 
foraged down to the river’s edge at night. 

The six specimens secured (nos. 10611-10616) represent the fol- 
lowing localities, all on the California side: five miles south of Needles, 
opposite The Needles, twenty miles north of Picacho, and Pilot Knob. 

The weights of three adult males were 16, 18, and 21 pounds; of 
two females, 18 and 20 pounds. 


bo 
On 
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1914] Grinnell: Mammals and Birds of the Colorado Valley 


Vulpes macrotis arsipus Elliot 
Desert Kit Fox 


This fox appeared to be a characteristic element in the purely desert 
fauna. The five specimens taken were all trapped on the desert mesas 
back from the river. In fact, no evidence at all was forthcoming to 
show that kit foxes ever visit the river or even the bottomlands. Their 
presence was generally detected in sandy tracts, especially around 
colonies of Dipodmys deserti. It is to be inferred that this rodent 
constitutes the main food-supply of the !nt fox in localities where 
both oceur. 

The accompanying table shows the data appertaining to the speci- 
mens caught. Although a xerophilous species, examples from opposite 
sides of the river do not differ appreciably, in either cranial or external 
characters. 

The two females taken at Pilot Knob showed evidence of having 
recently suckled young, though this stage was well passed. 


LIST AND MEASUREMENTS IN MILLIMETERS OF VULPES MACROTIS ARSIPUS 
FROM THE COLORADO VALLEY 


Weight Tail Hind 
Mus. no. Sex Locality Date Ibs. Length vertebrae foot Ear 
10581 Q Mellen, Ariz. side Feb. 26 2114, 670 260 110 80 
10582. 2 Ehrenberg, Ariz. side Mar. 27 3 725 290 120 85 
10583. § Opposite Cibola, Calif. side Apr. 5 = 740 290 115 92 
10584 Q Near Pilot Knob, Calif. side May 10 344 720 280 115 — 
10585 @Q Near Pilot Knob, Calif side May 15 3 724 297 108 86 


Urocyon cinereoargenteus scotti Mearns 
Arizona Gray Fox 

The Colorado River bottom and nearby desert proved to be well 
populated with a form of gray fox. Specimens were taken, as shown 
in the accompanying table, on both sides of the river. Besides the 
localities represented by specimens, gray foxes were also seen on the 
California side near Pilot Knob; so that the species doubtless occurs 
continuously the whole length of the portion of the river explored 
by us. 

Our Colorado River material added a subspecies to the known 
mammal fauna of the state. Urocyon c. scotti is well distinguished 
from U. c. californicus, of the San Diego district, on the basis of 


[ Vou. 12 


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1914] = Grinnell: Mammals and Birds of the Colorado Valley 257 


both external and cranial characters. Scotti, as exemplified by the 
Colorado River series, has a longer tail, higher ear and paler colora- 
tion; the rufous tinges are less bright; the white endings of the hairs 
on sides of body and tail and on top of head are more extensive, giving 
a grizzled effect in the desert race. The skull of scotti is of relatively 
lighter build; the teeth are smaller, being more slender and hence 
sharper; the auditory bullae are more inflated, that is, relatively. higher 
and steeper-sided; the rostrum is conspicuously narrower. For 
measurements of californicus see Dixon (1910, p. 304) and for dis- 
cussion of relationships see Grinnell and Swarth (1913, p. 373). 


Mephitis estor Merriam 


Arizona Striped Skunk 


A common species in the riparian strips along both sides of the 
river. Data pertaining to the eight specimens obtained is given in 
the accompanying table. This animal evidently does not stray out on 
to the open desert, being probably kept to the bottomlands by daily 
need for water; nor has it the physical ability to cover much distance 
in a night’s travel. Those caught were in mesquite, screwbean, willow 
or arrowweed tracts. Tracks were often seen on the mud at the water’s 
edge. 

A male, no. 10576, weighed three pounds, two ounces. The female 
taken near Pilot Knob May 12 contained four foetuses. 

This species of Mephitis is distinetly smaller than MW. occidentalis. 
Although in all of the six skins of estor from the Colorado Valley 
there is much more white than in occidentalis, the amount varies indiv- 
idually to a large extent. In five out of the six there is a white 
pectoral patch, and in four of these there are additional flecks of 
white mid-ventrally. The longer hairs of the tail are in each case 
fully white; but a thick growth of relatively short hairs clothing the 
tail is black terminally, white at base. As the season advances it 
looks as though the long white hairs of the tail tend to be shed, 
leaving the tail of a darker tone, until the black even predominates. 

The mid-frontal white streak varies in amount, but averages greater 
than in occidentalis. The dorsal white area (creamy white) is 
unbroken in no. 10575; in no. 10574 there are tufts of black in the 
mid-dorsal line over the rump; in the other four skins there is a 
well-defined black stripe separating the white into two lateral stripes 


258 University of California Publications in Zoology [ Vou. 12 


which join on the shoulders, and continue forward to cover the whole 
back part of the head behind an abrupt line of demareation joining 
the ears. (See pl. 13, fig. 21.) 

The question suggests itself whether or not geographic variation 
in proportion of white and black (as in Mephitis estor of the desert 
as compared with M. occidentalis of the Pacifie slope of California) is 
correlated with the maximum condition of efficiency from the stand- 
point of utility. Granted that the sole purpose of the skunk’s con- 
trasted markings is to offer a signal of warning, then the maximum 
of efficiency will be reached when the proportion of black and white 
is such as to bring the greatest degree of conspicuity amid the average 
of the natural surroundings. 

The surroundings on the desert, even in the riparian thickets, are 
far lighter-toned both night and day, than they are in the humid coast 
region. It would seem, therefore, that to secure the greatest con- 
spicuity the increased proportion of white must be provided for the 
darker surroundings, and conversely the increased amount of black 


must be presented in the lighter surroundings. This is the reverse of 
the case in fact. Hence it looks as though the warning theory does 
not gain substantial support from this direction. 

The problem of the significance of animal coloration promises 
important results when attacked upon a basis of the facts to be 
observed in geographic variation. 


LIST AND MEASUREMENTS IN MILLIMETERS OF MEPHITIS ESTOR FROM THE 
COLORADO VALLEY 


Nature of Total Tail Hind 
Mus. no. Sex material Locality Date length vertebrae foot Ear 
10574 @ Skinand Calif. side, 5 mi. south Feb. 23 665 300 65 15 
skeleton of Needles 
10575 2 Skinand Calif. side, Cheme- Mar. 9 625 300 70 25 
skeleton huevis Valley 
10576 =g¢ Skinand Arizona side, foot of Mar. 5 642 330 70 23 
skull The Needles 
10577. = g ~=Skullonly Arizona side, Ehrenberg Mar. 26 625 300 68 18 
12648 og Skeleton Arizona side, Ehren- Mar. 24 610 288 71 aly 
only berg 
10578 =g Skinand Calif. side, opposite Apr. 6 525 300 70 25 
skull Cibola 
10579 =@ Skin and Calif. side, 20 mi. N. Apr. 12 615 322 69 20 
skull of Picacho 
10580 92 Skinand Calif. side, near May 12 595 280 64 22 


skull Pilot Knob 


1914] Grinnell: Mammals and Birds of the Colurado Valley 259 


Spilogale arizonae arizonae Mearns 
Arizona Spotted Skunk 


We were told of the presence of ‘‘hydrophobia’’ skunks among 
The Needles, Arizona. Mr. Stephens saw tracks of one in the river 
bottom on the California side five miles below Needles. The species 
is certainly not common, or our traps would have given more indica- 
tion of its presence. The only specimen procured by our party was 
trapped in the arrowweed belt within one hundred yards of the river, 
on the California side at our last station, close to Pilot Knob. It is 
an adult male (no. 10573) with following external measurements 
by collector (Stephens) : length 440, tail vertebrae 170, hind foot 46, 
ear 22, weight 18 oz. The cranium measures: basilar length 49.8, 
zygomatic breadth 36.1, mastoid breadth 30.9, interorbital width 14.4, 
height of brain-case (as measured by Howell, 1906, p. 37) 17.8. 

I sent this skin-with-skull to the Bureau of Biological Survey, 
Washington, where Mr. A. H. Howell determined it to be ‘‘arizonae 
but not typical.’’ He further considered it ‘‘as grading toward 


5) 


martirensis,’’ which, from its geographical location, is quite to be 
expected (see Howell, 1906, pp. 29, 30, pl. 1). The present record 


is the basis of the first assignment of arizonae to Californian territory. 


Taxidea taxus berlandieri Baird 
Mexican Badger 


While no badgers were secured by our party directly, I purchased 
a flat skin, with skull, of a young one at the Barber Ranch twenty 
miles above Picacho. This had been killed on the California side in 
the autumn of 1909. Badgers were reported as frequently seen in 
the vicinity of the Draper ranch, eighteen miles north of Picacho, and 
also on the Arizona side in the neighborhood of Cibola. 

The skin secured (no. 10603) as compared with one of about the 
same age from Ventura County, California (no. 7078), is very much 
paler. The individual hairs have longer white tips, and a mid-dorsal 
white stripe extends from top of nose to rump. In the Ventura County 
specimen there is a white stripe from tip of nose to between shoulders, 
only. The skull of no. 10603 is smaller, doubtless due to younger age, 
but it appears to be also of relatively lighter build. 


260 University of California Publications in Zoology [ Vou. 12 


Procyon pallidus Merriam 
Pallid Coon 


An abundant representative of the riparian association everywhere. 
The accompanying list shows localities of capture from Needles to 
Pilot Knob, and also from one of the Colorado’s distributaries, New 
River, in the delta region near Imperial, California. 

At practically every place where we had occasion to examine the 
muddy margin of the river or its lateral sloughs, the conspicuous foot- 
prints of coons were to be seen. One coon was caught in an unbaited 
trap set for beaver beneath the surface of the water, at the river’s edge. 

The chief food of the coons was evidently fish. At a drying-up 
overflow pond, the shallow water of which was crowded with catfish, 
a profusion of the tracks of herons and coons showed a marked com- 
munity of interests on the part of these two fishers. 

No young-of-the-year came to notice. The female caught May 11 
near Pilot Knob contained four embryos. The one caught near 
Imperial May 10 contained five embryos. The weight of the female 
no. 10610, was 16 pounds. Two fat males, nos. 10606, 10607, weighed 
1814 and 18 pounds respectively. A lean male, no. 10609, weighed 
but 13 pounds. The very large old male (no. 7153), from near 
Imperial, weighed 20 pounds. 

In the original description of Procyon pallidus (Merriam, 1900, 
p. 151), both external and cranial characters of this form are given. 
The cranial peculiarities mentioned, as compared with Procyon psora, 
do not appear to hold in our material. There is much variation in the 
respects mentioned, as will be understood from the table of measure- 
ments. In coloration alone, however, there is abundant basis for the 
recognition of pallidus. 

The seven skins from the Colorado Valley, including Imperial 
‘Valley, although somewhat variable in coloration, are in mass effect 
conspicuously different from the coons of the Pacific Coast region. A 
large series of skins of the latter are available for comparison; and 
even selecting extremes from the pallidus and psora-pacifica series, 
no overlapping of characters can be found in the material at hand. 

The general pallor of pallidus as compared with psora is due to: 
(1) The far less amount of black on the individual hairs everywhere 
except on the nose and cheeks. This is the converse of saying that 
much more of each hair is white. Along the sides, on the ears and 
on the light intervals between the tail-rings the hairs are pure white 
with no black tippings. (2) The black areas are less in extent. The 


1914] Grinnell: Mammals and Birds of the Colorado Valley 261 


seven or eight black tail bars are narrower, thus making the light 
intervals broader; the whole tail is thus very much lighter colored. 
(3) The ‘‘black’’ color of the hairs is in many places not so intense, 
being, particularly on the tail and head, of a deep vandyke brown. 
Fading evidently brings a browning of the dark hairs to some extent. 
(4) The under fur is very much paler in color. Dorsally it is vandyke 
brown instead of bistre; on the sides and belly it grades through cin- 
namon to pale elay color. In the facial region the typical coon pattern 
is present, but the white markings are not so conspicuously contrasted. 
This is due to the less intense black, and to the fact that the pileum 
is scarcely darker than the rest of the back. The post-mental area 
and the median nasal stripe are rather pale vandyke brown. Only the 
transocular black patch is retained in nearly its usual distinctness. 
The whiteness of the backs of the ears is a notable feature; and there 
is altogether lacking the black patch behind the ears usually so well 
marked in psora. 

In a general way pallidus thus differs from psora in very much 
paler coloration. The two specimens from the vicinity of Imperial are 
topotypes of pallidus, and accord closely with the brief description of 
the form as given by Merriam. These two topotypes are the palest 
of our pallidus, but it will be observed from the list that they were 
secured in May. As might be expected from this circumstance, they 
had evidently been subjected to much more wear than especially our 
Needles and Mellen specimens, which are somewhat darker. It is easy 
to see that the cutting off of the black tips of the long hairs in the 
latter, particularly on the mid-dorsal region, would result in a decided 
paling in the general color. 

Eliminating this factor of wear, and the slight amount of fading 
which very probably occurs (even though these nocturnal animals 
may not be exposed to intense sunshine to any large extent as they 
probably spend the day in drift piles or thickets rather than in trees), 
I cannot see that those specimens from farthest up the river are any 
paler than those from farther down the river, or than the topotypes. 

It would not be reasonable, therefore, to hold that the apparently 
darker animals farthest up the river from the assumed center of differ- 
entiation (Colorado delta) indicate intergradation with darker coons 
to the northward, even though this might be expected. It is even pos- 
sible that the cliff-confined river above forms an effectual barrier to 
the north, so that there has been little chance of mingling of coon 
strains from elsewhere. Pallidus may be hemmed in by ordinarily 
uncrossable barriers, and hence has attained a clear-cut specific identity. 


[Vou. 12 


University of California Publications in Zoology 


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1914] Grinnell: Mammals and Birds of the Colorado Valley 263 


Corynorhinus macrotis pallescens Miller 
Pale Lump-nosed Bat 


At Riverside Mountain, California, March 18, three bats were found 
at the end of a sloping drift in the Steece copper mine. They were 
clinging to the rock wall, and at once took flight, attempting to pass 
us towards the opening of the tunnel. Two were secured, nos. 10694, 
10695. The fur of these has a slight reddish east, which is doubtless 
wholly adventitious, due to the fine, sticky red dust with which the walls 
of the mine were covered. Both were females, one of them containing 
a single embryo. 


Antrozous pallidus pallidus (LeConte) 
Desert Pallid Bat 


On several nights in April, large light-colored bats were moment- 
arily observed flitting about in the moonlight close over our beds. 
At times the flutter of their wings was clearly audible. But they 
seldom appeared until it was too dark to shoot. On April 20, on the 
California side, eight miles east of Picacho, a specimen was shot at 
late dusk as it flew among willows across a patch of open sky. This, 
the only example of the species obtained (no. 10696), was a female 
containing two embryos. 


Myotis occultus Hollister 
Hollister Bat 


This very distinct species was only recently described (Hollister, 
1909, p. 43) from two specimens taken May 14 and 15, 1905, on the 
California side of the river in the bottomlands ten miles above Needles. 
The fact that our own expedition failed to detect the presence of this 
bat until the first week in May would point towards its late spring 
arrival in the region generally. We obtained six specimens, as listed 
in the accompanying table of measurements. The first was shot at 
late dusk close to the river bank between files of cottonwoods, in just 
the same association as those taken by Hollister. At our second locality 
of capture, the remaining five specimens were shot over the water in 
a back eddy of the river. Here these bats arrived in considerable num- 
bers at early dusk to drink, flitting down to the water’s surface and 
dipping several times before flying off among the willows and ecotton- 
woods. We used a boat in shooting and retrieving the specimens 
obtained. 


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1914] Grinnell: Mammals and Birds of the Colorado Valley 265 


I sent one example to the Bureau of Biological Survey, where Mr. 
A. H. Howell made the specific determination here employed. Our 
series bears out closely the characters assigned in the original descrip- 
tion (Hollister, 1909, p. 43), and they are in all these respects sur- 
prisingly uniform. The very broad and flat-topped rostrum and 
brainease constitutes a character for discrimination from all other 
species of Myotis in California except orinomus. The proportions 
generally are peculiar (see table of measurements). 


/ : 

line 
Fig. H. Right upper dental series Fig. I. Right upper dental series 
of Myotis occultus, no. 10702, 2; mid- of Myotis occultus, no. 10706, g; mid- 
dle upper premolar absent. 4. dle upper premolar present. X 4. 


Variation in general proportions from 
those of preceding figure possibly due 
to age. 


An interesting fact pointed out by Hollister is the variability in 
a feature usually considered of much more fundamental importance 
than the external characters employed in distinguishing members of 
the genus, namely, the presence or absence of the middle upper pre- 
molar (pm*). In one of Hollister’s specimens this tooth was present, 
in the other absent; in three of ours it is present, in three it is wanting. 
Thus fifty per cent of the individuals so far collected lack the tooth 
in question, certainly a remarkable aberrancy from the norm in the 
genus Myotis (see Miller, 1907, p. 201), and denoting a tendency to 
specialization in this member of the genus, along a line regularly 
shown in other closely related genera (see figs. H, I). 


Myotis californicus pallidus Stephens 
Stephens Little Pallid Bat 


The accompanying table shows certain circumstances of capture of 
this species. Although obtained at but the two localities, Mellen, on 
the Arizona side, and opposite The Needles, on the California side, 


266 University of California Publications in Zoology [Vou 12 


I was fairly sure that I saw the same species at other localities along 
down the river. Those obtained were all shot at late dusk, considerably 
later in the evening than most of the appearances of Pipistrellus 
hesperus. Instead of flying high, against the sky, as in the case of 
the latter species, MW. c. pallidus was almost always foraging low over 
the bushes of the second bottom, or along shallow washes between 
clumps of mesquite, seldom appearing above the sky-line. The move- 
ments of flight were peculiar also. 


LIST AND MEASUREMENTS IN MILLIMETERS OF MYOTIS CALIFORNICUS 
PALLIDUS TAKEN IN 1910 ON THE COLORADO RIVER 


Tail Hind 

Mus. no. Sex Locality Date Length vertebrae foot 
10698 Jb Mellen, Ariz. Feb. 26 75 38 5 
10699 3 Mellen, Ariz. Feb. 28 77 38 6 
10700 3 Opposite The Needles, Calif. Mar. 1 81 40 6 
10701 2 Opposite The Needles, Calif. Mar. 3 75 35 6 


The four specimens obtained are uniform among themselves and 
with a topotype specimen of Myotis californicus pallidus (no. 7350) 
from Vallecito, on the western side of the Colorado desert in extreme 
eastern San Diego County. All agree closely with the description 
of M. c. pallidus (Stephens, 1900, p. 153). An additional feature, 
as compared with Myotis californicus californicus from Monterey, 
California, is the smaller skull of pallidus, with decidedly smaller 
brainease, less inflated in the parietal region. 


Myotis velifer (J. A. Allen) 
Cave Bat 


Not obtained by our party ; but there are in the Museum three skins- 
with-skulls (nos. 7762-7764) taken by Charles Camp at Needles July 
16 and 18, 1909. Mr. Camp states that this species was roosting in 
numbers in an old storehouse from which they were routed out and 
shot. One of the specimens was forwarded to the Bureau of Biological 
Survey, Washington, where the above determination was confirmed 
by A. H. Howell. I do not find a previously recorded occurrence of 
this species for California. 


1914] Grinnell: Mammals and Birds of the Colorado Valley 267 


Pipistrellus hesperus hesperus (H. Allen) 
Canon Bat 


The most abundant representative of the order Chiroptera 
observed during the period of our work. Seen abroad at dusk as early 
in the season as February 23, when the nights were still so cold that 
ice formed in suitable places. Numerous at Mellen, February 23 to 28, 
and swarming in the vicinity of The Needles March 1 to 3. Thence- 
forth seen at nearly every station all the way down the river. One 
thing was conspicuously noticeable in regard to occurrence, namely, 
that this bat varied directly in degree of abundance with nearness 
to cliffs, or hillsides with outcroppings of fractured rock. In other 
words, this species probably dwelt exclusively in the rocks during the 
day, from which it emerged at early dusk to forage out over the river 
bottom in the near vicinity. 

Individuals were often seen before the sunlight had yet left the 
eastern hilltops. On one occasion, as we were floating down the river 
near Picacho, a Pipistrellus appeared in flight in the glaring forenoon 
sunshine, dipped down to the surface of the water, where it touched, 
and thence flitted back to a crevice in the nearby cliff. 

Seventy-four specimens of this species were shot (nos. 10382-10423, 
10746-10777), 42 being preserved as skins and 32 as alcoholics. 
Highteen out of this series are from the California side of the river 
near Pilot Knob, only about six miles due west of old Fort Yuma, and 
are thus practically topotypes of Pipistrellus hesperus. 

In reviewing the Museum’s entire collection of Pipistrellus from 
California, it becomes clearly apparent that while there is but one 
species represented, there are two appreciably different subspecies of 
this species, one a pale-colored form occupying the arid desert regions 
from Owens Valley and the vicinity of Walker Pass southeast to 
the Mexican line, the other a darker-colored form occurring on the 
Pacifie slope of southern California (in the San Diegan district) and 
in certain parts of the San Joaquin and Sacramento valleys. 

It appears to the writer that we have here two races well worthy 
of recognition in nomenclature. The name hesperus was based on the 
desert form. A name is apparently available for the Pacific race in 
the Vesperugo merriami of Dobson (1886, p. 124). This has always 
been synonymized under Pipistrellus hesperus. While the habitat is 
given by Dobson as ‘‘North America (Locust Grove, State of New 
York) ’’, an error was committed, for the type really came from Red 
Bluff, Tehama County, California (fide Miller, 1897, p. 31). Dobson 


268 University of California Publications in Zoology | Vou. 12 


received his specimen from C. H. Merriam, whose residence at that 
time was Locust Grove, New York. 

The case seems to be clear, and I propose that the Pacifie slope race 
be called Pipistrellus hesperus merriami (Dobson), the type locality 
of which is thus Red Bluff, California. The characters of this form, 
as compared with P. h. hesperus, le in the darker, distinctly browner 
tone of coloration both above and below, and in somewhat larger size 
throughout. The Museum has specimens, unequivocally of this form, 
from the following localities, all within the state of California: Marys- 
ville Buttes, Sutter County; Raymond, Madera County; Cuyama Val- 
ley, Santa Barbara County; Fort Tejon, Kern County; San Fran- 
cisquito Canon, northern Los Angeles County; vicinity of Pasadena; 
Escondido, San Diego County. 


Eptesicus fuscus (Beauvois) 
Large Brown Bat 

One shot at dusk on the California side near Pilot Knob, May 6. 
Other bats supposed to be the same species were seen flying down 
the river high overhead the same evening. <A strong west wind was 
blowing at the time. The species was not seen at any other place. 

The specimen secured (no. 10697) is an adult female. It appears 
to differ in small size and extreme paleness from the average of the 
species from California. It about equals in the latter respect the 
palest out of a series of ninety brown bats from the Pacifie slope of 
California. The color dorsally is uniform isabella color, ventrally pale 
wood brown. Measurements: length 107 mm., tail vertebrae 44, foot 9, 
forearm 42.5, longest finger 72, ear (dry) 12. The skull, too, is appre- 
ciably small. 

A general inspection of the Museum’s series of this species from 
California points towards the existence within the state of at least 
three geographic races based on size and depth of color. But so much 
of the total area is unrepresented by specimens that systematic analysis 
at this time seems inadvisable. 


Nyctinomus mexicanus Saussure 
Mexican Free-tailed Bat 
We were fairly certain of seeing this bat at almost every station, 
as a rule flying high and often squeaking loudly. Only three specimens 
were secured (nos. 10690-10692) the first two at Mellen, February 26, 
the third in Chemehuevis Valley, March 11. 


1914] Grinnell: Mammals and Birds of the Colorado Valley 269 


Macrotus californicus Baird 
California Leaf-nosed Bat 
One of the rooms of an adobe ruin on the Arizona side, ten miles 
below Cibola, showed considerable bat excrement on the floor. No bats 
could be found in the thatch above, so the place was visited in the 
evening. No bats appeared in the vicinity until all daylight had faded. 
Then two were seen flying about the ruins; and by means of a lantern 
and butterfly net, one was caught in the room referred to. This was 
probably used by the bats merely to repair to while eating the moths 
caught outside; wings of the latter were strewn upon the floor. 
The specimen obtained (no. 10693) is an adult male; the date 
April 8. 


Transmitted June 13, 1918. 


LITERATURE CITED 


ADAMS, C. C. 
1905.. The postglacial dispersal of the North American biota. Biol. Bull., 
9, 53-71. 
AMERICAN ORNITHOLOGISTS’ UNION CommMiTTER, J. A. ALLEN, Chairman and 
Editor. 
1910. Cheek-list of North American birds. Ed. 38, revised (New York, 
American Ornithologists’ Union), 4380 pp., 2 maps. 
BaILey, V. 
1902. Synopsis of the North American species of Sigmodon. Proe. Biol. 
Soe. Wash., 15, 101-116. 
Bairp, 8. F. 
1861. ‘‘List of Birds colleeted on the Colorado Expedition’’ in ‘‘Zoology*’ 
in Ives, Report upon the Colorado River of the West, pp. 5-6 (of 
Part V). 
BENDIRE, C. E. 
1892. Life-histories of North American birds with special reference to their 
breeding habits and eggs. U. 8S. Nation. Mus., special bulletin, 
1, pp. vili+ 446, pls. 12. 
BRENINGER, G. F. 
1898. Hybridization of flickers. Osprey, 3, 13. 
BREWSTER, W. 
1883. On a collection of birds lately made by Mr. F. Stephens in Arizona. 
Bull. Nutt. Orn. Club, 8, 21-36. 
1887. Three new forms of North Ameriean birds. Auk, 4, 145-149. 
Brown, H. 
1908.- Arizona bird notes. Auk, 20, 43-50. 
1904. The elf owl in California. Condor, 6, 45-47. 
1906. The water turkey and tree ducks near Tucson, Arizona. Auk, 23, 
217, 218. 


270 University of California Publications in Zoology | Vou. 12 


Cooprr, J. G. 

1861. New California animals. Proce. Calif. Acad. Sci., 2, 118-123. 

1868. Some recent additions to the fauna of California. Proe. Calif. Acad. 
Sei., 4, 3-13. 

1869. The naturalist in California. Amer. Nat., 3, 470-481. 

1870a. The fauna of California and its geographical distribution. Proce. 
Calif. Acad. Sei., 4, 61-81. 

1870b. Ornithology of California. Volume I. Land birds. Edited by S. F. 
Baird, from the Manuseript and Notes of J. G. Cooper. xi+592, 
many text figs. 

Cougs, E. 

1866a. From Arizona to the Pacific. Ibis (2), 2, 259-275. 

1866b. List of the birds of Fort Whipple, Arizona; with which are incor- 
porated all other species ascertained to inhabit the Territory; with 
brief critical and field notes, descriptions of new species, ete. 
Proce. Acad. Nat. Sci., Phila., 1866, 39-100. 

1903. Key to North American birds. Fifth edition, 1, xli+1—535 pp., 2, 
vi+537-1152 pp. 

Court, E. J. 

1908. Treganza Blue Heron. Auk, 25, 291-296. 
Daccert, F. S. 

1902. Winter observations on the Colorado desert. Condor, 4, 37-39. 
Drxon, J. 

1910. A northern coast form of the California gray fox. Univ. Calif. Publ. 
Zool., 5, 308-305. 

Dosson, G. E. 

1886. Description of a new species of Vesperugo from North America. 

Ann. Mag. Nat. Hist., (5), 18, 124-125. 
Dwicut, J., JR. 

1900. The sequence of plumages and moults of the passerine birds of New 

York. Ann. N. Y. Acad. Sci., 13, 73-360, pls. 1-7. 
Euuiot, D. G. 

1905. A check-list of mammals of the North American continent, the West 
Indies and the neighboring seas. Field Columbian Mus., Zool. Ser., 
6, v + 761 pp., frontispiece. 

GILBERT, C, H., and Scorre.p, N. B. 
1898. Notes on a collection of fishes from the Colorado basin in Arizona. 
Proc. U. 8. Nation. Mus., 20, 487-499, pls. 36-39. 
GILMAN, M. F. 
1909. Nesting notes on the Lucy warbler. Condor, 11, 166-168. 
GOLDMAN, E, A. 

1910. Revision of the wood rats of the genus Neotoma. U.S. Dept. Agric., 
Bur. Biol. Surv., North Amer. Fauna, 31, 124 pp., 8 pls., 14 figs. 
in text. 

GRINNELL, J. 

1901. The Pacifie Coast yellowthroats. Condor, 3, 65, 66. 

1904, The ashy kinglet. Condor, 6, 25. 

1909a. Three new song sparrows from California. Univ. Calif. Publ. Zool., 
5, 265-269. 

1909b. A new cowbird of the genus Molothrus, with a note on the probable 
genetic relationships of the North American forms. Univ. Calif. 
Publ. Zool., 5, 275-281, 1 fig. in text. 


1914] Grinnell: Mammals and Birds of the Colorado Valley 271 


1910a. Two heretofore unnamed wrens of the genus Yhryomanes. Univ. 
Calif. Publ. Zool., 5, 307-309. 

1910b. Birds of the 1908 Alexander Alaska expedition, with a note on the 
avifaunal relationships of the Prince William Sound district. 
Univ. Calif. Publ. Zool., 5, 361-428, pls. 32-34. 

191la. Description of a new spotted towhee from the Great Basin. Univ. 
Calif. Publ. Zool., 7, 309-311. 

1911b. A new blue grosbeak from California. Proe. Biol. Soe. Wash., 24, 
163, 164. 

1911e. The linnet of the Hawaiian Islands: a problem in speciation. Univ. 
Calif. Publ. Zool., 7, 179-195. 

1912. The two pocket gophers of the region contiguous to the lower Colo- 
rado river, in California and Arizona. Univ. Calif. Publ. Zool., 
10, 171-178, pl. 5. 

GRINNELL, J., and Swarru, H. 8. 

1913. An account of the birds and mammals of the San Jacinto area of 
southern California, with remarks upon the behavior of geographic 
races on the margins of their habitats. Univ. Calif. Publ. Zool., 
10, 197-406, pls. 6-10. 

HENSHAW, H. W. 

1884. Description of a new song sparrow from the southern border of the 

United States. Auk, 1, 228, 224. 
HOouuister, N. 

1909. Two new bats from the southwestern United States. Proce. Biol. 

Soc. Wash., 22, 43, 44. 
Howe, A. H. 

1906. Revision of the skunks of the genus Spilogale. U.S. Dept. Agric., 

Bur. Biol. Surv., North Amer. Fauna, 26, 55 pp., 10 pls. 
MacDoueat, D. T. 

1908. Botanical features of North American deserts. Carnegie Inst. Wash., 

99, 1-111, 62 pls. 


Mearns, E. A. 

1892. <A study of the sparrow hawks (subgenus Tinnunculus) of America, 
with especial reference to the continental species (Falco sparverius 
Linn.). Auk, 9, 252-270. 

1896. Preliminary diagnoses of new mammals from the Mexican border of 
the United States. Proc. U. S. Nation. Mus., 18, 443-447. 

1897. Preliminary diagnoses of new mammals of the genera Lynaz, Urocyon, 
Spilogale, and Mephitis, from the Mexican boundary line. Proe. 
U. S. Nation. Mus., 20, 457-461. 

1907. Mammals of the Mexican boundary of the United States: a descrip- 
tive catalogue 01 the species of mammals occurring in that region; 
with a general summary of the natural history, and a list of trees. 
U.S. Nation. Mus. Bull., 56, Part 1; xv +530 pp., 13 pls., 126 figs. 
in text. 

Merriam, C. H. 

1892. The geographic distribution of life in North America with special 
reference to the mammalia. Proce. Biol. Soe. Wash., 7, 1-64, map. 

1894. Laws of temperature control of the geographic distribution of terres- 
trial animals and plants. Nation. Geog. Mag., 6, 229-238, 3 pls. 


(maps). 


272 University of California Publications in Zoology [ Vou. 12 


1899. Results of a biological survey of Mount Shasta, northern California. 
U. S. Dept. Agrie., Div. Biol. Surv., North Amer. Fauna, 16, 
1-179, 5 pls., 46 figs. in text. 

1900. Descriptions of two new mammals from California. Proc. Biol. Soe. 
Wash., 13, 151, 152. 

1901. Preliminary revision of the pumas (Felis concolor group). Proe. 
Wash. Acad. Sei., 3, 577-600. 

1903. Eight new mammals from the United States. Proc. Biol. Soc. Wash., 
16, 73-78. 

Minter, G. 8., JR. 

1897. Revision of the North American bats of the family Vespertilionidae. 
U. 8. Dept. Agrice., Div. Biol. Surv., North Amer. Fauna, 13, 140 
pp.-, 3 pls., 40 figs. in text. 

1907. The families and genera of bats. U.S. Nation. Mus. Bull., 57, pp. 
Xvii + 282, 14 pls., 49 figs. in text. 

Morcom, G. F. 

1887. Notes on the birds of soutnern California and southwestern Arizona. 

The Ridgway Ornithological Club Bull., 2, 36-57. 


NE.son, E. W. 
1900. Deseription of a new subspecies of Meleagris gallopavo and proposed 
changes in the nomenclature of certain North American birds. 
Auk, 17, 120-126. 


NEWELL, F. H. 


1904. Second annual report of the reclamation service, 1902-8. U.S. House 


Document no. 44, vol. 57 (Washington, Government Printing 
Office), pp. 550, pls. 69, figs. 56. 


OBERHOLSER, H. C. 

1898. <A revision of the wrens of the genus Thryomanes Sclater. Proce. 
U.S. Nation. Mus., 21, 421-449. 

1904. A revision of the American great horned owls. Proce. U. 8. Nation. 
Mus., 27, 177-192. 

1911. A revision of the forms of the ladder-backed woodpecker (Dryobates 
scalaris [Wagler]). Proe. U. S. Nation. Mus., 41, 139-159, pl. 12. 

1912. A revision of the forms of the great blue heron (Ardea herodias 
Linnaeus). Proce. U. S. Nation. Mus., 43, 531-559. 


Oscoop, W. H. 
1900. Revision of the pocket mice of the genus Perognathus. U. S. Dept. 
Agrie., Div. Biol. Surv., North Amer. Fauna, 18, 74 pp., 4 pls., 
15 figs. in text. 
1907. Four new pocket mice. Proce. Biol. Soe. Wash., 20, pp. 19, 20. 
1909. Revision of the mice of the American genus Peromyscus. U.S. Dept. 
Agriec., Bur. Biol. Surv., North Amer. Fauna, 28, 285 pp., 8 pls., 
12 figs. in text. 
Rimveway, R. 
1887. A manual of North American birds. (Philadelphia, J. B. Lippincott 
Company), xi+631 pp., frontispiece, 124 pis. 
1901. The birds of North and Middle America. U. 8. Nation. Mus. Bull., 
50, part I, xxx+715 pp., 20 pls. 
1902. Idem., part Il, xx +8384 pp., 22 pls. 
1904. Idem., part III, xx +801 pp., 19 pls. 


1914] Grinnell: Mammals and Birds of the Colorado Valley PAB} 


RuTHVEN, A. G. 

1907. A collection of reptiles and amphibians from southern New Mexico 
and Arizona. Bull. Amer. Mus. Nat. Hist., 23, pp. 483-6038, 22 figs. 
in text. 

SpaLpine, V. M. 

1909, Distribution and movements of desert plants. Carnegie Inst. Wash., 

113, 144 pp., 31 pls. 
STEPHENS, F. 

1900. Deseriptions of two new mammals from southern California. Proe. 
Biol. Soe. Wash., 13, 158. 

1903. Bird notes from eastern California and western Arizona. Condor, 5, 
75-78, 100-105. 

1906. California mammals (San Diego, West Coast Publishing Co.), 351 
pp-, frontispiece, many figs. in text. 

Stone, W., and Ryoaps, 8. N. 

1905. On a eolleetion of birds and mammals from the Colorado Delta, Lower 

California. Proe. Acad. Nat. Sei. Phila., 1905, 676-690. 
Swartu, H. 8. 

1905. Summer birds of the Papago Indian Reservation and of the Santa 
Rita Mountains, Arizona. Condor, 7, 22-28, 47-50, 77-81. 

1910. Two new owls from Arizona, with description of the juvenal plumage 
of Strix occidentalis occidentalis (Xantus). Univ. Calif. Publ. 
Zool., 7, 1. 

1912. Report on a collection of birds and mammals from Vancouver 
Island. Univ. Calif. Publ. Zool., 10, 1-124, pls. 1-4. 

TROTTER, S. 
1912. The faunal divisions of eastern North America in relation to vegeta- 
tion. Journ. Acad. Nat. Sei. Phila. (2), 15, 207-218. 
Van RosseM, A. 
1911. Winter birds of the Salton Sea region. Condor, 13, 129-137. 
WoopDHOUSE, 8S. W. 

1853. ‘‘Birds’’ in Sitgreaves, ‘‘Report of an expedition down the Zuni 

and Colorado rivers,’’ pp. 58-105, pls. 1-6. 


PLATE 3 


Map of Needles-to-Yuma portion of lower Colorado River, showing stations 
from which collecting was carried on by the 1910 expedition from the California 
Museum of Vertebrate Zoology. (See itinerary, pp. 53-57.) 


[274] 


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Fig. 2. Looking up the Colorado River from Mellen, Arizona. At this date, 
February 27, 1910, the river was at a low stage, leaving many mud bars 
uncovered. These bars were resorted to as forage grounds by herons and 
ravens. ; 

Fig. 8. California shore near Pilot Knob, showing dense mass of cane 
(Phragmites communis), partly submerged, and in part hanging over the bank 
into the water. The log held fast by the tangle of cane was the favorite resort 
of muskrats (Ondatra zibethica pallida), two of these animals being caught 
at this particular place. This cane is a conspicuous riparian element on per- 
manent banks from the vicinity of Picacho to the Mexican line. Resident 
birds showing marked preference for these cane thickets were Melospiza melodia 
saltonis and Geothlypis trichas seirpicola. Photograph taken May 15, 1910. 


[276] 


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Fig. 4. Looking north over the flood plain of the Chemehuevis Valley, 
California side. At this date (March 10, 1910) only the eottonwoods had eome 
into leaf. This is the typical willow-cottonwood association of the riparian 
belt as found in all the broad valleys. The component plants are willows, of 
two species, cottonwood, guatemote, and screw bean. Some of the latter show 
in the foreground because of the mistletoe clumps in their as yet leafless 
branches. At the time of taking this picture birds of the winter visitant 
eategory were plentiful (see text, p. 71). 


Vig. 5. The arrow-weed association, the typical element in which is the 
arrow-weed (Pluchea sericea). This plant forms an almost continuous growth 
over the river flood plain outwardly adjacent to the willow-cottonwood asso- 
ciation, Animal life was poorly represented in this association as compared 
with any other of the region, Those birds and mammals found therein 
appeared nearly all to traverse it only incidentally, in passing between the 
mesquite and willow associations. Photograph taken near Pilot Knob, May 
11, 1910, 


[278] 


[GRINNELL] PLATE 5 


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PLATE 6 


Fig. 6. Portions of quail-brush (in foreground) and mesquite associations, 
paralleling each other and situated next outwardly from the arrow-weed asso- 
ciation. Photograph taken on the Arizona side about one mile above Mellen, 
February 27, 1910. On this date the deciduous mesquites were still leafless, the 
dark patches being masses of the mistletoe (Phoradendron californicum). The 
latter parasitic plant produces an almost perennial and abundant crop of 
berries which form a staple food supply for many species of birds, notably 
the phainopepla, western bluebird, western robin, and mockingbird. The 
quail-brush (Atriplex lentiformis), because of its stoutly interlacing and spiny 
branches, forms an ideal refuge for such animals as the cottontail rabbit and 
desert quail. Abert towhees are permanent inhabitants of this belt as well 
as of the adjacent one on each side, while the winter-visiting Zonotrichias 
make it their headquarters. 


Fig. 7. Mesquite (Prosopis juliflora) in full leaf and fruit, the latter the 
bean-like pods. The mesquite marks a distinct association, the outermost one 
of the riparian set of associations. Both the foliage and the fruit constitute 
important food sourees for many of the animals of the region, either directly 
or indirectly. Among birds, characteristic permanent residents are the crissal 
thrasher and Abert towhee. The Colorado river wood rat (Neotoma albigula 
venusta) is a characteristic mammal. Photograph taken near Pilot Knob, May 
11, 1910. 


[280] 


UNIV, CALIF. PUBL. ZOOL. VOL, 12 [GRINNELL] PLATE 6 


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Fig. 8. The salt-bush association, on second-bottom above the reach of 
the highest overflow. The large plant in the center of the picture is the 
creosote bush (Larrea divaricata) which in places invades the second bottom 
nearly or quite to the edge of the mesquite and grows to larger size in such 
places than on the desert mesa. The prevailing low, light-colored shrub, is 
the salt-bush (Atriplex polycarpa). Winter visiting birds of this association 
were: Nevada sage sparrow, Brewer sparrow and desert Bewick wren; mammals 
caught at this point were Dipodomys merriami and Perognathus penicillatus. 
Photograph taken one mile above Mellen, Arizona, February 27, 1910. 


Fig. 9. Typical wash association, the catelaw (Acacia greggii) being the 
plant most constantly present. Thickets of catclaw are to be seen in the right 
foreground, while large ironwood and palo verde trees are to be seen in the 
middle distance. The distant hill slopes are dotted with creosote bushes, while 
Atriplea polycarpa margins the wash in the immediate foreground. Resident 
birds of this wash association were: verdin, plumbeous gnateatcher, and cactus 
wren. Photograph taken March 10, 1910, on California side, near lower end 
of Chemehuevis Valley. 


PLATE 8 


Fig. 10. Ironwood tree (Olneya tesota) photographed March 10, 1910, in the 
wash pictured in the previous plate. This individual, an unusually large one, 
was 90 inches in circumference of trunk two feet above the ground, 31 feet 
in extreme height, and with a foliage expanse of 50 feet. The thorny branches 
afforded protection to several nests, old and new, of the verdin. The blossoms 
of this plant, which appear in May, attract numerous hummingbirds. 


Fig. 11. Palo verde tree: (Parkinsonia torreyana) of unusual size. At the 
time of blossoming, in April, this tree is resorted to by many migrating birds, 
both for the flower nectar and the insects. Like other elements in the wash 
association the palo verde is frequented by verdins, plumbeous gnatcatchers 
and cactus wrens. Photograph taken February 27, 1910, on the Arizona side 
above Mellen, near the mouth of the Sacramento wash. 


UNIV, CALIF, PUBL ZOOL. VOI 2 


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PLATE 9 


Fig. 12. Giant cactus (Cereus giganteus) on California side four miles 
north of Potholes. A palo verde stands immediately beyond, its trunk being 
nearly hidden by that of the cactus. The extreme height of the latter was 
28 feet. Openings may be seen in the upper branches. Two of these were 
inhabited by a pair each, respectively, of the Gila woodpecker and ash-throated 
flycatcher. Photograph taken April 28, 1910. 


Fig. 13. Giant cactus on California side four miles above Potholes, photo- 
graphed April 28, 1910. In a hole in this cactus was’ found a brood of gilded 
flickers (just beneath short left-hand branch), and three feet higher up in a 
cavity opening on the opposite side of the trunk was a saguaro screech owl. 


- [286] 


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PLATE 10 


Fig. 14. Looking due south from Mellen, Arizona, and toward the group 
of spire-pointed hills known as ‘‘The Needles.’’? The Colorado River in the 
right distance. Typical rocky mesa in the immediate foreground, the scattering 
plants being creosote bushes. The desert mesa is here seen to abut closely upon 
the river, leaving only very narrow riparian strips. Mammals trapped on the 
mesa at this point were Perognathus intermedius and Ammospermophilus harrisi. 
Photograph taken February 28, 1910. 


Fig. 15. Photograph taken March 7, 1910, on the Arizona side, from upper 
slope of The Needles. Channel of the Colorado River at extreme left. The 
chief vegetation on the steep rocky slopes is the creosote bush and Encelia 
farinosa. The latter reappears so persistently upon such ground that its name 
has been selected to apply to the association marked by its presence. Mammals 
trapped on this slope were Neotoma intermedia desertorum, Perognathus inter- 
medius and Ammospermophilus harrisi. 


[288] 


UNIV, CALIF. PUBL. ZOOL. VOL. 12 [GRINNELL] PLATE 10 


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Fig. 16. Group of burrows of the large kangaroo rat (Dipodomys deserti). 
Tracks of the animals may be seen in the soft aeolian sand, here accumulated 
to considerable depth. The dessicated remains of a brief-lived annual vegeta- 
tion may be seen on the sand between the creosote and the salt-bushes. Other 
species of mammals trapped in this, a variation of the salt-bush association, 
were: Citellus tereticaudus, Peromyscus eremicus, and Perognathus penicillatus. 
Photograph taken about one mile north of Mellen, Arizona, February 27, 1910. 


Fig. 17. Mouth of burrow of the large kangaroo rat (Dipodomys deserti), 
showing parallel imprints of the hind feet and the tail in the soft sand. Photo- 
graph taken above Mellen, Arizona, February 27, 1910. 


[290] 


UNIV. CALIF. PUBL. ZOOL. VOL. 12 (GRINNELL] PLATE II 


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Fig. 18. Burrow of the Harris ground squirrel (Ammospermophilus harrisi) 
beneath creosote bush on desert mesa near Mellen, Arizona. The wind-worn 
pebbles of the mesa surface are here well shown, the loose sand being con- 
tinually removed by the prevailing winds. Photograph taken February 28, 
1910. 


Fig. 19. Ironwood tree almost completely killed by the rising of the water 
level in the soil at the outer edge of second bottom. A nest of the Lucy 
warbler (Vermivora luciae) was situated 35 inches from the ground in a cavity 
in the side of the trunk. Photograph taken April 12, 1910, near the Draper 
ranch, on the California side eighteen miles north of Picacho. 


a td 


PLATE 13 


Fig. 20. Nest of the Lucy warbler (Vermivora luciae) in crevice on side of 
trunk of partly dead ironwood shown in plate 12, figure 19. This nest contained 
three eggs. Photographed April 12, 1910, on the California side, eighteen miles 
above Picacho. 


Fig. 21. Selected specimens of Mephitis estor from the Colorado Valley: 
at left, no. 10575, Chemehuevis Valley, California side; middle, no. 10574, five 
miles south of Needles, California side; at right, no. 10579, twenty miles above 
Picacho, California side. The variation shown is individual. If the contrasted 
black and white markings are of warning signficance, hence of adaptive advan- 
tage to the species, why should the desert skunks have proportionally much 
more white than skunks from the humid northwest coast belt of the United 
States? (See text, p. 256.) 


[294] 


‘SUT 


CG 


Ad ‘SI1VO ‘AINN 


mks! 


€| alv1d [WaNNIYD] 


’ UNIVERSITY OF CALIFORNIA PUBLICATIONS—(Continued) 


5. On the Skeletal Morphology of Gonyaular catenata (Levander), by 
Charles Atwood Kofoid. Pp. 287-294, plate 18. 
6. Dinoflagellata of the San Diego Region, V. On Spiraulax, a New Genus 
of the Peridinida, by Charles Atwood Kofoid. Pp. 295-300, plate 19, 
Nos. 4, 5, and 6 in one cover. September, 1911 oo.c..0 ccc 
7. Notes on Some Cephalopods in the Collection of the University of Cali- 
fornia, by S. S. Berry. Pp. 301-310, plates 20-21. September, 1911. 
8, On a Self-Closing Plankton Net. for Horizontal Towing, by Charles 
Atwood Kofoid. Pp. 311-348, plates 22-25, 
9. On an Improved Form of Self-closing Water-bucket for Plankton In- 
vestigations, by Charles Atwood Kofoid. Pp. 349-352. 
Nos. 8 and 9 in one cover. Nevember, 1911 20002020 
Index, pp. 353-357. 


1, The Horned Lizards of California and Nevada of the Genera Phryno- 
oe and Anota, by Harold C. Bryant. Pp, 1-84, plates 1-9. Decem- 
PSR Gk BEN Rata SOS Be A Sik AE AACE ad OS, MN ae Mea RES ISS Oe ha 
On a Lymphoid Structure Lying Over the Myelencephalon of Lepisos- 
teus, by Asa C. Chandler. Pp. 85-104, plates 10-12. December, 1911. 
3. Studies on Early Stages of Development in Rats and Mice, No. 3, by 
E. L. Mark and J. A. Long. The Living Eggs of Rats and Mice with 
a Description of Apparatus for Obtaining and Observing Them (Pre- 
liminary paper), by J. A. Long. Pp. 105-136, plates 13-17, February, 
1912 


to 


» 


. The Marine Biological Station of San Diego, Its History, Present Con- 
ditions, Achievements, and Aims, by Wm..E. Ritter, Pp. 137-248, 
plates 18-24, and 2 maps. March, 1902 22... c.ccecccel i ccccecteeenesineneceenrene 

5. Oxygen and Polarity in Tubularia, by Harry Beal Torrey. Pp. 249- 

DUE MAS EOS ee aR a kh Anta ts aaa ca ean 

6. The Occurrence and Veriical Distribution of the Copepoda of the San 

Diego Region, with particular reference to Nineteen Species, by Cal- 
vin O. Esterly, Pp. 253-340, 7 text-figures.. July, 1912.02... 
7. Observations on the Suckling Period in the Guinea-Pig, by J. Marion 
Read. “Pp. 341-351. “September, 1912: 23.2222 cn 

8. Haeckel’s Sethocephalus eucecryphalus (Radiolaria), a Marine Ciliate, 

by Charles Atwood Kofoid. Pp. 353-357. September, 1912 —.......... 
Index, pp. 359-365. 


(Contributions from the Museum of Vertebrate Zoology.) 


1, Report on a Collection of Birds and Mammals from Vancouver Island, 
by Harry 8S. Swarth. Pp. 1-124, plates 1-4. February, 1912 ........... 
2. A New Cony from the Vicinity of Mount Whitney, by Joseph Grinnell. 
yy get 217 Biase Fo 8 a fata I i 7 pie eo as np OSE cn Sr a ap 
8. The Mole of ‘Southern California, by J. Grinnell and H. S. Swarth. 
Pp. 131-136, 2 text-figures. 
4, Myotis orinomus Elliott, a Bat New to California, by J. Grinnell and 
H. S, Swarth. Pp. 137-142, 2 text-figures. 
Nos. 3 and 4 in one cover. April, 1912 2... eeccel cise ccceceesteeeeenneeene 
5. The Bighorn of the Sierra apy adie by Joseph Grinnell. Pp. 143-153, 
A CORETBUT ORV SIMAY, 20S Dooce aaah sca ah tes wn eaten dung tnedectnostontencsecaw 
6. A New Perognathus from the San Joaquin Valley, California, by 
Walter P. Taylor. Pp. 155-166, 1 text-figure. 
7. The Beaver of West Central California, by Walter P. Taylor. Pp. 
167-169. 
: Nos. 6 and 7 in one cover, May, 1912-200... 2 ccc ceee eee ecentecnnecen eee 
8. The Two Pocket Gophers of the Region Contiguous to the Lower Colo- 
rado River, in California and Arizona, by Joseph Grinnell. Pp, 171- 
bbe 2 aibers Birks este be 9 a2 pens ats ale ee Van, ae eR A SE Sine, eer pacer eee 
9. The Species of the Mammalian Genus Sorex of West-Central Cali- 
fornia, with a note on the Vertebrate Palustrine Faunas of the 
Region, by Joseph Grinnell. Pp, 179-195, figs. 1-6. March, 1913 ........ 
10, An Account of the Birds and Mammals of the San Jacinto Area, of 
Southern California, with Remarks Upon the Behavior of Geographic 
~ Races on the Margins of Their Habitats, by J. Grinnell and H. 8. 
Swarth. Pp. 197-406, pls. 6-10. October, 1913 2.2... occa eeenenenaneee 
Index, pp. 407-417. 


1.50 
10 


40 


70 


1,00 
05 


1.00 
-10 
05 


1,00 


10 


15 


15 


15 


UNIVERSITY OF CALIFORNIA PUBLICATIONS— (Continued) , 


Vol. 11.° 1. Birds in Relation to a Grasshopper Outbreak in California, by Harold - 

Cc. Bryant. -Pp. 1-20. November, 1912 2.0.20. ecto 

2. On the Structure and Relationships of Dinosphaera palustris (Lhemm.), 

by Charles Atwood Kofoid and Josephine Rigden Michener. Pp. 21. cio e 

23°; Decomper, 21972 er Se he ee eh ae Cee Bh 

8. A Study of Epithelioma Contagiosum of the Common Fowl, by taean 

Clifford D. Sweet. Pp. 29-51. January, BAS IK Sete ainaletanehiee tag ripe 'i nom saa 3 i 

4. The Control of Pigment Formation in Amphibian Larvae, by Myrtle — 
B. Johnson. Pp. 53-88, plate 1. March, 1918 2... .cc..-2 cies eenceeceeeee 

5. Sagitta californica, u.sp., fromthe San Diego Region, including 

Remarks on Its Variation and Distribution, by Ellis L. Michael. 

Pp. 89-126, plate, 2. Sune OTS io ike es ae Ae ie _ 

6. Pycnogonida from the Coast of California, with Description of Two. 2 

New Species, by H. V. M, Hall. Pp. 127-142, plates 3-4. August, 1913. _ 

7. Observations on Isolated Living Pigment Cells from the Larvae of 

Amphibians. by S. J. Holmes, Pp. 143-154, plates 5-6. - 

8. Behavior of Ectodermic Epithelium of Tadpoles when Cultivated in * 

Plasma, by S. J. Holmes. Pp. 155-172, plates 7-8. ; 
Nos. 7 and 8 in one cover, September, 5 22S Se = OUR Ro ATE ie ‘i 

9. On Some Californian Schizopoda, by H. J. Hansen. Pp: 173-180, pi 8 9. aut 

NOVEM Der, LOLS eS as yap asco te ecco nwt Caper Oa iN gi a ee be ce : 

10. Fourth Zaxonomic Report on the Copepoda of the San Diego ‘Region, ~ 

by Calvin O. Esterly. Pp. 181-196, pls. 10-12. November, 1913 -.... 4 

11. The Behavior of Leeches with Especial Reference to Its Modifiability, 

A.The General Reactions of the Leeches Dina microstoma Moore and — 

Glossiphonia stagnalis Linnaeus; B,:Modifiability in the Behavior of 

the Leech Dina microstoma Moore, by Wilson Gee. Pp. 197- 305, 13. 

text fenres: “December, 1913 ir ae Reese ek, ree eee 

12. The Structure of the Ocelli of Polyorchis penicillata, by Etta Viola 

Little. -Pp..307-328, plates 13-15. February, 1914 2c. a a 

18. Modifications and Adaptations to Functions in the Feathers of. Circus 

hudsonius, by Asa C. Chandler. Pp. 329-376, plates 16-20, March, ~ 

BV: pet ia aap eins nee soar ne EL an My JM Lie ED Ria nck ee Seek SS 

14. A Determination of the Economic Status of the Western Meadowlark 

(Sturnella negjectay in California, by Harold Child Bryant. Pp. 377-_ 

510, plates 21-24, 5 text figures. February, 1914 ..2.80 coe 5 


Vol. 12. 1. A Study of a Collection of Geese of the Branta canadensis Group from ~ 
the San Joaquin Valley, California, by Harry 8. Swarth.: Pp. 1 ee 
plates 1-2, 8 text figs. November, 19130-20222. -....ccicecc. ice ecctecelidenceccteee ‘ 

2. Nocturnal Wanderings of the California Pocket Gopher, by Harold U2 
Bryant. Pp. 25-29, 1 text fig. November, 1913 2.0.00. 
3. The Reptiles-of the San Jacinto Area of Southern California, by Sarah 
Rogers Atsatt.. Pp. 31-50.- November, 1913 2.222.020 A 
4,-An Account of the Mammals and Birds of the Lower Colorado Valley, 
with Especial Reference to the Distributional Problems Presented, - 
by Joseph Grinnell. Pp. 51-294, plates 3-13, 9 text figs, March, 1914. 2.40 


te te es ee 


ee I no ee ee ee 


a 


a 5S 


UNIVERSITY OF CALIFORNIA PUBLICATIONS 
1N 


ZOOLOGY 
Vol. 12, Nos. 5 and 6, pp. 295-300 April 15, 1914 


APLODONTIA CHRYSEOLA, A NEW MOUNTAIN 
BEAVER FROM THE TRINITY REGION 
OF NORTHERN CALIFORNIA 


BY 
LOUISE KELLOGG 


A PREVIOUSLY UNDESCRIBED APLODONTIA 
FROM THE MIDDLE NORTH COAST 
OF CALIFORNIA 


BY 
WALTER P. TAYLOR 


1914 


UNIVERSITY OF CALIFORNIA PRESS (2° MAY @ 
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Vol. 7. (Contributions from the Museum of Vertebrate. Zoology.) 
1. Two New Owls from Arizona, with Description of the Juvenal Plum- 
age of Strix occidentalis occidentalis (Xantus), by Harry S. Swarth. 
Bp eB Se Mays LOL ee a cy ed ees Nee ah 
2. Birds and Mammals of the 1909 Alexander Alaska Expedition; by. 
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of the Region, by Walter P. Taylor. Pp. 319-436, Suge 7-12, 
By Neha tt tx page Op Bex ania pe a Re tele AS SES Uap ac an HR emda a ae Sb a 
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Vol, 8-1, The Vértical Distribution of Hucalanus Hon iuias: in the San Diego 

Region during 1909, by Calvin O. Esterly. Pp. 1-7... May, 1911 .. 

2. New and Rare Fishes from Southern California, by Edwin Chapin - 
Starks and William M. Mann. Pp. 9-19, 2 text-figures. July, 1911. 

8. Classification and Vertical Distribution of the Chaetognatha of the San 
Diego Region, Including Redescriptions of Some Doubtful Species of 
the Group, by Ellis I. Michael. Pp. 21-186, pls. 1-8. December, 1911.- 

4, Dinofiagellata of the San Diego Region, IV. The Genus Gonyaulaz, with 
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and Specific Characters, by Charles Atwocd mptsies Pp. 187-286, 
plates 9-17. 


UNIVERSITY OF CALIFORNIA PUBLICATIONS 
IN 
ZOOLOGY 
Vol. 12, No. 5, pp. 295-296 April 15, 1914 


APLODONTIA CHRYSEOLA, A NEW MOUNTAIN 
BEAVER FROM THE TRINITY REGION 
OF NORTHERN CALIFORNIA 


BY 


LOUISE KELLOGG 
(Contribution from the Museum of Vertebrate Zoology of the University of California) 


Discovery of the well-marked new form of mountain beaver here 
described was one of the results of field work carried on in the Trinity 
region of northern California by Miss Annie M. Alexander and the 
writer during the summer of 1911. 


Aplodontia chryseola, new species 


Trinity Mountain Beaver 

Type: Male adult, no. 13328, Mus. Vert. Zool.; Jackson Lake, 
Siskiyou County, California, altitude 5900 feet; June 22, 1911; col- 
lected by A. M. Alexander; original number 1441. 

Spreciric CHARACTERS: Coloration, both above and below, varying 
about ochraceous-buff or buffy golden; rostrum short as compared 
with that of Aplodontia californica (Peters) ; auditory tubes long in 
proportion to size of skull. 

Mareriau: The basis for this diagnosis consists of nine specimens, 
nos. 13324-13332, taken at the following localities, all in that portion 
of the Trinity Mountain region known as the Salmon Mountains: 
Jackson Lake; South Fork of Salmon River; Wildeat Creek; head of 
Grizzly Creek. 

Remarks: In size this new species of Aplodontia is nearest A. 
californica (Peters) (—A. major Merriam), the form occupying the 


bal 


s 


MAY 6 IO]a 


eee 
Agaoriiali lisae~ 
Woes t tyre 


ty 


~ 


G, 


296 University of California Publications in Zoology ‘Vou. 12 


Sierra Nevada Mountains, but it differs materially from the latter 
both in certain cranial characters and in its golden coloration. In the 
latter respect, however, it closely resembles the small Aplodontia phaea 
Merriam, found in Marin County, California. Specimens taken on 
the slopes of Mount Shasta seem to be typical of A. californica, show- 
ing no gradation towards A. chryseola. 

The general coloration of Aplodontia chryseola, on the back, is 
fulvous or ochraceous, thickly sprinkled with black. The dense under- 
fur is black at the base in fresh pelage, wearing to slate, and tipped 
with ochraceous-buff. The long hair is a mixture of gleaming 
ochraceous-buff and black, giving a pecuhar bright effect, of gold and 
black. The sides are ochraceous-buff with fewer black hairs. The 
underparts are plumbeous, wearing to lighter gray, and _ heavily 
sprinkled with ochraceous-buff, especially around the throat and 
cheeks. The tip of the nose is seal brown, face drab, whiskers mainly 
white. 

A further discussion of the Trinity mountain beaver, as well as an 
account of its habits, is contained in the writer’s paper on the mam- 
mals of the Trinity Mountain region now in preparation. In order 
to facilitate further work on the Aplodontiidae, in progress both in 
the Department of Palaeontology and in the Museum of Vertebrate 
Zoology, of the University of California, immediate publication of 
this description is deemed advisable. 


Transmitted January 24, 1914. 


UNIVERSITY OF CALIFORNIA PUBLICATIONS 
IN 
ZOOLOGY 
Vol. 12, No. 6, pp. 297-300 April 15, 1914 


A PREVIOUSLY UNDESCRIBED APLODONTIA 
FROM THE MIDDLE NORTH COAST 
OF CALIFORNIA 


BY 


WALTER P. TAYLOR 
(Contribution from the Museum of Vertebrate Zoology of the University of California) 


The recent discovery by field parties working in the interests of 
the California Museum of Vertebrate Zoology, of two new forms of 
Aplodontia, is a testimony both to the restricted habitats of the animals 
in this genus and to the incompleteness of our knowledge of even 
so small and well-worked an area as the State of California. Addi- 
tional attention is directed to these points by the fact that one of the 
new forms is the most strikingly marked species yet found within the 
state. 

Material in the collection of the Museum indicates the existence 
in the vicinity of Humboldt Bay, California, of a third distinct new 
form, most closely related to Aplodontia chryseola Kellogg, of the 
Trinity Mountains. The status of this form cannot certainly be de- 
termined, however, without more specimens. 


Aplodontia nigra, new species 
Point Arena Mountain Beaver 


Type: Male adult, no. 20320, Mus. Vert. Zool.; Point Arena, 
Mendocino County, California; July 10, 1913; collected by C. L. 
Camp; original number 1003. 

DiaGNostic CHaracters: Most closely related to Aplodontia phaea 
Merriam, but nasal outline swelling at the sides anteriorly, the broad- 
est portion of the nasals tending to be just posterior to their anterior 


298 University of California Publications in Zoology ‘Vou. 12 


ends, contrary to the condition in phaea; nasals uniformly broader 
than in phaea; interpterygoid fossa broader. Coloration dorsally 
shiny black instead of ochraceous-buff; ventrally warm buff instead 
of ochraceous-buff or light ochraceous-buff (Ridgway’s Color 
Standards and Nomenclature, 1912, used as color guide). 

Marertat: Twenty specimens of Aplodontia phaea; four speci- 
mens of Aplodontia mgra. It should be noted that both series, with 
the exception of two specimens of A. phaea (nos. 8973, 8974). were 
taken during the summer season, so are strictly comparable as to 
pelage. 

DescripTIVvE Remarks: Coloration.—The black-and-gray dorsal 
appearance of Aplodontia nigra is unique among known California 
aplodonts. This character marks the new species off sharply from A. 
phaea, to which its size and cranial characters show it to be most 
closely related. The prevailing color tone seen in a dorsal view is, in 
nigra, Shiny black; in phaea, ochraceous-buft. The dorsal coloration 
is modified somewhat in nigra by the dark plumbeous bases of the 
hairs showing through. Some of the dorsal hairs are tipped with 
buffy. These are so few, however, that the general dorsal aspect is 
shiny black only faintly sprinkled with grayish. In phaea, the 
ochraceous-buff of the dorsal surface varies toward light ochraceous- 
buff. The general coloration is modified by the showing through of the 
dark plumbeous bases of the hairs and also by the admixture of 
numerous black hairs. There is also a sprinkling of hairs tipped with 
light buff. The result of the mixture of these variously marked hairs 
is a grizzled ochraceous-buff appearance. 

Face in A. nigra is dark quaker drab; near pale quaker drab in 
A. phaea. 

Sides paler than back in A. nigra, there being fewer black hairs, 
and more buffy-tipped ones. In A. phaea the sides are nearly the 
same as the back, grading into the coloration of the underparts. 

There is much more blackish externally on both fore and hind feet 
in A. nigra, as well as on the rump and tail. In A. phaea these parts 
tend to be browner. The feet in A. nigra are blackish, in phaea 
brownish. 

A. nigra has basal portions of hairs ventrally varying between 
plumbeous, deep plumbeous, and dark plumbeous, and upper portions 
of the same hairs warm buff. Phaea has hairs ventrally a paler shade 
on their basal portions, varying between plumbeous and cinereous, 
and their outer portions ochraceous-buff or ight ochraceous-buft. 


1914] Taylor: Aplodontia 299 


Coloration is remarkably uniform in the series of phaeca, showing 
only a very narrow range of individual variation. Of the four speci- 
mens of nigra, three are young. In coloration these differ only slightly 
from the adult, and present the diagnostic characters of the species 
as clearly. 

It should, perhaps, be emphasized that coloration serves unques- 
tionably to differentiate A. nigra from any other species of the same 
genus found in California. 

Cranial characters.—See table of measurements, following. Only 
one specimen of Aplodontia nigra (no. 20320) is strictly comparable 
with the series of four adult male specimens of Aplodontia phaea 
listed in the table. The three other specimens show the open sutures 
and immature characteristics of youth. 

Three characters stand out as specifically distinetive: the width of 
the interpterygoid fossa, and the outline and breadth of the nasal 
bones. A. nigra (no. 20320) has the interpterygoid fossa thirty per- 
cent broader than it is in the average of phaea, at least as shown in the 
table of measurements. 

The outline of the nasals is different. A. nigra has this outline 
dilated anteriorly, the broadest part of the nasals being about six 
millimeters posterior to their anterior ends. In phaea the tendency 
is for the broadest part of the nasals to be at the anterior points at 
which nasals and premaxillae join. A. nigra, no. 20321, approxi- 
mates the condition in phaea, while A. phaea, no. 20309, tends toward 
the relation in nigra. The breadth of the nasals is definitely greater 
in the Point Arena form than in phaea, however, the youngest speci- 
men of the former exceeding in this respect all the adult males of the 
latter measured. 

In other cranial characters A. nigra is nearly identical with A. 
phaca, clearly exhibiting the comparatively close relationship of the 
two coast forms. 

The Point Arena Aplodontia may be separated from any other 
species represented in our collections on the basis of eranial as well 
as external characters. Perhaps the most convenient is the length of 
the incisive foramen, which is less in the erania of the two coast forms 
than in comparable specimens of A. rufa, or in the species heretofore 
deseribed from California. Size also is a differentiative characteristic 
when nigra is compared with rufa or with Californian species. 

It is perhaps worthy of note that the smallest, darkest forms of 
Aplodontia are found along the western coast of the United States. 


300 University of California Publications in Zoology Vou. 12 


Aplodontia pacifica Merriam, described from Newport, mouth of 
Yaquina Bay, Oregon, is the darkest species of mountain beaver known 
heretofore, and is the smallest species as yet characterized. 


Transmitted January 22, 1914. 


TABLE OF CRANIAL MEASUREMENTS OF Aplodontia phaea AND A. nigra 


All measurements in millimeters 


~ 
a3 
2 
ae 
= = = 
<= a r Sei 
) 2 2 be Soe 
a5 a6 2 7) 2o 
Sa = & = =] Se 
ais we a 5 
I =o Sin Nee of 4 Bok 
ae = oe nz H Se 
=F a = 2S ei 3 on 
ae == De Rey Tes = oeg 
Za Sa aie oo EPI Z Sa 
3 aa oa ins oS a 235 
B= mea H's Sie} He faa) ess 


Aplodontia phaea 
20309 § 6 mi. W. Inverness, 
Marin Co., Calif... 23.6 98 415 7.3 42 15.8 59.0 26.8 
8974 g Lagunitas, Marin 
Cot, Calif, Sees Bees CUS ieee 6.2 46 15.3 55.4 27.6 
20311 2 6 mi. W. Inverness, 
Marin Co., Calif... 22.1 98 443 65 44 160 569 28.1 | 
20305 2 6 mi. W. Inverness, 
Marin Co., Calif... 21.8 9:8 45.0) 16:5) 4. A738) 58:55 2956 


Aplodontia nigra 
20320 g Point Arena, Men- 

docino Co., Calif... 23.5 11.0 468 68 56 17.5 569 30.8 
20321 ¢ Point Arena, Men- 

docino Co., Calif... 21.9 10.00 456 55 48 154 52.9 291 
20319 g Point Arena, Men- 

docino Co., Calif... 22.2 10.9 491 5.0 49 15.3 55.0 27:8 
20318 g Point Arena, Men- 

docino Co., Calif... 22.7 10.5 462 5.2 5.7 145 52.9 27.4 


1 Most anterior point on nasal bones to most posterior point. 
2 Greatest width of nasals, across both of them. 


3 With cranium resting on its dorsal surface, rostrum pointing away from 
the worker, the greatest length of the foramen on the right side. 


4 Taken at expansion of interpterygoid fossa immediately back of hard palate. 


5 Most lateral point on foramen ovale to the point farthest laterally (with 
reference to the skull) on zygomatie side of auditory tube. 


Vol, 9. 


UNIVERSITY OF CALIFORNIA PUBLICATIONS—(Continued) 


5, On the Skeletal Morphology of Gonyaulax catenata (Levander), by 
Charles Atwood Kofoid. Pp. 287-294, plate 18. 
6. Dinoflagellata of the San Diego Region, V. On Spiraular, a New Genus 
of the Peridinida, by Charles Atwood Kofoid. Pp. 295-300, plate 19. 
Nos. 4, 5, and 6 in ome cover. September, 1911 200000 
. Notes on Some Cephalopods in the Collection of the University of Cali- 
fornia, by 8. 8S. Berry. Pp. 301-310, plates 20-21, September, 1911. 
. On-@ Self-Closing Plankton Net for Horizontal Towing, by Charles 
Atwood Kofoid. Pp. $11-348, plates 22-25. 
9. On an Improved Form of Self-closing Water-bucket for Plankton In- 
vestigations, by Charles Atwood Kofoid: Pp. 349-352. 
Nos: 8 and 9 in one cover. November, 197) 2... cceclcccccceeee 250 
Index, pp. 353-357. 


on 


1. The Horned Lizards of California and Nevada of the Genera Phryno- 
Ss and Anota, by Harold C. Bryant, Pp. 1-84, plates 1-9, Decem- 

[eS a ew Sieree Serie Roe chic hee ae ie yf A cy, eR ean ORR Ay ca ERAS Ry ee 

2, On a Lymphoid Structure Lying Over the Myelencephalon of Lepisos- 
teus, by Asa C, Chandler. Pp. 85-104, plates 10-12. December, 1911, 

8. Studies on Barly Stages of Development in Rats and Mice, No. 3, by 
E. L. Mark and J. A. Long. The Living Eggs of Rats and Mice with 


4,. The Marine Bi 
ditions, Achievements, and Aims, by Wm. E. Ritter, Pp. 137-248, 
plates 18-24, and 2 maps, March, 1912 occ. ease cceccceceenee Pe 
5, Oxygen and Polarity in Tubularia, by Harry Beal Torrey. Pp. 249- 
Doves Way OAS 10 ow an tai Ney te ed) et COE Me free EO 
6. The Occurrence and Vertical Distribution of the Copepoda of the San 
Diego Region, with particular reference to Nineteen Species, by Cal- 
vin O. Esterly. Pp. 253-340, 7 text-figures. July, 1912 0. 
7, Observations on the Suckling Period in the Guinea-Pig, by J. Marion 
Read. Pp. 841-351. September, 1912-2... eh. ee 
8. Haeckel’s Sethocephalus eucecryphalus (Radiolaria), a Marine Ciliate, 
by Charles Atwood Kofoid. Pp. 353-357. September, 1912 ........... 
Index, pp. 359-365. 


Vol. 10. _ (Contributions from the Museum of Vertebrate Zoology.) 


1. Report on a Collection of Birds and Mammals from Vancouver Island, 
by Harry S. Swarth. Pp. 1-124, plates 1-4. February, 1912 2.0. 
2, A New Cony from the Vicinity of Mount Whitney, by Joseph Grinnell. 
EPs 125-4200 TRNUALY oho ter hoe SE ee ee Oe ae 
8. The Mole of Southern California, by J. Grinnell and H. S, Swarth. 
Pp. 131-136, 2 text-figures. 
4, Myotis orinomus Elliott, a Bat New to California, by J. Grinnell and 
H. S. Swarth. Pp. 137-142, 2 text-figures. 
Nos. 3 and 4 in one cover. ~ April, 1912 22... cece ceeeee eee 
5. The Bighorn of the Sierra Nevada, by Joseph Grinnell. Pp. 143-153, 
AS CORP ONY OS. SIMERY, 240 be eon a ee a epee arene ate 
6. A New Perognathus from the San Joaquin Valley, California, by 
Walter P, Taylor. Pp. 155-166, 1 text-figure. 
7. The Beaver of West Central California, by Walter P. Taylor. Pp. 
167-169. 
Nos. 6 and 7 in one cover. May, 1912 ......ccc.ccce cece cence 
8, The Two Pocket Gophers of the Region Contiguous to the Lower Colo- 
rado River, in California and Arizona, by Joseph Grinnell. Pp, 171- 
SELES pi Uae Oe Bt 2c ooh cant oe ag ease eh erent Ne SS Eh a eae 
9. The Species of the Mammalian Genus Sorex of West-Central Cali- 
fornia, with a note.on the Vertebrate Palustrine Faunas of the 
Region, by Joseph Grinnell. Pp. 179-195, figs. 1-6. March, 1913 ........ 
10. An Account of the Birds and Mammals of the San Jacinto Area of 
Southern California, with Remarks Upon the Behavior of Geographic 
Races on the Margins of Their Habitats, by J. Grinnell and H. 8, 
Swarth. Pp. 197-406, pls. 6-10. October, 1913 .2n..e coe eee 
Index, pp. 407-417. 


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Vol. 18. 


UNIVERSITY OF CALIFORNIA PUBLICATIONS— (Continued) 


1. Birds in Relation to a Grasshopper Outbreak in California, by Harold ~ 


C. Bryant. Pp. 1-20: “November, 1912 2.c.cc.cs cece ibe neces 

2. On the Structure and Relationships of Dinosphaera palustris (Lemm.), 
by Charles Atwood Kofoid and Josephine Rigden Michener. Pp. 21- 
ZS, 32-DOcami Der TO ee eae eat Bt geg eee tometer 
A Study of Epithelioma Contagiosum of the Common Fowl, by 
Clifford D. Sweet. Pp. 29-51. January, 1913 2.222 


om fe 


Remarks. on Its Variation and Distribution, by Ellis L. Michael. 
Pp, 89-126, plate: 2.’ Tike, 4901S. 2 es tk eee ne : 
6. Pycnogonida from the Coast of California, with Description of. Two 
New Species, by H. V. M. Hall. Pp. 127-142, plates 3-4. August, 1913. 
7, Observations on Isolated Living Pigment Cells from the Larvae of 
: Amphibians, by S. J. Holmes. Pp. 143-154, plates 5-6. 
8. Behavior of Ectodermic Epithelium of Tadpoles when Cultivated in 
Plasma, by S. J. Holmes. Pp. 155-172, plates 7-8. 
Nos, 7 and 8 in one cover. September, 1913 -........0.....-. 
9. On Some Californian Schizopoda, by H. J. Hansen, Pp, 173-180, pl. 9. 
November, pak: 3 hy eee ane neem: SR geen ys aL eat ea ee Re are 
10: Fourth Taxonomic Report on the Copepoda of the San Diego Region, 
by Calvin O. Esterly. Pp. 181-196, pls. 10-12. November, 1913 ._.... 
11. The Behavior of Leeches with Especial Reference to Its Modifiability, 
A. The General Reactions of the Leeches Dina microstoma Moore and 
Glossiphonia stagnalis Linnaeus; B. Modifiability in the Behavior of 
“the Leech Dina microstoma Moore, by Wilson Gee. Pp, 197-305, 13 
text figures. December, 1913 
12. The Structure of the.Ocelli of Polyorchis pentcillata, by Etta Viola 


Little. Pp. 307-328, plates 13-15. February, 1914 20... ee 


13. Modgifications and Adaptations to Functions in the Feathers of Circus 
hudsonius, by Asa C, Chandler. Pp. 329-376, plates 16-20. March, 
SANE: See ea RES SS ae ale gl Wom NA Ene, aye eT eben ooeee ni - NES 

14. A Determination of the Economic Status of the Western Meadowlark 


(Sturnella neglecta) in California, by Harold Child Bryant. Pp. 377-. 


“ 510, plates 21-24, 5 text figures, February, 1914 22 
15; Parasynaptic Stages.in the Testis of Aneides lugubris (Hallowell), by 
Harry James Snook and J. A. Long. Pp. 511-528, plates 25-26, 1 text 

fie SADA VOTE eee See chan) Se SS ate Soe eee ge ee ea ws 


1. A Study of a Collection of Geese of the Branta canadensis Group from” 


the San Joaquin Valley, California, by Harry S. Swarth. Pp. 1-24, 
plates 1-2, 8 text figs. November, 1913. .....i...22t tec eee 
Nocturnal Wanderings of the California Pocket Gopher, by Harold C. 
Bryant. . Pp. 25-29, 1 text fig. Nowember, 1913. . 2.22.22 
The Reptiles of the San Jacinto Area of Southern California, by Sarak 
Rogers Atsatt. Pp. 81-50. November, 1913 ©2222. oi... eee 
4. An Account of the Mammals and Birds of the Lower Colorado Valley, 
with Especial Reference to the Distributional Problems Presented, 
by Joseph Grinnell. Pp. 51-294, plates 3-13, 9 text figs. March, 1914. 
. Aplodontia chryseola, a New Mountain Beaver from the Trinity Region 
of Northern California, by Louise Kellogg. Pp. 295-296. 


pon 


ao 


California, by Walter P. Taylor. Pp. 297-300. 


Nos. 5. and 6 in one cover. April, 19140 1...222.2.. tee . 
7; A Second Species of the Mammalian Genus Microdipodops from Cali-. 


fornia, by Joseph Grinnell. Pp. 301-304. April, 1914 202. 


1. The Schizopoda of the San Diego Region, by Calvin O. Esterly. Pp. ~ 


1-20,. plates’ ?-23°- April, 1914 23S oe Re Be eck wea 
2. A Study of the Occurrence and Manner of Distribution of the Cteno- 
phora of the San Diego Region, by Calvin O. Esterly. Pp. 21-38. 


Aprils TOU Ae OS So ees Bee See Ne EEE Oey Se Re seta ee x te 
A New Self-Regulating Paraffin Bath, by C. W. Woodworth. eis 89- = 


Moa 


42. 2 text-fiptres,= Aprils 191s ota ef aca tvee cose pnaconetaaleuabonsckege 


A Previously Undescribed Aplodontia from the Middle North Coast of- 


. The Control of Pigment Formation in Amphibian Larvae, by Myrtle a yes 
E. Johnson. Pp. 53-88, plate 1. March, 1913 - Se 
. Sagitta californica, n.sp., from the San Diego “Region, including © 


Se 
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yaaa f ARR Re Nn Te Sgeaay 


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; 1. Two New Owls from Arizona, with Description of the Juvenal Plum- . ~~ 
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BE ges NS «ge 1 Sy aie ha 8 etek les bs batten rae NSAI Vapi Rn apn SY 

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5. The Modesto Song Sparrow, by Joseph Grinnell. Pp. 197-199. Feb-— 

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Taylor: Pp..205-307.2duney AO ooo eh ol ee et 
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Description of a New Hairy Woodpecker from Southeastern Alaska, by 
H. 8. Swarth. Pp. 318-318. October, 19V0 - woe. eceeeeeee 
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County, Nevada, with a Discussion of Some of the Faunal Features 
of the Region, by Walter P. Taylor. Pp. 319-436, plates. 7-12. 
FQDrUsTry OY ee Ee daa states done anc cacy dak obtains ceases 

Index, pp. 437-446. 


Vol. 8 1. The Vertical Distribution of Hucalanus elongatus in the San Diego 
Region during 1909, by Calvin O. Esterly. Pp. 1-7. May, 1911 -....... 
2. New and Rare Fishes from Southern California, by Edwin Chapin 
Starks and William M. Mann. Pp. 9-19, 2 text-figures. -July, 1911. 
8. Classification and Vertical Distribution of the Chaetognatha of the San 
Diego Region, Including Redescriptions of Some Doubtful Species of — 
the Group, by Ellis L. Michael. Pp. 21-186, pls..1-8. December, 1911. - 
4, Dinoflagellata of the San Diego Region, IV. The Genus Gonyaulaz, with 
Notes on Its Skeletal Morphology and a Discussion of Jts Generic 
and Specific Characters, by Charles Atwood Kofoid. Pp. 187-286, 
plates 9-17. 


Penis 


UNIVERSITY OF CALIFORNIA PUBLICATIONS 
IN 
ZOOLOGY 
Vol. 12, No. 7, pp. 301-304 April 15, 1914 


A SECOND SPECIES OF THE MAMMALIAN 
GENUS MICRODIPODOPS FROM 
CALIFORNIA 


BY 


JOSEPH GRINNELL 


(Contribution from the Museum of Vertebrate Zoology of the University of California) 


The remarkable rodent genus Microdipodops came to the attention 
of naturalists only about twenty-three years ago. So far as at present 
known, it occupies a more restricted area than any other genus of 
Sonoran mammal, namely, the central and northern parts of Nevada, 
and the adjacent extreme southeastern portion of Oregon and eastern 
edge of California. This genus may thus be inferred to be an essen- 
tially Great Basin product. Even within this limited region geo- 
graphic speciation is strongly in evidence, so that four species have 
already been distinguished by name. The existence of any represent- 
ative of the genus within the state of California was first discovered 
by Dr. Walter K. Fisher, who, in August, 1900, sent to the United 
States Biological Survey some specimens obtained by him in Sierra 
Valley, Plumas County, near the town of Vinton. One of these speci- 
mens became the type of Microdipodops californicus C. H. Merriam 
(1901, p. 128). This has continued until the present time the only 
record of the occurrence of the genus in California. 

In July, 1912, two collectors from the California Museum of Verte- 
brate Zoology, Messrs. Chas. D. Holliger and Norman Stern, trapped 
for mammals in the vicinity of Benton, Mono County, California. As 
one result of their work there, Wicrodipodops was found to be rather 
commonly represented, and a series of ten specimens (nos. 17031- 
17040) was secured. These prove to represent a species altogether 
distinct from M. californicus, and also different from the other three 
species of the genus characterized to date. 


AK sonia a Ins tity ep 
Se 
\ MAY @ 1914 
XA a7 


Sonal 


a\\ 


AR ye 


302 University of California Publications in Zoology  'Vou. 12 


The previously deseribed forms of Microdipodops are: M. mega- 
cephalus (C. H. Merriam, 1891, p. 116), type locality Halleck, Elko 
County, Nevada; MW. megacephalus oregonus (C. H. Merriam, 1901, p. 
127), type locality Alvord Desert, Harney County, Oregon; M. pallidus 
(C. H. Merriam, 1901, p. 127), type loeality Carson Sink, near Still- 
water, Churchill County, Nevada; M. californicus (C. H. Merriam, 
1901, p. 128), type locality Sierra Valley, near Vinton, Plumas County, 


California. 


Microdipodops polionotus, new species 
Mono Kangaroo Mouse 


Type.—Male adult, no. 17031, Mus. Vert. Zool.; MeKeever’s Ranch, 
two miles south of Benton Station, Mono County, California, altitude 
5200 feet; July 10, 1912. Collected by C. D. Holliger; orig. no. 184. 

Diagnostic Characters—Nearest like Microdipodops pallidus in 
general appearance, but coloration paler, decidedly more ashy in tone, 
and white areas emphasized in brilliancy; pelage notably shorter, less 
lax and fluffy ; general size less, and tail decidedly shorter ; skull similar 
to that in pallidus and megacephalus, but auditory capsules slightly 
less inflated, particularly less protuberant behind. 

Coloration of Type.—Above, eartridge buff (of Ridgway’s Color 
Standards and Color Nomenclature, 1912), with the hairs minutely 
black-tipped especially on the rump; sides of face and rump least 
buffy, presenting a distinetly ashy tone; underfur a pale tint of gray, 
te 
gradually dusky-hued towards tip; lower surface of tail and feet, 
white ; whole lower surface of body brilliant white, abruptly contrasted 
with dorsal color along sides; some of whiskers black, and others, the 
longer posterior ones, white; ears like back in tint, but with a silvery 
white upper rim, a conspicuous snowy white spot at inferior base, and 


number dorsal side of tail cartridge buff proximally, becoming 


an even larger white patch just behind. 

Measurements —Of type, total length 145 millimeters, tail verte- 
brae 80, hind foot 24, ear from crown 9; average of seven adults, total 
length 148, tail vertebrae 82, hind foot 23.7, ear from crown 7.6. 

Comparative Remarks.—The new species differs from all those pre- 
viously described in extreme pallor of coloration. The contrast with 
californicus is particularly great, the dorsal color in that form being 
strongly brownish, approaching tawny-olive in fused tone. Polionotus 


1914] Grinnell: Microdipodops 303 


is ashy gray, very faintly tinged with buff. This tone of coloration, 
on the insides of the ears alone, serves to characterize every individual 
in our series of polionotus. The white around the ears is particularly 
conspicuous as compared with that in all the forms except californicus, 
in which species the darker surrounding color makes up for the lesser 
amount of white in producing the effect of contrast. 

There is a notable difference in quantity and texture of pelage 
among the different species. It is possible that some of the variation 
in this respect is due to season, but it cannot all be. Our skins of 
poliontus, taken in July, exhibit much longer pelage than is shown 
by specimens of californicus taken in August, and yet not so long and 
especially not so silky and lax as shown by examples of pallidus taken 
in May. 

The body size of all the species appears to be about the same, but 
there is some variation in tail length. Polionotus has a lesser tail 
length than pallidus as given by Merriam, resembling most nearly 
megacephalus in this particular. 

The skulls of megacephalus and pallidus are essentially alike ae- 
cording to Merriam. The skulls of polionotus differ from examples 
at hand of those just named, in noticeably smaller auditory capsules. 
The inflation is decidedly less, so that when looked at from behind the 
skulls of polionotus do not stand so high, due to less inflation of bullae ; 
when looked at from above the mastoid inflation is seen to be less pro- 
tuberant both laterally and behind; the notch between the bulging 
capsules behind is thus not so deep. The rostra are not of the relatively 
slender build shown in californicus. 

Habitat—The entire series of Microdipodops polionotus was ob- 
tained on a sandy, sagebrush flat, on the McKeever Ranch, two miles 
south of the railroad station of Benton, Mono County, California. The 
altitude of this flat is close to 5200 feet, and in common with the rest 
of the immediately adjacent region the type locality lies in an ex- 
tremely arid faunal division of the Upper Sonoran Zone. The dates 
of capture are July 10, 11, and 12. Three of the animals are obviously 
young, one of these being not over half grown. 

Acknowledgments.—The present study was put upon a satisfactory 
basis through the privilege accorded the writer by Mr. Henry W. 
Henshaw, Chief of the Bureau of Biological Survey, Washington, of 
examining specimens contained in the portion of the National collection 
under his charge. This borrowed material consisted of skins-with- 
skulls of each of the four previously deseribed forms, as follows: 


304 University of California Publications in Zoology (Vou. 12 


Microdipodops megacephalus, five topotypes; M. m. oregonus, two topo- 
types; MW. pallidus, two specimens from the general type area; M. cali- 
fornicus, five topotypes. 


Transmitted February 11, 1914. 


LITERATURE CITED 


Merriam, C. H. 


1891. Description of a new genus and species of dwarf kangaroo rat from 
Nevada (Microdipodops megacephalus). U. S. Dept. Agric., Div. 
Orn. and Mam., N. Amer. Fauna, 5, 115-117. 

1901. Descriptions of three new kangaroo mice of the genus Microdipodops. 
Proce. Biol. Soe. Wash., 14, 127-128. 


ay, UNIVERSITY OF CALIFORNIA PUBLICATIONS— (Continued) 


6b. Ou the Skeletal Morphology of Gonyaulax catenata (Levander), by 
Charles Atwood Kofoid. Pp. 287-294, plate 18, - 
6. Dinoflagellata of the San Diego Region, V. On Spiraulaz, a New Genus 
of the Peridinida, by Charles Atwood Kofoid. Pp. 295-300, plate 19. 
Nos. 4, 5, and 6 in one cover. “September, 1911 00.0.0... eee 
7. Notes on Some Cephalopods in the Collection of the University of Cali- 
fornia, by 5. 8S, Berry. Pp. 301-310, plates 20-21. September, 1911. 
8. On a Self-Closing Plankton Net for Horizontal Towing, by Charles 
Atwood Kofoid. Pp. 311-348, plates 22-25. 
9. On an Improved Form of Belf-closing Water-bucket for Plankton In- 
vestigations, by Charles Atwood Kofoid. Pp. 349-352. 
Nos, 8 and 9 in one cover. Nevember, 1911 222.0022. ci.cceccttccteceeeeneee 
index, pp. 353-357. 


Vol. 9. 1. The Horned Lizards of California and Nevada of the Genera Phryno- 

: soma and Anota, by Harold C. Bryant. Pp. 1-84, plates 1-9. Decem- 
ST SEO EL oa eS Se rah co seek eed gic en nce epee Re 

. On a Lymphoid Structure Lying Over the Myelencephaion of Lepisos- 

teus, by Asa C. Chandler, Pp. 85-104, plates 10-12. December, 1911. 

. Studies on Early Stages of Development in Rats and Mice, No. 3, by 

EH... Mark and J..A. Long, The Living Eggs of Rats and Mice with 

a Description of Apparatus for Obtaining and Observing Them (Pre- 
liminary paper), by J. A. Long. Pp. 105-136, plates 13-17. February, 

1912 


eo Nn 


4, The Marine Biological Station of San Diego, Its History, Present Con- 
ditions, Achievements, and Aims, by Wm. E. Ritter, Pp. 137-248, 
plates 18-24, and. 2 maps... March, 1912 -ono.c.cclecc levees ceeeee 

. Oxygen and Polarity in Tubularia, by Harry Beal Torrey. Pp. 249- 

SPORE NR Y, EOL eS Seay Sec ira New Me eae ae SMa eG en at Ota s 

6; The Occurrence and Vertical Distribution of the Copepoda of the San 

Diego Region, with particular reference to Nineteen Species, by Cal- 
vin O. Esterly. Pp. 253-340, 7 text-figures. July, 1912 - 

7. Observations on the Suckling Period in the Guinea-Pig, by a “Marion 

Read. Pp. 341-351. September, 1912 22 esse lee eet cence ee nees 

8. Haeckel’s Sethocephalus eucecryphalus (Radiolaria), a Marine Ciliate, 

by Charles Atwood Kofoid. Pp. 353-357. September, 1912 -.....200... 
Index, pp. 359-365. 


on 


Vol. 10. (Contributions from the Museum of Vertebrate Zoology.) 
1, Report on a Collection of Birds and Mammals from Vancouver Island, 
by Harry S. Swarth.- Pp. '1-124, plates 1-4. February, 1912 ~........... 
2. A New Cony from the Vicinity of Mount Whitney, by Joseph Grinnell. 
Pps 325-1205"  Fanuanyy Lot ek ee oe eee ee 
3. The Mole of Southern California, by J. Grinnell and H. S. Swarth. 
Pp..131-136, 2 text-figures. 
4, Myotis orinomus Elliott, a° Bat New to California, by J. Grinnell and 
H.S. Swarth.. Pp. 187-142, 2 text-figures. 
Nos. 3 and 4 in one-cover.. April, 1912 2c geececel ice cee cee en cece enee 
5. The Bighorn of the Sierra Nevada, by Joseph Grinnell. Pp, 143-153, 
PS OCCKETOULON: Hn NIB Ye LO oe a an Snag ee eon tama sees date 
6. A New Perognathus from the San Joaquin Valley, California, by 
Walter P, Taylor. Pp:-155-166, 1 text-figure. 
7. The Beaver of West Central California, by. Walter P. Taylor. Pp. 
167-169. 
Nos. 6 and 7 in one cover, May, 1912-2... ceciccccceeceeseceeecceeseene 
8. ‘The Two Pocket Gophers of the Region Contiguous to the Lower Colo- 
rado River, in California and Arizona, by Joseph Grinnell. Pp. 171- 
me LEAS PRES Bh ec Sg ee Se page RRC IS Sas MR Rp mt ne en neuer: Cam) eee 
9. The Species of the Mammalian Genus Sorex of West-Central Cali- 
fornia, with a note on the Vertebrate Palustrine Faunas of the 
Region, by Joseph Grinnell. Pp, 179-195, figs. 1-6. March, 1913 ..._... 
“10, An Account of the Birds and Mammals of the San Jacinto Area of 
Southern California, with Remarks Upon the Behavior of Geographic 
Races on the Margins of Their Habitats, by J. Grinnell and H, 8. 
Swarth. Pp, 197-406, pls. 6-10. October, 1913 22-22 nse. nce 
Index, pp. 407-417. 


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Vol. 12. 


Vol. 13. 


UNIVERSITY OF CALIFORNIA PUBLICATIONS—(Continued) 


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14. 


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rs Pycnogonida from the Coast of California, with Desc 


. Birds in Relation to a Grasshopper Outbreak in California, by Harold 


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. The Control of Pigment Formation in Amphibian Larvae, by ‘Myrtle 


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EG WOME SR, OT a a cas ean han cent eg ese ed 


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Glossiphonia stagnalis Linnaeus; B. Modifiability in the Behavior of . 


the Leech Dina microstoma Moore, by Wilson Gee. Pp. 197-305, 13 


text fignres.: “December, 1813-58 3 a ee é 
. The Structure of the Ocelli of Polyorchis penicillata, by Etta Viola 


Little. Pp. 307-328, plates 18-15.. February, 1914 22.0.0 


. Modifications and Adaptations to Functions in the Feathers of Cirews 


hudsonius, by Asa.C. Chandler. Pp. 329-376, plates 16-20. March, 


BBS 2 SS AS OER SOR LS APR ef Recipe alee Ba eee Se yk EL ARES eI oC yet ys ; 


A Determination of the Economic Status of the Western Meadowlark 
(Sturnella neglecta) in California, by Harold Child Bryant. Pp. 377- 
510, plates 21-24, 5 text figures. February, 1914 <2... .o2 

Parasynaptic Stages in the Testis of Aneides lugubris (Hallowell), by 
Harry James Snook and J. A. Long. Pp,.511-528, plates 25-26, 1 text 
HE. ETE OEE sess SE Bes al ter cepa pees 


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. Nocturnal Wanderings of the California Pocket Gopher, by Harold C. 


Bryant. Pp. 25-29, 1 text fig. Nowember, 1913 2222.2... 
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UNIVERSITY OF CALIFORNIA PUBLICATIONS 
IN 
ZOOLOGY 
Vol. 12, No. 8, pp. 305-310, plate 14 October 31, 1914 


DISTRIBUTION OF RIVER OTTERS IN 
CALIFORNIA, 


WITH DESCRIPTION OF A NEW SUBSPECIES 


BY 
JOSEPH GRINNELL 


(Contribution from the Museum of Vertebrate Zoology of the University of California) 


River otters (genus Lutra) are still known to oecur at irregular 
intervals in the streams of northern and central California. The 
southernmost ascertained point of occurrence in the coast belt is a 
creek flowing into one of the heads of Drake Bay, near Point Reyes, 
Marin County. In the great Sacramento-San Joaquin Valley there 
are definite reports of otters from various streams and sloughs south 
as far as near Lane Bridge, north of Fresno, in that portion of the 
San Joaquin River forming the boundary between Fresno and 
Madera counties. There are rumors of occurrence still farther south, 
namely in certain streams making down from the high southern 
Sierra Nevada; but there is as yet no acceptably authenticated 
instance. There are no records at hand from the coast belt south of 
San Francisco Bay and none from the San Diegan district. 

Otters are stated to be ‘‘occasionally caught in the Colorado River,’’ 
along the southeastern border of California (Stephens, 1906, p. 234). 
The writer just cited refers to the Colorado River animal under the 
name Lutra canadensis sonora Rhoads, apparently assuming its 
identity with the form deseribed from a tributary of the Gila River, 
in Yavapai County, Arizona. This assumption is probably correct, 
and the form sonora should not have been omitted, as it was, from 
my distributional list of the mammals of California (Grinnell, 1913, 
p. 297). However, I am unable at this time to add any corroborative 


306 University of Califorma Publications in Zoology — [Vou. 12 


evidence either as to the occurrence of otters in the Colorado River, 
or as to the status of the form there represented. It is practically 
certain that decided differences exist between the race occurring to the 
west beyond the broad expanse of waterless desert and that of the 
Colorado basin. Rhoads’ description of sonora comes near to provid- 
ing clear proof of this, although he had evidently not had the oppor- 
tunity to make comparisons with the Californian animal. Unfortun- 
ately, the present writer has no specimens of sonora for examination 

Returning again to the river otter of central and northern Cali- 
fornia, there are in the California Museum of Vertebrate Zoology 
seven specimens from this area, as follows: one (no. 4975), skin only, 
from John’s Camp, McCloud River, Shasta County, secured by E. L. 
Furlong; one, skull only (no. 12653), from Price Creek, tributary to 
Eel River, Humboldt County, obtained by F. Stephens; two, skulls 
only (nos. 19153, 19154), and two, skins with skulls (nos. 19098, 
19152), from Cuddeback, on tributary of the Eel River, Humboldt 
County, all taken by H. E. Wilder; and one, skin with complete 
skeleton (no. 20775), from Grizzly Island, Solano County, secured by 
Miss A. M. Alexander. In studying this Californian material the 
writer has had access to twenty-one other specimens from Alaska, two 
from Queen Charlotte Islands, British Columbia, one from Vancouver 
Island, British Columbia, and two from Klamath County, Oregon, all 
this material being also contained in this Museum; and three additional 
skulls from Oregon, two from MeKenzie River, Lane County, and 
one from Bend, Crook County, kindly loaned the writer from the 
Oregon State Fish and Game Office by Mr. Stanley G. Jewett. 

It is believed that the above specified material, in conjunction with 
the published measurements, plates, and descriptions by Rhoads (1898, 
pp. 423-439, pls. 24, 25) suffices for determining the systematic 
status of the California river otter. After an appropriate examina- 
tion of the facts in the case I am led to conelude that we have here 
an additional distinct subspecies, which requires naming. 


Lutra canadensis brevipilosus, subsp. nov. 
California River Otter 
Type.—Female adult, skin and complete skeleton, no. 20775, Mus. 
Vert. Zool.; Grizzly Island, Solano County, California; January 26, 
1914; secured from a local trapper by Miss Annie M. Alexander, and 
presented by her to the Museum. 


1914] Grinnell: Distribution of River Otters in California 307 


Diagnostic Characters.—Similar to L. c. pacifica and L. c. peri- 
clyzomae, but general size greater, pelage shorter, coloration paler, 
and proportions of skull different, the cranium for one thing being 
narrower and higher. 

Description of Type—Weight before skinning, 16 pounds, 10 
ounces. Total length, 1158 millimeters; caudal vertebrae, 447; hind 
foot, 123.5; height of ear from crown, 20; ear from notch, 23.8. 
Length of hair: on middle of back, 21; top of head between ears, 11; 
top of tail along median line half-way toward tip, 20.5; middle of 
belly, 16.4. Length of fur: on middle of back, 13.8; top of head, 8.1; 
top of tail, 9; belly, 9.8. Coloration (nomenclature, that of Ridgway’s 
Color Standards, 1912): above bister, with hairs distinetly paler 
tipped, giving a grizzled effect, gradually paling around sides to 
Saccardo’s umber on lower surface of body and tail; this further 
paling anteriorly to avellaneous on throat and to tilleul-buff on chin 
and upper lip; tops of fore and hind feet snuff brown; whiskers chiefly 
whitish. Skull small (see table of measurements) ; rostral portion 
relatively both narrow and shallow; braincase narrow and high: ratio 
of height of skull at bulla to mastoid breadth 64 per cent; dentition 
hight. 

Remarks.—Lutra c. pacifica Rhoads (1898, p. 429), type from Lake 
Keechelus, Kittitas County, Washington, is evidently a large race, very 
similar to L. c. periclyzomae Elliot (1905, p. 80), type from Queen 
Charlotte Islands, British Columbia. The differences between these 
two must be very slight. In fact Heller (1909, p. 262), after examin- 
ing good series in the national collections at Washington, was able to 
find but one ‘‘reliable character’’ to distinguish periclyzomae from 
pacifica, namely ‘‘the extreme flatness of the audital bullae.’’ The 
bullae of brevipilosus are much smaller, but relatively somewhat more 
elevated, than in British Columbian examples of periclyzomae. 

The type of brevipilosus is extreme in all skull characters (see 
pl. 14), so that it probably represents the farthest southern divergence 
of the Pacific Coast series of forms. The Humboldt County speci- 
mens are very similar, as shown in the table of measurements here- 
with given of Californian skulls. The five Oregon skulls are somewhat 
intermediate towards the pacifica-periclyzomae style; but because of 
their small size I should apply the name brevipilosus to them, along 
with all the Californian specimens. Characters of pelage and color 
are likely to prove intermediate also, though this surmise is practically 
worthless in absence of skins from Oregon and Washington. The three 


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1914] Grinnell: Distribution of River Otters in California 309 


other skins from California are almost identical in these respects with 
the type of brevipilosus. All are full-pelaged winter skins. 

As for name, the only synonym apparently requiring consideration 
is the ‘‘Lutra californica Gray,’’? and this seems to have been dis- 
posed of with finality by Thomas (1889, p. 198), as applying to a 
South American form of remote relationship. 

The type locality of brevipilosus is in the lowland area at the con- 
fluence of the Sacramento and San Joaquin rivers. In the ineluded 
Suisun marshes are many sloughs in which the water varies, accord- 
ing to the tide and the stage of water in the rivers, from salt to nearly 
fresh, averaging brackish. From information obtained by Miss 
Alexander, as well as from other sources, river otters would appear 
to be even at the present time common along these channels. At least 
six are reported to have been killed in the vicinity the past winter. 
Good evidence is at hand that otters have occurred recently across the 
Suisun marshes as far west as Cordelia Slough. 


Transmitted June 27, 1914. 


LITERATURE CITED 


Exuior, D. G. 
1905. Descriptions of three apparently new species of mammals. Proce. 
Biol. Soe. Wash., 18, 79-82. 
GRINNELL, J. 
1913. <A distributional list of the mammals of California. Proe. Calif. Acad. 
Sci., 3, 265-390, pls. 15, 16. 
HELLER, E. 
1909. The mammals, in ‘‘ Birds and mammals of the 1907 Alexander expedition 
to southeastern Alaska.’’ Univ. Calif. Publ. Zool., 5, 245-264, pl. 
26, 2 figs. in text. 
Ruwoaps, 8. N. 
1898. Contributions to a revision of the North American beavers, otters and 
fishers. Trans. Amer. Philos. Soce., n.s., 19, 417-439, pls. 21-25. 
STEPHENS, F. 
1906. California mammals (San Diego, West Coast Publishing Co.), 351 pp., 
frontispiece, many figs. in text. 
THomaS, O. 
1889. Preliminary notes on the characters and synonymy of the different 
species of otter. Proc. Zool. Soe. London, 1889, 190-200. 


PLATE 14 


Q adult, no. 20775, Mus. Vert. Zool., Grizzly Island, Solano County, Calif 
x dl 


[310] 


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1, Report on a Collection of Birds and Mammals from Vancouver Island, 
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A, The General Reactions of the Leeches Dina microstoma Moore and 
Glossiphonia stagnalis Linnaeus; B, Modifiability in the Behavior of 
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-Vol. 14. 


UNIVERSITY OF CALIFORNIA PUBLICATIONS — (Continued) 


12. The Structure of the Ocelli-of Polyorchis penicillata, by Etta Viola 


Little.” Pp. 307-328, plates 13-15. February, 191400. ccesctanem a 
13. Modifications and Adaptations to Functions in the Feathers of Circus ae 

hudsonius, by Asa C. Chandler. Pp. 329-376, plates 16-20. March, - 

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3, The Reptiles of the San Jacinto Area of Southern California, by § 2 
Rogers Atsatt. Pp. 31-50. November, 1918 0.00.02 tte 2 


4. An Account of the Mammals and Birds of the Lower Colorado Valley, - 


with Especial Reference to the Distributional Problems Presented, 
by Joseph Grinnell. Pp. 51-294, plates 83-13, 9 text figs. March, 1914, 

5. Aplodontia chryseola, a New Mountain Beaver from the Trinity Region 
of Northern California, by Louise Kellogg. Pp. 295-296... 


6. A Previously Undeseribed Aplodontia from the Middle North Coast of 


California, by Walter P. Taylor. Pp. 297-300. 


Nos. 5 and 6 in one cover, April, 1914702 aoe ) 


7. A Second Species of the Mammalian Genus Microdipodops from Cali- 
fornia, by Joseph Grinnell, Pp. 301-304. April, 1914 2.020. 
8. Distribution of River Otters in-California, with Description of a New 


Subspecies, by Joseph Grinnell, Pp. 305-310, plate 14. October, 1914 ~ 
1. The Schizopoda of the San Diego Region, by Calvin 0. Esterly. Pp. is 


1-20;-plates 4-275 Apmil, 1914 oo se ee Or ee a 


2A Study of the Occurrence and Manner of Distribution of the Cteno- : 
phora of the San Diego Region, by Calvin O. Esterly. Pp. 21-38. ~ 


mW cya BORG! 3 U: Sepa er ep oeesale ap MOEN aa ME Baer ect Ts Make ee arene 
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BQN ees a a eh RS i OS ES ers See eee 
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Frog, by S. J. Holmes. Pp. 167-174, plate 10. August, 1914 


8. Polychaetous Annelids of the Pacific Coast in the Collections of the 


Zoological Museum of the University of California, by Aaron i.” 
Treadwell. Pp. 175-234, plates 11-12. 

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B. Bigelow. Pp. 239-241, September, 1914 srrseeceencentnnscsscnrecencerspsognecsece 


1. A Report upon the Physical Conditions in San Francisco Bay, Based - 
upon the Operations of the United States Fisheries Steamer “Alba 
tross’’ during the Years 1912 and 1913, by F. B, Sumner, G. dD. 
Louderback, W. L. Schmitt, E. C. Jounston. Pp. 1-198, plates 1-13, 
20 text SEE: Suly, 1914 oe ee ees 


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UNIVERSITY OF CALIFORNIA PUBLICATIONS 
IN 


ZOOLOGY 
Vol. 12, No. 9, pp. 311-316 November 21, 1914 


FOUR NEW POCKET GOPHERS FROM 
CALIFORNIA 


BY 


JOSEPH GRINNELL 


(Contribution from the Museum of Vertebrate Zoology of the University of California) 


The pocket gophers (genus Thomomys) offer an interesting prob- 
lem in distribution and speciation. Reduced power of locomotion 
appears to have accentuated the action of barriers. Particularly in 
California, where topographical and climatic conditions are so varied, 
has differentiation of species proceeded to an extreme degree. 

For the past seven years, particular attention has been paid by 
the staff of the California Museum of Vertebrate Zoology to obtaining 
material in this group, with the result that we now have 1749 speci- 
mens from localities within the state. But even this amount of 
material is far from sufficient for exhaustive and accurate treatment. 
There is, however, sufficient ground for placing on record at this time 
characterizations of certain new species and subspecies. 

During a recent visit to this museum, Mr. Vernon Bailey, of the 
United States Department of Agriculture, went over our gophers for 
the purpose of gathering distributional data contributory to a revision 
of the North America gophers which he now has under way. The 
writer was privileged to work over part of the material with Mr. 
Bailey, and submit to him various questions. The validity of the 
supposed new forms was discussed, and during these discussions some 
of the writer’s impressions were emended or corrected. The writer 
takes this opportunity to thank Mr. Bailey for his friendly and helpful 
suggestions in these regards. 


312 University of California Publications in Zoology — | Vou. 12 


Thomomys monticola premaxillaris, new subspecies 


Yolla Bolly Gopher 


Type.—J adult, no. 20242, Mus. Vert. Zool.; two miles south of 
South Yolla Bolly Mountain, altitude about 7500 feet, in Tehama 
County, California; August 6, 1913; colleeted by G. F. Ferris; original 
no. 166. 

Diagnosis—A member of the monticola series of gophers; palest 
of the forms known from California. Feet and ears small; pre- 
maxillary tongues extending far back of posterior ends of nasals; 
interparietal relatively broad antero-posteriorly. 

Material—Twenty-five specimens (nos. 20223-20247), from three 
localities in Tehama County, in the vicinity of South Yolla Bolly 
Mountain: two miles south of South Yolla Bolly, about 7500 feet 
altitude; four miles south of South Yolla Bolly, about 6000 feet alti- 
tude; Mount Linn (South Yolla Bolly of residents of the region), 
about 7600 feet altitude. All these localities are in Canadian or high 
Transition Zone; semi-arid in faunal condition; and on gravelly 
mountain slopes in coniferous forest association. 

Measurements.—Of type (old adult male) : Total length, 215 mm.; 
tail, 59; hind foot, 27; oceipito-nasal length of cranium, 37.5; zygo- 
matic width, 20.8; mastoid width, 18.6; height of brainease at bullae, 
11.1. The hind foot averages, in thirteen adults of premawillaris, 
26.8 mm. 

Comparison.—From near topotypes of Thomomys monticola monti- 
cola Allen, from the central Sierra Nevada, in Eldorado County, 
California, the new form differs in paler coloration (above close to 
ochraceous-tawny [of Ridgway, 1912], below hght ochraceous-buft) , 
in smaller ear, shorter hind foot, in slightly shorter and narrower 
rostrum of skull, in greater development of temporal ridges and in 
closer and more nearly parallel approximation of these, in extension 
of premaxillary tongues far back of posterior ends of nasals, and in 
shape of interparietal which is on an average relatively much broader 
antero-posteriorly. From topotypes of 7’. monticola pinetorum Mer- 
riam, premacillaris differs in slightly paler coloration, in shape of 
interparietal, which averages very slightly broader antero-posteriorly, 
and in all other respects as from 7. m. monticola. It may be remarked 
that the differences distinguishing pinetorum and monticola are at best 


extremely slight. 


1914] Grinnell: Four New Pocket Gophers from California 313 


Thomomys diaboli, new species 
Diablo Gopher 


Type—® adult (contained five embryos), no. 14165, Mus. Vert. 
Zool.; Sweeney’s Ranch, in hills of Diablo Range twenty-two miles 
south of Los Banos, Merced County, California; April 2, 1911; col 
lected by C. H. Richardson and H. A. Carr; original no. 108. 

Diagnosis —A member of the bottae series of gophers; smallest. of 
all the forms so far known from west-central California; nearest in 
color to Thomomys angularis angularis Merriam, but shehtly darker 
brown, especially beneath; other characters: small ears and feet, very 
weak and narrow rostrum, small teeth, moderately projecting incisors, 
and short nasals. 

Material. 
type locality, as above; one (no. 16676) from top of divide on wagon 


Seven specimens (nos. 14160-14165, 14696) from the 


road, 3000 feet altitude, in the Temblor Range, eleven miles west- 
northwest of McKittrick, Kern County, California. The latter speci- 
men shows slight peculiarities of color and cranium, but is much 
nearer diaboli than any other form. In both places the zone is Upper 
Sonoran; rather arid; hillside juniper association. 

Measurements—Of type (adult female): total length, 180 mm.; 
tail, 60; hind foot, 25; occipito-nasal length of cranium, 32.9; zygo- 
matie width, 22.0; mastoid width, 17.9; height of braincase at bullae, 
11.6; length of nasals, 10.3. Total length of six adults: 180 to 193, 
average 189. 

Comparisons—F rom Thomomys bottae bottae (EKydoux and Ger- 
vais), of the San Francisco Bay region, the new form differs in very 
much smaller size, less blackish, more reddish, coloration, relatively 
smaller feet, smaller teeth, and weaker rostrum. From topotypes of 
T. angularis angularis Merriam, diaboli differs in much smaller size, 
relatively smaller feet and ears, shghtly darker and browner colora- 
tion, much weaker rostrum, narrower incisors and less angular skull. 
From 7. leucodon navus Merriam, of the Sacramento Valley, diaboli 
differs in slightly darker coloration, smaller size, much smaller molar 
teeth, smaller auditory bullae, much shorter nasals and narrower 
rostrum. From 7. nigricans nigricans Rhoads, of the coast ranges of 
San Diego County, diaboli differs in slightly paler coloration, and 
notably in cranial characters: the nasals are much shorter, the incisors 
project far more, and the braincase is broader and more inflated 


314 University of California Publications in Zoology [ Von. 12 


parietally. Diaboli is much like topotypes of 7. mewa Merriam in 
size and coloration, though not quite so reddish in corresponding 
pelage, but differs in longer tail, and cranially in more bulging brain- 
case, shorter nasals, and much more projecting incisors. 


Thomomys infrapallidus, new species 


Carrizo Plain Gopher 


Type—<¢ old adult, no. 14181, Mus. Vert. Zool.; seven miles south- 
east of Simmler, Carrizo Plain, San Luis Obispo County, California; 
May 25, 1911; collected by H. S. Swarth; original no. 9138. 

Diagnosis —A member of the bottae series of gophers; palest of 
all the forms of this series so far as known from west-central Cali- 
fornia; nearest in color to Thomonys angularis pascalis Merriam, but 
decidedly paler beneath; other characters: relatively large feet, long 
fore claws, long tail, narrow and high braincase, moderately spreading 
zygomatic arches, moderately projecting incisors, small molar teeth, 
and very small auditory bullae. 

Material—Nine specimens (nos. 14179-14187, Mus. Vert. Zool.), 
all from Carrizo Plain, San Luis Obispo County, California: six from 
seven miles southeast of Simmler, and three from five miles north of 
Painted Rock; all collected in May, 1911, by H. S. Swarth. The 
altitude of the Carrizo Plain varies from 1900 to 2500 feet. Faunally 
and zonally it may be considered very arid, high Lower Sonoran. <A 
sparse prairie vegetation grows on ground that is more or less strongly 
alkaline over most of the area. 

Measurements.—Of type (old adult male) : total length, 248 mm. ; 
tail, 76; hind foot 34; oecipito-nasal length of cranium, 43.0; zygo- 
matic width, 29.0; mastoid width, 22.8; height of brainease at bullae, 
14.0. 

Comparisons.—F rom topotypes of Thomomys bottae pallescens 
Rhoads, the new form differs at a glance in much smaller auditory 
bullae, smaller and slenderer teeth, and more anteriorly projecting 
incisors. From 7. bottae bottae (EKydoux and Gervais), infrapallidus 
differs in very much paler, more clayey, coloration, smaller general 
size, larger feet, longer tail, much smaller teeth, and smaller bullae. 
From topotypes of 7. angularis angularis Merriam, infrapallidus 
differs in smaller size, paler coloration, especially below, narrower 


1914] Grinnell: Four New Pocket Gophers from California 315 


braincase, and less squarely spreading zygomatic arches. From topo- 
types of 7. a. pascalis Merriam, infrapallidus differs in paler colora- 
tion ventrally, larger feet, higher braincase, more projecting incisors 
and smaller auditory bullae. 


Thomomys nigricans puertae, new subspecies 
La Puerta Gopher 


Type—g young adult, no. 7511, Mus. Vert. Zool.; La Puerta 
(Mason’s Ranch), eastern San Diego County, California; May 31, 
1909; collected by F. Stephens; original no. 1974. 

Diagnosis—A pale desert-slope race of Thomomys nigricans 
nigricans Rhoads; apparently identical in size and cranial characters 
with that race; pelage mid-dorsally and around ears with far less 
admixture of black; general tone of coloration both above and below, 
ochraceous-tawny (of Ridgway, 1912), brightest on sides, and shghtly 
subdued with dusky down middle of back; spot of sooty behind ear; 
face dusky. 

Material—The Museum contains forty-three specimens labelled La 
Puerta (nos. 7510-7519, 7582-7586, 16636-16654, 18822-18830). 
These are all of Mr. Stephens’s collecting. According to his field 
notebook, the majority were caught in cultivated land on Mason’s 
Ranch, which is located at the lower end of La Puerta Valley. Others, 
however, were taken on surrounding hillsides. Mason’s Ranch is five 
miles west of Vallecito, on the old overland (Butterfield) stage-road. 
La Puerta Valley is evidently arid Lower Sonoran. Along the course 
of the stream arrowweed (Pluchea) is a common plant, while there 
is an occasional mesquite. The valley floor is mostly covered with 
large creosote bushes (Larrea). 

Comparisons.—The series of La Puerta gophers is notably variable 
in color-tone. Picked specimens are fully as dark as the average of 
topotypes of nigricans, from Witch Creek, on the west side of the 
Cuyamaca Mountains. La Puerta is well down on the desert or east 
slope of the same general mountain mass, and less than ten miles 
away, so that it is quite possible that a continual invasion of true 
nigricans down on the east slope brings good examples of that form 
into the same valley with the more remotely derived and hence differ- 
entiated type here named puertae. Upon this theory the series at 
hand may be divided into bright-colored and dark-colored sections, 
the former meriting the name puertac, the latter nigricans. 


316 University of Califorma Publications in Zoology [ Vou. 12 


It is obvious that puwertae is a local, arid Lower Sonoran race of 
nigricans, the latter belonging to the closely-adjacent semi-humid 
Upper Sonoran of the various coast ranges of San Diego County, 
mostly on their western slopes. Typical pwertae, as compared with 
typical nigricans, shows, in addition to the characters given above, a 
tail of uniform pale tawny color both above and below, imstead of 
dusky above at base. From 7. cabezonae Merriam, possibly also a 
subspecies of nigricans, puertae differs in more tawny coloration above 
(this obtaining in small young as well as in adults), invasion of 
tawny over under surface, more depressed rostral region of cranium, 
narrower interorbital constriction and braincase, and more widely 
spreading zygomata. From 7. bottae pallescens Rhoads, puertac 
differs in smaller size, paler, more tawny coloration, lighter dentition, 
weaker and more depressed rostrum and smaller auditory bullae. 


Transmitted September 24, 1914. 


UNIVERSITY OF CALIFORNIA PUBLICATIONS— (Continued) 


4. The Marine Biological Station of San Diego, Its History, Present Con- 
ditions, Achievements, and Aims, by Wm. E. Ritter, Pp. 137-248, 


plates 18-24, and 2 maps. March, 191202... .cccsccccccesoecescsSenceencetsexpeotecesez $1.00 
5. Oxygen and Polarity in 7ubularia, by Harry Beal Torrey. Pp. 249- 
: PAT Bese SS oka hf Vase na Ea CTR SONS REID Bont 5S ons Tne eR a Pan Uo 05 
6. The Occurrence and Vertical Distribution of the Copepoda of the San 

Diego Region, with particular reference to Nineteen Species, by Cal- 

vin O. Esterly. Pp. 253-340, 7 text-figures, July, 1912 2.22.22 0 2 1.00 
7. Observations on the Suckling Period in the Guinea-Pig, by J. Marion 

Read. Pp. 341-361. September, 1912 osc cccc neces snncedecee 10 
8, Haeckel’s Sethocephalus eucecryphalus (Radiolaria), a Marine Ciliate, 

by Charles Atwood Kofoid.- Pp. 353-357. September, 1912 .......:...... .05 


Index, pp. 359-365. : 


(Contributions from the Museum of Vertebrate Zoology.) 
1. Report on a Collection of Birds and Mammals from Vancouver Island, 
by Harry S. Swarth. Pp. 1-124, plates 1-4. February, 1912 0.02.0... 1,00 
2. A New Cony from the Vicinity of Mount Whitney, by Joseph Grinnell. 
Pp. cious 29 saniary, LOUD S- oe  e e a0. 
3. The Mole of Southern California, by J. Grinnell and H. 8, Swarth. 
Pp. 131-136, 2 text-figures. 
4, Myotis orinomus Elliott, a Bat New to California, by J. Grinnell and 
H. S. Swarth. Pp. 137-142, 2 text-figures. 


Nos. 3 and 4 in one cover. April, 1912 2.20... pers | 
. the Bighorn of the Sierra Nevada, by Joseph Grinnell. Pp, 143-153, 
dv text-fisures: May, 1922 too, ee. ee Sh Ne 10 


. A New Perognathus from the San Joaquin Valley, California, by 
Walter P. Taylor, Pp. 155-166, 1 text-figure. 
~ 7 The Beaver of West Central California, by Walter P. Taylor. Pp, 
167-169, 
Nos. 6:and 7in-onhe ‘cover, = May;..1912.< 55) eo ee me 15. 
The Two Pocket Gophers of the Region Contiguous to the Lower Colo- 
tado River, in California and Arizona, by Joseph Grinnell. Pp. 171- 
B Ughs Bag 118 2) gti B22 B75 ene teeta TY cr SRE Se are Re RO eS 15 
The Species of the Mammalian Genus Sorex of West-Central Cali- 
fornia, with a note on the Vertebrate Palustrine Faunas of the 
Region, by Joseph Grinnell. Pp, 179-195, figs. 1-6. March, 1913 _..... 15 
An Account of the Birds and Mammals of the San Jacinto Area of 
Southern California, with Remarks Upon the Behavior of Geographic 
Races on the Margins of Their Habitats, by J. Grinnell and H, 8, 
Swarth.- Pp. 197-406, pls, 6-10.° October, 1913 W022 2.00 
Index, pp. 407-417. 


Birds in Relation to a Grasshopper Outbreak in California, by Harold 


Cc. Bryant. Pp. 1-20: _November, 1912 20 
On the Structure and Relationships of Dinosphaere palustris (Lemm.), 

by Charles Atwood Kofoid and Josephine Rigden Michener. Pp. 2i- 

OS MRIBS BY Torys ae) seeks £2 bia ieee ont CS ear cep ee edhe SE ap tata ts ay Se oe a 10 
A’ Study of Epithelioma Contagiosum of the Common Fowl, by 

Clifford D. Sweet. Pp. 29-51. January, 1913 o..00.2. ice ceeeceeeeti eee 25 
The Control of Pigment Formation in Amphibian Larvae, by Myrtle 

E. Johnson. Pp. 53-88, plate 1.. March, 1913... 22a 35 


Sagitta californica, u.sp., from the San Diego Region, including 
Remarks on Its Variation and Distribution, by Ellis L. Michael, 
Ppee89-1 26, pla be 2 EE ha aa os le Sages nace ene nore 35 

6. Pycnogonida from the Coast cf California, with Description of Two 

New Species, by:H. V. M. Hall. Pp, 127-142, plates 3-4. August, 1913.  .20 

7. Observations on Isclated Living Pigment Cells from the Larvae of 

Amphibians by S, J. Holmes, Pp. 143-154. plates 5-6. 
8. Behavior of Ectodermic Epithelium of Tadpoles when Cultivated im 
Plasma, by S. J. Holmes. Pp. 155-172, plates: 7-8, 


Nos. 7 and 8 in one cover. September, 1913 0228.22 see 30 
: 9. On Some Californian Schizopeda, by H. J. Hansen. Pp. 173-180, pl. 9. 
Sy : SS Pepa} 4 fot at Os ae NR aS Sa BN NRcaaseae aRaNICSe NNT 10 
i= “10. Fourth * axonomic Report on the Copepoda of the San Diego Region, 
by Calvin O. Esterly. Pp. 181-196, pls..10-12. November, 1913 ....... 15 


11. The Behavior of Leeches with Especial Reference to Its Modifiability, 
$ A. The General Reactions of the Leeches Dina microstoma Moore and 
% Glossiphonia stagnalis Linnaeus; B. Modifiability in the Behavior of 
the Leech Dina microstoma Moore, by Wilson Gee. Pp. 197-305, 13 

_ text figures. “December, 1913 on. senn cna ace nape hee aeons tdem ee -. 1,00 


UNIVERSITY OF CALIFORNIA PUBLICATIONS— (Continued) 


12. The Structure of the Ocelli of Polyorchis penicillata, by Etta Viola 
Little. Pp. 307-328, plates 13-15. February, 1914 ........20 
13. Modifications and Adaptations to Functions in the Feathers of Circus _ 
hudsonius, by Asa ©. Chandler. Pp. 329-376, plates 16-20. March, 
b A Te SO ecard AG ap gE Pee neg Se ain AME pa ME a a ER a hin a OL A AS 
14. A Determination of the Econoniic Status of the Western Meadowlark 
(Sturnella neglecta) in California, by Harold Child Bryant. Pp. 377- ~ 
510, plates 21-24, 5 text figures. February, 1914 ...2.e2- ic lecceeeeees meni 
15. Parasynaptic Stages in the Testis of Ancides lugubris (Hallowell), by ~~~ 
Harry dames Snook and J. A. Long. Pp. 511-528, plates 25-26, 1 text 
fig. April, 1914 


Vol. 12. 1. A’ Study of a Collection of Geese of the Branta canadensis Group from 

the San Joaquin Valley, California, by Harry S. Swarth. Pp. 1-24, 

plates 1-2; 8 text figs. November, 1913 -2..c2 on. oik ela eae ‘ 

Nocturnal Wanderings of the California Pocket Gopher, by Harold 0. 

Bryant, Pp. 25-29, 1 text fig. November, 1913 20000002. 

The Reptiles of the San Jacinto Area of Southern California, by Sarah ~ 

Rogers Atsatt. Pp. 81-50. November,°1913) 2.0.0.2 ee eke ecceecececceneee 

4, An Account of the Mammals and Birds of the Lower Colorado Valley, 

with Especial Reference to the Distributional Problems. Presented, 

by Joseph Grinnell. Pp. 51-294, plates 3-13, 9 text figs. March, 1914, 

Aplodontia chryseola, a New Mountain Beaver from the Trinity Region 
of Northern California, by Louise Kellogg. Pp. 295-296, 

. A Previously Undescribed Aplodontia from the Middle North Coast of 

California, by Walter P. Taylor. Pp. 297-300. 

Nos. 5 and 6 in one cover. April, 1914 o.oo. cee cece cecceteneceneeene 

7. A Second Species of the Mammalian Genus Microdipodops from Cali- 

fornia, by Joseph Grinnell. Pp, 301-304.. April, 1914 wu 

. Distribution of River Otters in California, with Description of a New 

Subspecies, by Joseph Grinnell... Pp. 305-310, plate 14. October, 1914 

. Four New. Pocket Gophers from California, by Joseph Grinnell. Pp. 

811-316. November, 1914 


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Ets re “9 
Lad cs i tech ees A ARE Aa “PE 


oOo 2 


Vol, 18. 1. The Schizopoda of the San Diego Region, by Calvin O. Esterly. Pp. 
1220. “plates: 1-2) = Aprilys OF4- 25 S27 ves paca eRe co yn oop aceon, cae einer 

2. A Study of the Occurrence and Manner ‘of Distribution of the Cteno- 
phora of the San Diego Region, by Calvin 0: Esterly. Pp. 21-38, 

WN rhs t Bhat Ib We tee Are ees OUR ah ay BE Rane nena ERS NU eR RE I 

3. A New Self-Regulating Paraffin Bath, by C. W. Woodworth. Pp. 39. 
AD o-text-feures,, \April, TOTS sven, cee os. kak os ohana ae 

4: Diplodinium ecaudatum, with an Account of Its Neuromotor Apparatus, 
by Robert G, Sharp. Pp. 43-122, plates 3-7, 4 text figures, May, 
LOU ee ee oP Sack Se Teas Gara as Satan ere cme aa Me sea MEER 

5. The Vertical Distribution and Movements of the Schizopoda of the 
San Diego Region, by Calvin O. Esterly. Pn. 123-145. May, 1914...... 

6. The Anatomy of Heterodontus Francisci. I. The Exoskeleton, by J. 


Frank Daniel. Pp, 147-166, plates 8-9, 4 text figures. May 23, 
1914 


7. The Movernents and Reactions of the Isolated Melanophores of the 
Frog, by-S. J. Holmes. Pp. 167-174, plate 1C. August, 1914 002. 
8. Polychaetous Annhelids of the Pacific Coast in the Collections of the 
Zoological Museum of the University of California, by Aaron L. 
Treadwell. Pp, 175-234, plates 11-12. 
9. New Syllidae from San Francisco Bay (collected by the U.&. S. 
** Albatross’’), by Aaron L. Treadwell. Pp. 235-238, 7 text figures. 
Nos. 8 and 9 in’one cover. October, 1914 oo... loe cei ice eceteactcneee : 
10. Note on the Medusan Genus Stomolophus, from San Diego, by Henry 
B. Bigelow. Pp. 239-241. September, 1914 


ee ee oe ee en ag 


Vol. 14. 1. A Report upon the Physical Conditions in San Francisco Bay, Based 
upoh the Operations of the United States Fisheries Steamer ‘‘Alba- 
tross’’ during the Years 1912 and 1913, by F. B. Sumner, G, D. 
Louderback, W. L. Schmitt, E. C, Johnston, Pp. 1-198, plates 1-19; So 
20 text figures. Duly, 1914 ooo... ccc ccc eed ecteceee Rake Te Sed paae eee Saray ns 


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UNIVERSITY OF CALIFORNIA PUBLICATIONS 


IN 
ZOOLOGY 
Vol. 12, No. 10, pp. 317-321 December 4, 1914 


4 ORE 


THREE NEW RACES OF VESPERTILIONID 
BATS FROM CALIFORNIA 


ie BY 

¥ 

: HILDA WOOD GRINNELL 
: 


ax 
spe UNIVERSITY OF CALIFORNIA PRESS 
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UNIVERSITY OF CALIFORNIA PUBLICATIONS 
IN 


ZOOLOGY 
Vol. 12, No. 10, pp. 317-320 December 4, 1914 


THREE NEW RACKS OF VESPERTILIONID 
BATS FROM CALIFORNIA 


BY 
HILDA WOOD GRINNELL 


(Contribution from the Museum of Vertebrate Zoology of the University of California) 


During the past six years there has been accumulated in the 
Museum of Vertebrate Zoology a collection of eight hundred and 
thirteen specimens of bats from within the boundaries of California. 
Although material is still lacking from many important localities 
within the state, there is now at hand in several instances a sufficient 
number of specimens to show clearly the existence of certain hitherto 
unnamed races. Three of these new races are described below. In 
these descriptions all measurements are given in millimeters; total 
length, tail vertebrae and foot measured in the flesh by the collector. 
Color deseriptions are based upon Ridgway’s Color Standards and 
Nomenclature (1912). 

The writer’s thanks are due to Mr. Henry W. Henshaw, Chief of 
the Bureau of Biological Survey, United States Department of Agri- 
culture, and to Mr. John Rowley, Curator of Mammals, California 
Academy of Sciences, for the loan of pertinent material used in 


comparisons. 


Myotis californicus quercinus, new subspecies 
Oak Foliage Bat 
Type—Female, adult; no. 6939, Mus. Vert. Zool.; Seven Oaks, 
5000 feet altitude, San Bernardino Mountains, San Bernardino County, 
California; July 8, 1905; collected by J. Grinnell; original no. 1120. 
Diagnosis—Similar to Myotis californicus californicus (Audubon 
and Bachman) and Myotis californicus pallidus Stephens, but inter- 
mediate in color between these two forms. Prevailing tone of color 


on back, cinnamon. 


318 University of California Publications in Zoology [ Vou. 12 


Description —Ears, feet and fur as in M. c. californicus. Mem- 
branes and bases of hairs everywhere as in californicus. On the back 
the terminal portions of the fur are glossy cinnamon, and this color 
extends down onto the sides. The terminal portions of the hairs be- 
low are light buff in color, rather than buffy-brown as in californicus, 
or pale cartridge-buff as in pallidus. 

Measurements.—A series of ten examples of WM. c. quercinus from 
southern California averages in millimeters as follows: Total length, 
81.6 (77.0-83.0) ; tail vertebrae, 36.8 (31.0-41.0) ; tibia, 14.1 (12.5- 
15.0) ; foot, 6.0 (4.0-8.0) ; forearm, 31.9 (31.0-33.2); greatest length 
of eranium, 13.1 (12.9-13.8); zygomatic breadth, 7.7 (7.4-8.0); 
breadth of brainease, 6.8 (6.6-7.0); interorbital constriction, 3.0 
(2:9=3:3)). 

Specimens examined.—The writer has examined twenty-two speci- 
mens of Myotis californicus quercinus from the following localities 
in California: San Diego County—Cuyamaea, 2, Julian, 5; Santa Cruz 
Island—Friar’s Harbor, 3; San Bernardino Mountains, San Bernar- 
dino County—Seven Oaks, 2, Bear Lake, 1, South Fork Santa Ana 
River, 2; San Jacinto Mountains, Riverside County—Kenworthy, 1, 
Sehain’s Ranch, 1; Tulare County—Trout Creek, 2; Ventura County— 
Matilija, 1, Mount Pinos, 2. 

Remarks.—The three specimens listed from Santa Cruz Island, 
while slightly darker than typical quercinus, are still nearer to this 
form than to MW. c. californicus. 

Distribution.—The range of M. c. quercinus, as so far worked out, 
occupies portions of the San Diegan faunal division of southern Cali- 
fornia, and the Santa Barbara Islands. The life-zone is high Upper 
Sonoran and low Transition. The bats appear at late twilight and 
are usually observed flitting close about the foliage of scrub, golden, 
and black oaks. 


Myotis yumanensis sociabilis, new subspecies 
Tejon Bat 


Type—Female, adult; no. 5158, Mus. Vert. Zool.; Old Fort Tejon, 
3200 feet altitude, Kern County, California; July 23, 1904; collected 
by J. Grinnell; original no. 715. 

Diagnosis —Similar in general characters to Myotis ywmanensis 
yumanensis (H. Allen) and Myotis ywmanensis saturatus Miller, but 


intermediate in color between these two forms. 


1914] Grinnell: Three New Races of Vespertilionid Bats 319 


Description—The fur is distributed as in topotypes of M. y. 
yumanensis. On middle of back it averages about six millimeters in 
length. Hairs everywhere clove brown at base; distal half of fur on 
dorsal surface wood brown; fur below light buff, with darker bases 
of hairs showing through. On throat, sides and chin the color varies 
toward warm buff; ears olive brown; feet, wings and tail-membranes 
clove brown. The young are darker and grayer throughout, entirely 
lacking the buffy tint of the adults. 

Measurements.—A series of five adult males of M. y. sociabilis 
averages in millimeters as follows: Total length, 81.4 (75.0-87.0) ; 
tail vertebrae, 33.3 (30.0—-37.0) ; tibia, 15.2 (15.0-16.0) ; foot, 8.1 (7.0- 
10.0); forearm, 34.6 (32.9-35.3); greatest length of eranium, 13.7 
(13.5-14.4) ; zygomatic breadth, 8.4 (8.0-8.6) ; breadth of braincase, 
7.1 (6.77.6) ; interorbital constriction, 3.8 (3.5-3.9). 

Ten adult females from Old Fort Tejon, Kern County, average in 
millimeters as follows: Total length, 81.9 (76.0-85.0) ; tail vertebrae, 
36.6 (34.0-37.0) ; tibia, 14.7 (13.5-16.0) ; foot, 8.9 (8.0-10.0) ; fore- 
arm, 34.2 (33.7-35.0) ; greatest length of cranium, 13.8 (13.4—14.2) ; 
zygomatic breadth, 8.1 (7.8-8.3) ; breadth of braincase, 7.2 (6.7-7.3) ; 
interorbital constriction, 3.7 (3.6-4.0). 

Specimens examined.—Total number sixty-nine, from the follow- 
ing localities in California: Butte County—Chambers Ravine, four 
miles north of Oroville, 1; Glenn County—Winslow, five miles west 
of Fruto, 1; Kern County—Old Fort Tejon, 61, Buttonwillow, 1 
(Calif. Acad. Sci.) ; San Bernardino County—Bluff Lake, 7500 feet 
altitude, 3, Bear Lake, 6700 feet altitude, 1, South Fork Santa Ana 
River, 8500 feet altitude, 1. 

Remarks.—Specimens of M. y. sociabilis from the San Bernardino 
Mountains show strong superficial resemblance to the smaller indi- 
viduals among a series of Myotis longicrus (True) from the same 
locality. The longer tibia of the latter species, however, together with 
the slightly greater size of skull and the more elevated occipital region, 
serves to allocate individuals. 

Distribution —The distribution of this bat cannot be stated with 
confidence without much further field-work. It appears to occupy an 
intermediate geographic position between that of M. y. ywmanensis 
and M. y. satwratus, namely the semi-arid Transition and Sonoran 
zones in California west and north of the southeastern deserts. 


320 University of California Publications in Zoology [ Vo. 12 


Corynorhinus macrotis intermedius, new subspecies 
Intermediate Lump-nosed Bat 


Type—Female, adult; no. 7753, Mus. Vert. Zool.; Auburn, 1300 
feet altitude, Placer County, California; July 31, 1909; collected by 
Dr. J. C. Hawver; original no. 2387, J. Grinnell. 

Diagnosis—Similar in general characters to Corynorhinus macrotis 
pallescens Miller and Corynorhinus macrotis townsendi (Cooper), but 
intermediate in color between these two forms. 

Description.—As compared with pallescens, intermedius is some- 
what larger in general size; ten examples of the latter form from 
Auburn, Placer County, average 102 millimeters in length, while ten 
specimens of pallescens from the San Jacinto region average but 97.2 
in the same dimension. In color intermedius is natal brown above; 
below, wood brown; membranes bone brown. 

Measurements.—A series of ten specimens from west central Cali- 
fornia averages in millimeters as follows: Total length, 102 (97.0- 
108.0) ; tail vertebrae, 48.6 (45.0-55.5) ; tibia, 19.7 (18.7-21.0) ; foot, 
9.8 (9.0-12.5); forearm, 42.0 (40.2-43.6); greatest length of skull, 
16.2 (15.4-17.1). 

Specimens examined.—Total number, thirty-two, from the follow- 
ing localities in California: Placer County—Auburn, 23, Pioneer Cave, 
3; Santa Catalina Island—Johnson Harbor, 1; Napa or Sonoma 
County—Mount Veeder, 1 (U. S. Biol. Surv.) ; Siskiyou County— 
Happy Camp, 1 (U.S. Biol. Surv.) ; San Benito County—Bear Valley, 
2 (U.S. Biol. Surv.), Hernandez, 1 (Calif. Acad. Sei.). 

Distribution —The evidence at hand indicates that this bat occupies 


a geographic position intermediate between that of pallescens and that 
of townsendi, namely the semi-arid and semi-humid portions of the 
Upper Sonoran zone in California west of the desert divides. 


Transmitted October 6, 1914. 


UNIVERSITY OF CALIFORNIA PUBLICATIONS—(Continued) 


4, The Marine Biological Station of San Diego, Ita History, Present Con- 
ditions, Achievements, and Aims, by Wm. E. Ritter, Pp. 137-248, 
Plates 18-24, and 2 maps, March, 1912 20......c..ccccccctecceecoceee seers ete eees 

5. Oxygen and Polarity in Tubularia, by Harry Beal Torrey. Pp. 249- 
COLT TAY Sd ONS ar OSS es ie a eae eR NM ae ae 

6. The Occurrence and Vertical Distribution of the Copepoda of the San 
Diego Region, with particular reference to Nineteen Species, by Cal- 
vin O. Esterly. Pp. 253-340, 7 text-figures. July, 1912 2.00.0. 

7. Observations on the Suckling Period in the Guinea-Pig, by J. Marion 
Read. Pp. 341-351. “September, 1912 20. 

8. Hacciel’s Sethocephalus eucecryphalus (Radiolaria), a Marine Ciliate, 
by Charles Atwood Kofoid. Pp. 353-357. September, 1912 20.02.20... 

Index, pp. 359-365. 


Vol. 10. (Contributions from the Museum of Vertebrate Zoology.) 


1, Report on a Collection of Birds and Mammals from Vancouver Island, 
ze by Harry 8, Swarth. Pp. 1-124, plates 1-4, February, 1912 ............. 
x 2. A New Cony from the Vicinity of Mount Whitney, by Joseph Grinnell. 

Bp ep 29. = a anyary, 1002 8 ee a a 
3. The Mole of Southern California, by J. Grinnell and H, 8, Swarth. 
nes Pp. 131-136, 2 text-figures. 
; 4. Myotis orinomus Elliott, a Bat New to California, by J. Grinnell and 
H. 8S. Swarth. Pp. 137-142, 2 text-figures. 
Nos. 3 and 4 in one cover. April, 1912 ok ccceeectacecene 

. the Bighorn of the Sierra Nevada, by Joseph Grinnell. Pp. 143-153, 

4 stext-Benres; | May, LOL 2a a ea ie Ce ee 

6. A New Perognathus from the San Joaquin Valley, California, by 

Walter P. Taylor. Pp. 155-166, 1 text-figure. 


f 
o 


7. The Beaver of West Central California, by Walter P. Taylor. Pp. 
“ : 167-169. 
4 Nos. 6 and 7 in one cover. May, 1912 oct ee teenie 


8. The Two Pocket Gophers of the Region Contiguous to the Lower Colo- 
tado River, in California and Arizona, by Joseph Grinnell. Pp, 171- 
U7 Be STUNG POU DE reer oh pcesatin Cece Nera kareena oad 

The Species of the Mammalian Genus Sorex of West-Central Cali- 
fornia, with a note on the Vertebrate Palustrine Faunas of the 
Region, by Joseph Grinnell. Pp. 179-195, figs. 1-6. March, 1913.1... 

10. An Account of the Birds and Mammals of the San Jacinto Area of 

Southern California, with Remarks Upon the Behavior of Geographic 

Races on the Margins of Their Habitats, by J. Grinnell and H. 8. 

Swarth. Pp. 197-406, pis, 6-10. October, 1918 22.002. 
Index, pp. 407-417. 


. Vol. 11. 1. Birds in Relation to a Grasshopper Outbreak in California, by Harold 

C. Bryant. Pp. 1-20.. November, 1912 22.0 n0. ccs een ete nene 

2. On the Structure and Relationships of Dinosphaera palustris (Lemm.), 

by Charles Atwood Kofoid and Josephine Rigden Michener. Pp. 21- 

Pa x VOCE DCPs) BON as. wach ee ea cs See SE a cava ue ste ee ON bs 

3. A Study of Epithelioma Contagiosum of the Common Fowl, by 

= Clifford D. Sweet. Pp. 29-51. January, 1913 ~.2..eeeeee 

. The Control of Pigment Formation in Amphibian Larvae, by Myrtle 

E. Johnson. Pp. 53-88, plate 1. March, 1913 22.2. 

Sa Sagitta californica, n.sp., from the San. Diego Region, including 

S Remarks on Its Variation and Distribution, by Ellis L, Michael. 

‘ Pp. 89-12G plate: 2: “Tune; 191s a eh a eres 

6. Pycnogonida from the Coast cf California, with Description of Two 

New Species, by H. V. M. Hall. Pp. 127-142, plates 3-4. August, 1913. 

7. Observations on Isolated Living Pigment Cells from the Larvae of 

Amphibians, by 8. J. Holmes. Pp, 143-154, plates 5-6. 

8, Behavior of Ectodermic Epithelium of Tadpoles when Cultivated in 
3 Plasma, by 8. J. Holmes. Pp. 155-172, plates 7-8. 

ae Nos. 7 and 8 in one cover. September, 1913 -..2.22...ee see. 

5 9, On Some Californian Schizopeda, by H. J. Hansen. Pp. 173-180, pl. 9. 


¥ 


x 
oe 


Shale 


AN OWOMTDEN OTS ssc Sires ease ccc wnat cetacean sar ace sane v aru oueclsaernes 
= 10. Fourth Taxonomic Report on the Copepoda of the San Diego Region, 
eae ~ ‘by Calvin 0. Esterly, Pp. 181-196, pls. 10-12, November, 1913 ..._.... 
aa 11. The Behavior of Leeches with Especial Reference to Its Modifiability, 

A, The General Reactions of the Leeches Dina microstoma Moore and 

Glossiphonia stagnalis Linnaeus; B. Modifiability in the Behavior of 

the Leech Dina microstoma Moore, by Wilson Gee. Pp. 197-305, 13 

toxt-figures; “December, 1918-7 ae - cactctasewsen ctacectois ae 


1.00 


16 


15 


16 


2.00 


1.00 


UNIVERSITY OF CALIFORNIA. PUBLICATIONS— (Continued) 


12. The Structure of the Ocelli of Polyorchis penicillata, by Etta Viola 
Little. Pp. 807-328, plates 13-15. February, 1914 ..0000.2.c-eeccctecececeeeeetee : 

13. Modifications and Adaptations to Functions in the Feathers of Circus 
hudsonius, by Asa C. Chandler. Pp. 329-376, plates 16-20.- March, 
BB A Ss Sa aber Olea i sm Ry “cme Serge Ps rE rah Maho peas) lanes 

14. A Determination of the Economic Status of the Western Meadowlark 
(Sturnella neglecta) in California, by Harold Child Bryant. Pp. $77- ~~ 
510, plates 21-24, 5 text figures. February, 1914 222i... cecstececceeteoeee % 

15. Parasynaptie Stages in the Testis of Aneides lugubris (Hallowell), by 


Harry James Snook and J, A. Long. Pp. 511-528, plates 25-26, 1 text — 
fig April, TOE nae ee rc aes SSI a ee ce Sage megucoaana OES 
Vol, 12. 1. A Study of a Collection of Geese of the Branta canadensis Group from 
the San Joaquin Valley, California, by Harry 8. Swarth. Fp. 1-24, | 
plates 1-2, 8 text figs. November, 1913 2.0... ..ic.eecee ee cpetee cine eee cerceee EE Bi 
2. Nocturnal Wanderings of the California Pocket Gopher, by Harold C, 
Bryant. Pp. 25-29, 1 text fig. November, 1913 2222.00. om By 
3%. The Reptiles of the San Jacinto Area of Southern California, by Sarah { 
Rogers Atsatt. Pp. 31-50. November, 1913 ....0.000....c2. te 


ee 
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SNH 
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4. An Account of the Mammals and Birds of the Lower Colorado Valley, 
with Especial Reference to the Distributional Problems Presented, 
by Joseph Grinnell. Pp. 51-294, plates 3-13, 9 text figs. March, 1914. 
5. Aplodontia chryseola, a New Mountain Beaver from the Trinity Region ; 
of Northern California, by Louise Kellogg, Pp. 295-296. 
6. A Previously Undescribed Aplodontia from the Middle North Coast of 
California, by Walter P. Taylor. Pp. 297-300, 
Nos. 5 and 6. in one cover, April, 1914 22.0 eee _ 05 
7. A Second Species of the Mammalian Genus Microdipodops from Cali- es 
fornia, by Joseph Grinnell. . Pp, 301-304. April, 1914 00u..cc2. = 
8. Distribution of River Otters in California, with Description of a New 
Subspecies, by Joseph Grinnell. Pp. 305-310, plate 14. October, 1914 
9, Four New Pocket Gophers from California, by Joseph Grinnell. Pp. 
311-316, SINoVentber, TOL arr yes ioe and oa cane tanta 
10. Three New Races of Vespertilionid Bats from California, by Hilda 
Wood Grinnell, Pp. 317-321. December, 1914-2... tec cet 


Vol. 18. 1. The Schizopoda of the San Diego Region, by Calvin O. Esterly. Pp. — 
1-20, plates':1-2> \ April, 10145 eee a a ee eae 
2. A Study of the Occurrence and Manner of Distribution of the Cteno- 
phora of the San Diego Region, by Calvin O. Esterly. Pp. 21-38, — 
PN 58 Wega 9 6: Spe iiees i a cine Sela Se cet A A RL meee SU das Leet has aE <F 
3. A New Self-Regulating Paraffin Bath, by C. W. Woodworth. Pp. 39- 
42,2 text-tigures.: “April, (1914-25. eo oe ee, a eae 
4. Diplodinium ecaudatum, with an Account of Its Neuromotor ‘Apparatus, 5 
by Robert G. Sharp. Pp. 43-122, plates 3-7, 4 text figures. May, — 
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: ~ ZOOLOGY 
‘ vol. 12, No. i, pp- 321-328, 1text-Agure -- January. 20, 1915 


FROM THE INNER NORTHERN COAST 
BELT OF CALIFORNIA 


<i BY 
JOSEPH GRINNELL 
fies Le 
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IN 


ZOOLOGY 
Vol. 12, No. 11, pp. 321-325, 1 text-figure January 20, 1915 


EUTAMIAS SONOMAE, A NEW CHIPMUNK 
FROM THE INNER NORTHERN COAST 
BELT OF CALIFORNIA 


BY 
JOSEPH GRINNELL 


(Contribution from the Museum of Vertebrate Zoology of the University of California)~- 


The considerable series of chipmunks (genus Hutamias) in the 
California Museum of Vertebrate Zoology, accumulated from the coast 
district of California north of San Francisco Bay, consists of three 
distinet species. Two of these, Hutamias hindsi (Gray) and Eutamias 
townsendi ochrogenys Merriam, are already well known. The third is 
herewith described, and the opportunity is taken to give the distri- 
bution of all three species, as shown by the series of specimens at hand. 
In this connection, there have been available for comparison, through 
the courtesy of Mr. Henry W. Henshaw and Mr. Vernon Bailey, of 
the United States Biological Survey, forty-six additional specimens of 
Eutamias hindsi, from the national collection at Washington. Color 
names are taken from Ridgway’s (1912) Color Standards and Nomen- 


clature. 


Eutamias sonomae, new species 
Sonoma Chipmunk 

Type.— adult, in full summer pelage ; no. 20825, Mus. Vert. Zool. ; 
one mile west of Guerneville, Sonoma County, California; July 12, 
1913; collected by J. and H. W. Grinnell; orig. no. 2250. 

Diagnosis—A member of the townsendi group of chipmunks (see 
Merriam, 1897, p. 194) ; nearest like Hutamias hindsi, from which it 
differs in greater size, relatively longer tail, longer ears, less deeply 
ferruginous tone of coloration dorsally, whiter lower surface, the three 
dorsal black stripes narrower, outermost light stripes whiter, post- 


322 University of Califorma Publications in Zoology [ Vou. 12 


auricular spots larger and whiter, top of head and rump grayer in 
tone, and tail hairs distinctly tipped with gray. 

Description —Adult summer, or post-breeding, pelage (from type) : 
Under parts from chin to base of tail, white, with creamy tinge mid- 
ventrally; insides of thighs and arms, lght ochraceous-buff; upper 
surface of feet, dull ochraceous-tawny; sides and general suffusion 
over back, bright cinnamon-rufous, approaching apricot-orange in tone 
along the lower sides; narrow black median stripe from between ears 
to rump, fading out near base of tail; rest of dorsal stripes much 
shorter, not extending forward of shoulders; median dorsal stripe bor- 
dered on each side by a gray line, this much obscured with ferrugin- 
ous; outwardly of gray stripe on each side is a black stripe dulled 
by intermixture of ferruginous hairs; succeeding this outwardly is a 
conspicuous clear ashy-white stripe on each side, and outwardly to 
this a much shorter stripe of dusky along the upper edge of the 
otherwise bright cinnamon-rufous side. There are thus three black 
stripes and four light stripes, the outermost pair of which is con- 
spicuously whitish (much more clearly so than in hindsi). Rump and 
flanks, grizzled Brussels brown; tail beneath, bright Sanford’s brown, 
outwardly margined with black and ashy white; tail dorsally black, 
with a little of the ochraceous-orange bases of the hairs showing 
through, and grizzled with the ashy-white hair tippings. Ears lightly 
lined inwardly with hazel hairs; front half of outer surface of ear, 
dull cinnamon-rufous, posterior half light mouse gray; triangular spot 
behind each ear, clear white, slightly tinged with ashy, but much more 
conspicuous than in hindsi; top of head grizzled grayish brown; head 
stripes more brightly contrasted with each other than in either hindsi 
or ochrogenys; stripes from nose to base of ear, above and below eye, 
white, slightly dulled’ with black and ferruginons hairs; a dusky stripe 
above each whitish supra-ocular stripe, becoming blacker anteriorly 
and meeting its fellow on tip of nose; a black stripe, dulled by fer- 
ruginous, from eye to base of ear; a similarly colored but narrower 
stripe beneath the whitish subocular stripe; a dull brownish suffusion 
between eye and snout; whiskers black. Measurements of type: total 
length, 260 mm.; tail vertebrae, 112; hind foot, 36; ear from crown, 
17. Thirty adult specimens of sonomae average, as measured by field 
collectors: total length, 251 mm.; tail vertebrae, 113; ratio tail to 
body, 82 per cent. Eighteen adults of hindsi average: total length, 
232 mm.; tail vertebrae, 102; ratio tail to body, 78 per cent. Cranial 
differences are not apparent. Breeding (worn winter) pelage: pattern 


1915] Grinnell: A New Chipmunk 323 


the same as above, but much duller in tones of color and hence with 
less of contrast; compared with corresponding stage in hindsi, grayer, 
less reddish; lower surface white instead of buffy; brown of tail much 
less reddish in tone, more ochraceous; dimensions diagnostic. 
Distribution—Of Eutamias sono- 
mae there are 68 specimens in the 
Museum of Vertebrate Zoology, repre- 
senting localities as follows: 5 miles 
south of Kunz, Trinity County, 1; 
Line Creek, 5500 ft. alt., 1 mile east 
of Castle Peak, near Trinity County 
line, Mendocino County, 1; 3 miles 
south of Covelo, Mendocino County, 
9; 3 miles west of summit of Sanhe- 
drin Mountain, 4500-6000 ft. alt., 
Mendocino County, 22;  Lierly’s 
Raneh, 2340 ft. alt., 4 miles south 
of Sanhedrin Mountain, Mendocino 
County, 2; 6 miles north of Willits, 
1400 ft. alt., Mendocino County, 2; 
Rumsey, 500 ft. alt., Yolo County, 3; 
8 miles west of Vacaville, Solano 


County, 5; 1 mile west of Guerneville, 


Sonoma County, 6; 7 miles west of 
Cazadero, Sonoma County, 15; Free- a E. t. ochrogenys 
stone, Sonoma County, 2. 4 E. sonomae 
Of Eutamias townsendi ochrogenys e E. hindsi 

there are 129 specimens in this Mu- 
seum, distributed as follows: Trini- 
dad, Humboldt County, 4; Areata, Fig. 1. Map of portion of north- 
Humboldt County, 1; Freshwater, ester Ca ormay ner Eromuoan 


Humboldt County, 2; Eureka, Hum- 


Francisco Bay, showing record 
stations and approximate ranges 
boldt County, 7; Fair Oaks, Hum- of Eutamias townsendi ochrogenys, 
boldt County, 17; C@uddeback Hum. Lutamias sonomae, and Hutamias 
: i 4 hindsi. 

boldt County, 5; Rockport, Mendocino 
County, 1; Mendocino City, Mendocino County, 22; Sherwood, Men- 
docino County, 23; 5 miles north of Willits, Mendocino County, 1; 
Gualala, Mendocino County, 23; 
Sonoma County, 12; Freestone, Sonoma County, 11. 

Of Eutamias hindsi there are available, from the collections of this 


tl 


miles west of Cazadero, 900 ft. alt., 


324 University of California Publications in Zoology | Vou. 12 


Museum and of the United States Biological Survey, 62 specimens, as 
follows, all from Marin County: Vicinity of Point Reyes, 3 to 5 
miles west of Inverness, 12; Inverness, 5; Olema, 15; Lagunitas, 2; 
‘‘Nicasio’’ (probably in near vicinity of San Geronimo), 26; Mail- 
hard, 2. 

Relationships and Ecology—Eutamias sonomae shows itself to be- 
long primarily to the Upper Sonoran and lower Transition zones, 
chiefly in the chaparral association, though locally invading the mar- 
gins of the forest. The species further belongs to what has been called 
the Clear Lake subfauna, namely, the aggregation of animal species 
occupying the semi-humid inner coast ranges and intervening valleys, 
lying between the narrow coastal Redwood fauna on the west and the 
Sacramento fauna on the east, and extending from the vicinity of San 
Pablo and Suisun bays north into western Trinity County. Hutamias 
hindsi has similar zonal and associational restriction, but belongs to a 
more humid area, namely, that oceupied by the Marin subfauna. This 
is a section of the coast belt proper, lying entirely within Marin County, 
and extending from Point Reyes eastwardly to include the slopes and 
foothills of Mount Tamalpais. Eutamias townsendi ochrogenys be- 
longs to the Transition and Boreal zones in their narrow, humid-coast, 
faunal divisions. It is a dweller in chaparral and forest, and has not 
been found south of Freestone, Sonoma County. At the latter point 
sonomae and ochrogenys have been taken in the same lines of traps, as 
also at a point about seven miles west of Cazadero, Sonoma County. 
The ranges of these two chipmunks thus not only meet, but actually 
overlap to a slight extent. On the other hand, the range of hindsi is 
separated from the ranges of both ochrogenys and sonomae by a belt 
of country apparently unfit associationally for the existence of any 
one of this group of chipmunks. Hindsi is thus quite isolated from 
any of its relatives (see fig. 1). 

It may be remarked that hindsi appears to be much nearer in agere- 
gate of characters to ochrogenys than is sonomae. The latter is thus 
an extreme departure from the townsendi stock. Sonomae reminds 
one in several respects of Hutamias quadrimaculatus of the western 
flank of the central Sierra Nevada, and the species just named may 
well be a still further manifestation in the fownsendi series. 

Eutamias merriami pricei, of the Santa Cruz subfaunal area, be- 
longs also to the townsendi group. It resembles sonomae in general 
aspect, but differs constantly and markedly in smaller ear, in dull 
gray instead of white postauricular spot, in light gray instead of 


1915] Grinnell: A New Chipmunk 325 


whitish outer side stripe, and in grayer, less brightly reddish, general 
tone of coloration dorsally. In the latter respect price? differs still 
more strongly from hindsi. The ranges of the two chipmunks last 
mentioned are separated by San Francisco Bay and the Golden Gate, 
together of course with adjacent strips of territory associationally 
unfit for habitation by chipmunks. 


Transmitted October 31, 1914. 


LITERATURE CITED 


MERRIAM, C. H. 
1897. Notes on the chipmunks of the genus Hutamias occurring west of the 
east base of the Cascade-Sierra system, with deseriptions of new 
forms. Proc. Biol. Soe. Wash., 11, pp. 189-212. 


esr ,,rlt (‘a lS 


UNIVERSITY OF CALIFORNIA PUBLICATIONS— (Continued) 


4,.The Marine Biological Station of San Diego, Its History, Present Con- 
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5. Oxygen and Polarity in Tubularia, by Harry Beal Torrey. Pp. 249- 


ae 
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‘ 
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. Vol. 10. (Contributions from the Museum of Vertebrate Zoology.) 

ne 1. Report on a Collection of Birds and Mammals from Vancouver Island, 
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<c 


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the Leech Dina microstoma Moore, by Wilson Gee. Pp. 197-305, 13 
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PAMUATY, TOLD ee eR ee Be SRN Be ent eT lia epee 


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6. The Anatomy of Heterodontus Francisci. I. The Exoskeleton, by J. 
Frank Daniel. Pp, 147-166, plates 8-9, 4 text figures, May 23, 
OTA Fr Oe Ace Eo eae Ns MO AAS Sn Eee ee ae ee es 
7. The Movements and Reactions of the Isolated Melanophores of the 
Frog, by S. J. Holmes. Pp, 167-174, plate 10. August, 1914 22.0... 1 
8, Polychaetous Annelids of the Pacific Coast in the Collections of the 
Zoological Museum of the University of California, by Aaron L. 
Treadwell. Pp. 175-234, plates 11-12. 
9. New Syllidae from San Francisco Bay (collected by the U.S. S. 
** Albatross’’), by Aaron L. Treadwell. Pp. 235-238, 7 text figures. 
Nos..8 and 9 in one cover. October, 1914 c2.n.. cc cence leceeccteeetemeeeeeeee 
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B. Bigelow. Pp. 239-241. September, 1914 .2...2.e2.tt eel cccce ceeee Ss 


1. A Report upon the Physical Conditions in San Francisco Bay, Based 
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tross’’. during the Years 1912 and 1913, by F. B. Sumner, G. D. 
Louderback, W. L. Schmitt, E. C. Johnston. Pp. 1-198, plates 1-153, 
20. text figured, Sully); 1904 ss re a cca senernetctiin ig 


3 
thomas sah fn it Moi glance fence saguiees ae SH 


. 
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ie UNIVERSITY OF CALIFORNIA PUBLICATIONS 

~ ; IN 

os ZOOLOGY 

. Vol. 12, No. 12, pp. 327-334 April 2, 1915 
ps Rag Ss ORES RA SO SERRE I 9 AO REN SE Me RS IS SEE 
ed 


BATRACHOSEPS MAJOR AND BUFO COGNATUS 
* CALIFORNICUS, NEW AMPHIBIA FROM 
3 SOUTHERN CALIFORNIA 


BY 


CHARLES LEWIS CAMP 


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BERKELEY 


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UNIVERSITY OF CALIFORNIA PUBLICATIONS 
IN 


ZOOLOGY 
Vol. 12, No. 12, pp. 327-334 April 2, 1915 


BATRACHOSEPS MAJOR AND BUFO COGNATUS 
CALIFORNICUS, NEW AMPHIBIA FROM 


SOUTHERN CALIFORNIA gcenien Institgp 
“es i 


BY aa MAY * 1935 


CHARLES LEWIS CAMP 
Nay 


(Contribution from the Museum of Vertebrate Zoology of the University of California) 


; Wy 
ional Muse 


During the author’s examination of the amphibia from southern 
California contained in the collection of the Museum of Vertebrate 
Zoology, two forms have been distinguished which seem to deserve 
recognition under new names. 

The occurrence of Bufo cognatus almost to the Pacifie seaboard 
in California is of considerable interest, the previous westernmost 
published record being from the Colorado River. The detection of 
a new species of Batrachoseps in southern California is not to be 
wondered at, considering the obseurity of the descriptions in the 
literature relating to this genus. This new salamander is fairly 
common in the western part of the San Gabriel Valley, especially in 
the district immediately surrounding Pasadena. 


Batrachoseps major, new species 
Garden Salamander 

Type.—Adult; no. 611, Mus. Vert. Zool. ; Sierra Madre, 1000 feet 
altitude, Los Angeles County, California; March 14, 1909; collected 
by C. L. Camp; orig. no. 218. 

Diagnosis—A large, pale, long-limbed Batrachoseps with light 
yellow underparts. Costal folds 18, rarely 17 or 19. 

Material—Twenty-nine specimens from Pasadena and Sierra 
Madre, California; twelve from the latter and seventeen from the 
former locality ; nos. 611, 954-956, 4566-4586, Mus. Vert. Zool. 


328 University of California Publications in Zoology — [Vou. 12 


Comparisons.—This species is intermediate in many of its char- 
acters between Batrachoseps pacificus from the northern Channel 
Islands and Batrachoseps attenuatus of the Pacifie Coast district. 
Size larger than any of the other species of Batrachoseps, not quite 
so slender as attenuatus; head wider; tail shorter and limbs longer 
than in attenuatus; head narrower, tail longer and limbs shorter than 
in pacificus. Van Denburgh (1905, p. 8) states that the number of 
costal grooves in B. pacificus is 17, rarely 16 or 18; this would make 
the number of costal folds in that species 16, rarely 15 or 17. 

To avoid confusion, the system of enumerating the costal folds as 
here employed should be set forth. For obvious reasons the costal 
grooves cannot be so accurately ascertained as the folds. The limbs 
may start in a groove or on a fold and so there may be in many eases 
fractional folds. These are never considered, only complete folds 
bounded on each side by grooves being taken into account. This makes 
the number of grooves always one more than the number of folds. 

The costal folds in B. major number 18, rarely 17 or 19, the 
number of folds in B. attenwatus (from points in southern and middle 
California) is 18 or 19, rarely 17 or 20, the number in B. caudatus 
(Hassler Harbor, probably on Annette Island, southeastern Alaska) 
from the record of Cope (1889, p. 126) is 20 (21 grooves). 

B. pacificus, with a wide head, short tail and body, longer limbs 
and few costal folds, stands nearest the Plethodon group of sala- 
manders, while B. caudatus, the most specialized member of its genus, 
with a narrow head, shorter legs, slender body and long tail, lies, both 
structurally and geographically, at the opposite end of the scale. 
B. attenuatus is intermediate; and B. major, representing an appar- 
ently restricted local race, is between pacificus and attenuatus in its 
proportional measurements (see table, p. 330). 

The coloration of the present species resembles that of pacificus 
more than that of attenuatus. It is distinet from attenwatus by 
reason of its pale color, especially of the ventral parts, which are 
yellow and never gray except in narrow transverse areas between the 
limbs. The dorsal surfaces are slightly paler than in B. pacificus. 
In Van Denburgh’s redeseription (1905, p. 8) of pacificus no mention 
is made of darker areas between the limbs on the ventral surface such 
as exist In major. 

Description of type-——Body and tail elongate, cylindric, annulated ; 
tail conical at tip, stout throughout its length, longer than body; head 
wider than neck, flat above, narrower than body in abdominal region ; 


1915] Camp: New Amphibia from Southern California 329 


fore and hind limbs do not meet when pressed to sides of body; 
digits rudimentary, four on both front and hind feet; nostrils sep- 
arated by nearly twice their distance from the orbits, not terminal, 
connected with upper lip by thick-edged grooves; gular fold not 
plainly marked ; body divided into eighteen folds, or segments, between 
the front and hind limbs; skin very smooth and shiny; openings of 
small pores barely visible on head region. Color in alcohol (from 
Ridgway’s Color Standards, 1912) light neutral gray above; sides, 
lower parts, upper lip, palms and soles near cream buff; slightly 
darker on under surface of tail and on ventral surface of body 
transversely between limbs. 

Variations.—In the series of twenty-five specimens at hand the 
costal folds number 17 in four, 18 in seventeen, and 19 in three. In 
one specimen there are 17 folds on one side and 18 on the other side 
of the body. In one specimen the outside (fourth) digit of the left 
front foot is lacking. 

Distribution—This large, light-colored Batrachoseps has been 
taken on south Euclid Avenue beneath boards in a yard, and also in 
a cellar, in Pasadena, California. The type was found in the neigh- 
boring town of Sierra Madre under a broken piece of cement side- 
walk, and others taken in the same vicinity were captured in piles 
of damp lumber and in post-holes. Two were taken in August, 
1905, several feet beneath the surface of the ground in loose gravel 
in a ravine bottom. The localities of capture lie in the upper edge 
of the Lower Sonoran life-zone (mesa oak association) and below the 
range of Batrachoseps attenuatus. The latter species appears to 
inhabit the Upper Sonoran zone (maple-sycamore association) in 
the mountain cafions, possibly getting out into the valleys occasionally 
along water courses. Both species are entirely terrestrial and both 
appear to estivate during the drier months, being then seldom found 


above ground. 


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1915] Camp: New Amphibia from Southern California 331 


Bufo cognatus californicus, new subspecies 
Arroyo Toad 


Type—Female, adult; no. 4364, Mus. Vert. Zool.; Santa Paula, 
800 feet altitude, Ventura County, California; May 22, 1912; collected 
by C. L. Camp; orig. no. 551. 

Diagnosis —A toad with divergent head crests, a nasal boss, short, 
shghtly divergent parotoids and with an internal, cutting tubercle on 
hind foot; femur short as in Bufo cognatus cognatus, the Great Plains 
toad. Size medium; parotoids wide; coloration nearly uniform, with- 
out large spots; no vertebral streak; external tubercle on hind foot 
small and rounded, not provided with a cutting edge. 

Material—Two alcoholic specimens: the type; and adult female, 
no. 767, Mus. Vert. Zool.; Tujunga Wash, near Sunland, Los Angeles 
County, California; April 1, 1904; collected by J. Grinnell. 

Comparisons—This Bufo is distinet from the common toad 
(Bufo halophilus) of the Los Angeles region, differing from it: in 
the possession of both transverse and longitudinal cranial crests, in 
the smaller size, in the absence of a vertebral stripe, in the thick 
head, and in the more even character of the tubereulation of the 
back. It is clearly most closely related to Bufo cognatus cognatus 
Say (1828, 2, p. 190) of Arizona and the Great Plains, east of the 
Rockies. 

The type locality of the latter is the Arkansas River in Colorado, 
probably between the present site of La Junta, Colorado, and the 
Colorado-Kansas boundary. Bufo cognatus ranges over a large part 
of the Great Plains district east of the Rocky Mountains, in Colorado, 
Kansas, Arkansas, New Mexico, Texas, Nebraska, Montana, and South 
Dakota. West of the Rockies it occurs in northern Mexico, Arizona, 
and, in southeastern California, along the Colorado River and in the 
Salton Basin. 

The two specimens here included in the subspecies californicus 
are the first representatives of this species known from the Pacific 
coast of California. They differ from a series at hand of Bufo 
cognatus taken on the Colorado River, in California and Arizona, in 
the slightly longer hind foot, in the lack of an external cutting 
tubercle, in the width of the parotoids, which are 10 to 20 per cent 
broader in californicus, and in the type of coloration, which is uniform 
and without a trace of the large green spots, so pronounced a feature 
of the coloration of most of the Colorado River examples. 


332 University of California Publications in Zoology — [Vou. 12 

From Bufo lentiginosus woodhousti, of the Lower Colorado River 
district and farther east, the new toad differs decidedly in its smaller 
size, short hind legs, divergent cranial crests, and in the presence of 
a bony nasal elevation. 

Description of type.—Size medium; hind legs very short, femur 
almost entirely enclosed in skin of abdomen; head short and thick; 
nasal region elevated into a bony protuberance; longitudinal cranial 
crests more or less united across median region, and slightly divergent ; 
transverse crests divided by width of median groove; parotoids oval, 
slightly divergent and very broad; inner tubercle of hind foot with 
a sharp edge; outer tubercle very small, rounded and without cutting 
edge; eyelids and back evenly tuberculated; tympanum oval, shorter 
than diameter of eye; a dozen or more large whitish tubercles below 
and just posterior to the tympanum. The color when taken was 
light olive-green above, with two light patches about six millimeters 
in diameter immediately behind the parotoids. Upper lip below the 
eye barred vertically with paler tone; light areas present on eyelids 
and anterior ends of parotoids. No vertebral stripe present; under- 
parts creamy white, unspotted. Bases of several of the tubercules on 
the back encircled by small black rigs, and no large greenish spots 
‘in the dorsal coloration. 

Distribution—The type was found on a lawn in the middle of 
town, about eight o’clock in the evening. The Santa Paula district 
is about sixty miles east of the San Fernando Valley (where the other 
specimen of this subspecies was found), and a mountain spur about 
one thousand feet in elevation separates the two localities of capture. 
Both localities lie within the San Diegan faunal area in the Lower 
Sonoran life-zone, as the latter has been delimited in California. 
Both localities are of semi-arid character, and the streams are inter- 
mittent in the arroyos near which each of these toads was captured. 


333 


€ 
e 


rom Southern Califorma 


j 


1 


D 


New Amph 


. 


MEASUREMENTS IN MILLIMETERS OF THE Two SUBSPECIES OF Bufo cognatus 


B g 4g 

ep bo ro b oa x 
a a ee Gee es 
= Zz 3 Zz S I ws SS 
Mus. Ce ee ee ae ee 
oo: Sex Locality? Date a | | q B iq Sige Bae, 

Bufo cognatus californicus 
4364 9 Santa Paula, Ventura Co. May 22, 1912 64.5 19.2 23.55 844 263 41.5 105 7.8 
767 2 ‘Tujunga Valley, Los Angeles Co. Apr. 1, 1904 Sip) ISON S22TGie BSt8 eb .Ole e419) aed: 
Bufo cognatus cognatus 

1139 g Needles, San Bernardino Co. July 16, 1909 60.2 18.0 22.7 804 25.0 38.7 8.8 5.5 
4361 ¢ Needles, San Bernardino Co. July 16, 1909 59.1 17.9 21.8 75.5 23.6 36.3 8.9 4.9 
4358 9 Needles, San Bernardino Co. July 16, 1909 60.3 19.0 23.8 81.8 24.7 40.1 9.6 5.2 
4360 2 Needles, San Bernardino Co. July 16, 1909 6255-1914" 122'6 780) 25.18 38:9 9.0 5.5 
4363. @ Needles, San Bernardino Co. July 16, 1909 62.5 19.1 23.0 81.6 25.9 38.5 9.2 4.6 
1137 @ Needles, San Bernardino Co. July 15, 1909 Feb. le OQie3) GIS -28u 36:6 8.5 5.2 
1134. @ Needles, San Bernardino Co. July 16, 1909 63.7 20.0 22:9 78.89 25.5 38.07 9.8 5.6 
457 2 Mecca, Riverside Co. Apr. 21, 1908 64.9 19.4 240 79.4 26.0 37.8 95 6.1 
210701 g Ft. Huachuea, Arizona July, 1893 60.0 17.7 21.6 5 Phill BN alls} Ges) 
210651 9 Animas Valley (Colorado?) Sept. 9, 1895? 89.8 223 28.8 96.6 318 495 121 7.2 
25621 ¢@ Red River, Arkansas 88.5 25.0 31.5 118.1 338.6 544 145 81 


Camp 


1915] 


1From collections of United States National Museum. 
?In California unless otherwise stated. 
8 Tip of snout to posterior border of pubic symphysis. 


*From pubie symphysis. 
* Including tarsus. 


B34 University of California Publications in Zoology — [Vou. 12 


LITERATURE CITED 
Corr, E. D. 
1889. The batrachia of North America. U.S. Nat. Mus., Bull., 34, 525 pp., 
86 pls., 119 figs. in text. 


Say, T. 

1823. In Long, Stephen H., Account of an expedition from Pittsburgh to 
the Rocky Mountains performed in the years 1819 and ’20 by order 
of the Hon. J. C. Calhoun, See’y of War; under the command of 
Major Stephen H. Long from the notes of Major Long, Mr. T. Say, 
and other gentlemen of the exploring party. Compiled by Edwin 
James, botanist and geologist for the expedition. (Philadelphia), 
2 vols. [+ atlas], 503 + 442. 


VAN DENBURGH, J. 
1905. The reptiles and amphibians of the islands of the Pacifie Coast of 
North America from the Farallons to Cape San Lueas and the 
Revilla Gigedos. Proce. Calif. Acad. Sei. (3), 4, 1-40, pls. 1-7. 


Transmitted February 9, 1916. 


UNIVERSITY OF CALIFORNIA PUBLICATIONS—(Continued) 


Vol, 10. (Contributions from the Museum of Vertebrate Zoology.) 


3. The Mole of Southern California, by J. Grinnell and H. S. Swarth. 
Pp. 131-136, 2 text-figures. 
4. Myotis orinumus Elliott, a Bat New to California, by J. Grinnell and 
Dan Hi. 8. Swarth. Pp. 187-142, 2 text-figures. 

. Nos, 3 and 4.in one cover. April, 1912 occ ccc ccccsccccccocessceecceeceaece 
te 5. The Bighorn of the Sierra Nevada, by Joseph Grinnell. Pp. 143-153, 
A VOSPUAUTOR “DA Yh od Olean ied Stee cca Ragan Seeger 
6. A New Perognathus from the San Joaquin Valley, California, by 

Walter P, Taylor. Pp. 155-166, 1 text-figure. 
fy Sag of West Central California, by Walter P. Taylor. Pp. 

-169. 

Nos, 6 and 7 in one cover. May, 1912 .......c2ccecccceceecccesetseseoeeseae 
8. The Two Pocket Gophers of the Region Contiguous to the Lower Colo- 
rado River, in California and Arizona, by Joseph Grinnell. Pp, 171- 
2 5 71s ett fats. Sark By by paw eo mane © renee a ae, oo ant SGN Sy Cua ASN ME 
ies 9. The Species of the Mammalian Genus Sorex of West-Central Cali- 
5 ; fornia, with a note on the Vertebrate Palustrine Faunas of the 
Region, by- Joseph Grinnell, Pp. 179-195, figs. 1-6. March, 19183 ........ 
10, An Account of the Birds and Mammals of the San Jacinto Area of 
Southern California, with Remarks Upon the Behavior of Geographic 
Races on the Margins of Their Habitats, by J. Grinnell and H. 8. 


Index, pp. 407-417. 


Vol. 11. 1. Birds in Relation to a Grasshopper Outbreak in California, by Harold 

Ox Bryant:.Pp: 1-20:>: November, 19120 3.02 ok na 

2. On the Structure and Relationships of Dinosphaera palustris (Lemm.), 

by Charles Atwood Kofoid and Josephine Rigden Michener. Pp. 21- 

ee FEOCeM DER, UBT aS Te OT A oe angpareaterbingte cg tape ceibenasn = 

8. A Study of Epithelioma Contagiosum of the Common Fowl, by 

_ Clifford D. Sweet. Pp, 29-51, January, 1913 2 lcci eceteee 

4, The Control of Pigment Formation in Amphibian Larvae, by Myrtle 

E. Johnson, Pp. 53-88, plate 1. March, 1913-20... eee ne 

5, Sagitta californica, u.sp., from the San Diego Region, including 

Remarks on Its Variation and Distribution, by Ellis L. Michael. 

Pp: 89-126, plate Bi TUS LOLS en A cass sacgncenunann vastly euadeasaweeanoee 

Pycnogonida from the Coast of California, with Description of Two 

New Species, by H. V. M. Hall. Pp. 127-142, plates 3-4: August, 1913. 

; 7. Observations on Isolated Living Pigment Cells from the Larvae of 
Se Amphibians, by 8. J. Holmes. Pp,.148-154, plates 5-6. 

: Behavior of Ectodermic Epithelium of Tadpoles when Cultivated in 

Plasma, by 8..J. Holmes. Pp. 165-172, plates 7-8. 

Nos. 7 and 8 in one cover. September, 1913 —..-.2-..2..---.--egeeeeeeeee 

9. On Some Californian Schizopoda, by H. J. Hansen. Pp. 173-180, pl. 9. 

‘ Ped syen bates Aga thes Ie es Rata get ten aes SR ESR ah Sree SOc pe Raburn ee tke ep 

Ber 10, Fourth ©:axonomic Report on the Copepoda of the San Diego Region, 

: by Calvin 0. Esterly. Pp. 181-196, pls. 10-12. November, 1913 _....... 

% -_>11. The Behavior of Leeches with Especial Reference to Its Modiflability, 

ah : A, The General Reactions of the Leeches Dina microstoma Moore and 

RA Glossiphonia stagnalis Linnaeus; B. Modifiability in the Behavior of 

the Leech Dina microstoma Moore, by Wilson Gee. Pp. 197-305, 13 

text. figures, Deceniber,: 1913 xe rae naar 


Sy) 


Swarth. Pp. 197-406, pls. 6-10. October, 1913 0.2 " 


10 


15 


6 


UNIVERSITY OF CALIFORNIA PUBLICATIONS—(Continued) 


12. The Structure of the Ocelli of Polyorchis penicillata, by Etta Viola 


Little. Pp. 307-328, plates 13-15. February, 1914 0.0... eeeeeeeee 
18. Modifications and Adaptations to Functions in the Feathers of Circus 


hudsonius, by Asa C. Chandler. Pp. 329-376, plates 16-20. March,. 


BP OY cece cg ie Gs Beem eae ene RRM ee OE CS RR Packs By 3 Nene any Lbs SCNT A 
14. A Determination of the Economic Status of the Western Meadowlark 
(Sturnella neglecta) in California, by Harold Child Bryant. Pp. 377- 
510, plates 21-24, 5 text figures. February, 1914 o22oo.2.tn2 2c. nesssceeens 
Parasynaptic Stages in the Testis of Aneides lugubris (Hallowell), by 
Harry James Snook and J. A. Long. Pp. 511-528, plates 25-26, 1 text 
fig ADT -TO TA pas Sas EO eS a oka ee cacao 


Vol. 12. 1. A Study of a Collection of Geese of the Branta canadensis Group from 
the San Joaquin Valley, California, by Harry S. Swarth. Pp. 1-24, 
plates 1-2, 8 text figs. November, 1913 2 .ci cic. cece le eceeeecencceomeene 

2. Nocturnal Wanderings of the California Pocket Gopher, by Harold 6. 
Bryant. Pp. 25-29, 1 text fig. November, 1913 02... cccc tect eceneeeecsneee 
$8. The Reptiles of the San Jacinto Area of Southern California, by Sarah 
Rogers Atsatt. Pp. 31-50. November, 1913 oo2.2.....cccececclicceeceetecereccarneee 
4. An Account of the Mammals and Birds of the Lower Colorado Valley, 
with Especial Reference to the Distributional Problems Presented, 
by Joseph Grinnell. Pp. 51-294, plates 3-13, 9 text figs. March, 1914, 


15 


5. 


of Northern California, by Louise Kellogg. Pp. 295-296. 
6. A Previously Undescribed Aplodontia from the Middle North Coast of 
California, by Walter P. Taylor. Pp. 297-300. 
Nos. 5 and 6 in one cover, April, 1914 cc.e... elec tceenteecennseeseeee 
A Second Species of the Mammalian Genus Microdtpodops from Cali- 
fornia, by Joseph Grinnell. Pp. 301-304. April, 1914 2c. 
8. Distribution of River Otters in California, with Description of a New 
Subspecies, by Joseph Grinnell. Pp. 305-310, plate 14. October, 1914 
9. Four New Pocket Gophers from California, by Joseph Grinnell. Pp. 
S11-816;.:: November): 19) 45 5 See ee a me Sm 


ra 


10. Three New Races of Vespertilionid Bats from California, by Hilda 


Wood Grinnell, Pp. 317-321. December, 1914-2022 ..ccccccccesk eee tene 


11. Eutamias sonomae, a New Chipmunk from the Inner Northern Coast 


Belt of California, by Joseph Grinnell. Pp. 321-325, 1 text figure. 
DANUALY, AOU BR i cca air Sar AE Sa 
12. Batrachoseps major and Bufo cognatus  californicus, New Amphibia 
from Southern California, by Charles Lewis Camp. Pp. 327-334. 
April TOL eas Ae Nh eee AO Th ee ca Ap needa Raa 


Vol. 19. 1. The Schizopoda of the San Diego Region, by Calvin O. Hsterly. Pp. 
1-29;-plates 1-2. April, 1914 te 


2. A Study of the Occurrence and Manner of 
phora of the San Diego Region, by Calvin O. Esterly. Pp. 21-38. 
We) 3 eee hh Sete eee ee een, cg Ang a oe eae RN NPS Ds Spal AOE pec Rae Qeeala te 

8. A New Self-Regulating Paraffin Bath, by C. W.. Woodworth. Pp. 39- 

42; 2-textfigures. April, 1014 nc shen oT ee ee, 

Diplodinium ecaudatum, with an Account of Its Neuromotor Apparatus, 

by Robert G. Sharp. Pp. 43-122, plates 3-7, 4 text figures. May, 


4 


6. The Vertical Distribution and Movements of the Schizopoda of the 
San Diego Region, by Calvin O. Esterly.. Pp. 123-145. May, 1914...... 
6. The Anatomy of Heterodontus Francisci. I. The Exoskeleton, by J. 
Frank Daniel. Pp. 147-166, plates 8-9, 4 text figures. May 23, 


01 I LA Nae AE ct a BOW Ete Seo NMI SBE INY OF Ge Bares SMR AB Bie cain m. Shae tp ey Be Ts 
7. The Movements and Reactions of the Isolated Melanophores of the 
Frog, by S. J. Holmes. Pp. 167-174; plate 10. August, 1914 0.02... 


8. Polychaetous Annelids of the Pacific Coast in the Collections of the 
Zoological Museum of the University of California, by Aaron L. 
Treadwell, Pp. 175-234, plates 11-12. 

9. New Syllidae from San Francisco Bay (collected by the U.S. S. 
‘‘Albatross’’), by Aaron L. Treadwell. Pp. 235-238, 7 text figures. 

Nos, 8 and 9 in one cover. October, 1904 ooo... cece tee ceecceeenensenene 

10. Note on the Medusan Genus Stomolophus, from San Diego, by Henry 

B. Bigelow. Pp. 239-241. September, 1914 ooi..etlc ee yeeee | peeeenee 

Vol. 14. 1. A Report upon the Physical Conditions in San Francisco Bay, Based 
upon the Operations of the United States Fisheries Steamer ‘‘Alba- 

tross’’ during the Years 1912 and 1913, by F. B. Sumner, G. D. 

Louderback, W. L. Schmitt, E. CO. Johnston. ‘Pp. 1-198, plates 1-13, 

20 text figures. July, 1914 co... ccs ccccinccccccecceece ees cedenece igheSue Bb Seas agora ft 


Aplodontia chryseola, a New Mountain Beaver from the Trinity Region - 


05 


2.25 


-UNIVERSITY OF CALIFORNIA PUBLICATIONS - 
IN 
ZOOLOGY 


Vol. 12, No. 13, pp. 335-398, plates 15-18 
Vol. 12, No. 14, pp. 399-410 


Issued January 27, 1916 


13. REPORT UPON MAMMALS AND BIRDS 
FOUND IN PORTIONS OF TRINITY, 
SISKIYOU AND SHASTA 
COUNTIES, CALIFORNIA 


BY 
LOUISE KELLOGG 


14. AN ANALYSIS OF THE VERTEBRATE 
FAUNA OF THE TRINITY REGION OF 
NORTHERN CALIFORNIA 


BY 
JOSEPH GRINNELL 


, s { 
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PUD BPS SLAB ao Soa ccgs cas eae sara sn ine laces ener See 


UNIVERSITY OF CALIFORNIA PUBLICATIONS 
IN 


ZOOLOGY 


Vol. 12, No. 13, pp. 335-398, plates 15-18 Issued January 27, 1916 


REPORT UPON MAMMALS AND BIRDS FOUND 
IN PORTIONS OF TRINITY, SISKIYOU AND 
SHASTA COUNTIES, CALIFORNIA, WITH 
DESCRIPTION OF A NEW DIPODOMYS 


BY 


LOUISE KELLOGG 


. 


(Contribution from the Museum of Vertebrate Zoology of the University of California) 


aes \ 
: rs ~&tion V 
CONTENTS SS 
PAGE 
ATRISIEGTS (3 GATT Uy) Teese mrss ete ee eee ke nes coe ee SOE et Sass cocoa ace be sou ce be coe oct Soc tacmucewaptaeliace 335 
MPG LIV GAGs 0 secs c cee ace se cee cea sne conn e a Sedona e stneecaneaaesnccsecvesecuee socussdcecte SacbessoaccieteccteesSolecdentueescosse 336 
SYOMVEXCH DEY areca dorccsclae eee A Sa e oe rr oe ee ere > ee 336 
Description of route Spooks 
(CHEE) THE aR ALS ee ya ONES ee 350 
Generaleaccounitswot thenmamimal s! 22<-22-2)2.< -c2a-. ce oe coc oence ace acanseecencune-beekaeeoapenceene 351 
List of the birds with annotations - ee OH) 
ID ahiG TERT) “COMET | oe seedecoccec cere eae eee eee ee eee ee eee hae ete eee nee 389 


INTRODUCTION 


The Trinity, Salmon and Scott mountains form a chain lying in 
an intermediate position between the Sierra Nevada and the Coast 
Ranges, on the east and west, respectively, and merging through 
the Siskiyou Mountains at the north into the Cascade Range. The 
river systems which drain this chain of mountains occupy valleys 
somewhat similar to those which furrow the western flank of the 
Sierra Nevada. The streams all flow ultimately in a_ westerly 
direction and finally meet the Klamath River, which forms their 
common outlet to the seacoast. There is thus offered an interest- 
ing problem in animal distribution; for the faunas of the Sierra 
Nevada to the east, the Coast belt to the west, and the Cascades 


336 Unwersity of California Publications in Zoology [Vow 12 


to the north, are decidedly different from one another. The fauna 
of the Trinity region could reasonably be expected to have resulted 
from an intermingling of forms from all three directions. 

With a view to obtaining exact information pertaining to the 
fauna and flora of this region, embracing a part of northeastern 
Trinity County, southwestern Siskiyou County and a corner of 
northwestern Shasta County, Miss Annie M. Alexander organized 
and financed two trips, one in February and March, 1911, to Helena, 
Trinity County, and another during the summer months of the 
same year, over the wider territory indicated in detail in the 
following itinerary and on the map (fig. A, page 337). The specimens 
obtained, numbering 449 birds, and 976 mammals, have been pre- 
sented by Miss Alexander to the California Museum of Vertebrate 
Zoology. The present report is based upon these specimens and is 
supplemented from the field notes of the collectors. 


ITINERARY 


SCHEDULE 


Helena, Trinity County, February 11 to February 26, 1911. 

Tower House, Shasta County, February 28 to March 8. 

Mayten, Siskiyou County, June 3 to June 6. 

Scott River, 6 miles northwest of Callahan, Siskiyou County, 
June 7 to June 14. 

Jackson Lake, Siskiyou County, June 15 to June 27. 

Wildeat Peak, Siskiyou County, June 27 to June 30. 

North Fork of Coffee Creek, Trinity County, July 1 to July 9. 

Saloon Creek Divide, Siskiyou County, July 9 to July 10. 

South Fork of Salmon River, Siskiyou County, July 12 to July 17. 

Summerville, Siskiyou County, July 17 to July 19. 

Hunters’ Camp, Trinity County, July 19 to July 20. 

Head of Grizzly Creek, Trinity County, July 20 to July 2 

Head of Rush Creek, Siskiyou County, July 26 to August 2. 

Kangaroo Creek, Siskiyou County, August 3 to August 5. 

Head of Bear Creek, Trinity County, August 5 to August 17. 

Castle Lake, Siskiyou County, August 18 to August 23. 


= 
o. 


| 1916] Kellogg: Mammals and Birds of 
} 


goat 
aS 
Rr ems 
Bye 


“s 
= 
¢ a) 


Ean 
ee 
Ean lM, Ml Rees Cat PRE « m5 
; te EM 


i SM MN eek = 
| Sy Jack, 


A gill lll 3 
Fame MN Za, 9A 
; = as 


ss alld, att 


QCecilville 


2 We 
= 2 

me : 
orn OSummerville” 


ne. 
haere a 
uth Fork 3Saimon River 
z ere noe 
CULE 
Iyer, 
Chee Z 
Tn 


3 gle. UM, 
tS AIAN 
’ 2S MAINS 
 y cy We 3 Ta 4 
eo Wty, A awe 


WN Finds 


my” “ay 


North Fork 


La range Mine 


é tA 
e = if rl Win tlle 

F lhe Mae MN 

1Q Weavervi Fe aa 


Seay, 
27 MS iy 
Zan Me 


; Ww 
RY als 


TN 


wi 


Fig. A. Map of portions of Siskiyou, Trinity and Shasta counties, show- 
ing routes (broken lines) traversed by Miss Annie M. Alexander and Miss 


Louise Kellogg in their zoological explorations of the region. 


338 University of California Publications in Zoology (Vow. 12 


DESCRIPTION OF ROUTE 


As a preliminary to the work of the summer, Miss Alexander 
and the writer spent a month, from February 10 to March 8, 
1911, at Helena, on the North Fork of the Trinity River, in Trin- 
ity County, and at Tower House, in Shasta County. 

We went by stage from Redding, Shasta County, following the 
Sacramento River for about three miles, then up Shasta Creek and 
over the Shasta divide, elevated 1800 feet above sea level, down into 
the valley of Clear Creek to Tower House and French Gulch, and 
from there over the Deadwood divide, 4550 feet by aneroid, to 
Lewiston, on the Trinity River. 

The east slope of the Shasta divide presents a scene of desola- 
tion as a result of the killing of the trees by fumes from the 
smelters. On the west side the vegetation is more flourishing, but 
miles of timber and chaparral have been demolished by forest fires. 
When we crossed the Deadwood divide, February 9, there was much 
snow near the summit. We saw very few birds, only a flock of chick- 
adees and kinglets, and a few quail. Mammals were even less in 
evidence, although in places we saw a few tracks in the snow near 
the road. Only one chipmunk and one gray squirrel were actually 
seen on the entire stage trip through to Weaverville. 

We spent the night at Lewiston and went on the next day to 
Weaverville, following the Trinity River for about three miles, 
crossing it and climbing over the shoulder of Brown Mountain, and 
down into Weaverville, arriving there February 10. Consultation 
with some of the people of the town seemed to point to Helena, on 
the North Fork of the Trinity, as being a good place for trapping, 
and further desirable because George Knowles, considered to be 
one of the best trappers in the county, was staying there. It is 
an eighteen mile drive up West Weaver Creek, then over the Oregon 
Gulch Mountain, 1050 feet, and down the gulch to Junction City 
on the Trinity River. This Oregon Gulch is being filled up at the 
rapid rate of twelve feet a month by debris from the La Grange 
hydraulic mine. The water of the Trinity River itself is turbid 
from the many mines scattered along its course. From Junction 
City we turned up Cafon Creek for a short distance, then doubled 
back to the Trinity and followed it down to where the North Fork 
empties into the main stream. The water of North Fork was beau- 
tifully clear in contrast to that of the muddy main river, and its 


1916] Kellogg: Mammals and Birds of Northern California 339 


narrow caiion leading up to a mass of snow-covered peaks gave 
one the impression of having entered a really wild and rugged 
country. Below North Fork, or Helena, as the small settlement 
is called, the Trinity River enters a deep rocky canon. 

HELENA is a postoffice town consisting of a hotel, store and 
couple of houses, situated at the junction of the Trinity River and 
its North Fork. The narrow cation of the North Fork opens out 
enough to make a little farming possible; but the wooded hills rise 
close on every side, with higher mountains in sight just beyond. 
The principal trees and shrubs noted in the vicinity were: digger 
pine, Douglas fir, oaks, ceanothus and poison oak. This is consid- 
ered a good trapping country for such fur-bearing mammals as 
fox, coon, skunk, civet cat, and, to a less extent, fisher. Larger 
mammals such as black-tailed deer, wild-cat and mountain lion were 
reported common. The life-zone at Helena is Upper Sonoran, with 
many Transition elements intruding from the closely surrounding 
area of pure Transition. 

We spent two weeks collecting at Helena, and then returned 
by way of Weaverville back along the stage line as far as Tower 
House, where we stayed for ten days, returning then to Oakland 
March 9. 

Tower House is a hotel resort at the lower end of Clear Creek 
Valley, eighteen miles from Redding. A violent storm which raged 
during most of our stay kept us from seeing much of the surround- 
ing country, but it appeared to be a place typical of the worked- 
out mining region so prevalent in Shasta County. The interests 
of the people center in the cultivation of small farms and in the 
cutting of wood. The tree stand is of digger pine and oak, and 
the chaparral consists of deer brush and manzanita. The zone may 
be considered Upper Sonoran. 

We made our second start from Oakland on the evening of 
June 3, with John Howard as assistant. We left the train at Edge- 
wood, Siskiyou County, and went by team to Mayten, twelve miles 
to the northward in Shasta Valley. 

Mayven is perhaps better known by the name of Big Spring, 
which is more specifically applied to a wet meadow several acres in 
extent in the center of Shasta Valley. The spring itself forms 
one of the sources of the Shasta River. A dam at the head of the 
spring backs the water up, forming a small lake, along the edge 


340 University of California Publications in Zoology (Vox. 12 


of which tules grow abundantly. The people told us that ducks 
occur here plentifully during the winter and that a few stragglers 
remain and nest. The water is used for irrigation, but the rocky 
nature of the soil in this part of the valley makes agriculture diffi- 
cult. Bird and mammal life centers close around the spring and 
where the land is cultivated. We had hoped to obtain here, at the 
type locality, specimens of Reithrodontomys megalotis klamathensis 
and might have had better success if we had trapped in a hayfield. 
As it was, near the spring, we did not get any. One farmer re- 
ported kangaroo rats as being abundant in the upper part of the 
valley where the soil is better and farming is carried on to a greater 
extent, but we saw no sign of them ourselves. Sage-brush grows 
on the rocky, uncultivated ground, and the scattering trees are 
juniper and small yellow pine. The region is distinctly Upper 
Sonoran in zone. We stayed here only two days, and then went 
to Gazelle, a small town on the line of the railroad and the first 
station north from Edgewood. While staying there over the night 
of June 6 so as to get the stage to Callahan, we set a few gopher 
traps in an alfalfa field close to the town, securing specimens of 
Thomomys leucodon navus. Our observations were naturally lm- 
ited, but indications pointed to the same zonal position as Mayten, 
namely Upper Sonoran. 

We left Gazelle the morning of June 7 by stage for Callahan, 
twenty-five miles to the west and at the south end of Scott River 
Valley. We passed over a divide of about five thousand feet alti- 
tude and reached Callahan a little after noon. As the immediate 
vicinity of the town did not look favorable for collecting we drove 
about six miles down the valley and made camp on a small slough 
tributary to the Scott River. According to report there was still 
too much snow in the mountains for us to attempt to go up to a 
high altitude, so we put in the time from June 8 to 14 collecting 
at this camp. 

Scorr River VaAuuery is a fertile stretch some twenty-four miles 
long and varying in width from one to six miles. It is hemmed in 
on the east by low rocky hills covered with sage-brush and ceano- 
thus, with a scattering growth of yellow pine and oak (see pl. 15, 
fiz. 1). On the western side the hills are somewhat more rugged 
and heavily timbered, and numerous small streams make their way 
down narrow eafions into the main river. The town of Callahan 


1916] Kellogg: Mammals and Birds of Northern California 341 


lies at the extreme southern point of the valley, where the river 
makes its entrance from a canon about a mile wide and six miles 
long. Beyond this hes the main valley, but we did not go farther 
north than our camp, which was situated just where the valley 
begins to widen. The altitude is 3000 feet. 

The water from the river is used extensively for irrigation. 
We were told that about fifty years ago the whole stream was 
diverted to the eastern side of the valley where it now runs, per- 
haps to facilitate in some way the use of the water. Our camp 
was situated in the old river-bed; the sandy bottom-land, undis- 
turbed, has gradually become covered with a dense growth of cotton- 
woods and willows. Zonally the valley is mainly Upper Sonoran, 
but there are a number of Transition elements mixed in. Of par- 
ticular interest was the discovery of Dipodomys, which finds here 
an ideal home in the sand of the river-bed. Birds, of the stream- 
side category, were numerous. 


As the weather had become very warm we decided to go up 
to Jackson Lake on the eastern slope of the Salmon Mountains. 
The move was made on June 15, and this may be considered our 
first base camp on the main line of the proposed fieldwork. We 
secured the services of Mr. John Baker as guide and hunter, with 
five pack animals to be used in transporting our outfit from camp 
to camp. 

Jackson Lake is a nearly cireular bit of water, at an altitude 
of 6000 feet, lying in an amphitheater of rocky peaks. The highest 
of these, situated on the north side of the lake, is called Wildeat 
Peak. The only break in the rocky wall is where Jackson Creek 
flows out from the lake to the east down a narrow valley, green 
with meadows and alder thickets. A thick stand of yellow pine, 
fir, spruce, hemlock, tamrac pine, and some sugar pine, fills in this 
open side of the lake and extends back up along the side of Wild- 
cat Peak almost to its barren summit. On our arrival, June 15, 
there was still much snow around the lake, especially on the rugged 
south side, with a few patches in the woods. There is a small 
pebbly beach on the north side of the lake, but for the most part 
the rocks descend sheer into the water. Several small streams 
enter the lake on the south, east, and north. We had fortunately 
arrived early enough to avoid any running of cattle or sheep into the 
meadows, and as the timber around the lake has not been cut, it was 


342 University of California Publications in Zoology (Vor. 12 


as natural and unsullied a collecting ground as one would wish to 
find. The variety of mammal life confirmed this idea thoroughly, 
for the locality yielded the greatest number of species of any of our 
camping places, with the exception of Helena. Birds were not numer- 
ous as to individuals except for chickadees and juncos. 

Judging from the mammalian fauna and the trees the region 
is in the Canadian division of the Boreal zone. Among the mam- 
mals taken were: golden-mantled ground squirrel, flying squirrel, 
mountain beaver, bushy-tailed wood rat, red-backed mouse, mink 
and marten. 

Witpcat PEAK was visited as a side trip of three days, June 
27 to 30. This is considered the highest point near the Lake, 
having an elevation of 7200 feet. We accomplished almost nothing 
in the collecting line while there, both because of the apparent 
scarcity of small mammals and birds and on account of a severe 
storm of rain and sleet. Our camp was about 300 feet below 
the summit, at the upper limit of the red fir growth. There were 
a few scattering firs above, and some white-barked pines; but the 
top of the ridge is very narrow and rocky, falling off abruptly 
to the north. From the summit one obtains a comprehensive view 
of the Seott and Salmon mountains to the south and west, and of 
Mount Shasta to the east. There was almost no life at this altitude; 
but the fact was due probably not so much to the height as to 
the barren, rocky nature of the ground. But five species of small 
mammals were taken, one an Aplodontia secured in a meadow below 
the peak on the north side. Birds were as scarce as mammals. The 
trees indicate that the peak rises barely into the Hudsonian zone. 

Our next move, July 1, was to the North Fork of Coffee Creek, 
almost due south across Saloon Creek divide, 6850 feet, in Trinity 
County. 

Norra Fork or Corree CREEK is a good-sized stream making 
its way precipitately down a narrow, well-wooded canon. There 
are numerous small meadows where creeks make into the main 
stream, the banks of which are densely clothed with alders. The 
timber is mainly white fir, Douglas fir, yellow, sugar, and silver 
pine, cedar, and a few cottonwoods. This growth does not extend 
far above the creek bed on the north side, but gives way to a dense 
growth of deer oak and white-flowered ceanothus which was in full 
bloom. Our camp was at the junction of the North Fork of Coffee 
Creek with Granite Creek, at an elevation of 4500 feet, with only 


1916] Kellogg: Mammals and Birds of Northern California 343 


1500 feet difference between it and Jackson Lake, but the differ- 
ence in temperature and the character of the surroundings was 
very marked. Here we were in full summer, while there it was 
spring. Just above our camp was a deer lick well known to our 
euide, and he told great tales of the deer he had seen in that lick. 
The zone may be considered Transition, but this diagnosis was 
based more upon the vegetation than upon the animal life, for 
we found here several Boreal mammals which thus ranged well 
down into the Transition, Zapus for example. 

SaLoon CREEK Divin—e.—On July 9, Miss Alexander and I made 
a trip back to Saloon Creek Divide, staying one night and putting 
in the afternoon and morning in collecting. This divide, of 6850 
feet altitude, forms part of the line between Trinity and Siskiyou 
counties. On the northern, Siskiyou, side, which is almost devoid 
of trees, a descent of about 500 feet brings one to a small stream 
fed from the snow banks of the divide (see pl. 15, fig. 2). Here we 
had seen numerous holes of the golden-mantled ground squirrel, 
and it was to collect some of these that we made the trip up from 
the North Fork. The southern side is equally steep, but covered 
thickly with ceanothus brush. We camped on the north side in a 
grove of red fir and devoted our collecting to ground squirrels and 
chipmunks. This was the first time we had ever found such a 
colony of the former (Callospermophilus). Their burrows were 
under every rock, as well as out in the open, and we could see 
many of the animals running about or sunning themselves on the 
rocks. We expected to find Microtus and Zapus in the alders along 
the stream, but the ground had been so beaten down by cattle that 
the smaller mammals had evidently not thriven. 


After twelve days on the North Fork, we started out again, fol- 
lowing up the creek and then striking across a 6100-foot divide 
and down gradually to the main Coffee Creek. We camped the 
night of July 11 about a mile above its junction with Union Creek. 
The next day we proceeded up the creek to the Salmon Flats, large 
mountain meadows which form a low divide between Trinity and 
Siskiyou counties. The ascent was a gradual one and the vegeta- 
tion abundant at the divide itself. Cottonwoods attain an immense 
size in the moist creek bottomland, and willows form dense thickets 
on either side of the stream. From the divide we went up the 
South Fork of the Salmon River about two miles and camped in 


344 University of California Publications in Zoology [Vow 12 


a small grove of pines by the river, where we stayed from July 12 
to 17. Here we found ourselves again in Siskiyou County. 

SoutH Fork or THE SALMON River heads in a semicircle of high 
peaks of about 7500 feet altitude. Our camp was at an altitude 
of 5000 feet, on the edge of a fine meadow of white clover inter- 
spersed here and there with large patches of false hellebore. South- 
west of us, looming up across the river, was a high rocky peak 
with a good bit of snow on it, and beyond that was a sharp- 
pointed peak of solid rock with precipitous sides, probably about 
8000 feet in height. There is no heavy timber on either side of 
the valley, but more on the east side than the west, where the 
mountains are rocky and covered mostly with chaparral. The trees 
around us. were yellow, sugar, and tamrac pine, white fir, and 
Douglas fir. The banks of the river were brushy with alders and 
willows. We had evidently again reached the Canadian zone, and 
a few new birds were added to the inevitable juncos and chicka- 
dees in the way of kinglets, both ruby-crowned and golden-crowned, 
Lincoln sparrows, and creepers. 

SUMMERVILLE, eleven miles down the river, was our next objective 
point. This is the name given to a series of mines and farms 
along the main Salmon River, 2000 feet below our last camp. 
The change from firs and tamrac pines to oak, manzanita and scat- 
tered madrone was very marked. The country looked dry and 
unattractive after the higher mountain region; but the land can 
be made to produce well under irrigation, as we saw on the farm of 
Mr. Jack Hinz at whose place we stopped. His land les on a 
bench some distance above the river, back of which rise the hills, 
while between them and the stream stretches a strip of glaring 
rock and sand, the remains of former hydraulic mining. Across the 
river the mountains rise steeply to a height of 5600 feet, covered 
below with black oak, madrone, Douglas fir and sugar pine, and 
higher up with a chaparral of chinquapin, white ceanothus and 
manzanita, which runs to the top of the ridge. The zone at Sum- 
merville may be considered high Upper Sonoran, with close invest- 
ment on all sides by Transition. 

At Summerville we added to our party Mr. Jack Hinz, at whose 
ranch we camped for a night, and who proved familiar enough 
with the country to be able to follow up an old government trail 
leading to the head of Grizzly Creek. Here we would be within 
reach of the highest peak in the Salmon Mountains. It took us two 


1916] Kellogg: Mammals and Birds of Northern California 345 


days from Summerville to reach the head of Grizzly Creek, the 
night of July 19 being spent at Hunters’ Camp. 

Hunters’ Camp, used by hunters and cattle men, is situated 
about a hundred feet down on the southern slope of the divide 
between Salmon River and Grizzly Creek, in a grove of white fir. 
The slope down to Grizzly Creek is very steep and the underbrush 
of chinquapin and ceanothus dense. There is a good spring coming out 
just below the camp, and its downward course is marked by dense 
alder clumps. Mr. Hinz says he often comes up on this ridge during 
the winter to trap, and has caught several fisher here. 

The trail up Grizzly Creek from Hunter’s Camp had been blazed 
years ago by a government surveying party which had made the 
ascent of a peak next to that known as Thompson Peak, and con- 
sidered the highest point of the Salmon Range. We were unable 
to ascertain the name of this peak from any of the people in the 
region, and it is not given on the maps, but is so close to Thomp- 
son Peak and so nearly of the same height that for the sake of 
convenience it also may be called Thompson Peak. 


GrizzLy CREEK has as its source a fair-sized lake lying in a 
rocky bowl between the mountains, and fed by great snow banks. 
The outlet of the lake is a waterfall which makes a sheer leap of 
about fifty feet over a rocky wall, and then by a succession of 
smaller drops descends into Grizzly Cafon. Our camp was just 
at the base of this precipice in a clump of red fir, white pine, and 
hemlock, flanked by large open meadows and willow thickets. 


In scenic beauty this spot surpassed any of our other camping 
places (see pl. 17, fig. 5), but collecting was rather arduous on 
account of the ruggedness of the ground and the steep slopes, 
either up or down; for the cafion is hemmed in on all sides, except 
in the direction of its outlet, by rocky walls. Small mammals were 
not particularly abundant, and Mr. Hinz attributed the lack of 
marten and fisher sign to the absence this year of suitable food, 
especially chipmunks. Aplodontia sign was abundant in a small 
canon where a stream and alders provided their favorite habitat 
(see pl. 16, fig. 3), and Zapus and flying squirrels were secured. One 
deer was shot high up on the crest of the mountain a thousand 
feet above camp. Save for elusive thrushes which sang in the 
underbrush, birds were rather scarce right around our camp. The 
density of the tree growth probably accounted for this. In the 


346 University of California Publications in Zoology (Vor. 12 


more open places and meadows the birds were about as plentiful 
as at other camps. This locality is in the Canadian zone. 

Our aneroid registered 7900 feet at the top of Thompson Peak, 
which is a pointed mass of rock dropping off steeply on all sides 
and surmounted by a government monument. Except for the last 
hundred feet or so up this point, the ascent of the mountain was 
not difficult, mainly over great stretches of granite rock and snow 
slides until we reached a granite ridge which forms the main 
approach to the peak. About half way up we obtained a good view 
of the lake which is the source of Grizzly Creek. There was ice still 
in it and we were told that the snow bank between the two peaks, 
from which it is fed, never entirely melts. On the eastern slope 
of the ridge were wind-blown specimens of white-bark and foxtail 
pines forming a heavy growth, while on the western slope tongues 
of hemlock ran up the mountain sides. 

The view from the summit well repaid us for the climb. On the 
east was Mount Shasta; directly below us to the south, two small 
lakes which constitute the source of the Stewart Fork of the 
Trinity River. Rattlesnake Creek also heads off to the south, sep- 
arated from Stewart Fork by a jagged crest of rocks. The water- 
shed of the Rattlesnake is fan-shaped, the granite smooth as if 
planed by glaciers, but covered with a scattering growth of pines 
and hemlocks. To the south and west we saw the Trinity Moun- 
tains and the Coast range; in fact, as far as the eye could reach, 
there were mountains. We were told that Mount Hood can be seen 
from here on a clear day, but this sounds like an exaggeration. 
We collected specimens of red and white heather and other plants 
of the Hudsonian zone. 

We put in four days collecting on Grizzly Creek and then 
returned to Summerville. Another night, that of July 25, at Hinz’s 
ranch and we were off up Rush Creek to its head, where we camped 
in a large meadow. 

Rusu Creek heads in broad, open meadows with occasional strips 
of alders (see pl. 17, fig. 6). Patches of red fir skirt the outer 
edges of the meadows and extend up to the divide, which rises on 
either side of the lowest point of the saddle. Our camp was at an 
altitude of 6400 feet, with the divide some thousand feet higher. 
The outlook to the south and west was quite comprehensive and 
we could see directly across to the two Thompson Peaks with the 
snow bank between. A side ecafon to the south of our camp con- 


1916] Kellogg: Mammals and Birds of Northern California 347 


tained a small lake surrounded by trees and proved a favorite 
haunt of birds. The meadow was variegated with flowers, helle- 
bore, painted cup, and larkspur. The cattlemen say that larkspur 
and wild parsley are fatal to cattle and they never keep their stock 
more than one night in this region. They claim horses are not 
affected. Flying squirrels were abundant and easily trapped in 
the higher groves of red fir, and we also secured several marten 
here. Mr. Baker shot a magnificent buck on a high ridge to the 
south. Zapus lived among the alders along the stream. The Lincoln 
sparrow was nesting here in a clump of hellebore. The zone may be 
therefore considered Canadian. 

On August 2 we made a forced march back to Callahan, over 
the Rush Creek Divide, around the heads of Taylor Creek and one 
of the branches of Coffee Creek, down into the East Fork of the 
Salmon River, up again until we struck the Scott River watershed, 
and thence down into Callahan. 

KANGAROO CREEK, a tributary of Scott River, then became our 
base of operations. Our camp here was situated at some deserted 
mining cabins about eight miles northeast from Callahan and 
several hundred feet higher, at an altitude of 3300 feet. We 
were disappointed to find that hydraule mining had been earried 
on in the creek, thus spoiling all the natural aspect of the place. 
Also there was almost no water except from a small spring near 
the buildings, so that any hope of finding many small mammals was 
vain. The side-hills were dry and unattractive, and birds conse- 
quently scarce. 

The thing of chief interest about the place was the mixture of 
zones, for we here found round-tailed wood rats and bushy-tailed 
ones inhabiting the same cabins. Also there were golden-mantled 
ground squirrels, which, with the bushy-tailed wood rats, must 
have been at the lowest line of their distribution. The place being 
shut in, as it is, in the canon, is probably somewhat colder in winter 
than at Callahan, although there is so little difference in altitude 
and general conditions; this, combined with heavier timber, may 
account for the presence of the two high-zone mammals named. 
The tree stand is chiefly yellow pine. The locality may be consid- 
ered prevailingly Transition. 

The collecting at Kangaroo Creek was so disappointing that 
after two days’ work we were glad to start for Bear Creek, one 
of the northern tributaries of the Trinity River, and at the base of 


348 Unwersity of Califorma Publications in Zoology (Vou. 12 


Mount Eddy. Here we remained from August 5 to 17, and, besides 
making the ascent of Mount Eddy, we spent a day at Toad Lake, 
one of the sources of the Sacramento River. 

BEAR CREEK is the second tributary from the head of Trinity 
River. We camped almost at its head, altitude about 6000 feet, in 
an attractive meadow, dotted with clumps of tamrac pine, alder, and 
willow, and supporting a heavy growth of grass and rank hellebore. 
Several acres had been fenced in by the forest rangers so that they 
could have a place in which to pasture their horses, that had not 
been trampled by the hordes of cattle. The trail from Kangaroo 
Creek, for the most part through sparsely wooded hills and open 
rocky stretches, had not prepared our minds for anything so green 
and inviting as this stopping-place proved to be, so we were very 
agreeably disappointed in it, and found the collecting excellent. 

The side-hills were covered with a growth of deer-oak and 
manzanita, white, and red fir, cedar, and yellow pine.’ Of the 
cedar there were some especially fine old trees. There was an 
abundance of water coming down in small streams on both sides 
of the main ereek, and that element always makes for plentiful 
bird and mammal life. The list of mammals ineluded the golden- 
mantled ground squirrel, two species of chipmunks, flying squirrel, 
snowshoe rabbit, Zapus, and marten. Birds were plentiful, both 
near camp and in the woods, among them being Lincoln sparrow, 
Lewis woodpecker, ruby-crowned kinglet, and creeper. The locality 
may be considered as prevalently Canadian in zone. 

Mount Eppy is a bare, cone-shaped peak, 9151 feet in elevation, 
next to Shasta the highest point in the region. But, because of 
the lack of snow and its ruggedness, it does not give one the im- 
pression of great height. We made the ascent August 14 by way 
of Deadfall Cafon and experienced no hard climbing at all, but 
found it undesirable to stay long on the summit on account of the 
violent wind that was blowing. The southwest side of the peak is 
well timbered with foxtail pime, and a few straggling white-bark 
pines reach almost to the barren summit, which is covered with 
loose shale rock. There was a small bank of snow on the north 
side. We got a good view of the Salmon Mountains, and counted 
eight ridges between us and Lassen Butte. Mount Shasta looked 
stupendous, but the view toward the Sacramento Valley was un- 
satisfactory on account of the haze. We saw some nuterackers and 
vireos near a small lake about 500 feet below the summit, and 


1916] Kellogg: Mammals and Birds of Northern California 349 


almost at the top I heard a chipmunk. We collected a number of 
botanical specimens, mostly species of the Canadian and Hudsonian 
zones. 

We spent August 12 at Toad Lake, Siskiyou County, which is 
across the divide, south, from Bear Creek. In making the ascent to 
the divide we passed through a forest of young silver pine, and 
on the summit saw a much-branched white-bark pine. The lake 
is circular, several acres in extent, and has an underground outlet 
which is the source of one of the western branches of the Sacramento 
River. On the eastern side is a tundra-like marsh with tamrac pines 
erowing along the edge. Other trees around the lake are silver pine, 
red and white fir, Jeffrey pine, and hemlock. The south wall of the 
basin has no timber and is very rocky. 

Castte Lake—On August 17 we left Bear Creek and, after 
crossing the divide at its head, followed the North Fork of the 
Sacramento in to Sisson. John Baker left us here, and the next day 
we secured a wagon and went to Castle Lake, a favorite summer 
camping place for the people of Sisson. This lake lies at an alti- 
tude of 5434 feet, about twelve miles southwest of Sisson. There 
are a few trees left in their natural state around the lake and for 
a mile or so down the cafion from it, but over all the rest of the 
country between it and Sisson the timber either has been cut or is 
in process of being cut, and I believe this in part accounts for the 
strange mixture of life-zones that we found there. The eastern 
slope of the lake is sparsely wooded with white fir, and tamrac, 
yellow and silver pine, trees belonging to the Canadian zone, while 
the western side is a brushy hill covered with chaparral of plum, 
currant, ceanothus, manzanita, and spiraea. The south side is a 
wall of granite and broken rock slides, precipitous and forbidding. 
It was on these rock slides that we caught a bushy-tailed wood rat, 
and a dark bit of fir woods produced a flying squirrel. Golden- 
mantled ground squirrels lived on the dry side-hill, and the com- 
mon ground squirrel of the lowlands (Citellus douglasii) was taken 
where the creek leaves the lake. Two unusual forms also taken here 
were Hvotomys and the least weasel. 

Considered from the zonal point of view, this seemed about the 
strangest association of mammals that one could encounter. It is 
probable that in altitude and original state the locality belonged 
to the Canadian zone. The cutting of the timber raised the tem- 
perature and lessened the fall of rain and snow so that animals 


350 Unwersity of California Publications in Zoology (Vow. 12 


of the Transition zone gradually worked up from the valley; and 
yet the higher zone forms are not yet altogether crowded out. 
It is only a question of time, it would seem, before the locality will 
become more purely Transition. At any rate, Castle Lake proved 
an interesting collecting point and made a good finish for our trip, 
which terminated at Sisson on August 23. 


CHECK LIST OF THE MAMMALS 


Scapanus latimanus latimanus (Bachman). 
Neurotrichus gibbsi major Merriam. 

Sorex vagrans amoenus (Merriam). 

Sorex montereyensis montereyensis Merriam. 
Myotis longicrus longicrus (True). 
Lasionycteris noctivagans (LeConte). 
Eptesicus fuscus fuscus (Beauvois). 

Ursus americanus Pallas. 

Canis lestes Merriam. 

10. Uroeyon cinereoargenteus townsendi Merriam. 
11. Bassariscus astutus raptor (Baird). 

12. Procyon psora pacifica Merriam. 

18. Martes caurina caurina (Merriam). 

14. Martes pennanti pacifica (Rhoads). 

15. Mustela muricus (Bangs). 

16. Mustela saturata (Merriam). 

17. Mustela vison energumenos (Bangs). 

18. Spilogale phenax phenax Merriam. 

19. Mephitis occidentalis occidentalis Baird. 

20. Felis oregonensis oregonensis Rafinesque. 

21. Lynx fasciatus Rafinesque. 

22. Reithrodentomys megalotis klamathensis Merriam. 


Wey ee) SSN Er) i ee WS) Sp J 


23. Peromyscus maniculatus gambelii (Baird). 
24, Peromyscus boylii boylii (Baird). 

25. Peromyscus truei gilberti (Allen). 

26. Neotoma fuscipes fuscipes Baird. 

27. Neotoma cinerea occidentalis Baird. 

28. Evotomys obscurus Merriam. 

29. Microtus montanus montanus (Peale). 


30. Microtus ealifornicus californicus (Peale). 
31. Microtus mordax mordax (Merriam). 

32. Thomomys leucodon navus Merriam. 

33. Thomomys monticola pinetorum Merriam. 

34. Dipodomys californicus trinitatis, subsp. nov. 
35. Zapus trinotatus alleni Elliot. 

36. Aplodontia chryseola Kellogg. 

37. Citellus douglasii (Richardson). 

38. Eutamias amoenus amoenus (Allen). 

39. Eutamias senex (Allen). 


1916] Kellogg: Mammals and Birds of Northern California 351 


40. Callospermophilus chrysodeirus trinitatis Merriam. 
41. Sciurus douglasii albolimbatus Allen. 

42. Sciurus griseus griseus Ord. 

43. Glaucomys sabrinus flaviventris Howell. 

44. Lepus washingtonii klamathensis Merriam. 

45. Lepus californicus californicus Gray. 

46. Sylvilagus bachmani ubericolor (Miller). 

47. Odocoileus columbianus columbianus (Richardson). 


GENERAL ACCOUNTS OF THE MAMMALS 


Scapanus latimanus latimanus (Bachman) 
Central California Mole 
One skull (no. 12996) was picked up at Tower House, and one 
skin-with-skull (no. 13798) was secured at Scott River. The latter, 
a male, measures as follows: total length, 165 millimeters; tail 
vetebrae, 35; hind foot, 21. 


Neurotrichus gibbsi major Merriam 
Large Shrew-Mole 

The two specimens (nos. 12908, 12909), which we obtained at 
Tower House, are placed under this subspecies since both in meas- 
urements and in the presence of an anterior cusp on the cingulum 
of the upper premolar they correspond to the description (Mer- 
riam, 1899, p. 88). There is considerable difference in elevation 
between the type locality of N. g. major, Carberry Ranch, Shasta 
County, altitude 4100 feet, and Tower House, altitude 1200 feet. 
The former locality is in the upper part of the Transition zone, the 
latter in high Upper Sonoran. 


Sorex vagrans amoenus (Merriam) 
Sierra Nevada Shrew 
Shrews were not common at any of the points where we col- 
lected. A series of sixteen of this form was secured (nos. 13780- 
13795), seven of them taken at Mayten, Siskiyou County, our lowest 
camp. Other localities where they were found are: Castle Lake, 
Siskiyou County, two; Salmon River, Siskiyou County, three; Rush 


352 University of California Publications in Zoology (Vou. 12 


Creek, Siskiyou County, three; and Bear Creek, Trinity County, 
one. 

Average measurements of seven specimens from Mayten are as 
follows: total length, 98.8 millimeters; tail vertebrae, 38.4; hind 
foot, 12. Average of nine from other localities: total length, 98.5; 
tail vertebrae, 38.4; hind foot, 12.3. 


Sorex montereyensis montereyensis Merriam 
Monterey Shrew 
Four specimens of this shrew were taken, three at Tower House, 
Shasta County (nos. 12906, 12907, 12997), one at Castle Lake, Sis- 
kiyou County (no. 13797), and one at Jackson Lake, Siskiyou 
County (no. 13796). Average measurements of four of these: total 
length, 112 millimeters; tail vertebrae, 48.7; hind foot, 13. 


Myotis longicrus longicrus (True) 
Long-legged Bat 

There is nothing quite so wasteful of ammunition as shooting 
at bats. At Kangaroo Creek and Castle Lake we used to spend 
the evenings until dark trying to hit the wavering, fluttering things. 
The time for shooting was limited because the bats would not come 
out until the light was nearly gone; this accounts in part for the 
few secured. Two specimens of the above species (nos. 13804, 
13805) were taken at Castle Lake, August 22. 


Lasionycteris noctivagans (Le Conte) 
Silver-haired Bat 
Two specimens were gotten at Kangaroo Creek, August 4 (nos. 
13802, 13803); these are evidently young, though they were well 
able to fly. 
Eptesicus fuscus fuscus (Beauvois) 
Large Brown Bat 


Two specimens taken at Kangaroo Creek, August 3 and 4 (nos. 
13799, 18800), and one at Castle Lake, August 19 (no. 13801). 


1916] Kellogg: Mammals and Birds of Northern California 353 


Ursus americanus Pallas 
Black Bear 

The region around Callahan, Siskiyou County, and from there 
westward through the mountains as far as we went, is one in 
which, according to common report, black bears are still fairly 
numerous. At Callahan we were told how men went out with 
dogs in the winter and without having gone any great distance 
would return bringing a bear. We saw some good-sized skins for 
sale there, and Miss Alexander purchased two (nos. 13765, 13766) 
in good winter pelage, but without skulls. 

At Jackson Lake we saw some old sign; but it was not until 


’ 


Hunters’ Camp was reached, on our way up Grizzly Creek, that 
we were told we were in the heart of the bear country. It cer- 
tainly began to look like it, when John Howard took his rifle and 
within a few minutes walk of camp saw a bear ambling along among 
the trees. It had not scented him, so was going leisurely and he 
had a good shot. The bear was left all night where it fell, with 
a coat thrown over it; for, according to the hunters, coyotes will not 
touch anything which has any article of clothing left around it. 
This bear, a female, was not a large individual, possibly two years 
old. It was saved as skin and complete skeleton (no. 13764). 

Subsequently, a skin and complete skeleton (no. 14712) taken 
near Callahan, October 7, 1911, was sent to the Museum by J. 
Baker. 


Canis lestes Merriam 


Mountain Coyote 

When we were at Helena, George Knowles reported seeing many 
tracks of coyotes, but he did not secure any specimens. However, 
he later sent in a skin and skull (no. 12876) from Hay Fork. 
There is evidence that the two do not belong to the same animal, 
since the skull showed considerable weathering, while the skin was 
fresh. At any rate, the same locality is doubtless represented. 

The skin shows no significant characters in either size or colora- 
tion; measurements (by collector): total length, 1040 millimeters ; 
tail vertebrae, 305; hind foot, 178; height of ear, 89. The skull, 
however, differs somewhat from typical Canis lestes, and if more 
material were available to bear out the points, it would appear that 
we had at last found indications of a northwest-coast race of coyote. 


354 University of California Publications in Zoology [Vou. 12 


Compared with a skull of Canis lestes, this skull shows only a very 
shght development of the deuterocone on P*, the teeth are ex- 
tremely massive and more crowded and the auditory bullae are 
larger and more inflated. Such characters would constitute good 
ground for specific differentiation if found repeated in several 
specimens. 

During the summer we secured only one coyote, too young to 
be of any value for comparison (no. 13763). Our guide, Mr. Baker, 
saw fresh tracks on our trip up Wildcat Peak, at Jackson Lake, and 
confidently set some steel traps, baited with grouse, but he caught 
nothing. For the most part we were not in country where we 
should expect to find them, it being too high and mountainous. 
During our stay at Helena, Knowles reported seeing many coyote 
tracks and said that they bothered him by digging out and springing 
his steel traps set for other animals. 


Urocyon cinereoargenteus townsendi Merriam 
Townsend Gray Fox 

At Helena foxes seemed to be abundant. Knowles secured seven 
in all (nos. 12879-12885), by trapping. The weights of these varied 
from 414 to 1014 pounds. We bought two skins, without skulls, 
from D. M. Corliss at French Gulch, Shasta County (nos. 12877, 
12878). In the Trinity and Salmon mountains we saw no sign 
of foxes. 

In the series obtained, and which are otherwise referable to 
this form, the white stripe on the hind leg is quite well defined, 
although Merriam, in his description of townsendi (1899, p. 103), 
says: ‘‘The white stripe on the hind foot of californicus has dis- 
appeared and is represented by a pale streak.’’ 


Bassariscus astutus raptor (Baird) 
California Ring-tailed Cat 
All four specimens (nos. 12886-12889) of this species were taken 
at Helena, Trinity County. We might have found them at Tower 
House, too, but the weather while we were there was stormy and 
we did not devote much time to the setting of steel traps. George 
Knowles trapped three of the civet cats, and Miss Alexander and I, 
after a good deal of maneuvering, managed to trap one. We had 


1916] Kellogg: Mammals and Birds of Northern California 355 


seen the tracks of a small animal in the sand along the river and, 
by taking an imprint of the foot of one of Knowles’ specimens as 
a pattern, had decided they pertained to a civet cat. We set a 
number 1 steel trap under a willow tree, hanging the bait from a 
branch. The first night the animal climbed the tree, got out on the 
branch and dragged the bait over a side branch toward it. The 
second night we hung the bait farther out and nearer the ground, 
and cut off the side branches. The additional effort proved successful. 

The largest specimen taken by Knowles (male, no. 12889) meas- 
ured: total length, 720 millimeters; tail vertebrae, 340; hind foot, 
70; ear, 45. Its weight was two pounds. 


Procyon psora pacifica Merriam 
Pacific Coon 

Our specimens (nos. 12890-12894), three from Helena, one from 
Hay Fork and one from Tower House, all belong to this dark form 
of Procyon (see Merriam, 1899, p. 107). The saying ‘“‘cunning as 
a coon’’ was exemplified in the case of one we finally trapped at 
Helena; but it was perhaps more because of our way of fixing the 
trap than of much cunning on the part of the animal that he was 
able to take our bait two nights in succession. We were trying a 
trap called ‘‘Stop-thief’’, which is supposed to catch the animal 
around the head, this being more humane than the common steel 
trap; but on account of its mechanism the animal had to step 
through it, so we arranged a cave of rocks with the bait inside and 
the trap at the entrance. The first morning after setting we found 
the remains of the bait, a saw-bill duck, about two yards away 
from the cave with the trap attached. It had been dragged through 
the entrance. The next morning the coon had torn the cave open 
from the opposite side. So we gave up the ‘‘Stop-thief’’, and read- 
ily caught the coon the following night in an ordinary steel trap, 
with suspended bait. 

We saw no signs of coons in the higher mountains of the Salmon 
and Trinity ranges. Neither were any tracks seen at Seott River. 


Martes caurina caurina (Merriam) 
Northwestern Pine Marten 


We secured seven specimens of marten (nos. 13767-13773), one 
at Jackson Lake, three at the head of Rush Creek, and three at the 


356 University of California Publications in Zoology (Vor. 12 


head of Bear Creek. I have designated them as Martes caurina, 
although they are not typical of that form as represented by speci- 
mens in the Museum collection from Vancouver Island, British Colum- 
bia. Our specimens show both summer and winter pelages and in 
both instances the markings of the throat and under surface are not 
an orange red, as in the Vancouver specimens, but more of a yel- 
low, and the whole body color is much paler. They also differ from 
the Vancouver specimens in having the metaconid of the lower car- 
nassial more distinct. A female taken at Crescent City is de- 
scribed by Merriam (1890, p. 27) as being of a uniform light seal 
brown with yellowish markings. It thus appears that that indi- 
vidual together with our specimens represents an extreme southern 
type of Martes caurina. 

Our no. 13772, male, in fresh fall pelage, has the center of 
the back raw umber shading to tawny olive on the sides; underfur 
wood brown. The underparts are tawny olive sprinkled with white 
hairs; markings of the throat, deep chrome. The ears are drab 
with a whitish edge; the nose vandyke brown. The front feet shade 
from Prouts brown to bistre. The tail is darker than the back; 
the brush seal brown. In another specimen, with worn pelage, the 
underfur of the back shows in patches vandyke and wood brown, 
and the throat markings have faded to maize yellow. 

Our first specimen was taken at Jackson Lake at the foot of a 
rock slide on the east side of the lake. It was caught only by the 
side pad of one front foot, but so securely that the trap held in 
spite of the fact that the animal had gotten down below the rock 
where the trap was set and had crawled into a hole. At Rush 
Creek we caught one in a trap set under a big log in a dense grove 
of firs on the sidehill near a small stream, and the other two, evi- 
dently a pair, close together, out of six traps we had set up the 
canon of a small stream leading into a lake. Here also the timber 
was dense. At Bear Creek we found martens in much the same situa- 
tion among the timber and near water and I had the pleasure of 
seeing one running along a log, but he saw me first and his disappear- 
ance was rapid. 

Martes pennanti pacifica (Rhoads) 
Pacific Fisher 

Knowles reported one day at Helena that a fisher had been 

around to all his traps, which were set on a ridge, and eaten the 


1916] Kellogg: Mammals and Birds of Northern California 357 


bait. He had to dig the traps out of the snow, which had fallen 
since his previous round, and set them again, confident that he 
would catch the fisher; and he succeeded. He told us this speci- 
men (no. 12901) showed a somewhat lighter tone of coloration than 
he had seen in other individuals. Before leaving Helena, Miss 
Alexander purchased a fisher skin (no. 12902) which had been 
taken about six miles from Helena in 1910. 

Our next information in regard to the presence of fisher came 
from Jack Hinz at Summerville, Siskiyou County, on the Salmon 
River. He said he had often trapped them on ridges near his 
place. The night that we spent on the divide between the Salmon 
River and Grizzly Creek, at Hunters’ Camp, Hinz set some traps 
for fisher, but his efforts here, and afterwards at the head of Grizzly 
Creek, were unsuccessful. He stated that the animals were much 
more difficult to catch in summer than in winter. The Museum 
collection has since then been enriched by four more specimens 
(nos. 16386, 16531, 16596, 19095), taken by Hinz near Cecilville, 
December 13, 1911, February 19, 1912, March 25, 1912, and Jan- 
uary 27, 1913, respectively. 

Description of the winter pelage follows: Above, from between 
ears to middle of back, buffy yellow shaded with black; an irregular 
black streak from middle of back to base of tail; sides cinnamon; 
general effect of tail black with background of vandyke brown; nose 
to eyes seal brown; underfur of head and neck vandyke brown, of 
back, hair brown; head grizzled with grayish white; throat blackish 
seal brown, darker on breast and belly; legs and feet black. In worn 
pelage: Above yellowish white to middle of back; black stripe more 
restricted; sides tawny olive; underparts lighter. 


MEASUREMENTS IN MILLIMETERS OF Martes pennanti pacifica FRomM NORTHERN 
CALIFORNIA 


o 
g s 
5 z 3 
Mus. _ ; = = = K 
no. Sex Locality 2 é a & 
129 O1NOn Elelenalmrimitys | COs nee srecseesceseenesene 830 340 90 35 
16386 9 Cecilville, Siskiyou Co..................... 864 375 88:5 50:5 
16531 ¢@ Cecilville, Siskiyou Co....................- 997 381 114 50.5 
16596 ¢ Cecilville, Siskiyou Co................... 991 368.5 120.5 50.5 
19095 @Q Cecilville, Siskiyou Co..................: 864 349. 101.5 38 


358 University of California Publications in Zoology Vou. 12 


Mustela muricus (Bangs) 
Sierra Least Weasel 


Of this tiny and apparently little known weasel we secured two 
specimens (nos. 13776, 13777), one at the head of Rush Creek, Sis- 
kiyou County, at an altitude of 6400 feet, and the other at Castle 
Lake, Siskiyou County, altitude 5434 feet. The first one was caught 
in a number 1 steel trap set under a log on the margin of a small 
lake. The one at Castle Lake came to oatmeal bait on a rat-trap set 
in a clump of firs where we were trapping for flying squirrels. 

The two specimens agree with the description of muricus by 
Bangs (1899, p. 71) except for the tail of one, which is nearly all 
white but with the tip dark and with the upper basal half of the 
same color as the back. As this specimen also shows spots of white 
on the nose, behind the ears and on the upper flanks, it may be 
inferred that the species turns white in winter and that this indi- 
vidual had not fully completed its summer molt. The date of cap- 
ture, July 28, however, is late for retention of even remnants of the 
winter pelage. The second specimen was taken August 20; close 
scrutiny of this one discloses a few white hairs in the tail, and two 
or three in the back. 

The measurements of the two specimens are as follows: No. 
13776, 2 , total length, 210 millimeters; tail vertebrae, 55; hind foot, 
27; no. 13777, 2, total length, 205; tail vertebrae, 51; hind foot, 27. 


Mustela saturata (Merriam) 
Siskiyou Weasel 

Two specimens taken at Jackson Lake (nos. 13778, 13779) have 
been referred to this species, although as far as known this is the 
first record of its occurrence south of the type locality, Siskiyou, 
Oregon. There are no white facial markings. At the corner of the 
mouth is a distinct brown spot considered as characteristic of the 
species, and in general coloration the specimens otherwise corre- 
spond with the first published description (Merriam, 1896, pp. 
21-22). 

Measurements of the specimens, both male, are as follows: No. 
13778, total length, 412 millimeters; tail vertebrae, 136; hind foot, 
50; no. 13779, total length, 403; tail vertebrae, 150; hind foot, 43. 
These average smaller than the two males from the Siskiyou Moun- 


1916] Kellogg: Mammals and Birds of Northern California 359 


tains, Oregon, the average measurements of which as given by Mer- 
riam are: total leneth, 423; tail vertebrae, 164; hind foot, 48. 


Mustela vison energumenos (Bangs) 
Pacific Mink 

Two specimens were taken by us, one at Jackson Lake, Siskiyou 
County (no. 13774), and one on Coffee Creek, Trinity County (no. 
13775), while three (nos. 12903-12905) were sent in later from Hay 
Fork, Trinity County, by George Knowles. The one secured at 
Jackson Lake was a female in extremely poor condition, thin, blind 
in one eye, and with an abnormal growth in the intestinal tract. 
The female caught on Coffee Creek, July 4, had borne young ones 
recently. Both of our specimens are darker than the ones from 
Hay Fork; the latter show a better defined mid-dorsal stripe. But 
the worn pelage of the two females precludes any accurate com- 
parison of their coloration with that of other material at hand. 


Spilogale phenax phenax Merriam 

California Spotted Skunk 
One specimen of this small skunk was taken at Helena (no. 
12900). The teeth are much worn and the lower canines look 
somewhat deformed. The inner cusp of the upper ecarnassial is 
almost lacking. On geographical grounds we might expect to find 
Spilogale phenax latifrons here; but the less prominent black areas 

of the specimen place it under phenax proper. 


Mephitis occidentalis occidentalis Baird 
Northern California Striped Skunk 

Three specimens (nos. 12895-12897) were secured while we were 
at Helena and two sent in later by George Knowles from Hay 
Fork (nos. 12898, 12899). The specimens average small in meas- 
urements, especially of the hind foot; only one approaches the 
measurement of the type in that regard, but the discrepancy may 
possibly be due to the method of measuring, so I have placed them 
without question under the above name. 


360 University of California Publications in Zoology (Vor. 12 


Felis oregonensis oregonensis Rafinesque 


Northwestern Cougar 


One specimen (skin and skull, no. 12871) of mountain lion was 
gotten by George Knowles at Hay Fork, Trinity County, and sent 
in after our return. 


Lynx fasciatus Rafinesque 
Barred Wildeat 


As the status of the various species of Lynx seems to be rather 
uncertain I have referred our specimens (four from Helena and 
Tower House, nos. 12872-12875) to the species first described. 
They certainly belong to the fasciatus group, if that name is to be 
applied to the more northern form of Lynz, rather than to the 
lighter-colored californicus from the south; but they are not as 
dark as some specimens in the Museum collection from Humboldt 
Bay. As no specimens of Lynx fasciatus pallescens are available 
for comparison I would hardly venture to identify ours with that 
form, even though Merriam (1899, p. 104) refers specimens taken 
on Mount Shasta to it. An adult male taken at Helena weighed 
nineteen pounds. 


Reithrodontomys megalotis klamathensis Merriam 


Klamath Harvest Mouse 


It was not our good fortune to secure the harvest mouse from 
Mayten, the type locality. Our camp there was rather unfortunately 
situated in a dry rocky pasture and we were not conveniently near 
to the hayfields where we might have found the species. But three 
specimens from Scott River (nos. 13360-13362), and five from 
Tower House (nos. 12788-12792) have been referred to this form 
because of skull characters. The large skull, heavy rostrum, wider 
brain-case and relatively smaller bullae serve to distinguish our 
specimens from R. m. longicaudus. But the coloration is practically 
as dark as in longicaudus, so that our specimens might best be 
considered intermediate, and nearest klamathensis. The hind foot 
of four male specimens averages but 17 millimeters, as against an 
average of 18.5 for two adults from the type locality of klamathensis, 
as given by Merriam (1899, p. 93). One male (no. 12788) meas- 
ures: total length, 147 millimeters; tail vertebrae, 77; hind foot, 18; 


1916] Kellogg: Mammals and Birds of Northern California 361 


ear, 11.5; but this is no older, as shown by the teeth, than some of 
the smaller ones. 


Nore: Since the above was written, Howell’s Revision of the American 
Harvest Mice (1914) has appeared, in which the name klamathensis is put 
into the synonymy of longicaudus. This ruling does not seem to properly 
dispose of the case, for the Museum of Verebrate Zoology contains material 
which points strongly towards the existence of a distinguishable race in 
northeastern California. This form does not appear to be merely an inter- 
mediate stage between longicaudus and megalotis, as Howell asserts. The 
cranial characters, as above specified, are too prominent to ignore, and, in 
combination with color, seem to be of diagnostic value. 


Peromyscus maniculatus gambelii (Baird) 
Gambel White-footed Mouse. 

This white-footed mouse was in evidence at all the camps visited 
during the summer, and was also taken at Helena. <A series of 
forty-one in all was secured (nos. 12745-12759, 13334-13359), rep- 
resenting the following localities: Helena, Tower House, Mayten, 
Seott River six miles northwest of Callahan, Jackson Lake, Wildeat 
Peak, north fork Coffee Creek, Saloon Creek divide, head of Grizzly 
Creek, head of Rush Creek, head of Bear Creek, Castle Lake. 
The specimens are variously intermediate in characters between P. 
m. gambelu and P. m. rubidus, averaging a trifle nearer the former. 


Peromyscus boylii boylii (Baird) 
Boyle White-footed Mouse 
Taken only at Helena and Tower House (nos. 12760-12779), and 
not found at any of the summer camps, although it was to have 
been expected at the localities of low altitude. 


Peromyscus truei gilberti (Allen) 
Gilbert White-footed Mouse 
With the exception of one specimen from Mayten, Siskiyou 
County (no. 13333), this mouse also is represented only from Helena 


and Tower House, where eight specimens were secured (nos. 12780- 
12787). 


Neotoma fuscipes fuscipes Baird 
Dusky-footed Wood Rat 


Most of our specimens of this animal came from Helena and 
Tower House, although the species was found to occur also at Seott 


362 University of California Publications in Zoology (Vor. 12 


River and Kangaroo Creek. Most of our summer camps were evi- 
dently above the level of their range. Nineteen specimens (nos. 
12812-12822, 13383-13390) were secured at the four localities named 
above. 


Neotoma cinerea occidentalis Baird 
Western Bushy-tailed Wood Rat 


The bushy-tailed wood rat was not common at any of our camps, 
except that on the North Fork of Coffee Creek. Here Miss Alex- 
ander found a colony in a deserted cabin and specimens were 
trapped without any particular difficulty. At Jackson Lake where 
we found them first we looked for fresh sign in the rock slides at 
the south and west sides of the lake without success; but there was 
plenty of old sign, and rat-traps baited with bacon produced seven 
in all. The altitude at Coffee Creek was rather low for the species, 
as was also that at Kangaroo Creek, near Scott River. At the 
latter place, however, we again found them inhabiting an old cabin 
and this time in company with their round-tailed kin. 

At Castle Lake we did some intensive trapping for what seemed 
to be the only individual at the Lake. We set a quantity of traps 
in rock slides at the end of the lake with no success until one morn- 
ing Miss Alexander found a hind foot left in one of her traps. 
Since this proved the presence of the species we kept up the trap- 
ping and on our last morning she captured the rest of the rat— 
unmistakably the same animal, for he was minus a hind foot. This 
we had saved and so, by fastening it in its proper place, were 
able to preserve a complete specimen. 

Twenty examples were secured (nos. 13363-13382), from the 
following localities: Jackson Lake, North Fork of Coffee Creek, head 
of Grizzly Creek, Kangaroo Creek, and Castle Lake. 


Evotomys obscurus Merriam 
Dusky Red-backed Mouse 
Owing to lack of material for comparison it is rather difficult to 
determine the systematic status of the two specimens of Hvotomys 
which we secured, one rather young male from Jackson Lake and one 
adult female from Castle Lake; but the two points in which these 
incline most strongly to the above-named species are the dull color 


1916] Kellogg: Mammals and Birds of Northern California 363 


and the arched dorsal line of the skull in contrast to the conditions 
in FE. mazama, which species is deseribed (Bailey, 1897, p. 132) as 
being bright-colored and with a notably flat-topped skull. Our 
specimens agree with obscurus also in the basilar length of the 
skull and in the zygomatic and mastoid breadth. In external 
measurements our examples do not agree with obscwrus so well, 
at least with the published measurements of the type of HL. obscurus; 
but in a species of which so few individuals have been taken some 
latitude may be allowed in this respect. Our specimens measure as 
follows: no. 18392,? adult, total length, 147 millimeters; tail verte- 
brae, 48; hind foot, 20. No. 138391, ¢, total length, 137; tail verte- 
brae, 40; hind foot, 19.5. The type specimen of obscurus measured 
(Merriam, 1897a, p. 72) total length, 155 millimeters; tail vertebrae, 
47; hind foot, 17. Thus we see that our two specimens average 
smaller in total length and that of tail vertebrae, but have a longer 
hind foot. In this latter respect they incline toward EH. mazama, 
in four specimens of which according to Merriam (1897a, p. 72) 
the hind foot measured 18.75 millimeters. 

Both of our specimens were taken in dark shady places among 
the pines, in traps baited with oatmeal. 


Microtus montanus montanus (Peale) 
Peale Meadow Mouse 


One specimen (no. 13404), from Mayten, Siskiyou County. 


Microtus californicus californicus (Peale) 
California Meadow Mouse 

This meadow mouse was only taken at our two lowest summer 
camps, Scott River and Mayten, and in winter at Helena and Tower 
House. It was not at all common at Scott River, which was rather 
a wet place, but the dry grass in the other localities was, as usual, 
full of their runways. Thirty-three specimens were secured: nos. 
13393-13404, 13405-13407, 12793-12811, from Helena, Tower House, 
Mayten and Scott River. The Helena series is perceptibly darker 
in tone of coloration than the specimens from elsewhere. 


Microtus mordax mordax (Merriam) 
Cantankerous Meadow Mouse 


A series of seventy-one specimens (nos. 13408-13476) of this 


364 Unwersity of California Publications in Zoology (Vor. 12 


long-tailed mountain meadow mouse was taken, the species proving 
common at all the higher camps except Castle Lake. We found 
them in damp meadows where they seemed to favor especially patches 
of hellebore with green grass growing between. No difficulty was 
experienced in trapping them where their runways and cuttings were 
much in evidence. Localities of capture are as follows: Jackson 
Lake, Saloon Creek divide, south fork Salmon River, head of 
Grizzly Creek, head of Rush Creek, head of Bear Creek, and Castle 
Lake. 


Thomomys leucodon navus Merriam 
Red Bluff Pocket Gopher 


Comparatively few specimens of this species were taken, the 
greatest number being twenty-six from Helena (nos. 12823-12848), 
with five from Tower House (nos. 12849-12853). During the sum- 
mer they were found at only one locality, Gazelle, Siskiyou Connty, 
where six specimens were taken (nos. 13477-13482). 

The leucodon group is a distinct one, easily distinguishable in 
skull characters from its nearest relative, laticeps, by the white- 
ness of the incisors and the fact that they project more strongly 
forward, and by the smallness of the molars. Considerable diffi- 
culty was experienced in deciding from the published description 
whether or not the subspecies navus is a well-marked form and if 
so, whether our specimens belonged to it or to lewcodon. Navus 
is deseribed* (Merriam, 1901a, p. 112) as bemg a much smaller 
form than leucodon; but a series of topotypes from Red Bluff, in 
the collection of the Museum of Vertebrate Zoology, shows some 
individuals so large as rather to eliminate size as a dependable 
point of distinction. Through the kindness of Mr. Henry W. Hen- 
shaw, Chief of the Biological Survey at Washington, some topo- 
types of lewcodon were loaned the writer for comparison, and be- 
tween the two sets of topotypes well-marked differences were seen 
to exist in skull characters. The comparison further indicated 
that the specimens from Helena and Tower House, which we have 
put under navus, are somewhat intergradient toward lewcodon. 

The coloration of specimens from Helena and Tower House in 
winter pelage is as follows: Above, general color bistre; lighter 
on sides, shading into color of underparts, which extends well up 
onto the sides. Underparts varying from creamy white to buff, 


1916] Kellogg: Mammals and Birds of Northern California 365 


the slate black bases of the hairs forming a distinet dark back- 
ground. Nose and region around mouth dark seal brown; throat 
patches, when present, and inside of pouches, white. 

Coloration of specimens from Gazelle, Siskiyou County, in sum- 
mer pelage is as follows: Middle of back fulvous brown, brighten- 
ing into fulvous on the sides; face slightly darker than back. Un- 
derparts tawny ochraceous, except the throat and sides of head, 
which are bright fulvous; patches at sides of mouth seal brown; 
throat patch and inside of pouches white. 


Thomomys monticola pinetorum Merriam 
Trinity Pocket Gopher 


This name seems to be applicable to a shghtly marked sub- 
species ranging west from Mount Shasta through the higher parts 
of the Trinity region. As far as coloration goes I cannot say that 
there is much difference between Thomomys monticola monticola 
and 7. m. pinetorum, although the latter may be considered a trifle 
brighter; but in skull characters, series of the two are for the most 
part distinguishable. The best skull character of monticola is the 
shape of the interparietal, which is about twice as broad as it is 
long; in pinetorum the interparietal is only shghtly broader than 
its leneth. One point emphasized in the original description of 
pinetorum (Merriam, 1899, p. 97) is that the zygomata are much 
more widely spreading than in monticola. Our specimens do not 
seem to bear this out; but they do show much more distinct tem- 
poral ridges, and this we may consider a diagnostic feature of the 
subspecies. 

A series of 182 specimens was procured (nos. 13483-13664), 
from the following localities: Scott River, Jackson Lake, Saloon 
Creek divide, North Fork of Coffee Creek, Coffee Creek, South Fork 
of Salmon River, head of Grizzly Creek, head of Rush Creek, head 
of Bear Creek, Castle Peak. A series of fourteen males from Rush 
Creek averages: total length, 210.5 millimeters; tail vertebrae, 58.7; 
hind foot, 28. 

The coloration of a specimen from Jackson Lake in worn winter 
pelage may be described as dark fulvous above; the colored ends 
of the hairs entirely worn off in places, leaving patches of slate 
black; underparts light ochraceous; nose and mouth parts seal 


366 Unwersity of California Publications in Zoology [Vou. 12 


brown; a black patch around the ear. Specimens in summer pelage 
are lighter fulvous above, with the dark bases of the hairs not as 
prominent; underparts varying from ochraceous to light buff; feet 
and tail whitish. Very few of the specimens are in good pelage, 
the majority being in ragged transition state from winter pelage 
to summer, or back again to winter, apparently, before the whole 
summer molt was completed. The impression gained was that 
gophers living in cultivated fields, for instance, of alfalfa, accom- 
plish their molts much more completely than animals living in 
wild land, which, except in summer, are harder put to it to obtain 
food. 

Our largest series of pinetorum was secured at Rush Creek. It 
was while trapping there that we decided that gophers are sensi- 
tive to smell, for traps set with bare hands were not so liable to 
catch anything as those set with gloved hands and the hole excavated 
with a trowel. Miss Alexander successfully tried baiting the traps 
by covering them and the entrance to the hole with a species of 
mint of which the gophers seemed to be fond. With these methods 
the animals were easily caught. At Jackson Lake we were par- 
ticularly struck by the winter workings of the gophers. The snow 
was just off the ground and there had been as yet no cattle around 
to trample down the innumerable serpentine ridges which covered 
the open ground and extended even among the shrubbery under 
the pine trees. These ridges, or ‘‘earth plugs’’, consist of the 
earth which in winter is pushed up into the snow from under- 
ground passages, and they prove the continued activity of the 
gophers even during cold weather. 


Dipodomys californicus trinitatis, new subspecies 
Trinity Kangaroo Rat 

Type-—Male adult; no. 12860, Mus. Vert. Zool.; Helena, Trinity 
County, California, altitude 1405 feet; February 18, 1911; collected 
by A. M. Alexander; original no. 1159. 

Subspecific characters —As compared with Dipodomys californi- 
cus californicus, coloration dark; thigh patches small; nasals long 
and heavy anteriorly. 

Coloration—Upperparts from between ears to band across 
thigh, bistre, suffused with dark buff yellow, brightening into 


1916] Kellogg: Mammals and Birds of Northern California 367 


orange buff on the sides, and somewhat lighter between eyes; thigh 
patches small, same color as back, darkening towards ankle; supra- 
orbital white spot distinct; black ring around eye; white spot on 
upper edge of ear; line over nose and at base of whiskers black; 
upper portion of feet white, beneath seal brown; upper and lower 
tail stripes seal brown, the upper merging into black some distance 
in front of the white pencil. Underparts pure white. 

Measurements—Of type, total length, 310 millimeters; tail 
vertebrae, 194; hind foot, 46; height of ear, 14. Length of nasals, 
measured along median line, average of 12 adult males, 14.7 milli- 
meters, average of 3 adult males of D. californicus, 13.5; average 
of 10 adult females of D. c. trinitatis, 14.3; average of 5 adult females 
of D. californicus, 13.5. Greatest anterior breadth of nasals, average 
of 12 adult males of D. c. trinitatis, 4.3; average of 3 adult males of 
D. californicus, 3.6 ; average of 10 adult females of D. c. trinitatis, 4.3 ; 
average of 5 adult females of D. californicus, 3.8. 

Discussion.—This new subspecies is based chiefly on the series of 
fifteen skins (nos. 12854-12868) obtained at Helena. These are 
in winter pelage, while the 28 specimens (nos. 13665-13692) from 
Scott River, Siskiyou County, are on account of the difference in 
season slightly lighter in average coloration. In these the orange 
buff of the sides is more pronounced, but the skull characters are 
similar. The series from Helena, compared with topotypes of 
californicus, from Ukiah, Mendocino County, is distinetly darker 
both in the general color and, more markedly still, in the color of 
the dark stripes on the tail. The white of the thighs appears 
restricted and in the dry skins is almost concealed. Such exterior 
differences, however, are not so convincing as points of difference 
that can be readily distinguished in the skull, and in the case of 
this subspecies the length of the nasals and their width anteriorly 
are well-marked characters. 

Another subspecies, Dipodomys californicus pallidulus (Bangs, 
1899, pp. 65-66), has been described, from Sites, Colusa County ; 
but a study of a series of topotypes of this form in the collection 
of the Museum of Vertebrate Zoology does not substantiate the 
subspecies. The characters given for pallidulus as compared with 
californicus are: ‘‘Color very much paler; thigh patches not dusky ; 

the black markings at base of whiskers and over nose 
nearly wanting ... .’’ A further statement (Elliot, 1901, p. 


368 University of California Publications in Zoology (Vor. 12 


431) that the tail is ‘‘above sepia, beneath white’’, has no founda- 
tion in fact, as the specimens show the regulation two dark stripes, 
one above and one below, with an intervening white stripe on each 
side. It is my impression that the specimens from Sites are a trifle 
lighter than those from Ukiah, especially in the face, but the shade 
of difference is so slight that I should call them both californicus and 
simply say that individuals from Sites show a tendency toward an 
extreme of light coloration for the species. 

Habitat—As far as the present geographical distribution of 
Dipodomys c. trintatis shows, the stock form may have originally 
worked up over the comparatively low divide from the head of the 
Sacramento Valley and then differentiated, because of approximate 
isolation, spreading in like manner to the north and east. George 
Knowles stated that kangaroo rats oceur at Hay Fork, Trinity 
County; but as far as our present material goes we have no other 
actual record than those from the two localities at which we col- 
lected specimens. 

At Helena we trapped for Dipodomys on the north side of the 
Trinity River near its junction with the north fork. For a short 
distance above this junction small sandy flats extend back from 
the banks of the river, due partly to hydraulic mining carried on 
in earlier days. With this exception the river runs through a nar- 
row, rocky canon, so that it would seem as if Dipodomys had here 
found a very limited area suited to its needs. The area of our 
trapping did not cover over half a mile, and was confined altogether 
to the north bank. We crossed the river at one point, about a mile 
above the junction, but although there was a sandy flat there and 
conditions seemed to be ideal for Dipodomys, we found no signs 
of them whatever. Where we did trap them, sign was plainly to 
be seen, consisting of burrows and the tracks of the animals. The 
prints of the hind feet showed at intervals of about twenty inches, 
directed to or from clumps of bushes, or around them. These 
bushes were manzanita and willow, and under them we set our 
traps. The holes were filled up with sand, evidently by the ani- 
mals. Trapping was more likely to be successful when done near 
these holes, even though they did not look as though recently used. 
Many of the Dipodomys secured had succeeded in filling their cheek 
pouches full of oatmeal before being caught. 

Our trapping area at Scott River was not much more extensive 


1916] Kellogg: Mammals and Birds of Northern Calcforma 369 


than that at Helena, and was again in sandy ground, in the old 
bed of the Scott River, but nearly a mile back from the present 
main stream which, according to report, had been at one time arti- 
ficially diverted to its present bed. Numerous small creeks and 
back waters of the river, are, however, on this side of the valley, 
used for irrigation. Some of the land is under cultivation, but a 
great deal consists of sand, overgrown with cottonwoods and willows 
and is used for pasture. The kangaroo rats here seemed to prefer 
the thicker brush for their headquarters, but Miss Alexander was 
most successful in securing specimens by setting the traps in a 
horse trail, to which the animals were no doubt attracted by the 
grain to be gathered from the droppings. 


Zapus trinotatus alleni Elliot 


Allen Jumping Mouse 

A series of sixty-three specimens of this species was obtained 
(nos. 13696-13758), from the following localities: Jackson Lake, 
North Fork of Coffee Creek, Saloon Creek divide, South Fork of 
Salmon River, head of Grizzly Creek, head of Rush Creek, head 
of Bear Creek. The series is remarkable for the freshness and uni- 
formity of the pelage and the fact that nearly all the specimens 
are adult. In some groups of mice, such as Microtus and Peromys- 
cus, nearly aS many young ones as adults are taken, but in our 
trapping we did not take a single very young specimen of Zapus 
in spite of the fact that some of the females showed signs of having 
bred. 

Another rather singular fact was that we never saw any signs 
of Zapus, such as runways, excrement or cuttings, and yet we 
caught some individuals at every camp where there were meadows 
or alders along running water, from an elevation of 4500 feet, on 
the North Fork of Coffee Creek, to our highest camp at Jackson 
Lake, 7200 feet. The species was most abundant at Coffee Creek, 
where we secured half of the whole series. We always found them 
in places where the vegetation was rank, generally, as just stated, 
under alders. 


Aplodontia chryseola Kellogg 
Trinity Mountain Beaver 
Ten specimens secured (nos. 13324-13332, 13806) from Jackson 


370 Unwersity of California Publications in Zoology (Vor. 12 


Lake, south fork of Salmon River, Wildcat Peak, and head of 
Grizzly Creek. Upon this series was based the description of Aplo- 
dontia chryseola as a new species showing nearest affinities with 
A. californica of the central Sierra Nevada (see Kellogg, 1914, p. 
295). This species proves to be not closely related to the coast 
form, A. phaea, as its golden brown coloration might seem to indi- 
cate. Cranial and other characters lead the writer to conclude 
that chryseola is derived from californica stock, the latter thus 
having invaded the territory westwardly towards the coast through 
the Trinity region. For cranial characters see table of measure- 
ments, page 371, and photographs, plate 18. 

Our first systematic search for signs of mountain beaver was 
begun at Jackson Lake, Siskiyou County. Here clumps of alder 
grew on sidehills from which trickled small streams of water. This 
seemed ideal ground for the animal, but the first real sign was 
found on drier ground where a runway was located between two 
such groups of alders. The tunnel, or runway, was finally traced 
direct from one alder patch to the other through a growth of wild 
plum, and measured seventeen meters in length. There were a dozen 
or more exposures of the runway. Although it seemed to be con- 
tinuous, the tunnel proved, upon examination, to be disconnected 
at about a third of the distance, as it took a decided turn down 
the hill, while the one in line with it tended diagonally up hill. 
Before the first section of tunnel took the downward turn there 
was a small side exit, showing that the animals foraged abroad 
into the rather open wild plum grove as well as into the alder 
thickets. There may have been underground connections between 
the two sections. The first tunnel ran back into the alders, and 
there was no visible exit within a radius of several yards. The 
tunnel throughout its exposed length was dry and looked to be in 
recent use. In hunting among the alders we found several old 
burrows which now serve as underground channels for small streams. 

Miss Alexander found other Aplodontia burrows north of Jack- 
son Lake in alders along one of the main tributaries of Jackson 
Creek. In places the ground was literally honeycombed by them. 
In such a multiplicity of runways it was hard to select just the 
right places for the traps. Fresh sign was found near an old fallen 
tree by the stream. The tree was about forty-five feet long and 
the main tunnel seemed to run lengthwise underneath it with exits 


371 


Mammals and Birds of Northern California 


. 
. 


Kellogg 


1916] 


TABLE OF CRANIAL MEASUREMENTS OF Aplodontia chryseola From NorrHeRN CALIFORNIA 
Oldest available specimen of each sex at top of column, youngest at bottom 


All measurements in millimeters 


o a) B eG 

3 xe, fe S ge 

nya as) Meee ote ge sg 

Bw a2 12) ais} Pa 2 a oF 
Hs es celts rates, NSH} He fq ges 

13331 Oo Headl Grizzly s@re yirinilb ya © Os geccrseenersees cere eres = 22.9 12.3 53.7 6.7 4,9 21.1 61.8 34.1 
13328 } JACks Onmlaa ey Sis 10 U a Oo eee cesses ee aera eres ees 24.6 13.0 52.8 7.3 5.3 19.8 60.9 32.5 
13326 Que dacksonla ke Sis lily Os CON cre ree ecceceeeesceeee ence eens 25.1 11.7 46.6 7.6 5:3 19.4 58.6 33.1 
13327 3 S, Hork Salmon’ Ri Siskiyou CO. 2 eiccccsceee eee 25.3 10.8 42.7 7.3 5.1 17.9 58.7 30.5 
13329 6 wJackson’ Gales Siskiyou COn cose csececenencecacessenreecnse 22.4 10.9 48.6 6.7 5.0 17.2 59.5 28.9 
13330 9 Wilds @ateR ely SIS Kay OU C0 iaceceese tens ree reeeeensere eee 24.4 10.8 44.2 6.7 4.9 18.0 57.9 31.1 
13324 Q Jackson Lake, Siskiyou Co. .... vs 26.1 IPA) 47.9 6.7 yal 19.5 59.1 apy 
fe) HeadiGrizzly Ore wiry, «© Ongescscecsecenerea scree eee 24.3 11,2 46.1 7.6 5.1 17.8 58.3 30.5 
Q TACKSONMWAKe GSTS Ici yi Olt © Op yeesecceceeeeeeeree see eceeee eee 25.1 10.7 42.6 6. 4.6 16. Diit 28.6 


1 Most anterior point on nasal bones to most posterior point. 

2 Greatest width of nasals, across both of them. 

8 With cranium resting on its dorsal surface, rostrum pointing away from the worker, the greatest length of the foramen 
on the right side. 

4 Taken at expansion of interpterygoid fossa immediately back of hard palate. 

5 Most lateral point on foramen ovale to the point farthest laterally (with reference to the skull) on zygomatic side of 
auditory tube. 


372 Unwersity of California Publications in Zoology (Vou. 12 


on either side, but chiefly on the water side. Two holes in the ceil- 
ing, from the neighborhood of which dirt had been scratched into 
a pile, exposed a passage way about fourteen inches in diameter. 
It was damp, but there was no running water in it, as was the 
case a few feet farther down in the tunnel. The fresh sign con- 
sisted in the trampled appearance of the exits, fresh cuttings of 
grass in them, and the broken leaves of plants at the entrance. 
Another runway was found with a passage leading up to a large 
hollow under a rock, the floor of which was strewn with alder cut- 
tings about a foot in length. The branches were generally cut 
diagonally, and one was as much as half an inch in diameter. The 
animals seem to well deserve their name of ‘‘mountain beaver’’. 


Miss Alexander caught three males in one limited area at Jack- 
son Lake, but on the whole the idea that these animals live in colo- 
nies did not seem to be correct. They may do so at certain times 
of the year, as during the breeding season and until after the young 
ones are full grown; but we never trapped more than one animal 
at a single set of burrows. 

On the Salmon River we found the mountain beavers making 
what we called ‘‘hay’’—large bunches of green plants of various 
kinds eut up and spread out as if to dry and to be used later. (See 
pl. 16, figs. 3, 4). On a sidehill down which ran several small 
streams among alders and poplars there was a collection of bur- 
rows more extensive even than any found at Jackson Lake. These 
were situated both near the stream and out in the more open, drier 
eround. The runways led under roots and over rocks, with open- 
ings at short intervals, and there were plain runs out into the 
brush where short branches ran from each main tunnel, somewhat 
like a gopher working, with a dump of earth at each terminus. 


Citellus douglasii (Richardson) 


Douglas Ground Squirrel 


Most of our camps were above the range of this ground squirrel. 
At Seott River it was common in the sandy ground along the 
numerous irrigation canals leading off from the river. We caught 
a good many more individuals than we wanted, especially of young 
ones. The weather was very warm and their greasy skins slipped 
quickly. The occurrence of the species at Castle Lake, however, 


1916] Kellogg: Mammals and Birds of Northern California 373 


made them seem a little more desirable; for here they were ap- 
parently above the limit of their usual range and were found both 
on the dry rocky hillside above the lake and in damper ground 
where the creek came out from the lake; altitude 5434 feet. I 
presume that until the timber between Sisson and Castle Lake was 
cut out the ground squirrels were probably not found there. They 
doubtless followed the steadily retreating timber-line which re- 
sulted in unnatural conditions of dryness for such an altitude. 
The specimens taken at Castle Lake were in the process of molt 
into fresh fall pelage. 

Eleven specimens were taken (nos. 13018-13028), from Mayten, 
Scott River, Summerville, six miles northwest of Callahan, and 
Castle Lake. 

Eutamias amoenus amoenus (Allen) 
Klamath Chipmunk 

This small chipmunk was encountered at every camp. Its 
greatest abundance was at Jackson Lake and on Rush and Bear 
creeks. Here the timber conditions seemed to be just right for 
them, and when other material was scarce we could always depend 
upon the chipmunk. Ninety-six specimens were obtained (nos. 13083- 
13177, 13280), from the following localities: Scott River, Jackson 
Lake, Wildeat Peak, Salmon Creek divide, South Fork of Salmon 
River, head of Grizzly Creek, head of Rush Creek, head of Bear 
Creek, head of Deadfall Creek, Castle Lake. 


Eutamias senex (Allen) 


Allen Chipmunk 

We found this larger chipmunk along with the smaller one at 
every camp, and collected a good-sized series, the bulk of it coming 
in at Jackson Lake, Bear Creek and Castle Lake. Our specimens 
differ somewhat from the description of senex in having the tail, 
below, cinnamon rufous, instead of orange; in this respect they 
incline toward EF. quadrimaculatus, a form of a lower zone. But 
their measurements and coloration otherwise identify them unques- 
tionably as senez. 

One hundred and two specimens were obtained (nos. 13178-13279), 
from the following localities: Scott River, Jackson Lake, Wildcat 
Peak, North Fork of Coffee Creek, South Fork of Salmon River, 


374 University of California Publications in Zoology  [Vot. 12 


Grizzly Creek, head of Grizzly Creek, head of Rush Creek, Kan- 
garoo Creek, head of Bear Creek, Castle Lake. 


Callospermophilus chrysodeirus trinitatis Merriam 
Trinity Golden-mantled Ground Squirrel 

The distribution of this form is given (Merriam, 1901b, p. 126) 
as ‘“‘the Siskiyou, Salmon, and Trinity mountains of northwestern 
California and southwestern Oregon’’. It is described as being 
“much larger and darker than chrysodeirus, and [it] never, so far 
as known, develops the golden mantle which covers the head and 
shoulders of that species’’. The average measurements of six speci- 
mens from the type locality is stated to be ‘‘total length, 283; tail 
vertebrae, 100; hind foot, 438”’. 

On geographical grounds one would expect the golden-mantled 
ground squirrels of the parts of Trinity and Siskiyou counties 
covered by our trip to come under this subspecies. Twelve of our 
specimens, males, from the various localities, average: total length, 
269.6 millimeters; tail vertebrae, 96; hind foot, 41.3. An average 
of four specimens, male, of C. chrysodeirus from Mount Shasta, is; 
total length, 263.7; tail vertebrae, 89; hind foot, 40.7. Seven 
specimens, male, from near Mount Whitney, east-central Cali- 
fornia, average: total length, 276; tail vertebrae, 94; hind foot, 41.4. 
Three specimens, male, from Independence Lake, Nevada County, 
California, average: total leneth, 258; tail vertebrae, 78; hind 
foot, 41.3. 

Of the three measurements, that of the hind foot seems to be 
the most constant, and judging from this feature as an index of 
size the specimens from these different regions should all be classed 
under the same species, namely, chrysodeirus. Moreover, many of 
our specimens show a well-developed golden color to the mantle, 
and, as already stated, trinitatis was said to lack this mantle color. 
On the whole, however, I should say that our series is slightly 
darker than those from Mount Whitney and Independence Lake, 
and this, even if other ascribed characters fail, might in itself 
justify the recognition of a form from the Trinity region. Further 
study of relevant material, with particular attention to changes 
of pelage with season, and a greater number of measurements of 
both body and cranium, may show that there are not sufficient 
grounds for recognition of a form trinitatis. 


1916] Kellogg: Mammals and Birds of Northern California 375 


This squirrel was found most abundant on Saloon Creek divide, 
where it was the predominating species of mammal, as also at the 
head of Bear Creek, eleven specimens being taken at each of these 
localities. Its occurrence at Kangaroo Creek must have been at 
about the lowest limit of its range. Fifty-four specimens were 
taken (nos. 13029-13082), from the following localities: Jackson 
Lake, Wildcat Peak, North Fork of Coffee Creek, Saloon Creek divide, 
South Fork of Salmon River, head of Grizzly Creek. head of Rush 
Creek, Kangaroo Creek, head of Bear Creek, and Castle Lake. 


Sciurus douglasii albolimbatus Allen 
Sierra Chickaree 

Twenty-one specimens were taken (nos. 12738-12740, 13280- 
13298) representing five localities: Helena, Jackson Lake, head of 
Rush Creek, Bear Creek, and Castle Lake. This shows quite a 
range of zone from the Transition of Helena to the Boreal of Jack- 
son Lake. Those from Bear Creek and Castle Lake, taken in 
August, show the lght ochraceous-buff beneath, of the summer 
pelage. 


Sciurus griseus griseus Ord 

California Gray Squirrel 
Helena, Trinity County, was the only locality at which we 
found gray squirrels, and because of the cold weather there in 


February they were not particularly in evidence. The seven speci- 
mens taken (nos. 12731-12737) bear dates from February 13 to 17. 


Glaucomys sabrinus flaviventris Howell 
Trinity Flying Squirrel 

We secured our first flying squirrel at Jackson Lake in a rat- 
trap baited with oatmeal and set near the head of our bed. For 
two nights some creature had startled us by running across our 
faces. When we found what the intruder was, we looked carefully 
in all the adjacent red firs for some trace of a nest; but our knowl- 
edge of the home and habits of Glawcomys received no increment by 
actual observation during our entire trip. 

Only three specimens were taken at Jackson Lake, where it may 
have been too early in the season for them to be out in numbers. 


376 University of California Publications in Zoology (Vor. 12 


Coffee Creek was too low for them, but we secured one at the 
head of Grizzly Creek; while the largest series from any one locality, 
fourteen, came from Rush Creek. Five from Bear Creek and two 
from Castle Lake make up the total of twenty-five (nos. 13299- 
13323). Their occurrence at Castle Lake was one of the remark- 
able features of that camp; but a thick grove of red fir on a side- 
hill showed that the squirrels as well as the trees may have been 
more wide-spread there previous to the time of timber cutting. 
We always selected a thick growth of firs in which to set the traps, 
and put them at the bases of the larger trees. We often wished 
we could see the animals in action, but they were as noiseless and 
mysterious as their fur is soft and silky. 

Our series shows an interesting variation in coloration and con- 
dition of pelage. As a whole, it seemed to answer quite closely 
to the original description of the form klamathensis (Merriam, 
1897), p. 225), and this name we had accordingly adopted. Recently 
our series was loaned to the United States Biological Survey where 
a study of the group is in progress. A preliminary paper (Howell, 
1915) announces the separation of the Trinity flying squirrel as a 
recognizable race, the new name flaviventris being applied to it. 
The type is no. 13,319, Mus. Vert. Zool.; head of Bear Creek, 6400 
feet altitude, Trinity County, California; collected August 13, 1911, 
by Annie M. Alexander; original number 1775. 


Lepus washingtonii klamathensis Merriam 
Oregon Showshoe Rabbit 


One male (no. 13759), presumably adult, and an immature 
female (no. 13760) were taken at Bear Creek, Trinity County, 
August 8 and 15, respectively. In coloration and cranial char- 
acters the adult specimen agrees closely with the latest published 
description of L. w. klamathensis (Nelson, 1909, p. 107). Hitherto, 
as far as known to the writer, this rabbit has been recorded only 
from the vicinity of Fort Klamath, Oregon, and from the central 
Sierra Nevada of California. Our adult example, in summer pelage, 
may be described as follows: mixed grizzled fulvous and black 
above; head and face of the same fulvous but with less black; pec- 
toral collar fulvous with a sprinkling of white; chin, throat, and 
belly white; a white stripe extending along the upper surface of 


| 
} 
| 


1916] Kellogg: Mammals and Birds of Northern California 377 


the hind leg on the inner side and including the toes. The skull 
shows the deeply grooved jugal, and small auditory bullae con- 
sidered distinctive of the species. The post-orbital processes are 
noticeably long. In total length this specimen measures less than 
the average given of three specimens from Fort Klamath; but the 
hind foot is even longer and this may be considered a more de- 
pendable index to actual size. 

Measurements of this specimen are: total length, 367 millimeters; 
tail vertebrae, 26; hind foot, 130; height of ear, 87. Average of 
3 specimens from Fort Klamath as given by Nelson (1909, p. 108) : 
total length, 414, tail vertebrae, 39; hind foot, 126; ear from 
notch, 64. 

We had been told by Mr. Hinz at Summerville that snowshoe 
rabbits were frequently seen in that region in the winter and that 
they were quite abundant on the south fork of the Salmon River. 
We had seen no sign of them at our camp there, and had decided 
that only the merest chance would furnish any specimens during 
the summer season. Our camp at Bear Creek was on the edge of 
a mountain meadow, and within two hundred yards of our tent 
was some higher dry ground covered with small brush through 
which ran innumerable cattle paths. Miss Alexander was making 
the rounds of her small traps one day about noon, when she scared 
up a rabbit in this brush. It ran around a bush and then sat per- 
fectly still in the middle of the trail. The creature was so motion- 
less and so nearly the same color as the dried grass of the trail 
that he could not be seen again until he moved and then he was 
gone in a flash. 

I set a number of small steel traps in the trails under the 
bushes and baited them with dried apples. The next morning the 
rabbit was in one of them. The young one was caught in a rat- 
trap set for squirrels among the trees bordering the meadow. The 
rest of this family were probably lying in the neighborhood, but 
further trapping failed to secure any more specimens. 


Lepus californicus californicus Gray 


California Jack Rabbit 


Scott River was the only camp at which we saw jack rabbits, 
and even here they were not especially abundant. They affected the 


378 Unwersity of California Publications in Zoology (Vor. 12 


adjacent hillsides. Two specimens were secured (nos. 18761, 13762). 
A third was sent in by Mr. Knowles from Hay Fork, Trinity 
County (no. 12869). 


Sylvilagus bachmani ubericolor (Miller) 
Redwood Brush Rabbit 
Mr. Knowles sent in a specimen (no. 12870) of this race of 
brush rabbit taken at Hay Fork, Trinity County, in March. We 
did not encounter it anywhere during the summer collecting. 


Odocoileus columbianus columbianus (Richardson) 


Columbian Black-tailed Deer 

The entire region which we covered, with the exception of Mayten 
and Scott Valley, is supposed to constitute an excellent deer coun- 
try; but our experience was that these animals had to be hunted 
for long and arduously and that big bucks were hard to get. The 
writer did not personally see a single deer during the whole sum- 
mer, but the other members of the party were all more fortunate. 
A total of eight specimens were secured (nos. 13013-13017, 12729, 
12730, 13807), from the following localities: North Fork of Coffee 
Creek, Summerville, head of Grizzly Creek, head of Rush Creek, 
Helena, Hay Fork, and Callahan. 

During our stay at Helena, in February, George Knowles re- 
ported seeing a band of twenty-four deer. That was at a time of 
the year when they would be herding together, and, too, on account 
of the scarcity of feed, they would then be coming down nearer to 
human habitations. At Jackson Lake, in June, some men camped 
below us had seen a doe drinking at the stream, and Miss Alexander 
jumped one in the brush; also there were many fresh tracks. In 
spite of several nights spent by the men at the lick, near which 
we camped, only one young buck was shot. They said a big 
buck started to come in one night, but it was moonlight and he 
saw them before they had a chance to shoot. Mr. Baker told us 
he thought the deer were then feeding by moonlight and lying close 
in the brush during the day so that they could not be scared up 
unless nearly stepped upon. 

As I have said, each animal secured meant a hard day’s tramp 
over miles of country. The best specimen obtained was at Rush 


1916] Kellogg: Mammals and Birds of Northern California 379 


Creek, a fine buck, with four points to the horns, which were in 
the velvet. Mr. Baker found a big hole in the body where the ani- 
mal had been shot before. He says deer often get off with bad 
wounds, the bullet finally becoming encysted, and the animal ap- 
parently suffering no permanent ill effects. The big bucks, he 
claimed, along towards the end of July, go off by themselves on 
the high ridges, only coming down for water. The does and young 
ones stay more in the canon bottoms. As is often the ease in other 
regions, there are popular theories as to there being several dif- 
ferent species of deer, for instance a long-legged kind and a short- 
legged one. But we failed to secure evidence that would go to 
show the presence of any true species other than Odocoileus colum- 
bianus. 


LIST OF THE BIRDS, WITH ANNOTATIONS 


Actitis macularius (Linnaeus). Spotted Sandpiper. Two adult 
skins (nos. 19120, 19121), from Jackson Lake, June 18 and 20. 

Oxyechus vociferus (Linnaeus). Killdeer. Observed June 4 
at Mayten. 

Oreortyx picta picta (Douglas). Mountain Quail. Nine speci- 
mens (nos. 17299-17302, 19122-19126), from Helena, head of Bear 
Creek and Castle Lake. Three of these, taken August 15, 16 and 
20, are juvenals, and in this plumage the characters distinguishing 
picta from plumifera are better shown than in the adult. These 
characters consist in a warmer brown suffusion dorsally and 
greater extent of black markings. 

Numerous at nearly all of the mountain stations. On July 8, 
on north fork of Coffee Creek, the writer caught sight of a weasel 
in pursuit of a mountain quail. The bird was clucking in a dis- 
tressed manner and evidently leading the enemy away from where 
her chicks were. When the weasel got her to a safe distance he 
ran back, jumped over a log, and was seen to make off with a 
small victim in his mouth. The whole episode did not occupy two 
minutes and occurred in a clearing in broad daylight. 

Lophortyx californica vallicola (Ridgway). Valley Quail. Two 
skins (nos. 17303, 19127), from Helena and Scott River. A nest 


380 University of California Publications in Zoology  [Vou. 12 


found at the latter point, June 8, contained seventeen eggs. Seven 
of these occupied the center, while the remaining ten were disposed 
in an outer row. Observed also at Tower House. 

Dendragapus obscurus fuliginosus (Ridgway). Sooty Grouse. 
Two skins (nos. 17304, 17305), taken at Hay Fork March 20, sent to 
the Museum by George Knowles. These are not typical fuliginosus, 
but show themselves to be a step nearer to this humid-coast form than 
the grouse of the localities enumerated under the next heading. The 
Trinity region evidently constitutes a part of the area of intergrada- 
tion between sverrae and fuliginosus proper. 

Dendragapus obscurus sierrae Chapman. Sierra Grouse. Hight 
specimens (nos. 19128-19135), from Callahan, north fork of Coffee 
Creek, Grizzly Creek, Summerville, head of Rush Creek and head 
of Bear Creek. One of these is a downy young, taken en July 8; 
three others are juveniles, taken July 18 and 25 and August 7. 
The females and young average very slightly browner dorsally and 
are more heavily marked on the breast than typical sierrae, thus 
indicating intergradation towards fuliginosus. 

Heard drumming at Jackson Lake the third week in June, and 
one shot on flanks of Wildcat Peak, June 27. Brood of half-grown 
young seen July 18, at Summerville, where these birds are said to 
visit the farmyard occasionally to share the feed with the chickens. 
A farmer on Kangaroo Creek complained that grouse were so 
numerous that they were injuring his young grain. At this season, 
August, the grouse come down off the hillsides and collect around 
the margins of clearings where they prove locally destructive to 
erops. Abundant at head of Bear Creek during second week of 
August. 

Zenaidura macroura marginella (Woodhouse). Western Mourn- 
ing Dove. . Nest and two eggs found June 5 at Mayten. A flock 
noted at Scott River, June 11. 

Accipiter velox (Wilson). Sharp-shinned Hawk. One imma- 
ture (no. 19136), from Castle Lake, August 19. 

Accipiter cooperi (Bonaparte). Cooper Hawk. One immature 
(no. 19137), from Kangaroo Creek, August 3. 

Astur atricapillus striatulus Ridgway. Western Goshawk. One 
adult (no. 17306), from Hay Fork, taken March 26 by George 
Knowles. 

Falco sparverius sparverius Linnaeus. Sparrow Hawk. One 


1916] Kellogg: Mammals and Birds of Northern California 381 


seen on Wildeat Peak, June 29, and a pair seen at Summerville, 
July 18. 

Dryobates villosus orius Oberholser. Modoe Hairy Woodpecker. 
Eleven specimens (nos. 17307, 19138-19147), from Helena, Scott 
River, North Fork of Coffee Creek, and head of Bear Creek. This 
series is on the whole darker on the breast and less spotted than 
typical orius, but not to such a degree as to be referred to harrisi, 
though intergradation in this direction is indicated. 

Dryobates pubescens gairdnert (Audubon). Gairdner Wood- 
pecker. Four skins (nos. 17308, 17309, 19148, 19149), from Helena, 
Tower House, and Scott River. These are not typical gairdneri, 
but tend towards turatv. 

Xenopicus albolarvatus albolarvatus (Cassin). Northern White- 
headed Woodpecker. Two skins (nos. 19150, 19151), from Kangaroo 
Creek, and head of Bear Creek, August 4 and 6. Seen also on 
North Fork of Coffee Creek, July 8. 

Sphyrapicus varius daggetti Grinnell. Sierra Red-breasted Sap- 
sucker. Nine specimens (nos. 17310, 173811, 19152-19158), from 
Helena, Scott River, head of Grizzly Creek, and head of Bear Creek. 
Six, from the last two localities, July 23 to August 16, are juvenals. 

Phloeotomus pileatus abieticola (Bangs). Northern Pileated 
Woodpecker. One skin (no. 17312), taken at Hay Fork by George 
Knowles, March 27. One seen at Jackson Lake, June 30, working 
on a dead pine. 

Melanerpes formicivorus bairdi Ridgway. California Woodpecker. 
Two skins (nos. 19159, 19160), from Seott River, where common. 

Asyndesmus lewisit Riley. Lewis Woodpecker. One skin (no. 
19161), from head of Bear Creek, August 11. <A family of the birds 
was here observed using as a rendezvous a dead tree standing in an 
open tract of chaparral. 

Colaptes cafer collaris Vigors. Red-shafted Flicker. Eight skins 
(nos. 17313, 17314, 19162-19167), from Helena, Tower House, Jack- 
son Lake, and head of Bear Creek. At Seott River, June 13, a 
bird was observed excavating a nest-cavity near the summit of a 
dead cottonwood. Bill-fulls of fine chips were repeatedly brought 
to the doorway, and allowed to scatter to the ground immediately 
about the tree. The bird finally emerged with an egg in its bill 
and flew off with it over the fields. The bird soon returned and 
resumed work upon the interior of the cavity, as evidenced from 


382 University of California Publications in Zoology (Vou. 12 


the continual tapping from within. It is to be inferred that the | 
ege was laid before the nest-cavity had reached its proper dimen- 
sions, and that the bird viewed the egg in the light of an obstruc- 
tion, to be gotten rid of at that stage of nest-building. The species 
was also observed at Kangaroo Creek, August 4. 

Selasphorus rufus (Gmelin). Rufous Hummingbird. Five 
specimens (nos. 19171, 19172), from South Fork of Salmon River, 
July 14, and nos. 19168, 19173, 19174, July 27, 29 and 30, from 
head of Rush Creek. 

Stellula calliope (Gould). Calliope Hummingbird. Three skins, 
no. 19175, from Wildeat Peak June 30; no. 19170 from South Fork 
of Salmon River, July 14, and no. 19169, from Castle Lake, August 20. 

Sayornis nigricans (Swainson). Black Phoebe. One skin, no. 
17315, from Tower House, March 2. 

Nuttallornis borealis (Swainson). Olive-sided Flycatcher. Two 
skins (nos. 19176, 19177), from head of Grizzly Creek, July 23. 

Myiochanes richardson richardson (Swainson). Western 
Wood Pewee. Five specimens (nos. 19178-19182), June 9 to July 13, 
from Scott River, Jackson Lake, and South Fork of Salmon River. 

Empidonax tralli trailli (Audubon). Traill Flycatcher. One 
skin, no. 19187, from Scott River, June 12. 

Empidonax hammondi (Xantus). Hammond Flycatcher. Adult 
male, no. 19183, from South Fork of Salmon River, July 15; juve- 
nile male, no. 19184, head of Grizzly Creek, July 23; juvenile male, 
no. 19186, Castle Lake, August 20. 

Empidonaz wrighti Baird. Wright Flycatcher. One juvenal, no. 
19185, from head of Bear Creek, August 14. 

Cyanocitta stelleri frontalis (Ridgway). Blue-fronted Jay. 
Nineteen skins, nos. 17316, 17317, 19188-19204, from Helena, Jack- 
son Lake, North Fork of Coffee Creek, head of Grizzly Creek, head 
of Rush Creek, head of Bear Creek, and Castle Lake. Some of 
these show indications of intergradation towards the coast form 
carbonacea. 

Aphelocoma californica californica (Vigors). California Jay. 
One skin, no. 17318, from Helena, February 24, where only two or 
three were noted. Observed also at Seott River, in June. 

Nucifraga columbiana (Wilson). Clarke Nuteracker. One 
specimen, no. 19205, from head of Bear Creek, August 12. Seen 
at Jackson Lake, June 23 and 24, and on Wildcat Peak, June 28. 


1916] Kellogg: Mammals and Birds of Northern California 383 


Agelwius phoeniceus nevadensis Grinnell. Nevada Red-winged 
Blackbird. Seven breeding adults, nos. 19206-19212, from Mayten, 
June 4 and 5. At this locality the species was common on marshy 
ground around Big Spring. For systematic treatment, see Grinnell 
1914, p. 107. 

Sturnella neglecta Audubon. Western Meadowlark. Observed 
June 4 at Mayten, and June 13 at Scott River. 

Icterus bullocki (Swainson). Bullock Oriole. Three skins, nos. 
19214-19216, from Mayten (June 4) and Scott River (June 11 
and 12). 

Euphagus cyanocephalus (Wagler). Brewer Blackbird. One 
skin, no. 19213, from Mayten, June 4. 

Hesperiphona vespertina montana Ridgway. Western Evening 
Grosbeak. One specimen, no. 17319, taken at Weaverville, February 
27 (see Kellogg, 1911, pp. 119-120). 

Carpodacus purpureus califormcus Baird. California Purple 
Finch. Five skins, nos. 17320, 19220-19223, from Helena (February 
24), South Fork of Salmon River (July 14), and Castle Lake 
(August 21). 

Carpodacus cassini Baird. Cassin Purple Finch. One skin, 
no. 19224, from Jackson Lake, June 18. 

Carpodacus mexicanus frontalis (Say). California Linnet. Two 
specimens, nos. 19225, 19226, from Mayten and Scott River, June 4 
and 11. Common at both of these stations. 

Astragalinus tristis salicamans (Grinnell). Willow Goldfinch. 
Two skins, nos. 19227, 19228, from Scott River, June 10 and 11. 

Astragalinus psaltria hesperophilus Oberholser. Green-backed 
Goldfinch. Three skins, nos. 19229-19231, from Scott River, 
June 11-13. 

Spinus pinus pinus (Wilson). Pine Siskin. Four specimens, nos. 
17321-17324, from Tower House, March 4 and 7. Seen also at 
Helena, in February. 

Chondestes grammacus strigatus Swainson. Western Lark Spar- 
row. Three skins, nos. 19232-19234, from Scott River, June 9-11. 

Passer domesticus Linnaeus. English Sparrow. Two specimens, 
nos. 17325, 17326, from Tower House, March 7. Seen also at 
Weaverville. 


384 University of California Publications in Zoology (Vou. 12 


Zonotrichia coronata (Pallas). Golden-crowned Sparrow. Three 
skins, nos. 17327-17329, from Helena (February 14 and 23), and 
Tower House (March 8). 

Spizella passerina arizonae Coues. Western Chipping Sparrow. 
Four specimens, nos. 19235-19238, from Scott River and South Fork 
of Salmon River, June 8 and 11, and July 13. Observed also at 
Kangaroo Creek, August 4. 

Junco oreganus thurbert Anthony. Sierra Juneo. Thirty-one 
specimens, nos. 17330-17338, 19239-19260, from the following loeal- 
ities: Helena, February 15 and 25; Tower House, March 1 and 2; 
Jackson Lake, June 17-26; north fork of Coffee Creek, July 2 and 6; 
South Fork of Salmon River, July 13 and 14; head of Grizzly Creek, 
July 21-31; Kangaroo Creek, August 4; head of Bear Creek, 
August 11 and 16; Castle Lake, August 19. This was the most 
wide-spread and abundant bird of the higher mountains during the 
summer. 

Melospiza melodia rufina (Bonaparte). Rusty Song Sparrow. 
Eight skins, nos. 17343-17350, from Helena (February 15-24), and 
from Tower House (March 1-7). For use of this name see Kellogg 
(1911, p: 120). 

Melospiza melodia fisherella Oberholser. Modoe Song Sparrow. 
Sixteen skins, nos. 17339-17342, 19261-19272, from Helena (February 
16), Tower House (March 2-7), Scott River (June 8-12), Mayten 
(June 4 and 6), Castle Lake (August 20). Those from the first 
two localities were recorded (Kellogg, 1911, p. 120) under the name 
merrilli. This form of song sparrow was common on the more or 
less marshy ground around the Big Spring at Mayten; numerous 
also along the stream at the station on Scott River. 

Melospiza lincolui lincolni (Audubon). Lincoln Sparrow. Eleven 
specimens, nos. 19273-19283, from Jackson Lake (June 21), South 
Fork of Salmon River (July 14), head of Rush Creek (July 27), 
head of Bear Creek (August 6-10). 

Passerella iliaca wnalaschcensis (Gmelin). Shumagin Fox Spar- 
row. One specimen, no. 17351, from Helena, February 17. 

Passerella iliaca meruloides (Vigors). Yakutat Fox Sparrow. 
One skin, no. 17352, from Tower House, March 8. 

Passerella iliaca megarhyncha Baird. Thick-billed Fox Sparrow. 
Five skins, nos. 17353-17355, 19284, 19285, from Tower House 
(March 2 and 7), and Castle Lake (August 19 and 21). The 


ie) 
oO 


1916] Kellogg: Mammals and Birds of Northern California 38: 


latter was the only locality where any form of fox sparrow was 
encountered during the summertime. 

Pipilo maculatus falcinellus Swarth. Sacramento Towhee. Ten 
specimens, nos. 17356-17358, 19286-19292, from Tower House 
(Mareh 1-5), Seott River (June 9-11), and Summerville (July 18). 
For systematic treatment, see Swarth (19138, pp. 168, 172). <A 
nest with young just hatching was found at Scott River, June 10. 
It was on the ground under a branch of a fallen yellow pine and 
was screened by a bunch of dead pine needles. 

Pipilo crissalis carolae McGregor. Northern Brown Towhee. 
Three skins, nos. 17359-17361, from Helena (February 24), and 
Tower House (March 1 and 7). For status of this form, see 
Grinnell (1912, p. 199). 

Oreospiza chlorura (Audubon). Green-tailed Towhee. One 
specimen, no. 19293, from head of Bear Creek, August 6. 

Zamelodia melanocephala capitalis (Baird). Pacific Black-headed 
Grosbeak. Three skins, nos. 19217-19219, from Scott ‘River, 
June 9-11. 

Passerina amoena (Say). Lazuli Bunting. Seven skins, nos. 
19294-19300, Seott River and Summerville, June 9 to July 18. 
Particularly plentiful at the latter locality. 

Piranga ludoviciana (Wilson). Western Tanager. Five skins, 
nos. 19301-19305, from Scott River, Jackson Lake, Summerville, and 
head of Rush Creek, taken in June and July. Seen at head of 
Bear Creek, August 11. 

Bombycilla garrula (Linnaeus). Bohemian Waxwing. Nine 
specimens, nos. 17362-17370, from Helena and Tower House, Feb- 
ruary 9 to Mareh 1. For further account, see Kellogg, 1911 
(pp. 120-121). 

Vireosylva gilva swainsoni (Baird). Western Warbling Vireo. 
Four skins, nos. 19306-19309, from Seott River and Jackson Lake 
(in June), and Kangaroo Creek (immature, August 4). 

Lanivireo solitarius cassini (Xantus). Cassin Vireo. Six skins, 
nos. 19310-19315, from Scott River (June 10), head of Grizzly 
Creek (July 23), and head of Bear Creek (August 13). 

Vireo huttoni huttoni Cassin. Hutton Vireo. One skin, no 17371, 
from Helena, February 20. 

Vermivora ruficapilla gutturalis (Ridgway). Calaveras Warbler. 
Ten specimens, nos. 19316-19325, from Jackson Lake, South Fork of 


386 Unwersity of California Publications in Zoology (Vor. 12 


Salmon River, head of Grizzly Creek, and head of Rush Creek 
(in June and July), and Castle Lake (August 20). 


Vermiwvora celata lutescens (Ridgway). Lutescent Warbler. 
Four skins, nos. 19326-19329, from head of Grizzly Creek (July 23), 
head of Rush Creek (July 29), head of Bear Creek (August 11), 
and Castle Lake (August 21). 


Dendroica aestiva brewsteri Grinnell. California Yellow Warbler. 
Seven skins, nos. 19330-19336, from Scott River, June 8-13. 


Dendroica auduboni auduboni (Townsend). Audubon Warbler. 
Thirteen skins, nos. 19337-19349, from Jackson Lake (June 17-25), 
South Fork of Salmon River (July 14), head of Grizzly Creek 
(July 22 and 23), head of Bear Creek (August 12 and 15). 

Dendroica nigrescens (Townsend). Black-throated Gray Warbler. 
Three skins, nos. 19350-19352, from Scott River (June 11), and 
Summerville (immatures, July 18 and 19). 


Dendroica townsend: (Townsend). Townsend Warbler. One 
immature male, in full fall plumage, no. 19353, from head of 
Bear Creek, August 10. Probably a migrant. 

Dendroica occidentalis (Townsend). Hermit Warbler. Three 
immatures, nos. 19354-19356, from head of Grizzly Creek (July 28), 
head of Rush Creek (July 28), head of Bear Creek (August 13). 


Geothlypis trichas occidentalis Brewster. Western Yellowthroat. 
Four breeding adults, nos. 19357-19360, from Mayten and Scott 
River, June 4-11. Common on the more or less marshy ground 
around the “‘Big Spring’’ at Mayten. Nest with four eggs found 
there in bunch of dry tules, June 5. Nest with four eggs found 
near the ground in a cornus bush, June 11, at Scott River. 


Icteria virens longicauda Lawrence. Long-tailed Chat. Four 
skins, nos. 19361-19364, from Scott River, June 12 and 13. Here 
the bushes along the creek seemed alive with different birds, largely 
because of the chat’s voluble imitations of his various neighbors. 


Wilsoma pusilla chryseola Ridgway. Golden Pileolated Warbler. 
Nine skins, nos. 19365-19373, from Jackson Lake (June 16-18), 
head of Grizzly Creek (July 21), and head of Bear Creek 
(August 6-16). 

Cinclus mexicanus wnicolor Bonaparte. American Dipper. One 
juvenile, no. 193874, North Fork of Coffee Creek, July 5. 


1916] Kellogg: Mammals and Birds of Northern California 387 


Thryomanes bewicki drymoecus Oberholser. San Joaquin Wren. 
Three adult skins, no. 17372, from Helena, February 23, and nos. 
19375, 19376, from Scott River, June 11. 

Troglodytes aedon parkmani Audubon. Western House Wren. 
Ten specimens, nos. 19377-19386, from Scott River, head of Grizzly 
Creek, Summerville, head of Bear Creek, and Castle Lake, June to 
August. 

Certhia familiaris zelotes Osgood. Sierra Creeper. Ten speci- 
mens, nos. 19387-19396, from North Fork of Coffee Creek, South Fork 
of Salmon River, head of Grizzly Creek, and head of Bear Creek, 
July and August. 

Sitta carolinensis aculeata Cassin. Slender-billed Nuthatch. 
Two skins, nos. 19397, 19398, from Seott River, June 13. 

Sitta canadensis Linnaeus. Red-breasted Nuthatch. Three 
skins, nos. 19399-19401, from head of Rush Creek (July 28), and 
head of Bear Creek (August 6 and 10). 

Bacolophus imornatus inornatus (Gambel). Plain Titmouse. 
Three skins, nos. 17373, 17374, from Tower House, Mareh 2, and 
no. 19402, from Scott River, June 9. 

Penthestes atricapillus occidentalis (Baird). Oregon Chickadee. 
Four skins, nos. 19403-19406, from Scott River, six miles north- 
west of Callahan, June 10 and 13. Two are adults and two juvenals 
not fully grown, one of each on each of the two dates. This is the 
first unquestionably authentic record of this species for California. 

Penthestes gambeli gambeli (Ridgway). Mountain Chickadee. 
Twenty-two skins, nos. 19407-19428, from Jackson Lake, south fork 
of Salmon River, head of Grizzly Creek, head of Rush Creek, Kan- 
garoo Creek, head of Bear Creek, and Castle Lake, June 16 to 
August 19. 

Penthestes rufescens rufescens (Townsend). Chestnut-sided 
Chickadee. One skin, no. 17375, from Helena, February 15. Appar- 
ently numerous at this time and place, but not seen anywhere in 
the Trinity region during the summer. 

Psaltriparus minimus miuumus (Townsend). Coast Bush-tit. 
Five skins, nos. 17390-17392, 19429, 19430, from Tower House, 
March 7, and Scott River, June 8 and 11. For systematic status, 
see Swarth (1914, p. 510). 

Chamaea fasciata henshawi Ridgway. Pallid Wren-tit. Fifteen 
specimens, nos. 17376-17389, from Helena and Tower House, Feb- 


388 University of California Publications in Zoology (Vow. 12 


ruary 20 to March 7, and no. 19431, from Scott River, June 11. 
Most numerous at Tower House. 

Regulus satrapa olwaceus Baird. Western Golden-crowned 
Kinglet. Seven skins, nos. 17432-17488, from South Fork of Salmon 
River (July 14 and 16), head of Grizzly Creek (July 21-23), and 
head of Rush Creek (July 27). 

Regulus calendula (Linnaeus). Ruby-crowned Kinglet. Four 
skins, no. 17393, from Helena (February 14), no. 17394, from Tower 
House (March 7), no. 19439, from South Fork of Salmon River 
(July 16), and no. 19440, from head of Bear Creek (August 18). 

Myadestes townsendi (Audubon). Townsend Solitaire. Five 
skins, nos. 17395, 17396, from Helena (February 14 and 23), no. 
17397, from Tower House (March 1), no. 19441, from North Fork 
of Coffee Creek (July 3), and no. 19442, from Kangaroo Creek 
(August 4). 

Hylocichla ustulata ustulata (Nuttall). Russet-backed Thrush. 
Two adult skins, nos. 19443, 19444, from Jackson Lake (June 25), 
and head of Grizzly Creek (July 23). Nest and four eggs (no. 
1097) taken at Jackson Lake, June 25. This was situated three 
feet above the ground on a sagging branch of a small fir, and was 
loosely constructed of dry fir twigs, dead ferns and moss. 

Hylocichla ustulata swainsont (Tschudi). Olive-backed Thrush. 
One adult male, no. 19447, from head of Grizzly Creek, 6000 feet 
altitude, July 22. 

Hylocichla guttata nanus (Audubon). Dwarf Hermit Thrush. 
Seven skins, nos. 17398-17404, from Helena and Tower House, 
February 13 to March 7. 

Hylocichla guttata slevint Grinnell. Monterey Hermit Thrush. 
Two specimens: no. 19445, female juvenal, head of Rush Creek, 
July 29; no. 19446, female immature, head of Bear Creek, 
August 12. 

Planesiicus nugratorius propinquus (Ridgway). Western Robin. 
Hight skins, nos. 17405, 17406, 19448-19453, from Helena (February 
24), Mayten (June 5), Jackson Lake (June 19 and 24), and head 
of Bear Creek (August 10 and 16). At the latter locality and 
dates robins had gathered in flocks, feeding in the ceanothus brush. 

Ixoreus naevius naevius (Gmelin). Varied Thrush. Three 
skins, nos. 17407-17409, from Helena and Tower House, February 13 
to March 7. 


1916] Kellogg: Mammals and Birds of Northern California 389 


Sialia mexicana occidentalis Townsend. Western Bluebird. 
Three specimens: nos. 17410, 17411, from Tower House, March 2; 
no. 19454, from Scott River, June 13. Common at Tower House 
in March, but notably scarce in summer throughout the Trinity 
region. 

Sialia currucoides (Bechstein). Mountain Bluebird. One adult 
male, no. 19455, from Wildcat Peak, June 28. 


Transmitted June 27, 1914. 


LITERATURE CITED 


BAILEY, V. 
1897. Revision of the American voles of the genus Hvotomys. Proce. 
Biol. Soc. Wash., 11, 113-138, pl. 3. 
Banes, O. 
1899. Descriptions of some new mammals from western North America. 
Proc. New England Zool. Club, 1, 65-72. 
Exxiot, D. G. 
1901. A synopsis of the mammals of North America and the adjacent 
seas. Field Columbian Mus., Zool, Ser., 2, pp. vx + 471, 49 pls., 
94 figs. in text. 
GRINNELL, J. 
1912. The northern brown towhee. Condor, 14, 199. 
1914. A new red-winged blackbird from the Great Basin. Proc. Biol. 
Soc. Wash., 27, 107-108. 
Howe tu, A. H. 
1914. Revision of the American harvest mice (genus Reithrodontomys). 
U. S. Dept. Agric., Bur. Biol. Surv., N. Amer. Fauna, 36, 97 pp., 
7 pls., 6 figs. in text. 
1915. Descriptions of a new genus and seven new races of flying 
squirrels. Proc. Biol. Soc. Wash., 28, 109-114. 
KELLoGe, L. 
1911. <A collection of winter birds from Trinity and Shasta counties, 
California. Condor, 13, 118-121. 
1914. Aplodontia chryseola, a new mountain beaver from the Trinity 
region of northern California. Univ. Calif. Publ. Zool., 12, 
295-300. 
Merriam, C. H. 
1890. Descriptions of twenty-six new species of North American mam- 
mals. U. S. Dept. Agric., Div. Orn. and Mamm., N. Amer. 
Fauna, 4, pp. v + 60, 3 pls., 3 figs. in text. 
1896. Synopsis of the weasels of North America. Jbid., 11, 44 pp., 
5 pls., 16 figs. in text. 


390 Unversity of California Publications in Zoology (Vou. 12 


1897a. Descriptions of two new red-backed mice (Hvotomys) from 
Oregon. Proc. Biol. Soc. Wash., 11, 71-72. 
1897b. Description of a new flying squirrel from Ft. Klamath, Oregon. 
Tbid., 11, 225. 
1899. Results of a biological survey of Mount Shasta, California. 
U. S. Dept. Agric., Div. Biol. Surv., N. Amer. Fauna, 16, 179 pp., 
5 pls., 46 figs. in text. 
190la. Descriptions of twenty-three new pocket gophers of the genus 
Thomomys. Proc. Biol. Soe. Wash., 14, 107-117. 
19010. Two new rodents from northwestern California. Jbid., 14, 
125-126. 
NELSON, E. W. 
1909. The rabbits of North America. U. S. Dept. Agric., Bureau 
Biol. Surv., N. Amer. Fauna, 29, 314 pp., 13 pls., 19 figs. in text. 
SwarrH, H. S: 
1913. A revision of the California forms of Pipilo maculatus Swain- 
son, with description of a new subspecies, Condor, 15, 167-175, 
1 map. 
1914. A revision of the bush-tits of California. Auk, 31, 499-526, 1 pl. 


PLATE 15. 


Fig. 1. View towards the east, of a portion of Scott River Valley, 
Siskiyou County, California, about six miles north of Callahan, altitude 
about 3,000 feet. Photograph taken June 13, 1911. The floor of the 
valley is timbered with yellow pine, cottonwood and willow. The hills 
shown in the picture were not visited, but looked much drier and were 
decidedly more sparsely timbered than the hills on the west side, at 
the base of which camp was located. These western hills were well 
covered with sage-brush, deer-brush and scattering groves of yellow pine. 
The sandy ground of an old creek bed proved to be good trapping 
ground for Dipodomys californicus trinitatis. Other small mammals of the 
lower lands were Peromyscus maniculatus gambelii, Butamias amoenus 
amoenus, Eutamias senex, and Citellus douglasii. On the sidehill Lepus cali- 
fornicus californicus was common. Dryobates pubescens gairdneri, As- 
tragalinus tristis salicamans, Astragalinus psaltria hesperophilus, Chon- 
destes grammacus strigatus, Melospiza melodia fisherella, Pipilo macu- 
latus falcinellus, Zamelodia melanocephala capitalis, Dendroica aestiva 
brewsteri, Icteria virens longicauda and Penthestes atricapillus occi- 
dentalig were birds which frequented the willow thickets along the small 
ereek beds. On the sidehill were seen Lophortyx californica vallicola, 
Melanerpes formicivorus bairdi and Piranga ludoviciana. The life-zone 
may be considered mixed Upper Sonoran and Transition, prevalently 
the former. 


Fig. 2. North side of Saloon Creek divide, Siskiyou County, Cali- 
fornia, altitude 6,275 feet. Photograph taken July 10, 1911. A scattering 
growth of silver pine and red fir on sidehill, and patches of alder along 
creek bottom. As a collecting station the locality was chosen chiefly 
because of the presence there of a colony of the golden-mantled ground 
squirrel (Callospermophilus chrysodeirus trinitatis). The small Klamath 
chipmunk (ELutamias amoenus amoenus) was almost as numerous and 
there was abundant sign of gophers (Thomomys monticola pinetorum). 
The life-zone here represented was the Canadian division of the Boreal. 


[ 392 ] 


PLATE 16. 


Fig. 3. Burrows of the Trinity mountain beaver (Aplodontia chrys- 
eola) near Hunters’ Camp, Grizzly Creek Canon, Trinity County, Califor- 
nia. Photograph taken July 19, 1911. The animals had taken advantage 
of the protection afforded by the tangled roots of a fallen tree. Although 
the ground was here dry, there was running water within a few yards 
to one side. 


Fig. 4. Mouth of burrow of Aplodontia chryseola, showing heap of 
freshly-cut vegetation. Photograph taken near Hunters’ Camp, Grizzly 
Creek Cafion, Trinity County, California, July 19, 1911. The presence 
in this locality, as well as at several other points, of cuttings of grass 
and leaves seems to establish the fact that these mammals gather and 
dry vegetable materials for subsequent use. 


[ 394 ] 


PLATE 17. 


Fig. 5. View taken July 28, 1911, looking towards head of Grizzly 
Creek, Trinity County, California, altitude about 6,000 feet. The timber 
is white pine, red fir, and hemlock, scattered through a meadow and 
reaching up on the rocky slopes of Thompson Peak, the summit in the 
background. A bank of perpetual snow feeds a lake from which a water- 
fall (shown in left center) descends to form Grizzly Creek. Small mam- 
mals were not numerous but included Microtus mordax mordazr, Zapus 
trinotatus alleni, and the two species of Hutamias, amoenus and senez. 
Aplodontia chryseola was fairly abundant on the west side of the canon, 
and a number of black-tailed deer were seen in the neighborhood. Birds 
noted were: Nuttallornis borealis, Cyanocitta stelleri frontalis, Junco 
oreganus thurberi, Lanivireo solitarius cassini, Vermivora rubricapilla gut- 
turalis, Vermivora celata lutescens, Dendroica auduboni auduboni, Den- 
droica occidentalis, Certhia familiaris zelotes, Penthestes gambeli gambeli, 
Reyulus satrapa olivaceus, Hylocichla ustulata ustulata, Hylocichla ustu- 
lata swainsoni (one specimen). The life-zone represented is evidently 
Boreal, in its Canadian and Hudsonian divisions. 


Fig. 6. View taken July 31, 1911, looking southwest from head of 
Rush Creek, Siskiyou County, California, towards the Salmon Mountains; 
altitude about 6,400 feet. A typical Boreal meadow of luxuriant grasses, 
with thickets of alder and patches of hellebore. Red firs skirt the edge 
of the meadow and, interspersed with hemlock, extend in dense stand 
to the top of the divide. Below the meadow the silver pine and cedar 
predominate. Small mammals taken in and near the meadow were: 
Sorer vagrans amoenus, Peromyscus maniculatus gambelii, Microtus mor- 
dax mordaz, Thomomys monticola pinetorum and Zapus trinotatus al- 
leni. Glaucomys sabrinus flaviventris was abundant in the groves of 
red fir, and in a side canon three marten (Martes caurina caurina) were 
secured. Birds were confined chiefly to the meadow and its vicinity, and 
included Oreortyx picta picta, Dendragapus obscurus sierrae, Cyanocitta 
stelleri frontalis, Junco oreganus thurberi, Melospiza lincolni lincolni, 
Dendroica occidentalis, Penthestes gambeli gambeli and Hylocichla gut- 
tata slevini. 


[ 396 ] 


oO 
G) 
( 


3] PLATE 17 


UNIV. CALIF, PUBL. ZOOL. VOL. 16 [KELL¢ 


PLATE 18. 


Skulls of Aplodontia; all x 0.62. 


7. A. chryseola Kellogg: ¢@, no. 18331, Mus. Wert. Zool., head of 


Grizzly Creek, Trinity County, California. 


. A. californica (Peters): ¢, no. 18663, Mus. Vert. Zool., Blue 
Canon, Placer County, California. 


. A. chryseola Kellogg: ¢@, type, no. 13328, Mus. Vert. Zool., Jack- 
son Lake, Siskiyou County, California. 

Note in A. chryseola (the two end skulls): the short rostrum, 
short and broad nasals, expanded frontal region, broad rostrum 
posteriorly, less squarish zygomatic arch, broad interorbital con- 
striction, and lengthened auditory tubes. 


[ 398 ] 


UNIVERSITY OF CALIFORNIA PUBLICATIONS 
IN 


ZOOLOGY 


Vol. 12, No. 14, pp. 399-410 Issued January 27, 1916 


AN ANALYSIS OF THE VERTEBRATE FAUNA 
OF THE TRINITY REGION OF NORTH- 
ERN CALIFORNIA 


BY 


JOSEPH GRINNELL 


(Contribution from the Museum of Vertebrate Zoology of the University of California) 


From the array of facts presented in the foregoing paper (Kel- 
logg, 1916), in conjunction with what is known of adjacent areas, 
it seems possible to draw some general inferences as to the composi- 
tion and derivation of the vertebrate fauna of the Trinity region. 
Faunas vary through a wide range in degree of distinctness, from 
that which is searecely different from the one or ones adjacent, to 
the fauna which is of marked peculiarity, showing throughout 
nearly all its elements totally distinct characters. An example of 
the latter would be the San Diegan fauna as compared with the 
adjacent Colorado Desert fauna (see Grinnell and Swarth, 1913, 
page 388). Of far less degree of difference, we may cite the Santa 
Cruz faunal division of the Humid Coast Belt as compared with 
the Marin division of the same belt. 

The difficulties arising in attempts to diagnose and properly 
classify faunas are surprisingly similar to those encountered in 
dealing with subspecies, species, genera and other systematic groups. 
Not only is there such wide variability in amounts of difference 
between faunas as to make the matter of rank often indeterminable, 
but exact geographic boundaries may be impossible to fix because of 
intergradation over intermediate territory of greater or less width. 
As in some cases of intergrading series of subspecies, the location of 
lines of separation may be subject only to arbitrary choice, so that 
where six faunas might be recognized by one student, only three 
would be thought properly distinguishable by another. 


400 University of California Publications in Zoology ‘Vou. 12 


The fauna of the Trinity region seems to be best dealt with, by 
considering its Boreal zonal elements, and its Sonoran (or Austral) 
zonal elements, separately. Adjacent Boreal faunas are those of: 
(1) the Sierra Nevada of east central California, (2) the Cascades 
of Oregon, (3) the Humboldt Bay division of the Humid Coast 
Belt. Adjacent Sonoran faunas are those of: (1) the Sacramento 
Valley, (2) the Modoe region of northeastern California, (3) the 
Humid Coast Belt. The following tabulations will serve to segre- 
gate the pertinent facts appropriately : 


Table 1. Boreal species of the Trinity region which occur also on both 
the Sierra Nevada and the Cascade Mountains, but not as regular members 
of the humid coast fauna. 


MAMMALS 


Canis lestes (there are indications that a separate form exists in the 
area towards the coast from Mount Shasta, of which the Trinity 
region is a part). 

Martes caurina caurina (subspecifie status in doubt; probably differ- 
ing from the Cascade animal and most nearly identical with a 
Sierran race). 

Martes pennanti pacifica (subject to same remark as above). 

Microtus mordax mordaz. 

Eutamias amoenus amoenus. 

Eutamias senex. 

Callospermophilus chrysodeirus trinitatis (very slightly different 
from C. c. chrysodeirus of the Sierras and Cascades). 


BIRDS 


Dryobates villosus orius (intergradient towards the humid coast form 
harrisi). 

Xenopicus albolarvatus albolarvatus. 

Asyndesmus lewisi (possibly not to be considered Boreal). 

Stellula calliope. 

Empidonax wrighti (possibly a straggler, merely). 

Nucifraga columbiana. 

Carpodacus cassini. 

Melospiza lincolni lincolni. 

Oreospiza chlorura. 

Vermivora ruficapilla gutturalis. 

Dendroica auduboni auduboni. 

Dendroica nigrescens (not strictly Boreal). 

Dendroica occidentalis. 

Certhia familiaris zelotes. 

Penthestes gambeli gambeli. 

Myadestes townsendi. 

Sialia currucoides. 


1916] Grinnell: Analysis of Trinity Fauna 401 


Table 2. Boreal species of the Trinity region which are identical with, 
or show closest affinities to, representatives on the Sierra Nevada, but not on 
the Cascades. 


MAMMALS 


Sorex vagrans amoenus (very slightly differentiated from S. v. 
vagrans, of the humid coast belt and the Cascades). 

Mustela muricus. 

Microtus montanus montanus (found only at Mayten; occurs also in 
the Klamath region at extreme southern end of Cascades). 

Thomomys monticola pinetorum (very slightly different from 7. m. 
monticola of the Sierra Nevada). 

Zapus trinotatus alleni. 

Aplodontia chryseola (decidedly different from A. californica of the 
Sierra Nevada). 

Sciurus douglasii albolimbatus. 

Glaucomys sabrinus flaviventris. 


BIRDS 


Dendragapus obscurus sierrae (intergradient towards the humid 
coast form fuliginosus). 

Sphyrapicus varius daggetti. 

Cyanocitta stelleri frontalis (slightly intergradient towards the coast 
form carbonacea). 

Junco oreganus thurberi. 

Passerella iliaca megarhyncha. 


Table 3. Boreal species of the Trinity region which are identical with, 
or show closest affinities to, representatives on the Cascade Mountains, but 
not on the main Sierra Nevada. 


MAMMALS 


Mustela saturata. 
Neotoma cinerea occidentalis (occurs also on northern Sierra Nevada 


and locally to the coast). 

Evotomys obscurus (also on extreme northern Sierra Nevada). 

Lepus washingtonii klamathensis (the snowshoe rabbit reported under 
this name from the central Sierra Nevada, is probably distinct 
from klamathensis). 


BIRDS 


(none ) 


Table 4. Boreal species of the Trinity region which are identical with, 
or show closest affinities to, representatives in the humid coast belt. 


MAMMALS 


Scapanus latimanus latimanus (wide ranging zonally to the south- 
ward through the coast ranges; not a typical humid coast form; 
group relationships not yet worked out satisfactorily). 


a 


University of California Publications in Zoology [Vou. 12 


Neurotrichus gibbsi major (status of races not worked out satisfac- 
torily; while the genus belongs to the northwest coast belt, there 
may prove to be a Sierran subspecies with which the Trinity 
animal belongs). 

Sorex montereyensis montereyensis (but slightly different from the 
Sierran form, S. m. mariposae). 

Lynx fasciatus. 

Odocoileus columbianus columbianus. 


BIRDS 


Oreortyx picta picta. 

Dendragapus obscurus fuliginosus (intergradient towards the Sierran 
form sierrae). 

Melospiza melodia rufina (in winter, only). 

Penthestes atricapillus occidentalis. 

Penthestes rufescens rufescens (in winter, only?). 

Hylocichla guttata slevini. 


Table 5. Boreal species of the Trinity region which appear to be 
endemic, that is, different from related forms in either the Sierra Nevada, 
the Cascades, or the humid coast belt. 


MAMMALS 


Thomomys monticola pinetorum (extends also to Mount Shasta). 
Aplodontia chryseola. 

Callospermophilus chrysodeirus trinitatis. 

Glaucomys sabrinus flaviventris. 


BIRDS 


(mone) 


Table 6. Sonoran species of the Trinity region which belong to variable 
groups and which are identical with, or show nearest affinities to, repre- 
sentatives in the Sacramento Valley. 


MAMMALS 


Bassariscus astutus raptor (extends also through the coast ranges to 
the southward and into Oregon on the north). 

Spilogale phenax phenax (wide-ranging through the coastal region to 
the south). 

Mephitis occidentalis occidentalis (wide-ranging through many faunas 
to the south). 

Peromyscus boylii boylii. 

Peromyscus truei gilberti. 

Microtus californicus californicus. 

Thomomys leucodon navus. 

Dipodomys californicus trinitatis (slightly different from the form 
D. c. californicus of the upper Sacramento Valley). 

Lepus californicus californicus (also ranges locally nearly or quite 
through the coast belt). 


1916] Grinnell: Analysis of Trinity Fauna 403 


BIRDS 


Lophortyx californica californica. 
Astragalinus tristis salicamans (also in humid coast belt). 
Astragalinus psaltria hesperophilus, 
Chondestes grammacus strigatus (occurs also in the Great Basin). 
Pipilo maculatus falcinellus. 
Pipilo crissalis carolae. 
Thryomanes bewicki drymoecus. 
Baeolophus inornatus inornatus. 
Chamaea fasciata henshawi. 
Table 7. Sonoran species of the Trinity region which belong to variable 
groups, and which are identical with, or show nearest affinities to, repre- 
sentatives in the Modoc region. 


MAMMALS 


Reithrodoniomys megalotis klamathensis. 


BIRDS 


Agelaius phoeniceus nevadensis (found at Mayten only, in Shasta 
Valley; so not really a bird of the Trinity region as explicitly 
defined). 

Melospiza melodia fisherella. 


Table 8. Sonoran species of the Trinity region which belong to variable 
groups and which are identical with, or show nearest affinities to, repre- 
sentatives in the humid coast belt. 


MAMMALS 


Neotoma fuscipes fuscipes (not exclusively a humid coast form; ranges 
across head of Sacramento Valley). 

Sylvilagus bachmani ubericolor (extends also interiorly across head of 
Sacramento Valley). 


BIRDS 


Dryobates pubescens gairdneri (tending towards the race twrati, of 
the Sacramento Valley and other faunas to the southward). 

Psaltriparus minimus minimus (vanges somewhat beyond the most 
restricted confines of the humid coast belt). 


Table 9. Sonoran species of the Trinity region which are apparently 
endemic, that is, different from related forms in any of the other faunas. 


MAMMALS 


Dipodomys californicus trinitatis. 


BIRDS 
(none) 


Table 10. Species which are either so widespread through all the faunas 
here concerned as to be non-significant in the present study, or which are 
of unknown status. 


404 


Unwersity of California Publications in Zoology (Vou. 12 


MAMMALS 


Myotis longicrus longicrus. 

Lasionycteris noctivagans. 

Eptesicus fuscus fuscus. 

Ursus americanus. 

Urocyon cinereoargenteus townsendi (merging into the races U. c. 
californicus and U. c. sequoversis immediately to the south of the 
Trinity region). 

Procyon psora pacifica. 

Mustela vison energumenos, 

Felis oregonensis oregonensis. 

Peromyscus maniculatus gambelii (slightly intermediate towards the 
humid coast form rubidus). 

Citellus douglasii. 

Sciurus griseus griseus (apparently non-differentiated in the three 
Boreal faunas here concerned! ). 


BIRDS 


Actitis macularius. 

Oxyechus vociferus. 

Zenaidura macroura marginella. 

Accipiter velox. 

Accipiter cooperi. 

Astur atricapillus striatulus. 

Falco sparverius sparverius. 

Phloeotomus pileatus abieticola. 

Melanerpes formicivorus bairdi. 

Colaptes cafer collaris (not approaching the northern humid coast 
form saturatior). 

Selasphorus rufus (probably a migrant, only). 

Sayornis nigricans (in winter, only, and not in the main Trinity 
region. 

Nuttallornis borealis. 

Myiochanes richardsoni richardsoni. 

Empidonaz trailli trailli. 

Empidonaz hammondi (not certainly breeding). 

Aphelocoma californica californica. 

Sturnella neglecta. 

Icterus bullocki. 

Euphagus cyanocephalus. 

Hesperiphona vespertina montana (probably only in winter). 

Carpodacus purpureus californicus. 

Carpodacus mexicanus frontalis. 

Spinus pinus pinus. 

Passer domesticus. 

Zonotrichia coronata (in winter, only). 

Spizella passerina arizonae. 

Passerella iliaca unalaschcensis (in winter, only). 

Passerella iliaca meruloides (in winter, only). 

Zamelodia melanocephala capitalis. 

Passerina amoena. 


1916] Grinnell: Analysis of Trinity Fauna 405 


Piranga ludoviciana. 

Bombycilla garrula (in winter, only). 
Vireosylva gilva swainsoni. 

Lanivireo solitarius cassini. 

Vireo huttoni huttoni. 

Vermivora celata lutescens. 

Dendroica aestiva brewsteri. 

Dendroica townsendi (probably a transient, only). 
Geothlypis trichas occidentalis. 

Icteria virens longicauda. 

Wilsonia pusilla chryseola. 

Cinclus mexicanus unicolor. 

Troglodytes aedon parkmani. 

Sitta carolinensis aculeata. 

Sitta canadensis. 

Regulus satrapa olivaceus. 

Regulus calendula. 

Hylocichla ustulata ustulata. 

Hylocichla ustulata swainsoni (possibly an exceptional occurrence). 
Hylocichla guttata nanus (in winter, only). 
Planesticus migratorius propinquus. 
Ixoreus naevius naevius (in winter, only). 
Sialia mexicana occidentalis. 


Kellogg (1916) lists 47 mammals and 95 birds from the Trinity 
region, a total of 142 vertebrate species. In spite of the very high 
grade of field-work carried on by Misses Alexander and Kellogg dur- 
ing the time of their exploration in the region, it is not for a 
moment to be inferred that this is the total number of mammal and 
bird species regularly occurring in the region. But it is the known 
fauna. A few included species may prove to be stragglers and not 
really to be considered in faunal analysis. Obviously the present 
assignment of species must be accepted only as provisional; at the 
same time it is probable that the general situation as regards pro- 
portional constituency of the various categories of species would 
remain very nearly the same as here indicated, even after such 
intensive and long-continued survey as might be devoted to the 
Trinity region in the future. Increments from future field-work 
would be likely to be distributed pretty evenly among the several 
categories so that the ratios would continue but little if any changed. 

As indicated in the foregoing tables, 65 species out of the total 
142 are either widespread through all the faunas here concerned 
or are of unknown status, in either case being removed from con- 
sideration in any attempt to establish differences between the faunas. 
This disposes at once of 46 percent of the Trinity fauna. , 


406 University of California Publications in Zoology (Vor. 12 


As is well known, the appertaining faunas of greatest contrast 
are those of the humid coast belt and the Sierra-Cascades. Twenty- 
four Boreal species, or 17 percent of the Trinity mammals and birds, 
occur also on both the Sierra Nevada and the Cascade Mountains, 
but not regularly as elements of the humid coast fauna. Also there 
are thirteen species, or 9 percent, of the Trinity boreal fauna which 
are identical with, or show closest affinities to, representatives on 
the Sierra Nevada, but not on the Cascades; and there are four 
species, or 3 percent, of the Trinity Boreal fauna which are identi- 
cal with, or show closest affinities to, representatives on the Cas- 
cade Mountains, but not on the main Sierra Nevada. Mount Shasta 
is here considered as properly included in the Sierran fauna. This 
mountain, in spite of its relative isolation, certainly shows only 
minor differences from the main central Sierras (see Merriam, 1899, 
pp. 71-81). 

It is thus to be seen that a total of 41 mammals and birds, or 28 
percent of its species, are shared by the Trinity fauna exclusively 
with that of both the Sierras and Cascades together. On the other 
hand, there are only eleven, or less than 8 percent, of the Trinity 
species which are shared exclusively with the closely adjacent humid 
coast fauna. As regards its Boreal elements, therefore, we may con- 
clude that the Trinity fauna is much more nearly allied to the 
Sierra-Cascades than to the humid coast belt. 

The fauna of the Trinity region is preponderantly Boreal; much 
the larger part of the territory lies in the Transition and Canadian 
life-zones. The strictly Soncran (Upper Sonoran) elements occur 
only in the valleys of low altitude. Of these there are three, of small 
area, situated in the interior of the region: Scott River, south fork 
of the Salmon River at Summerville, and Trinity River in the vicin- 
ity of Helena; and two on the borders: Shasta Valley, in which 
Mayten is situated, and the upper extremity of the Sacramento 
Valley in the vicinity of Tower House. Taking into account chiefly 
the Sonoran ‘‘islands’’, we find that the Trinity region possesses 
18 species in common with the Sacramento Valley. This is some- 
what less than 12 percent of the total mammal and bird fauna of 
the Trinity area. Three species, or little more than 2 percent, are 
shared with the Modoe region of extreme northeastern California 
(and which has its western confines in Shasta Valley); while only 
four Sonoran species, not quite 3 percent, oceur also in the humid 


coast belt. 


1916] Grinnell: Analysis of Trinity Fauna 407 


It would thus appear that, as regards its Sonoran fauna, the 
Trinity region allies itself strongly with the Sacramento fauna 
rather than with either of the other two named. 

As far as specimens have become available and carefully studied 
(Kellogg, 1916), only five endemic forms have been found, all 
mammals. Four of these are Boreal: Thomomys monticola pine- 
torum Merriam, Aplodontia chryseola Kellogg, Callospermophilus 
chrysodeirus trinitatis Merriam, and Glaucomys sabrinus flaviventris 
Howell; one is Sonoran: Dipodomys californicus trinitatis Kellogg. 
Only one of the five is well-marked, the Aplodontia, the other four 
being but faintly characterized. This indicates a differentiation 
center of weak power. 

It is apparent that the Trinity region shows but weak faunal 
individuality. It is not sharply set off, except on the west, nor does 
it contain notably distinct forms. This relative unimportance as a 
differentiation center is doubtless due to its small area and to the lack 
of efficient barriers which would prevent intercrossing with repre- 
sentatives in adjacent faunas. No very low zone is interposed, the 
Transition, merely, being continuous between the Boreal areas. The 
climatic features, too, are clearly not so pronouncedly different from 
those of the Sierra Nevada as to make up for the very short distance 
between the main Trinity mass and the Sierras. 

There must be a very abrupt line of demarcation between the 
Trinity fauna and the humid coast fauna, for we have very few forms 
venturing from the latter into the former, in spite of the very short 
distance. It is true that with a few birds we find intergrades in the 
Trinity region between typical Sierran and typical coast-belt forms ; 
for example, in the genera Dendragapus, Dryobates and Cyanocitta. 
But in by far the greater number of appropriate cases, in both birds 
and mammals, the Trinity representatives are unequivocally of in- 
terior forms, and this in spite of the fact that practically all of the 
Trinity drainage is west directly to the Pacifie and hes within a 
maximum distance of ninety miles of the coast. The climatic barrier, 
of abruptly and greatly increased humidity on the west, is evidently 
much more efficient in checking the spread of species than the various 
Transition-zone ‘‘gaps’’ between the mountain masses of the interior 
or the Transition-zone divides between the “‘islands’’ of Upper 
Sonoran. 

The whole idea of basing the efficacy of zonal ‘‘gaps’ 
summation of the species occurring on either side (see Merriam, 


’ 


on the 


408 University of California Publications in Zoology  {Vor. 12 


1899, pp. 69-86) must to the writer’s mind be reconsidered. Zone 
must be thought of in conjunction with conditions of varying humid- 
ity, associational features, and relative sizes of all the segregated 
areas concerned. Species may be delimited in their ranges by other 
factors than that of temperature. This remark is not, however, 
meant as implying a denial that temperature is the one factor most 
frequently limiting the ranges of species. A narrow Transition-zone 
cap between two Boreal areas, as that at Sisson, between mounts 
Eddy and Shasta, cannot serve as a particularly effective barrier 
between the Boreal species encountering it, unless it be coupled with, 
on opposite sides, marked differences in humidity or associational 
conditions, which, however, do not appear to exist in the present 
instance. 

The problem here arises, how to treat the Trinity region in faunal 
nomenclature. What is the criterion for nominal separation of 
faunas? What percentage of its species have to be peculiar, or what 
minimum ratio of differences obtain in comparison with the animal 
life of adjacent areas, to render the Trinity assemblage of forms 
worthy of separate recognition by name on our fanual maps? Mount 
Shasta has but two mammals and two birds (even these doubtful), 
which do not occur on the central Sierras; it is chiefly characterized 
by absence of forms (due to its small area). We are therefore led 
to include Mount Shasta with the Sierra Nevada in the ‘‘Sierra 
Nevadan Faunal Area’’. But we do not hesitate to recognize as 
distinct the Cascade fauna and the Sierra Nevadan fauna. And the 
Humid Coast fauna is conspicuously peculiar. 


The difficulties here encountered have been alluded to in a pre- 
ceding paragraph. The Trinity region in its vertebrate animal life 
resembles much more closely the Sierras than the Cascades; it is in 
a way intermediate, but has in addition some humid coast elements. 
The differences between the Trinity and Sierran faunas are con- 
cerned with only eleven percent of its species all told. Is this a 
sufficient amount of difference to warrant their recognition as dis- 
tinct faunal areas, as indicated on a current distributional map 
(Grinnell, 1913, pl. 16)? Possibly not, though the writer believes 
that for purposes of faunal analysis such fine distinctions may be 
useful. It will depend on circumstances, for, with wider problems 
in view, it might for the moment be better to lump the Trinity area 
in with the Sierra Nevada under a common designation. Expediency 
will fix the criterion for recognizing faunal division, and this may 


1916] Grinnell: Analysis of Trinity Fauna 409 


vary as circumstances vary, just as with the recognition of genera, 
families, etc. The term subfauna has been previously employed for 
the more slightly characterized assemblages of animal inhabitants, 
and this might be a useful term to adopt regularly in connection 
with such minor faunal divisions as that of the Trinity region. 

The exact location of boundaries for the Trinity subfauna is a 
matter of uncertainty. This uncertainty arises from two circum- 
stances: first, that, as experience elsewhere would lead one to expect, 
there are broad marginal belts of intermingling, rather than sharp 
lines of demarcation; and, second, lack as yet of exploration of much 
of the surrounding territory. No field-work has been done by anyone 
connected with the California Museum of Vertebrate Zoology on a 
line directly to the coast west from the Salmon Mountains. To the 
north, the Siskiyou Mountains, across the deep Klamath River canon, 
are so far known only as regards their birds (see Anderson and Grin- 
nell, 1903). The birds are indicative of close faunal similarity of 
the Siskiyou and Trinity mountain masses. To the south, the Yolla 
Bolly country presents some obvious peculiarities of its own, but 
here, again, no adequate study has been accorded the animal life, 
and conclusions are therefore best deferred. To the east, the Trinity 
area seems satisfactorily bounded by the upper Sacramento Valley 
and by Shasta Valley together with the interconnectant Sisson gap, 
extending between mounts Shasta and Eddy. 

Summary.—tThe collections of specimens thus far available from 
the Trinity region cannot be considered anywhere nearly complete; 
neither is there sufficient information at hand, published or otherwise, 
concerning the Cascades. Even with the best-worked of the faunas 
here concerned, that of the Sierra Nevada particularly in its Mount 
Shasta portion, much remains in doubt. The statistics here set forth 
must therefore be considered provisional. Nevertheless the general 
conclusions are believed closely to approximate the truth. 

The Trinity region as regards its Boreal fauna is clearly far more 
closely allied to the Sierra-Cascade fauna than to the Humid Coast 
fauna. It is nearer to the Sierran fauna than to the Cascade fauna; 
indeed it might with propriety be included in the Sierra Nevadan 
faunal area, ranking merely as a subfauna. 

The Trinity region contains some Sonoran 
of these is most closely similar to that of the Sacramento Valley ; 
there are a few Great Basin or Modoc forms, and but scanty evidence 


> 


‘islands’’. 


‘ 


The fauna 


of humid coast intrusion. 


410 University of California Publications in Zoology  [Vou. 12 


The Trinity region shows but very slight endemic individuality. 
It possesses but five distinguishable races or species of its own, four 
of which are Boreal and one Sonoran. Only one of these is well 
marked. 

The failure of the Trinity Mountains to have developed a mark- 
edly distinct fauna from that of the Sierra Nevada, may be ascribed 
to three conditions: (1) Absence of extreme, that is, practically 
insurmountable, barriers, such as a continuous body of water, or a 
strip of the Sonoran zone, or a belt of excessive aridity; (2) close 
similarity in those features of climate included in the term humidity, 
for zonal identity implies similar temperature conditions at least as 
to mean; (3) small area as compared with that of adjacent mountain 
masses which, because of the greater mass of their fauna, have 
exerted a dominating influence in the interacting processes of 
invasion. 


Transmitted June 29, 1914. 


LITERATURE CITED 


Anpverson, M. P., and GRINNELL, J. 
1903. Birds of the Siskiyou Mountains, California: a problem in dis- 
tribution. Proc. Acad. Nat. Sci. Phila., 1903, 4-15. 


GRINNELL, J. 

1913. A distributional list of the mammals of California. Proc. Calif. 

Acad. Sci., (4) 3, 265-390, pls. 15, 16. 
GRINNELL, J., and SwartTH, H. S. 

1913. An account of the birds and mammals of the San Jacinto area of 
southern California, with remarks upon the behavior of geo- 
graphic races on the margins of their habitats. Univ. Calif. Publ. 
Zool., 10, 197-406, pls. 6-10. 


Ketoae, L. 
1916. Report upon mammals and birds found in portions of Trinity, 
Siskiyou and Shasta counties, California, with description of 
a new Dipodomys. Ibid., 12, 335-398, pls. 15-18. 


Merriam, C. H. 
1899. Results of a biological survey of Mount Shasta, California. Usiss 
Dept. Agric., Div. Biol. Surv., N. Amer, Fauna, 16, 179 pp., 5 


pls., 46 figs. in text. 


UNIVERSITY OF CALIFORNIA PUBLICATIONS— (Continued) 


‘ZOOLOGY, Vol. 12 (Continued) 

18. Report upon Mammals and Birds found in Portions of Trinity, 
Siskiyou and Shasta Counties, California, by Louise Kellogg. Pp. 
335-398, plates 15-18. 

14, An Analysis of the Vertebrate Fauna of the Trinity Region of 
Northern California, by Joseph Grinnell. Pp. 399-410. 

Nos. 13 and 14 in one cover. January, 1916-2-22 22.02... ae 


5 


Vol. 13. 1. The Schizopoda of the San Diego Region, by Calvin O. Esterly. 

PD 20, splates: 1-25 A prs LOT Be es eg ee a paola 

« - 2. A Study of the Occurrence and Manner of Distribution of the 

ae Ctenophora of the San Diego Region, by Calvin.O. Esterly. Pp. 

BP-38, Aprile TOU isco ect ech ages RE cae ase eee seas ce acen oregano 

8. A New Self-Regulating Paraffin Bath, by C. W. Woodworth. . Pp. 

39-42-2: text figures: April, 19 EA aaa retoncencede canna etunn 

4. Diplodinium ecaudatum, with an Account of Its Neuromotor Ap- 

paratus, by Robert G. Sharp. Pp, 43-122, plates 3-7, 4 text figures. 

US page co Yeates Sass a RE ent Ae OR ce © ARPS ort Sa ROO RRE PA EA 

5. The Vertical Distribution and Movements of the Schizopoda of the 

San Diego Region, by Calvin O. Esterly.. Pp. 123-145. May, 1914 

6. The Anatomy of Heterodontus francisci. I. The Exoskeleton, by 

J. Frank Daniel. Pp. 147-166, plates 8-9, 4 text figures. May 23, 

EGTA RS iS RE aia a ee aE oh Bey cae 

= 7. The Movements and Reactions of the Isolated Melanophores of the 

Frog, by S. J. Holmes. Pp. 167-174, plate 10. August, 1914.......... 

8. Polychaetous-Annelids of the Pacific Coast in the Collections of the 

Zoological Museum of the University of California, by Aaron L. 
Treadwell. Pp. 175-234, plates 11-12. 

was 9; New Syllidae from San -Francisco Bay (collected by the U. S. 8. 

“4 ** Albatross’’), by Aaron L. Treadwell. Pp. 235-238, 7 text figures. 

‘ Nos. 8 and 9 in one cover. October, 1914. .....2...2.220..-:eee ence 

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Henry B. Bigelow. Pp. 239-241. September, 1914.....-....20- 


Vol. 14. 1. A Report upon the Physical Conditions in San Francisco Bay, Based 
upon the Operations of the United States Fisheries Steamer ‘‘Al- 
pbatross’’ during the Years 1912 and 1913, by F. B. Sumner, G. 
D. Louderback, W. L. Schmitt, E. C. Johnston. Pp. 1-198, plates 
1-13, 20 text figures. FULY,, POEs ch aS SI es 


- \ Vol. 15. 1. Hydrographic Plankton, and Dredging Records of the Scripps In- 
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1901 to 1912, compiled and arranged under the supervision of W. 
E. Ritter by Ellis L. Michael and George F. heap din Pp. 1-206, 
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Vol. 16. 1. An Outline of the Morphology and Life History of Orithidia lepto- 
coridis, sp. nov., by Irene McCulloch. Pp. 1-22, plates 1-4, 1 text 
figure. September, pS a pocee ails Cred i pesngs Ae, caNcaice te yb coap ets baa Sl eae Oe 

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30-54, plates 5-8, 1 figure in text. 

i Nos. 2 and 3 in one cover.~ November, 1915__........0...-2-22---e--- 

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. On a New Trichomonad Flagellate, Trichomitus parvus, from the 
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Nos. 6 and 7 in one cover. December, 1915. ...200.0...eo to eeceeeeteee 


as | 


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15 


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05 


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65 
05 


2.25 


2.25 


+30 
10 


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25 


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: UNIVERSITY OF CALIFORNIA PUBLICATIONS 


aks 200L0GY z Pele US Ay 
Vol. 12,No. 15, pp. 413-495 -  Mareh 20, 1916 
“THE STATUS OF THE BEAVERS OF 
‘ WESTERN NORTH AMERICA, WITH 
As. CONSIDERATION OF THE 
_ FACTORS IN THEIR 
: SPECIATION 
"WALTER P, ae ey 
i eareciein oF canes PRESS on ef onal Meee sc 
i Ge : 


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: ZOOLOGY 
Vol. 12, No. 15, pp. 413-495 March 20, 1916 


THE STATUS OF THE BEAVERS OF WESTERN 
NORTH AMERICA, WITH A CONSIDERATION 
OF THE FACTORS IN THEIR SPECIATION 
WALTER P. TAYLOR 


(Contribution from the Museum of Vertebrate Zoology of the University of California) 


CONTENTS 
PAGE 
SAPPIRTUEVOCMCULOIN petecc: cereete tcc ec ck cosas ace ee see epee aats pet tnenrts eee U es eee actaoe Unseen ston 414 
Pe EVOMONA EY CONSTOGTACLOMS) ccccecceece sc aneee-s arose ce-ceeees eee oc onweane ne ee cueaness 414 
SD etailsmotaenee tim Cnt yleee ese ween. tee teosedcsotoretutenmaamteoete neete ee tle eesueaeeeecee 415 
Baa Materialmand hacknOwded SM EMtS een msec ct en cee cetera ca erases ne san ate acer eed nese eats 416 
COS DN Kop co KF Yh EEN AY B72) 4 ee a Sa a aa nc Re ee 417 
D. Changes due to age in a single species (Castor canadensis leucodonta 
CORTE) Re ee ear ks ee Ae ee 418 
PABBIXGOUTM Al e CHATACLOMS) seen cee oe oe cc nee ssa cere oe cee cease ecceccecceceeeee 418 
Tha WIG SENS ANT SY 00 SY aU OYE Slo 2 ee nee 418 
Dy COPS TRGY ee Aa a oes na Ue oe) Eee a eee ee i 420 
BUTE Gorse ra eR AT ACE CTS ys occ oe a aces ees secon seamen eee cc eae pie Lewpeneueequeceeececluns 422 
lem ememalle Chane Cj eee we ress taces crete scns stn oes caer erence ae Bh sas EE eee se 422 
Pe SEES ST aN A Se ee ee ne ROE oP nee Pe ce ae nae Ua . 422 
(1) Milk dentition and the eruption of the teeth........ . 422 
(2) Some characteristics of the permanent dentition........ 423 
(3) The dental armature as a cutting and grinding 
EPPA ET 6A eee Be ae re Pen ee an ee aero eRe a Ri ean ES 426 
(4) Parallelism in Castor and Erethizon...........................-.- 426 
SoMDLMENSIONS Of ‘CRAM ay oc orc nns pees ses ec cc eec encase ree oes ee ccs ee ecceececesccenscceeseck 427 
Are CEO ON Mss) OS CS acess nec ercc ee na rerescneennneterseat oveuieiannanrnensnaswececcsenunerase 427 
E. Description of a new subspecies of beaver from the Cook Inlet region, 
DINE EUS eet oye oo Renae ee Sees a faa Sod cp ecco) nace reaentsy Meet pesaee ccwerceeecesdeuees 429 
Castor canadensis belugae, new subspecies...............--..--..-----------eeee 429 
F. Description of a new subspecies of beaver from eastern Shasta 
Chey Sistine ACE Te ot a a rR Se Ue te ee eee ae 433 


Castor subauratus shastensis, new subspecies.............0.00------e ee 433 


414 University of California Publications in Zoology  [Vou.12 


G. Comparisons of certain American beavers. ..-.------ ccc cceeeeeeeeeee ee. 437 
I. Castor canadensis phaeus Heller, from Admiralty Island, Alaska 437 
II. Castor canadensis leucodonta Gray, from Vancouver Island, 


British Columbia, 2252.22 2p oe eek ee ne 440 

Ill. Castor canadensis pacificus Rhoads, from the Mainland of Brit- 
ishiColumbiagan di swWashine tore eee ann nee 442 

IV. Castor subauratus subauratus Taylor, from the San Joaquin 
Walley, sCallitorniiay cnc aes a 446 

V. Castor canadensis frondator Mearns, from the Colorado and 
Sanh Pedromiverss. Mexico ses ee eee 454 
He Outlinevor the history cot thesDeavers sesame mee 457 
I. Summary of relationships of certain North American beavers............ 460 
I. Some difficulties to precise statement of relationship... 460 
II. Consideration of certain North American forms... 460 
J. Remarks on isolation and its relation to speciation... 462 
I. Consideration of some of the evidence... eee 462 


2. Evidence from certain other families of mammals... 
(GO "Soricidae:. 2 see ee ee 
(C2) Bro cyomid aley weer ee re 
(3) PiMustelidalcheeserce = ene 
(4) eMuridae: 22 ae eee ee 
(oD Zapodidae ses see ee eee eee 
(6) Aplodontiidae 
CAMO chotonidaee =a 
(8) iCervidaewme = eee ee 
(9) Summarization _.. 


ries 
b. Relation of the evidence to Wagner’s theory of 
migration and geographical isolation.............. 475 


d. Tentative suggestions regarding the manner in 
which geographic isolation acts in the process 


OL, Speciation -~2s. 2-2 a ee 482 

OSL 09101 ya nee a Pa a ec 487 

Ib, Literature: Cited 2.52 secu .c cca ctswctesce tence sete eee ee 490 
INTRODUCTION 


EVOLUTIONARY CONSIDERATIONS 


Problems concerning the causes and conditions of organic evolu- 
tion are numerous and many of them are as yet unsolved. Even a 
cursory examination of zoological literature for a number of years 
reveals the fact that one branch of investigation has held the center 
of the stage for a time, only to be displaced by another and this in 
turn by another. This tendeney toward successive popularity of 
different fields of work is for the most part good, of course, since it 


1916] Taylor: Beavers of Western North America 415 


leads to the discovery of new problems, the evolution of new theories, 
and the coordinated accumulation of new facts. Ill effects, how- 
ever, may sometimes be realized as a result of it. A field of investiga- 
tion the resources of which are by no means exhausted may be for- 
saken by those best fitted to prosecute researches therein on the 
ground that some other field looks more promising. 

The studies of chorology, that is to say, of the geographical dis- 
tribution of living forms, and of the relations of the living organism 
to its natural environment, hold, in the opinion of the writer, posi- 
tions in scientific interest subordinate to those to which their abundant 
and practically unexplored resources would seem appropriately to 
assign them. 

With the swing of the pendulum of scientific interest away from 
these fields, work on what is probably an important condition in 
polytypic evolution, namely, isolation, has practically ceased. The 
large place in organic evolution which may be filled by this condition 
has been emphasized by Wagner (1868), Dixon (1885), Romanes 
(1886), Gulick (1905), and Jordan (1905). 

Questions arise immediately: What is isolation? Has it any 
importance in organic evolution? Is it not all-important? Have 
not the mutation and Mendelian concepts of the last few years done 
away with the necessity for postulating it at all as a condition of 
evolution? If isolation is the fictitious result of speculative induc- 
tion, the sooner the concept is thrown overboard the better. If, on 
the other hand, it is of importance as a factor in the evolution of any 
group of living forms, it ought to receive broader recognition than 
it has heretofore. 

It is the intention to present here some of the facts which seem 
to demand consideration, and which are drawn, not only from the 
study of beavers, but also from the geographical distribution and 
relationships of certain other families of west American (chiefly 
Californian) mammals. It is the hope of the writer that he may be 
able to emphasize: The importance of the study of isolation and 
certain related problems; the pertinence and indispensability of 
evidence from zoogeography. 


DETAILS OF TREATMENT 


In the preparation of this paper the inadequacy of material has 
been sharply felt. Still, it has been possible to get together a greater 


416 Unversity of California Publications in Zoology (Vou. 12 


amount of critical material representative of western beavers than 
has probably ever before been available to any one worker. 

Ridgway’s Color Standards and Color Nomenclature (1912) has 
been used as a guide to color names. 

Overhar, as used in the following pages, refers to the long hairs 
making up the contour pelage, those which would be removed in the 
furrier’s process of plucking. The underfur is the short, soft hair 
which covers the skin closely and which remains in the plucked skin. 

For manner of taking cranial measurements see Taylor, 1911, 
p. 206. Special or exceptional methods of measuring are explained 
in the course of the paper. 


MATERIAL AND ACKNOWLEDGMENTS 


For the loan of material grateful acknowledgment is made to the 
following institutions: The United States National Museum through 
Mr. Richard Rathbun, Assistant Secretary, and Mr. Gerrit S. Miller, 
Jr., Curator, Division of Mammals; the Field Museum of Natural 
History through Mr. Wilfred H. Osgood, Assistant Curator of Mam- 
malogy and Ornithology; and the United States Department of 
Agriculture through Mr. H. W. Henshaw, Chief of the Bureau of 
Biological Survey. 

Considerable material representative of western beavers is con- 
tained in the collection of the Museum of Vertebrate Zoology of the 
University of California. The gathering of this material has been 
largely due to the interest of Miss Annie M. Alexander in the par- 
ticular problem. The specimens from the San Joaquin River, Cali- 
fornia, were obtained directly by her from a local trapper, and those 
from Vancouver Island and southeastern Alaska were collected on 
three expeditions from the Museum made possible through means 
furnished by her. Altogether 86 specimens of beavers, some repre- 
sented by skins and skulls, others by skulls alone or skins alone, and 
one by jaws only, have been available for study. 

The writer is also indebted to the following persons, who have 
very generously given of their time and interest in assisting through 
helpful criticism and suggestion: Professor Charles A. Kofoid, 
Professor Samuel J. Holmes, Professor J. Frank Daniel, Professor 
John C. Merriam, Dr. Harold C. Bryant, Mr. F. H. Holden, and 
especially Dr. Joseph Grinnell. 


1916] Taylor: Beavers of Western North America 417 


NOMENCLATURE 


Three species of beavers, with altogether twelve subspecies, have 
been described from North America. 

The American beaver was separated from the European by Kuhl 
(1820, p. 64) under the name of Castor canadensis, its type locality 
being Hudson Bay. 

The specific name americanus was applied to the American beaver 
by F. Cuvier (1821, not seen) but this name is antedated by Kuhl’s 
canadensis. Although the account on which Kuhl’s name is based 
is fragmentary, it includes a description of the animal. 

Gray (1869, p. 293) separated the beaver of the ‘‘northwest coast 
of America’’ as Castor canadensis leucodonta. This deseription was 
on the basis of specimens collected by Dr. Robert Brown. It is very 
probable that they were obtained on Vancouver Island (Osgood, 
1907, p. 47). 

The beaver of northern Mexico and the southern Rocky Mountain 
region was described by Mearns (1897, p. 502) under the name 
Castor canadensis frondator, its type locality being San Pedro River, 
Sonora, Mexico, near monument no. 98 of the Mexican boundary line. 

A year later two more races were described by Rhoads (1898, 
pp. 420 and 422 respectively): Castor canadensis carolinensis, type 
locality Dan River, near Danbury, Stokes County, North Carolina; 
and Castor canadensis pacificus, type locality Lake Kichelos or 
Kecheelus, Cascade Mountains, Kittitas County, Washington. 

The beaver of Texas was shown to be distinct by Bailey (1905, 
p. 122), and was described as Castor canadensis texensis, type locality 
Cummings Creek, Colorado County, Texas. 

Expeditions from the Museum of Vertebrate Zoology of the 
University of California found beavers on several of the islands of 
southeastern Alaska, although Admiralty Island is the only one 
which is so far represented by specimens. The race found on this 
island was described by Heller (1909, p. 250) as Castor canadensis 
phaeus, type locality Pleasant Bay, Admiralty Island, Alaska. 

The beaver inhabiting the Sacramento and San Joaquin valleys 
of California was recently characterized as a full species, Castor 
subauratus (Taylor, 1912, p. 167), type locality Grayson, Stanislaus 
County, San Joaquin River, California. 

The beaver of Newfoundland, like so many others of the mammals 
inhabiting that island, is apparently restricted to it alone. It was 


418 University of California Publications in Zoology {Vou. 12 


recently described by Bangs (1913, p. 513), under the name of 
Castor caecator. 

Two more subspecies of canadensis, presenting respectively a very 
pale desert coloration, and a dark, rich coloration, have been described 
by Bailey (1913, pp. 191-193). These are Castor canadensis meai- 
canus, type locality Ruidoso Creek, six miles below Ruidoso, New 
Mexico, and Castor canadensis michiganensis, type locality Tahqua- 
menaw River (five miles above falls), Luce County, Michigan. 

The beavers of the west coast recognized in this paper are as 
follows: 

Castor canadensis belugae, new subspecies (see p. 429), Cook 
Inlet region, base of Alaska Peninsula and probably Kenai Penin- 
sula, and southward west of Rocky Mountains to central British 
Columbia. 

Castor canadensis phaeus Heller, Admiralty Island, Alaska; prob- 
ably neighboring islands and mainland. 

Castor canadensis leucodonta Gray, Vancouver Island. 

Castor canadensis pacificus Rhoads, probably mainland of British 
Columbia, Washington and Oregon; precise limits of range unknown. 

Castor canadensis frondator Mearns, Colorado River drainage, 
and probably of broad distribution in southern Great Basin region. 

Castor subauratus subauratus Taylor, Sacramento, Feather, 
American, and San Joaquin rivers, California. 

Castor subauratus shastensis, new subspecies (see p. 433), east of 
Sierra Nevada Mountains, California; drainage of the Pit River. 


CHAance Due To AGE IN A SINGLE Species (Castor canadensis leuco- 
donta Gray) 


Beavers secured on Vancouver Island by the expedition of the 
Museum of Vertebrate Zoology in 1910 represent three generations, 
and so make possible an outline of the changes in certain character- 
istics due to age. 


EXTERNAL CHARACTERS 
MEASUREMENTS 
(See table, p. 419) 
Difficulties are immediately apparent when one attempts to set 
down laws of change of form with age, the most important of which 
are that (a) weights and measurements of the youngest beavers are 


419 


ved 


Taylor: Beavers of Western North Amerw 


1916] 


I. EXTERNAL MEASUREMENTS* OF Castor canadensis leucodonta FROM VANCOUVER ISLAND, 


BRITISH COLUMBIA 


Specimens arranged approximately in order of ag 


(All measurements in millimeters) 


oO 

ee: 
= 2 =) 
Museum B ie Ae 
number Sex LOCALITY i a q 
12109 ? Hall’s Ranch, Alberni Valley 9 -------- 00 e000 se 
12110 XS Hall’s Ranch, Alberni Valley ------- 00-2000 => 
12106 3 Hall’s Ranch, Alberni Valley 9 ------- 00 e200 = 
12105 fe) Hall’s Ranch, Alberni Valley 503 162 93 
12104 fe) Hall’s Ranch, Alberni Valley, 900 280 165 
12108 Q Hall’s Ranch, Alberni Valley 967 400 170 
12102 a Hall’s Ranch, Alberni Valley 932 350 173 
12103 fe) Hall’s Ranch, Alberni Valley 925 350 170 
12101 é Hall’s Ranch, Alberni Valley 1000 340 175 
12111 a Great Central Lake 990 395 175 
12107 fe) Hall’s Ranch, Alberni Valley 1157 450 200 


*For method of taking measurements see Taylor, 1911, pp. 206, 207. 
yEar from crown. 


e from top of table to bottom 


Weight in pounds 


Length sealed portion 
of tail (dry skin) 


ive} 
_ 
a> 


110.6 
116.0 
124.2 
213 
260 
225 
225 
237 
243 
270 


Width scaled portion 
of tail (dry skin) 


wooo & RW 
arworoan 
aw 


wo oO 
me Oo 


120.9 
108.5 
124.2 


Number of scale-rows in 


broadest part of tail 


oo 
eae 


wow ce ww ow ow wo 
OrRRrE TT Fe 


Ratio tail vertebrae 


to total length 


Ratio width scaled portion 


of tail to length 


420 University of California Publications in Zoology (Vor. 12 


lacking; (b) dimensions of the scaled portions of the tails were not 
usually taken in the field, and absolutely accurate measurement is 
impossible in dry skins on account of their shrinking and crinkling; 
(c) the irregularity of their arrangement makes it difficult to avoid 
error in counting the scale-rows transversely on the tail. 

Keeping these possibly modifying factors in mind, it is believed, 
however, that certain general propositions may be formulated and 
regarded as fairly dependable: 

(1) Weight and dimensions inerease with age. Growth con- 
tinues as in certain other mammals (for example, the gopher, 
Thomomys) practically through life. 

(2) The number of scale-rows on the tail is apparently the same 
in adults and in juvenals, the increase in size taking place through an 
augmentation in measurements of the individual scales. 

(3) The ratio of the length of the tail vertebrae to total length 
apparently increases with age (no. 12108 constitutes an apparent 
exception to this statement). 

(4) There is evident a tendeney for the tail to increase in width 
somewhat more rapidly than in length. 

There is great individual variation in the ratio of the width of 
tail to length. According to the table, the maximum of this ratio is 
51.0 percent, minimum 31.8, indicating a variation of 19.2 percent. 
The average of all the ratios is 41.3 percent. Although the animals 
presenting the highest ratios are adults, there is no very clear cor- 
relation between age and different proportional dimensions of tail. 

Two specimens (nos. 71830, 71833, loaned by the Biological Sur- 
vey), being younger than the youngest listed in the table, have ratios 
of 44 and 47 respectively. 


CoOLORATION AND PELAGE 


On the whole, the coloration of the juvenals is very much like 
that of the adults. There is a change toward a deepening in general 
coloration, and a slight differentiation of color areas with increased 
age. In the adults there tends to be a dark area (one obtains a 
general impression of seal brown or a little paler) in the middle of 
the back, with a lightening of coloration (varying from near hazel 
or cinnamon-buff to chestnut) on the sides of the face, the top of 
the head, the nape of the neck, the shoulders and the rump. The 
pelage of the young is softer and fluffier than that of the adults. 


1916 | Taylor: Beavers of Western North America 421 


No differences in relative amounts of underfur and overhair can be 
clearly correlated with age. No individual out of four juvenals at 
hand (nos. 12109, 12110, 12106, and 12105, taken June 20 to 27) 
has the overhair so worn as it is in certain adults (as nos. 12108 and 
12107, taken June 25). 

The coloration of the underfur changes little with age, varying 
above, in both old and young, from fuscous and fuscous-black to 
benzo and hair brown, and varying beneath about light drab and 
hght cinnamon-drab. One adult specimen (no. 12101) has the under- 
fur above an almost uniform drab. 

Two very young juvenals, loaned by the Biological Survey (nos. 
71830, 71833) are very similar to the juvenals mentioned above. 
Being younger, the hair of nos. 71830 and 71833 is noticeably shorter, 
softer, and fluffer. The only difference in coloration is a slightly 
darker general effect dorsally. 

Dorsal coloration.—There is in the older animals a distinct dark- 
ening in dorsal appearance, the color varying from cinnamon to chest- 
nut. This darkening is partly the result of darker coloration of 
individual hairs, and partly the result of the showing through to a 
greater degree of the dark underfur. The juvenals are pinkish 
cinnamon to cinnamon, sometimes a little darker dorsally. 

The forefeet are near warm sepia or mars brown in the young, 
while in the older ones they have a deeper shade and exhibit some- 
thing of a luster. 

The hind feet of the young are near mars brown, although it is 
very difficult to fix the tone, while those of the adults are browner, 
varying from near carob brown to near hazel. 

Ventral coloration—The juvenals have more of a golden luster 
ventrally along the sides of the belly than the adults. The coloration 
mid-ventrally varies about drab in the juvenals, with a tendency to 
be darker in the adults. The area just anterior of the tail ventrally 
varies in the adult between cinnamon-brown and chestnut or bay, 
while in the juvenals it varies between walnut brown and cinnamon- 
buff. 

Molt and range of individual variation—Adequate material for 
the study of the molting process in beavers is lacking. It seems 
probable that the molt is not regional, as it is in chipmunks and 
gophers, but general, the hair being renewed gradually all over the 
body. 


422 University of California Publications in Zoology (Vou. 12 


In some species there seems to be decided individual variation in 
coloration of pelage. Of the specimens from eastern Canada at 
hand, one (no. 4358, U. S. Nat. Mus., May 21) is in light pelage, 
while two (nos. 174525, 174526, U. S. Nat. Mus., Sept. 25) are in dark 
pelage. Among skins of Castor canadensis phaeus from Admiralty 
Island, one (no. 209, Mus. Vert. Zool., May 16) is very dark, while 
another (no. 210, Mus. Vert. Zool., June 1) is paler. Details of these 
differences appear in the tabulations of coloration in the following 
pages. 

Tal—The variation in measurements and proportions of tails is 
recorded in table I, p. 419. 

Scattered hairs, which grow from between the scales, appear 
in the tails of the juvenals, but are generally lacking in those of the 
adults. 


CRANIAL CHARACTERS 
(See table, opp. p. 426, and fig. B, p. 424) 


GENERAL CHANGE 


The skull is rounded in young animals, with frontals elevated, 
and interparietal region sloping. In adults it is more flattened, with 
frontals not elevated and interparietal region not sloping so much. 

As growth continues, the comparatively undifferentiated skull of 
the juvenal becomes adapted to the increased strains put upon it, the 
sutures tend to disappear, the bones harden, processes and ridges 
develop greatly, and there is an increase in size. 

Every bone changes somewhat in outline as the animal grows older, 
the most evident modifications being (a) loss by the frontals of their 
jardinier or vase-shape in outline as viewed dorsally, and their assump- 
tion of a fleur-de-lis shape, due to encroachment of temporal ridges 
anteriorly ; (b) narrowing and antero-posterior extension of the inter- 
parietal, giving it, in outline as viewed dorsally, an Indian-club 
rather than a subrectangular shape; (c) change in outline of the 
palatine as viewed ventrally, so that instead of being nearly an equi- 
lateral triangle it is isosceles; (d) widening of the foramen magnum 
proportionally to its height. 


TEETH 
MiLtk DENTITION AND THE ERUPTION OF THE TEETH 
The dental formula of the beaver is I1, C$, P1, M3x 2=— 20. 


1916] Taylor: Beavers of Western North America 423 


Milk premolars are brachydont, usually with three 


: ee well-developed roots, though in one specimen at 
\ / hand there are two roots only. Tooth eruption is 


as follows: milk premolar 4; molar 1; molar 2; 
Fig. A. Occlusal 


surface of P4, to 2 c 
show method of appearance of the teeth is the same on both jaws, 


molar 3; permanent premolar 4. The order of 


taking measure- the corresponding upper and lower teeth appear- 
ments. Approx- 


imately natural 
size. 


ing simultaneously. 


Il MEASUREMENTS* OF TEETH OF Castor canadensis leucodonta 
GRAY, FROM VANCOUVER ISLAND, BRITISH COLUMBIA 


(All measurements in millimeters) 


os 
Transversey Basilar 


Museum T 9 ’ | length 
number Sex p4 M1 M? Ms Le? 4 My Ms M 3 of crania 


12104 3 5.7 5.8 Bear 5.2 4.8 5.6 5.8 5.2 99.6 
12108 2 4.7 6.2 5.8 5.4 4.8 6.2 PA 5¥3) 100.6 
12103 Q a2 bet 5.3 5.2 5.0 5.9 5.8 ue) 103.9 
12102 é iy 6.0 5.8 5.4 5.2 6.2 6.4 5.7 105.8 
12111 é eth 7.4 6.7 6.0 6.7 7.7 7.4 a4 110.9 
12101 é 8.4 Tat 6.9 6.1 7.0 13) 7.0 6.2 111.9 
12107 fe) atl 7.4 6.5 6.2 6.8 7.6 7.0 6.2 122.6 

Longitudinal; Basilar 
Moot jc «PS Mi M2 M® PS Mp Mz, Mj csuatia 
12104 é 5.5 6.1 5.6 See 6.1 7.0 tee 6.8 99.6 
12108 fe} Dali 5.9 5.5. 5.3 6.1 7.4 eal 6.8 100.6 
12103 Q 5.4 6.0 5.4 beg 6.1 6.7 6.7 6.8 103.9 
12102 é 5.6 6.0 yard bea 6.6 6.9 7.0 7.2 105.8 
12111 é 7.4 6.4 6.3 6.1 8.9 7.6 7.9 eb, 110.9 
12101 as 7.6 6.5 6.7 6.5 9.0 8.0 8.1 13 111.9 
12107 fe) 8.2 6.9 6.2 5.9 9.0 eb) eo) 7.6 122.6 


*Bach measurement is taken three times and the results averaged to 
give the measurement here entered. 


7See figure A. 
SomME CHARACTERISTICS OF THE PERMANENT DENTITION 


The permanent dental armature exemplifies a high degree of 
hypsodonty. The oldest crania available to the writer have the pulp- 
cavities of the cheek-teeth almost completely closed. In order to 
ascertain whether there is a pronounced change in size of teeth with 
age, all the cheek-teeth on the left-hand side, above and below, of seven 
available skulls from Vancouver Island, were measured. The skulls 
themselves belonged to animals of different ages. In nos. 12104, 


University of California Publications in Zoology  [Vor.12 


424 


-Ixoidde soins 


“BIQUINIOD YSig ‘pues, JaAnooue, Tueq Ty ‘6 ‘LOTZT ‘ou :19AB9q INP PIO 
“BICUIN[OD YS ‘pur[sy Jaanoour, ‘TuIeqry ‘2 ‘ZOLZT ‘ou ‘[BUIUe IaplO 
“BICUIN[OD YS ‘purjs] JaAnoour, ‘TUIEqTV ‘P ‘90TZT ‘ou ‘[eullue Sunox 

‘OZIS [BIN}VU SpIIY}-OM} ATO] eur 


sos 


‘DIUOPOINA] SisuapoUDI LO]SDO Ut eSB YIM Sutranoo0 Sasueyo ay} JO aUIOS MoYsS 0} SSUIMBIP OUITINO “gq “SIq 


2 D 


SEZ? 


1916] Taylor: Beavers of Western North America 425 


12108, and 12103 the permanent premolar has hardly become fune- 
tional. No. 12107 is the skull of a comparatively old adult, as shown 
by its dimensions and degree of development. To reduce the margin 
of error, each dimension was carefully taken three times, and the 
results averaged, in case they did not exactly agree (see table II). 

It is apparent, from the data of the table of measurements, that 
all the cheek-teeth first increase in size with age, then undergo a 
slight absolute or proportional decrease. It should be noted that the 
initial increase is of greater magnitude than is the later decrease. 
The tooth pierces the gum, increases in size up to a certain point, 
then decreases slightly. Since the oldest skulls at hand, without 
exception, have the longest maxillary tooth-rows, the decrease in 
tooth dimension late in life is not sufficient to affect the validity of the 
‘Vength of the maxillary tooth-row’’ as a comparative measurement. 

The table of measurements indicates that there are no important 
differences in the relation of the antero-posterior to the transverse 
diameter of the teeth in crania of different ages. One notes that the 
relation of the longitudinal or antero-posterior diameter of the teeth 
to the transverse is more variable in the superior teeth than in the 
inferior, in which latter the antero-posterior diameter exceeds the 
transverse in nearly all eases. 

A number of possibly significant conclusions are derivable from 
the tables of measurements of the teeth of beavers of different species 
(tables II and IV): 

(1) All the superior molars measured have the transverse diam- 
eter equal to or exceeding the antero-posterior, except in the following 
cases: in no. 12104, Vancouver Island (M’, M*, M*) ; no. 12103, Van- 
couver Island (M?, M2) ; no. 12111, Vancouver Island (M®*) ; no. 12101, 
Vancouver Island (M*) ; no. 3672, Skagit River, Washington (M*) ; 
in nos. 209 and 210, Admiralty Island (M?’, M*). 

(2) All the superior premolars measured have the antero-pos- 
terior diameters equal to or exceeding the transverse, except in the 
following cases: nos. 12101, 12104, 12111, Vancouver Island; nos. 
174525, 174526, New Brunswick. 

(3) Inferior molars in lewcodonta have the antero-posterior diam- 
eter greater than the transverse (except M, in nos. 12111 and 12107, 
Vancouver Island). 

(4) All the inferior premolars measured have the antero-posterior 
diameter greater than the transverse. 


426 University of California Publications in Zoology (Vou. 12 
THE DENTAL ARMATURE AS A CUTTING AND GRINDING AGENCY 


The work of the beaver, as exemplified in the cutting of materials 
for lodges and dams, serves to illustrate the efficiency of the incisors 
as cutting agents. The beaver’s incisor is said to have been the 
hardest substance except flint known and used as a cutting tool by 
certain tribes of North American Indians. That the grinding por- 
tions of the beaver’s dental armature are no less efficient to perform 
the function required of them is evident when it is considered that 
bark, a substance requiring very powerful mastication, is the beaver’s 
principal food. Counting from front to back there are about 40 
transverse cutting blades on each maxillary tooth-row, making 80 
cutting blades for the upper teeth. A similar number obtains for 
the lower teeth. Only one side at a time can be opposed in the process 
of mastication, so that 40 blades above are brought against 40 blades 
below in the course of one chewing movement. If there is enough 
lateral motion during this movement, however, all 80 of the blades 
of the upper teeth may be ground against the 80 blades of the lower 
teeth. In the former case 80 cuts, in the latter 160, would be given 
to the mouthful of material. In case the beaver makes 100 chewing 
movements a minute, the number of cuts for that period would be in 
the former case 8000, in the latter 16,000. 

Unfortunately, the writer has never been privileged to ascertain 
from watching the animal in life what the characteristic jaw move- 
ments are, nor have references to the matter been found in the litera- 
ture examined. On the basis of the arrangement of the series of 
teeth with reference to one another it may be concluded, however, 
that there is an antero-posterior movement of probably 15 to 20 
millimeters magnitude. That there is lateral motion is equally cer- 
tain, although it must be much less than the antero-posterior. Hight 
millimeters appears to be about the maximum sidewise movement 
possible. 


PARALLELISM IN Castor AND Erethizon 


The general resemblance of the enamel pattern of the cheek-teeth 
obtaining between the genera Castor and Erethizon has been remarked 
by former workers. Another character, of interest in this connec- 
tion, is the condition of the palato-maxillary region, which is rounded 
instead of plane in both genera. Authorities on classification agree 
in referring the beavers to the sciuromorph section of the Rodentia, 


1916] Taylor: Beavers of Western North America 42 


while the porcupine belongs to the section Hystricomorpha. If the 


above-mentioned resemblances are not due to inheritance from a 
common ancestor, and the bulk of evidence 


conc b ann EI a, would seem to indicate that they are not, they 
| Uy | | \ Se illnatratea «a natawarthyw aaca af narallal da- 
IA 


§ 

E 

- s 

= 5 

cI s 

é 2 = S a 

E 3 2 - E Sa 

B 5 5 = : ar 

i £ E 3 3 Bs 

ee ow ae, ay) sg 

[a7 as) © to = a a 

Be) eee Seuss 

B ry ; 2 SS Ss 

RE? 34 58 88 2 4&3, 

a 3 ra R oe zi 

a ge ge) oe | 84 38 26 
number Sex LOC4: 9 me ae poe BE Be 

12109 2 Hallie Ranehy, sen ieee) | ob. guest | ane 

12110 2 Halles Ranch, ee imei del! A Woh ee mS At 
12106 é$ Hall’s Ranch, 36.7 23.2 57.8 20.7 17.6 
12105 rc) Hall’s Ranch, | __ 37.6 24.3 57.8 2255 anTe7 
12104 2 Hall’s Ranch, }6.8 40.4 19.5 57.6 DIESe TAU 
12108 Q Hall’s Ranch,?78 38.2 225 580 213 144 
12102 3 Hall’s Ranch, 36.2 36.1 PAULEY GIR ONG IGLOS 
12103 Q Hall’s Ranch,}5.9 37.3 21.7 56.9 20 eto e7 
12101 3 Hall’s Ranch,}5.5 314 243 56.6 20.8 12.9 
12111 $ Great Central}5.6 39.1 22.0 56.6 20.9 i1S}0) 
12107 Q Hall’s Ranch, }5.4 31.9 PAS A OES TS) 


~ *Ali measurements except as be 
Width of nasals: across both n 
Length of frontals: along interl 
Length of interparietal: along 1 
Length of nasals: measured, noOharrow tongue e i ; 
posterior outline of bone. 7 ere ds boskerterly cf msty 


LEHOS LO WILIIM ov iin. UL a@ iiiic 
drawn at right angles to the 
antero-posterior axis of the skull 


Fig. D. Lateral view of mandible, to at the narrowest part of the 
show method of taking  interorbital constriction. 


measurements. About one- F 
half natural size. Bailey (1905, p. 122) refers 


Museum 
number 


12109 
12110 
12106 
12105 
12104 
12108 
12102 
12108 
12101 
12111 
12107 


Sex 


+0 O3 Os 40 O3 40 40 410 DO +d + 


_ *All measurements except as below specified taken in accordance with methods given by Taylor, 1911, pp. 206, 207. 


Hall’s 
Hall’s 
Hall’s 
Hall’s 
Hall’s 
Hall’s 
Hall’s 
Hall’s 
Hall’s 
Great 
Hall’s 


LOCALITY 
Ranch, Alberni 
Ranch, Alberni 
Ranch, Alberni 
Ranch, Alberni 
Ranch, Alberni 
Ranch, Alberni 
Ranch, Alberni 
Ranch, Alberni 
Ranch, Alberni 
Central Lake...... 
Ranch, Alberni 


III. CRANIAL MEASUREMENTS* OF Castor canadensis leucodonta FROM VANCOUVER ISLAND, BRITISH COLUMBIA 


Specimens arranged approximately in order of age from top of table to bottom 
(All measurements in millimeters) 


Valley 
Valley 
Valley 
Valley 
Valley 
Valley 
Valley 


Basilar length of Hensel 


© Zygomatie width 


oO 


=> Mastoid width 


= 
=~ Interorbital constriction 
> 


17.8 
17.7 
18.0 
22.3 
24.8 
22.5 
22.5 
23.9 
24.4 
25.7 


Length of nasals 
(See fig. C) 


bo 


tn 
tS 
Cars 


Width of nasals 


Maxillary tooth-row 


Length of frontals 


22.7 
24.1 
24.1 
24.9 
40.2 
38.4 
38.2 
38.8 
35.1 
43.3 
39.1 


Length of interparietals 


angle 


Oo. to anterior surface of alveolus of 
(See fig. D) 


cn Greatest length of mandible: 


incisor. 


> & mandible to coronoid 


$3 SS Ventral surface of 


Most dorsal point on outline of 
foramen magnum to occipital crest 
(See fig. G) 


on median line. 


ss 
o 


Vertical diameter of foramen 
(See fig. E) 


magnum. 


Width of nasals: across both nasals at right angles to the antero-posterior axis of skull, so as to include the most lateral points on their outline. 


Length of frontals: along interlying suture, or where this is obscure, along median line of skull . 


Length of interparietal: along median line of skull. i < 
poueth of nasals: measured, not strictly along interlying suture, but so as to include the most anterior and most posterior points, except where an exceedingly narrow tongue extends posteriorly of main 
posterior outline of bone. 


Transverse width of 
foramen magnum 


Ratio maxillary tooth-row 


to basilar length 


Ratio length of frontals 


to basilar length 


Ratio length of interparietals 


to basilar length 


Ratio mastoid width to 
basilar length 


Ratio width of nasals to 


basilar length 


Ratio vertical diameter of foramen 


magnum to basilar length 


426 University of California Publica 


THE DENTAL ARMATURE AS A CUTTING A 


The work of the beaver, as exemplified i 
for lodges and dams, serves to illustrate th 
as cutting agents. The beaver’s incisor 


& to 0h ‘e 
ni 16. dee 
+ hra4 ( %o Giarnt” 
: ; , onne :eleten to Syme: 
: : lo +clisie 10braye 


en renee op 


obtaining between the genera Castor and Ere 
by former workers. Another character, of 
tion, is the condition of the palato-maxillary 
instead of plane in both genera. Authoriti 
in referring the beavers to the sciuromorph 


1916] Taylor: Beavers of Western North America 427 


while the porcupine belongs to the section Hystricomorpha. If the 
above-mentioned resemblances are not due to inheritance from a 
common ancestor, and the bulk of evidence 
would seem to indicate that they are not, they 
illustrate a noteworthy case of parallel de- 
velopment in these widely different sections 
of the Rodentia. It is doubtless something 
more than coincidence in this connection that 


the beaver and the poreupine are bark-feed- 


Fig. C. Rostrum, dor- ers. The rounded instead of plane conforma- 
sal aspect, to show 
method of taking 
measurement“Length similar complication of the enamel pattern of 
of nasals”’. About . 
one-half natural size, the cheek-teeth, appear to be direct adapta- 


tions: the first to the stripping of bark from 


tion of the palato-maxillary region, and the 


twigs and branches; the second to its effective mastication. 


DIMENSIONS OF CRANTA 


From the table of cranial characters (opp. p. 426) it appears that 
with age: (a) nearly every bone increases in size, each outside skull 
dimension becoming greater; (b) the ratio of the length of the inter- 
parietal to basilar length decreases (though no. 12101 departs rather 
widely from this rule); (c) the mastoid width increases at prac- 
tically the same rate as does the basilar length, although giving evi- 
dence of a slight tendency to increase less rapidly. 


TEMPORAL RIDGES 


The degree of approximation of the temporal ridges is in general 
indicative of age, although the rate of approximation may vary in 
different forms. In leucodonta the temporal ridges first come together 
posteriorly, then continuously in 
an anterior direction. In the 
oldest skulls the ridges form a 
sagittal crest,narrow posteriorly, 
broader anteriorly, which ex- 
tends to within 25 mm. of a line 
drawn at right angles to the 
antero-posterior axis of the skull 


Fig. D. Lateral view of mandible, to at the narrowest part of the 
show method of taking 


measurements. About one- - 
half natural size. Bailey (1905, p. 122) refers 


interorbital constriction. 


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1916] Taylor: Beavers of Western North America 429 


to the lyrate condition of temporal ridges as a character of Castor 
canadensis texensis. While it is probable that the character is of sub- 
specific value in the Texan form of beaver, 

as the fact that the material on which the de- 
scription was based was exceedingly limited 

in amount emphasizes the alternative possi- 


bility that it is a character due to age only. 

Fig. E. Outline of fora- 

men magnum, to 

show method of tak- fact that in the following forms the char- 
ing measurements. Agee Onc Fo ; 

acter is one the condition of which appar- 

About one-half nat- PP 

ural size. ently depends solely on age: Castor fiber, 

Castor canadensis phaeus, Castor canadensis 

belugae, Castor canadensis canadensis, Castor canadensis leucodonta, 


and Castor canadensis frondator. 


Significant in this connection is the further 


DESCRIPTION OF A NEw SUBSPECIES OF BEAVER FROM THE CooK INLET 
Reaion, ALASKA 


In the labor of allocation of specimens and determination of their 
status it soon became apparent that the skulls from Cook Inlet were 
somewhat different from those of any other race of beaver. HExamina- 
tion of additional comparable material confirms the differences first 
observed. 


Castor canadensis belugae, new subspecies 


Type—Skull only, ¢ youngish adult, no. 4224, Mus. Vert. Zool. ; 
Beluga River, Cook Inlet region, Alaska; ‘‘1907’’; collected by Jacob 
Seminoff ; orig. no. 2. 

Diagnostic characters—Perhaps nearest Castor canadensis leuco- 
donta Gray, but crania immediately distinguishable through the nar- 
rower blades of the hamular processes of the pterygoids in Castor 
canadensis belugae; bony ridge laterally on rostrum less strongly 
developed in belugae than in leucodonta; belugae with tendeney for 
maxillary tooth-row, and ratio of maxillary tooth-row to basilar 
leneth, to be greater. 

Belugae is similar to Castor canadensis canadensis Kuhl, but with 
nasals of different outline, the lateral swelling being more posteriorly 
placed; maxillary tooth-row and ratio of maxillary tooth-row to 
basilar length tending to be greater. 

External characters—Only one skin of the new form (no. 4347, 


430 Unversity of California Publications in Zoology Vou. 12 


Mus. Vert. Zool.) is at hand. For comparison of this with Castor 
canadensis leucodonta see below. A comparison with Castor cana- 
densis canadensis is given in table opposite p. 432. 

Range.—lt is impossible at this time to define precisely the limits 
of range of Castor canadensis belugae. Specimens from the follow- 

ing localities have been examined: Beluga 

SAD River (tributary to Cook Inlet from the 

north) ; Nenilehuk (sometimes spelled Ninil- 

Hach Odiiaeon ae chik [ Baker, 1906, p. 463], a village on east- 

ular process, show- ern shore of Cook Inlet, south of the mouth 

ing method of taking of the Kasilof River); Kasiliff (probably 
measurement. About et : é 

one-half natural size, Kasilof, a fishing village at the mouth of 

Kasilof River, Cook Inlet, according to 

Baker, 1906, p. 353); Snug Harbor, Alaska Peninsula (probably 

Snug Harbor on the western shore of Cook Inlet, near [liamna Peak 

[Baker, 1906, p. 586]); and the general vicinity of Stuart Lake, 

British Columbia. 

It is probable that the form occupies territory on the mainland 
from central British Columbia on the south to the Alaskan Moua- 
tains on the north. The main chain of the Rocky Mountains doubt- 
less bounds its range on the east, and the ocean, or possibly certain 
coast mountains on the west. 

This region is much interrupted topographically and it is not 
unlikely that adequate material would show considerable local differ- 
entiation, possibly the presence of well-marked subspecies, within its 
boundaries. Militating against this suggestion, however, is the 
observed similarity of specimens from such widely separated points 
as the Cook Inlet region, Alaska, and Stuart Lake, British Columbia. 


REMARKS 
ComMPARISON WitH Castor canadensis leucodonta GRaAy 

The new subspecies, belugae, is perhaps nearest leucodonta, of 
which examples from Vancouver Island are at hand, although it is 
intermediate between that form and canadensis from eastern Canada. 
Crania can be distinguished from leuwcodonta by the narrower blades 
of the hamular processes of the pterygoids in belugae (see fig. F, 
above) ; by the ridge laterally on the rostrum less strongly developed 
than in leucodonta; by the tendency in belugae for maxillary tooth- 
row to be longer, shown also in ratio of maxillary tooth-row to basilar 


length. 


1916] Taylor: Beavers of Western North America 431 


Fy QDs Fig. G. Outline of posterior portion of cranium 


e. Castor canadensis canadensis; inner line, no. 174526 9, U. S. Nat. 
Mus., Nepisiquit River, New Brunswick; outer line, no. 174525 ¢, 
U. S. Nat. Mus., same locality. 


V. CRANIAL MEASUREMENTS* OF Castor canadensis belugae FROM COOK INLET REGION, ALASKA, AND BRITISH 
Specimens arranged approximately in order of age from top of table to bottom 


(All measurements in millimeters) 


& 

°o 

4 

a 

2 

5 

Museum me 

num! Sex LOCALITY fea] 
4233 ? Si) CEREUS) as foe eran ek 100.1 
4226 é Beluga River, Cook Inlet region, Alaska......................--+ 106.5 
4229 é Nenilchuk, Cook Inlet region, Alaska..........0...22.....-21:0e0--0- 106.9 
4223 fe) Beluga River, Cook Inlet region, Alaska................0.........-. 108.5 
4230 é Nenilchuk, Cook Inlet region, Alaska....................2::::::0ce-eec00 111.4 
4221 é Beluga River, Cook Inlet region, Alaska 112.3 
4222 Q Beluga River, Cook Inlet region, Alaska... 112.4 
4227 é Beluga River, Cook Inlet region, Alaska... ESS 
4224 $ Beluga River, Cook Inlet region, Alaska... 117.2 
4219 $ Beluga River, Cook Inlet region, Alaska 116.0 
4232 ? masilitt cCook Inlet Tegion, AlaSla. .i..1..cc-c0ccencceccsnsoccenone 112.6 
4228 $ Beluga River, Cook Inlet region, Alaska 116.2 
4231 ? wis TERED oa ae 110.9 
4225 $ Beluga River, Cook Inlet region, Alaska... 122.5 
4220 3 Beluga River, Cook Inlet region, Alaska... 118.0 
4347 3 Snug Harbor, Alaska Peninsula, Alaska... am 109.9 
77159 ) 20 mi. E Stuart Lake, British Columbia.........---.00-------- 111.4 
T7147 $ 12 mi. N Stuart Lake, British Columbia... 116.2 
77157 P 17 mi. N Stuart Lake, British Columbia... 121.7 
T7158 3 30 mi. E Stuart Lake, British Columbia... 116.0 
17155 3 15 mi. N Stuart Lake British Columbia...... ie 119.4 
77150 9 20 mi. NE Stuart Lake, British Columbia 119.2 


*For manner of taking measurements see Taylor, 1911, pp. 206, 207, an 


Zygomatic width 


Mastoid width 


nterorbital constriction 


20.9 
23.2 
23.2 
24.0 
23.0 
23.6 
24,1 
24.3 
25.7 
26.5 
23.6 
24.7 
23.5 
24.7 
26.6 
20.9 
23.9 
24.7 
22.9 
22.7 
25.8 
22.7 


Length of nasals 


& tS o 
aes SEE fig. C) 


40.6 
43.9 
43.6 
44.6 
49.5 
47.1 
47.0 
44.0 
46.6 
45.8 
50.1 
47.9 
44.6 
42.8 
45.3 
47.0 
46.9 
46.2 
49.2 


|e 
‘© Width of nasals 
bo 


20.7 
19.8 
20.7 
22.8 
21.1 
22.2 
22.7 
21.8 
22.4 
20.7 
21.5 
22.0 
26.1 
24.1 
21.9 
23.7 
25.5 
23.5 
24.8 
26.3 
23.9 


Maxillary tooth-row 


28.6 
28.8 
29.1 
29.1 
29.9 
29.9 
28.7 
30.9 
31.4 
31.4 
29.7 
30.8 
30.1 
33.4 
30.7 
29.6 
29.6 
31.0 
29.9 
31.8 
32.8 
29.7 


pital crest 


point on outline of 
(See fig. G) 


oramen magnum to occi 
on median line, 


Most dorsal 


f 


(See fig. E) 


Vertical diameter of foramen 


magnum. 


13.3 


Transverse diameter of 
= foramen magnum 


COLUMBIA, CANADA 


Ratio width of nasals 
to basilar length 


Hee 
~ 2 © 
ar oOo 


19.1 
20.5 
18.8 
19.7 
18.9 
18.6 
19.3 
18.4 
18.3 
19.8 
21.3 
20.4 
19.9 
21.3 
21.9 
19.3 
21.4 
22.0 
20.0 


atio maxillary tooth-row 


basilar length 


me 


Ratio vertical diameter of foramen 


magnum to basilar length 


13.3 


430 University of California Publications in 


Mus. Vert. Zool.) is at hand. For comparison o 


canadensis leucodonta see below. A comparison 
densis canadensis is eiven in table onvosite p. 432. 


row to be longer, shown also in ratio of maxillary t 


length. 


1916] Taylor: Beavers of Western North America 431 


Fig. G. Outline of posterior portion of cranium 
of beaver, to illustrate method of 
measuring dorsal outline foramen 
magnum to inion, or most dorsal point 
on outline of foramen magnum to 
occipital crest on median line. About 
one-half natural size. 


Belugae, on the basis of the table of measurements, has longer 
antero-posterior diameter of P* than in lewcodonta, although speci- 
mens nos. 4347 from Cook Inlet and 12101 from Vancouver Island 
are nearly the same in this respect (see table IV, p. 428). Belugae 
generally has broader teeth transversely than in leucodonta (P* in 
specimen no. 12101, from Vancouver Island exceeds P* in no. 4347, 


from Cook Inlet). 
d eS 


\ 


Fig. H. Outline drawings of tails of different races of western beavers. 

Approximately one-sixth natural size. 

a. Castor subauratus subauratus, 2, no. 12654, Mus. Vert. Zool.; Gray- 
son, San Joaquin River, Stanislaus County, California. 

b. Castor canadensis leucodonta, 9, no. 12107, Mus. Vert. Zool.; 
Alberni, Vancouver Island, British Columbia. 

ec. Castor canadensis phaeus; solid line, no. 210, Mus. Vert. Zool., 9, 
Hasselborg Lake, Admiralty Island, Alaska; dotted line, no. 209, 
Mus. Vert. Zool., ¢, Pleasant Bay, Admiralty Island, Alaska. 

d. Castor canadensis frondator, 9, no. 20751, U. S. Nat. Mus.; San 
Pedro River, Sonora, Mexico. 

e. Castor canadensis canadensis; inner line, no. 174526 9, U. S. Nat. 
Mus., Nepisiquit River, New Brunswick; outer line, no. 174525 ¢, 
U. S. Nat. Mus., same locality. 


432 University of California Publications in Zoology (Vou. 12 


Comparable skins of the two subspecies are distinguishable as 
follows: the single skin of belugae with both overhair and underfur 
thicker than in leucodonta; belugae paler in coloration than average 
of leucodonta, particularly about base of tail, which is near cinna- 
mon-buff in belugae, varying from near hair brown to a shade 
between chocolate and bay in leucodonta; tails are similar in gen- 
eral outline. 


CoMPARISON WitH Castor canadensis canadensis KuHL 


Skulls of belugae comparable with canadensis as regards age 
have outline of nasals different; maxillary tooth-row longer in com- 
parable crania; ratio of the maxillary tooth-row to basilar length 
averaging greater. Belugac, according to the table of measurements, 
has teeth tending to be broader and longer than in canadensis. A 
detailed comparison of external characters of the Cook Inlet race and 
canadensis will be found in table VII, opposite. 

The following table illustrates differences in tail outline: 


VI. MEASUREMENTS AND RATIOS OF SCALED PoRTIONS OF TAILS 


All measurements in millimeters, and taken from dry skins 


Ratio 

Museum width to 

Subspecies— number Length Width length 
Castor canadensis belugae...... 4347 245 115 47.0 
Castor canadensis canadensis.. 4358 223 92 41.1 
Castor canadensis canadensis... 174525 265 116 43.4 
Castor canadensis canadensis.. 174526 260 108 41.3 


The tails of 4847 and 4358 were considerably crinkled and dried 
hard, so that their measurements are less dependable than those of 
the others. 


CoMPARISON witH Castor canadensis phaeus HELLER 


Separated from Castor canadensis phaeus Heller, inhabiting 
Admiralty and probably neighboring Alaskan islands, on following 
characters: belugae with broader interorbital constriction than in 
phaeus; nasals of different outline, these bones not tapering caudad 
as they do in phaeus; nasals with lesser degree of extension back 
of a line joining the points of the antorbital tubercles. While indi- 
vidual specimens show intergradation in one or two of these char- 
acters, aS might be anticipated, it is true that in the series at hand 
every specimen is clearly separable. Externally belugae is dis- 


roe Cc 


Vil. COMPARISON OF EXTERNAL CHARACTERS OF Castor canadensis phaeus HELLER, FROM ADMIRALTY ISLAND, ALASKA Ww 


IT i 
TAYLOR, FROM COOK INLET REGION, ALASKA, AND WITH (Castor canadensis canadensis KUHL, FROM EASTERN ane 


Points of Comparison 


mereral coloration ........-......--.---.---.-- 


SLAC ¢ 2.9 CCS: RI a 


oy AIS TPES a1 hie 


RPE GHG VOL BIGGS... 2--.-2 con 2ccts-cce een 


op $. Titi ee 


0 Ee ee 


EERIE 1s) 1 ALIOVC: <0. -<---2-.-<---------nes-anee- 


Forefeet 


WR a 


MPN EKal) OVETHAIT.........-----2---------------+-2° 


olansp inne sc ane eee 


image of tail beneath........-..-.--.----------- 


Ventral 


underfur 


Castor canadensis phaeus 
(two examples) 


No. 210 similar in coloration to no. 4347 
from Cook Inlet; no. 209 much darker; 
all the other specimens of phaeus similar 
to no. 209. 

No. 210 with tips of dorsal overhair between 
cinnamon and sayal brown; no. 209 vary- 
ing about seal brown. 

No. 210 benzo brown to bone brown; no. 
209 light seal brown to benzo brown. 

No. 210 with dark coloration of dorsal over- 
hair grading into the verona brown or 
warm sepia of the ventral overhair with 
slight intermediate lightening; no. 209 
cinnamon and sayal brown of dorsal over- 
hair grading toward the verona brown of 
the ventral overhair. 

No. 210 orange-cinnamon; no. 209 mikado 
brown. 


Nos. 210 and 209 cream-buff grading toward 
honey yellow. 

No. 210 near bister, becoming paler anteri- 
orly; no. 209 near warm sepia. 


No. 210 near cinnamon-drab; no. 209 cin- 


namon-drab to benzo brown. 


Lighter than in canadensis: No. 210 near 
mars brown, burnt umber or Hay’s 
brown; no. 209 deep brownish drab. 

Nos. 210 and 209 varying about verona 
brown and warm sepia. 


No. 210 varying about cinnamon-buff, 
slightly paler dorsally, slightly darker 
ventrally; no. 209, scarcely indicated 
pinkish buff in coloration. 

No. 210 hazel to chestnut; no. 209 varying 
about verona brown, warm sepia and 
mars brown. 


No. 210 shaft of hair pale drab-gray, term- 
inally approaching drab, general impres- 
sion of light drab given; no. 209 similar, 
but general impression nearer light cin- 
namon-drab. 

Broader than in canadensis. 


Castor canadensis belugae 
(one example) 


No. 4347 from Cook Inlet much like no. 210, 
somewhat paler; considerably lighter in 
coloration than the rest of the series of 
phaeus. 


No. 4347 cinnamon to ochraceous-tawny, so 
definitely lighter. 


No. 4347 a slightly paler tint of benzo or 
bone brown than in no, 210. 

No. 4347 with hair of sides approaching cin- 
namon and then grading into coloration 
of the underparts which is near army 
brown. 


No. 4347 similar to rest of upperparts, but 
shade very slightly lighter, near orange- 
cinnamon. 

No. 4347 cinnamon-buff and pinkish buff. 


No. 4347 between clay color and cinnamon- 
buff. 


No. 4347 pinkish buff mixed with a color 
near cinnamon-drab. 


No. 4347 varying about cinnamon-drab and 
benzo brown. 


No. 4347 lighter than in phaeus, nearest 
army brown, Rood’s brown, or benzo 
brown. 


No. 4347 with few long hairs near cinnamon- 
puff; no definite throat band. 


No. 4347 between pinkish buff and cinna- 
mon-buff. 


No. 4347 shaft of hair pale drab-gray, 
terminally ecru-drab to cinnamon-drab, 
general impression near ecru-drab. 


Narrower than in phaeus. 


CANADA, 


Castor canadensis canadensis 
(two examples) 


Similar to phaeus but no specimen as dark 
as no. 209. 


Hairs a mixture of blackish and ochraceous- 
tawny (nos. 174525, 174526), former the 
darker; general appearance near chocolate 
or bay. 

Near clove brown (nos. 174525, 174526). 


Overhair of sides tipped with cinnamon; 
broad area of intergradation between 
chocolate or bay dorsal coloration and 
warm sepia ventral coloration (nos. 
174525, 174526). 


Similar to rest of upperparts. 


Near pinkish cinnamon (no. 174526); near 
cinnamon-buff or pinkish buff (no. 174525). 

Cinnamon-drab to verona brown; hairs im- 
mediately at base of tail tipped with seal 
brown (no. 174525); chestnut to bay, 
hairs at immediate base of tail tipped 
with seal brown (no. 174526). 

General impression pale brownish drab (no. 
174525); approaching warm _ blackish 
brown (no. 174526). 

Near dark vinaceous-drab (no. 174525); 
similar but with tendency to be lighter 
(no. 174526). 

Bone brown, dark grayish brown, or dusky 
drab (no. 174525); a trifle paler, near 
dark vinaceous-drab or natal brown (no. 
174526). 

Definite throat band, the scattered lighter 


hairs being near cinnamon-buff (no. 
174525);  mearer  ochraceous-buff (no. 
174526). 


Liver or carob brown approaching orange- 
cinnamon laterally and anteriorly (no. 
174526); mixture of chestnut and cinna- 
mon-drab approaching pinkish buff lat- 
erally and anteriorly (no. 174525). 

Shaft of hair pale, nearest pale gull gray, 
tipped with drab-gray (nos. 174525, 
174526). 


Narrower than in phaeus. 


3 


1916] Taylor: Beavers of Western North America 433 


tinguished by slightly paler coloration. See table of color characters, 
table VII, opposite page 432. 


MATERIAL 


Twenty-two specimens, skulls only with one exception, partly 
from the collection of the Museum of Vertebrate Zoology, partly 
from that of the Biological Survey Collection in the U. S. National 
Museum: Beluga River, Cook Inlet region, Alaska, 10 (nos. 4219- 
4228, skulls only, Mus. Vert. Zool.) ; Nenilechuk or Ninilchik, east 
shore Cook Inlet, 2 (nos. 4229, 4230, skulls only, Mus. Vert. Zool.) ; 
Kasiliff (probably Kasilof), mouth of Kasilof River, Cook Inlet, 
1 (no. 4232, skull only, Mus. Vert. Zool.) ; Snug Harbor, probably 
west shore Cook Inlet near Iliamna Peak, 1 (no. 4347, skin and 
skull, Mus. Vert. Zool.) ; and probably from Cook Inlet, no data 
attached, 2 (nos. 4231, 4233, skulls only, Mus. Vert. Zool.) ; 12 miles 
north Stuart Lake, British Columbia, 1 (no. 77147, skull only, U. 8. 
Nat. Mus., Biol. Surv. coll.) ; 15 miles north Stuart Lake, 1 (no. 
77155, skull only, U. S. Nat. Mus., Biol. Surv. coll.) ; 17 miles north 
Stuart Lake, 1 (no. 77157, skull only, U. S. Nat. Mus., Biol. Surv. 
coll.) ; 20 miles east Stuart Lake, 2 (nos. 77150, 77159, skulls only, 
U. S. Nat. Mus., Biol. Surv. coll.) ; 30 miles east Stuart Lake, 1 
(no. 77158, skull only, U. S. Nat. Mus., Biol. Surv. coll.). 


DESCRIPTION OF A New SUBSPECIES OF BEAVER FROM EASTERN SHASTA 


CouNnTY, CALIFORNIA 


Certain crania of beavers obtained in California have recently 
been received from the Bureau of Biological Survey, of the United 
States Department of Agriculture. These belong to specimens col- 
lected at Cassel, on Hat Creek, a tributary of the Pit River, eastern 
Shasta County, California, which, while they are undoubtedly most 
closely related to Castor subauratus subauratus of the interior val- 
leys of California, do present constant differences therefrom. 


Castor subauratus shastensis, new subspecies 


Type—Skull only, 3 adult; no. 50978, U. S. Nat. Mus., Biol. 
Surv. coll.; Cassel [Hat Creek], Pit River, Shasta County, Cali- 
fornia; January 3, 1893; collected by H. E. Williams. 

Diagnostic characters—No skins of the new form are available. 


434 University of California Publications in Zoology (Vou. 12 


The outline of the nasals is distinctive. In Castor subauratus shas- 
tensis the nasals do not taper so regularly or rapidly posteriorly as 
in Castor subawratus subawratus. The lateral nasal outline is, in 
shastensis, invaded by the postero-medial portion of backward- 
extending tongue of the premaxilla. Consequently there is formed 
a “‘bay’’ in the lateral outline of the nasals posteriorly, which is 
lacking in subauratus. In one or two specimens of the latter there 
is a slight indication of such a relation, but the diagnostic value of 
the character as given, both in adult and young specimens of both 
species, holds in available material. The nasals tend to maintain 
their full breadth farther posteriorly in shastensis than in sub- 
auratus. The temporal ridges tend to form a distinct sagittal crest 
posteriorly and to show a higher degree of approximation anteri- 
orly in shastensis, in specimens of the age which in subauratus 
exhibits a weak development of the crest posteriorly and a lyrate 
arrangement of the ridges anteriorly. Not only is there developed 
a distinct sagittal crest, but also a much more distinet knob dorsally 
on the crest located about 29 millimeters anterior of the posterior 
border of the lambdoidal ridge. The size of the cranium and the 
condition of certain sutures, whether open or closed, constitute the 
chief bases for age determination. Specimen no. 51477 of shastensis, 
which is considerably younger than specimen no. 12654 of suwb- 
auratus, presents the same general arrangement of the temporal 
ridges. Shastensis no. 50979, which is certainly not older than 
specimen no. 12654 of subawratus, exhibits the characteristically 
distinet sagittal crest posteriorly and the higher degree of approx- 
imation anteriorly. The lambdoidal ridge is also more strongly 
developed in ecrania of nearly equal age. The use of these char- 
acters as subspecifically differentiatory might be unjustified without 
a mass of material, were they not correlated with others; for they 
undergo marked modification with age within the same subspecies. 
However, a different degree of development for the same age 
undoubtedly can be relied upon. In this case, although the available 
material is not sufficient to prove, for instance, that very old examples 
of subauratus would not show the distinet sagittal crest posteriorly, 
the higher degree of approximation of temporal ridges anteriorly, 
and the more strongly developed lambdoidal ridge, it does suffice to 
indicate a difference at least in rate of progress, and this difference 
is valid as a subspecific character. 

The interorbital constriction is broader in shastensis. This meas- 


1916] Taylor: Beavers of Western North America 435 


urement in shastensis no. 50977 exceeds that in subawratus no. 
12668, which has the same basilar length, by 8.9 percent. The 
difference holds throughout the series in specimens of comparable 
age. 

The fronto-maxillary suture, situated dorsally on skull between 
backward-extending tongue of premaxilla and malar, is longer in 
shastensis than in subauratus. This holds for all specimens, regard- 
less of age (see tables of measurements, pp. 486 and 449.) 

The interparietal is somewhat broader in all specimens of shas- 
tensis, old and young, than it is in any specimen of swbauwratus. 

Remarks——The new subspecies, while clearly marked off from 
the beaver of the San Joaquin Valley by a number of valid cranial 
characters, nevertheless finds in the golden beaver its closest ally. 
This is shown by the facts that: (1) It is nearly identical with 
Castor subauratus subauratus in many cranial dimensions; (2) its 
foramen magnum shows the same general proportions; (3) its 
process medially in the interpterygoid fossa is nearly identical 
with that in subauratus, being different in form from that of any 
other west American beaver. 

Cassel, Shasta County, California, the type locality of the new 
form, is situated on Hat Creek, a tributary of the Pit River, which 
is in turn a tributary of the Sacramento River. The two forms, 
Castor subauratus subauratus and Castor subauratus shastensis, are 
found in the same hydrographic basin, namely that draining into 
San Francisco Bay. It should be noted, however, that the type 
locality of the Shasta beaver is on the eastern slope of the main 
chain of the Sierra Nevada Mountains. The surrounding region is 
characterized by environmental conditions probably much more 
typical of the Great Basin faunal area than of those of the Sacra- 
mento Valley. It is entirely possible, if not probable, that the Pit 
River Narrows at present constitute a barrier not regularly crossed 
by beavers. The limits of the range of Castor subauratus shastensis 
are yet to be defined. There would seem to be a possibility that 
the beaver of the Great Basin will be found to be referable to it. 

Material.—Five specimens, skulls only, all loaned to the writer by 
the authorities in charge of the Biological Survey mammal collection, 
United States National Museum: Cassel, Hat Creek, Pit River, Shasta 
County, California (nos. 50976-50979, 51477). 


[Vot. 12 


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1916] Taylor: Beavers of Western North America 437 


CoMPARISONS OF CERTAIN AMERICAN BEAVERS 


CASTOR CANADENSIS PHAEUS HELLER, FROM ADMIRALTY ISLAND, 
ALASKA 


MATERIAL 


Six specimens of Castor canadensis phaeus, skins with skulls, 
all in collection of Museum of Vertebrate Zoology: Hasselborg 
Lake, Admiralty Island, Alaska, 4 (nos. 129, 185, 210, 211); Mole 
Harbor, Admiralty Island, Alaska, 1 (no. 128); Pleasant Bay, 
Admiralty Island, Alaska, 1 (no. 209). 

Three specimens of Castor canadensis canadensis, all from col- 
lection of Smithsonian Institution, United States National Museum: 
Nepisiquit River, New Brunswick, 2 (nos. 174525, 174526, skins with 
skulls); Moose River, Ontario, 1 (no. 4358, skin with cranium 
inside). 


CoMPARISON WitTH Castor canadensis canadensis KuHL, FRoM EASTERN 
CANADA 


General external characters.—Specimens compared: Castor cana- 
densis phaeus, nos. 128 ¢,129 6,185 8, 209 ¢, 210 ¢, and 211 ¢, 
Mus. Vert. Zool., from three localities on Admiralty Island, 
Alaska, May 16 to June 1, 1907; and Castor canadensis canadensis, 
no. 4858 ¢, May 21, 1860, and nos. 174525 ¢, 174526 ?, Septem- 
ber 25, 1911; all U. S. National Museum coll. from eastern Canada. 

In coloration the Admiralty Island skins as a series resemble 
the two specimens from New Brunswick, although the type of the 
Admiralty Island race (no. 209) is darker than either. No. 210 is 
a lighter example than the rest of the series of phaeus, with more 
cinnamon to sayal brown coloration dorsally. Canadensis no. 4858 
is paler than the other specimens representing the same subspecies 
(nos. 174525, 174526). 

The ratio of width of tail to its leneth in the two examples of 
phaeus (nos. 209, 210) is greater, as shown below, even than it is 
in the specimen of frondator at hand (no. 20751, U. 8S. Nat. Mus.), 
attaining 54.1 percent in no. 209. The widest part of the tail, how- 
ever, is located more proximally in phaeus than it is in frondator. 

For comparison of external characters, see table VII, opposite 
p. 482. 


438 University of Califorma Publications in Zoology (Vor. 12 


IX. MEASUREMENTS AND RATIOS OF SCALED PoRTIONS OF TAILS 


All measurements in millimeters, and taken in dry skins; see fig. H, p. 431 


Ratio 

Museum width to 

Subspecies— number Length Width length 
Wastor .¢; DWACUS eerecceeeeeen sete eeeeeee 209 240 130 54.1 
Gaston cy PHAGUS) se. 210 248 132 53.2 
Castor ¢c. canadensis -.-----....----- 174526 260 108 41.3 
Castoric) canadensis) = 174525 265 116 43.4 
Castor c. canadensis ............-...-- 4358 223 92 41.1 


General cranial characters——(See table of measurements, oppo- 
site.) Crania compared: Castor canadensis phaeus, nos. 209 2, and 
210 3, Mus. Vert. Zool., from Admiralty Island; and Castor cana- 
densis canadensis, nos. 174525 2, 174526 9, U. S. Nat. Mus., from 
New Brunswick. 


Interorbital constriction decidedly narrower in phaeus. Nasals 
in phaeus longer and narrower. Foramen magnum broader, the 
difference amounting to 13 and nearly 15 percent respectively in 
nos. 210 and 209, percentage taken on the basis of the average 
width in canadensis. Maxillary tooth-row longer in phaeus, conse- 
quently ratio of this measurement to basilar length greater. Teeth 
in phaeus with a tendency to be longer than in comparable skulls 
of canadensis. 

Process in middle of interpterygoid fossa shorter in phaeus. 
Hamular processes of pterygoid (see fig. F, p. 4830) broader-bladed in 
phaeus (no. 209, 4.1 mm.; 210, 4.1; no. 174525, 3.5; 174526, 3.1). 

Chief points of difference: Narrower interorbital constriction 
(in phaeus), nasals of different length and outline, broader foramen 
magnum, longer maxillary tooth-row, longer teeth. 


Compartson WitH Castor canadensis belugae TAYLOR, FROM THE CooK INLET 
REGION 

General external characters—Specimens compared: Castor can- 
adensis phaeus, nos. 128 3,129 ¢,185 ¢, 209 6,210 ¢, and 211 ¢, 
Mus. Vert. Zool., from three localities on Admiralty Island, Alaska; 
Castor canadensis belugae, no. 4847 6, Mus. Vert. Zool., Snug Har- 
bor, Alaska Peninsula. The specimens of phaeus were collected, as 
above stated, between May 16 and June 1. The single skin of 
belugae was secured June 14, 1904. 


1916] Taylor: Beavers of Western North America 439 


Specimens nos. 4347 and 210 are similar in coloration, the 
former being slightly paler. No. 209 is much darker than either, 
the rest of the series from Admiralty being nearest 209. 


phaeus. Hamular blades (see fig. F, p. 430) broader (no. 210, 4.1 
mm.; 209, 4.1. No. 4347, 2.5; 4225, 2.5; 4224, 3.1). Distance from 
most dorsal point on outline of foramen magnum to inion less 
(see fig. G, p. 431; no. 210, 19.4 mm.; 209, 17.1. No. 4847, 20.5 mm. ; 
4225, 22.4; 4224, 22.7). 


Museum 
number 


16383 
8988 
12654 
12101 
12111 
12107 
4220 
4347 
4225 
210 
209 
60354 
35946 
174525 
174526 
18525 
18526 
18527 


Os 40 40 03 40 03 O3 05 05 0 O5 40 O_O 10 10 


$ 


Castor 
Castor 
Castor 
Castor 
Castor 
Castor 
Castor 
Castor 
Castor 
Castor 
Castor 
Castor 
‘ Castor 
Castor 
Castor 
Castor 
Castor 
Castor 


X. COMPARATIVE MEASUREMENTS* OF ADULT SKULLS OF CERTAIN AMERICAN SPECIES OF BEAVER 
(All measurements in millimeters) 


SUBSPECIES—GENERAL LOCALITY 
subauratus subauratus (San Joaquin Valley, Calif.) 
subauratus subauratus (San Joaquin Valley, Calif.) 
subauratus subauratus (San Joaquin Valley, Calif.) 
canadensis leucodonta (Vancouver Id., B. C.)............ 
canadensis leucodonta (Vancouver Id., B. C.)............ 
canadensis leucodonta (Vancouver Id., B. C.).......-.... 
canadensis belugae (Cook Inlet Region, Alaska)...... 
canadensis belugae (Cook Inlet Region, Alaska)...... 
canadensis belugae (Cook Inlet Region, Alaska)...... 
canadensis phaeus (Admiralty Id., Alaska).............- 
canadensis phaeus (Admiralty Id., Alaska).............- 
canadensis frondator (Colorado River, Mex.).-.....-..-.- 
canadensis frondator (San Pedro River, Mex.)...........- 
canadensis canadensis (New Brunswick, Canada)... 
canadensis canadensis (New Brunswick, Canada).... 
canadensis michiganensis (Michigan) .......-.-.----..-----.- 
canadensis michiganensis (Michigan) ...........------------- 
canadensis michiganensis (Michigan) ......--....-.-----.--- 


Basilar length of Hensel 


120.1 
119.8 
126.3 
111.9 
110.9 
122.7 
118.0 
109.0 
122.5 
117.6 
114.7 
118.7 
ata liler¢ 
117.6 
115.2 


108.5 


Zygomatic width 


96.7 
94.5 
103.4 
94.7 
91.6 
101.0 
96.5 
90.3 
101.3 
97.1 
92.3 
96.8 
94.3 
95.2 
92.7 
91.7 
89.2 
87.9 


Mastoid width 


66.2 
69.4 
70.5 
63.4 
62.8 
69.5 
69.6 
61.7 
68.2 
64.2 
62.6 
65.3 
61.3 
63.0 
64.6 
60.4 
61.5 
60.9 


a Interorbital constriction 
~] 


25.6 
28.3 
23.9 
24.4 
25.7 
26.6 
20.9 
24.7 
20.8 
22.7 
25.0 
21.8 
24.5 
24.5 
24.3 
23.0 
23.7 


Length of nasals 
(See fig. C) 


*For manner of taking measurements see Taylor, 1911, pp. 206, 207, and table opp. p. 426 of present paper. 


Width of nasals 


26.6 


Maxillary tooth-row 


32.1 


ertical diameter of foramen 
(See fig. E) 


agnum. 


al) 


Transverse width of 
foramen magnum 


19.3 


angle 
of 


to anterior surface alveolus 
(See fig. D) 


Greatest length of mandible; 


incisor. 


105.7 


Ventral surface of mandible 


to coronoid 


Ratio width of nasals 
to basilar length 


Ratio maxillary tooth-row 
tm to basilar length 


26.0 


bo 
is 
os) 


25.5 
25.6 
25.4 
26.0 
26.9 
27.3 
27.6 
28.1 
27.5 
26.4 
24.8 
25.3 
24.9 
25.7 
24.8 


Ratio vertical diameter of foramen 


magnum to basilar length 


9.3 


438 University of California Publicatio 


IX. MEASUREMENTS AND RATIOS OF SCALED 


All measurements in millimeters, and taken in dr 


Mus. Vert. Zool., from three localities on Adr 
Castor canadensis belugae, no. 4847 3, Mus. 
bor, Alaska Peninsula. The specimens of ph 
above stated, between May 16 and June 1 
belugae was secured June 14, 1904. 


1916] Taylor: Beavers of Western North America 439 
Specimens nos. 43847 and 210 are similar in coloration, the 
former being slightly paler. No. 209 is much darker than either, 
the rest of the series from Admiralty being nearest 209. 
XI. MEASUREMENTS AND RATIOS OF SCALED PoRTIONS OF TAILS 


All measurements in millimeters, and taken in dry skins 


Ratio 
Museum width to 
Subspecies— number Length Width length 
Castor canadensis phaeus 209 240 130 54.1 
Castor canadensis phaeus 210 248 132 53.2 
Castor canadensis belugae 4347 245 115 47.0 


General cranial characters.—Crania compared: Castor canadensis 
belugae, nos. 4219-4233, 4347, Mus. Vert. Zool., all from the Cook 
Inlet region, Alaska; Castor canadensis phaeus, nos. 209 6, 210 6, 
Mus. Vert. Zool., from Admiralty Island. Specimens 209 and 210 
from Admiralty Island, and specimens 4220, 4225 and 4347 from 
Cook Inlet comparable as to age. 

Skulls of phaeus slightly smaller than the average of those of 
belugae. Interorbital constriction narrower in phaeus than in 
eighteen of the twenty-two specimens of belugae of all ages figured. 
Nasals longer, more tapering posteriorly. Sharp process medially 
in interpterygoid fossa reduced in phaeus. Foramen magnum 
averaging broader in phaeus. Vertical diameter of foramen mag- 
num in no. 209 exceeding that in any specimen from Cook Inlet, 
but itself exceeded by no. 77150 from Stuart Lake, British Colum- 
bia. Teeth narrower in phaeus, proportionally to their length. 
Maxillary tooth-row averaging longer, consequently ratio of max- 
illary tooth-row to basilar length greater in phaeus. No. 4233, the 
youngest cranium from Cook Inlet, furnishes the single exception 
to this rule. In this specimen the ratio of maxillary tooth-row to 
basilar length is greater than in no. 210. Coronoid process of 
mandible hghter in phaeus. lateral ridge on rostrum larger in 
phaeus. Hamular blades (see fig. F, p. 430) broader (no. 210, 4.1 
mm.; 209, 4.1. No. 4347, 2.5; 4225, 2.5; 4224, 3.1). Distance from 
most dorsal point on outline of foramen magnum to inion less 
(see fig. G, p. 481; no. 210, 19.4 mm.; 209, 17.1. No. 43847, 20.5 mm.; 
4225, 22.4; 4224, 22.7). 


440 University of California Publications in Zoology (Vou. 12 


CASTOR CANADENSIS LEUCODONTA GRAY, FROM VANCOUVER 
ISLAND, BRITISH COLUMBIA 


MATERIAL 


Sixteen specimens, skins with skulls, and skulls only, partly from 
the Museum of Vertebrate Zoology and partly from the United 
States National Museum, Biological Survey collection: Hall’s 
Ranch, Alberni Valley, Vancouver Island, British Columbia, 10 
(nos. 12101-12110, skins with skulls, Mus. Vert. Zool.) ; Great Cen- 
tral Lake, Vancouver Island, British Columbia, 1 (no. 12111, skin 
with skull, Mus. Vert. Zool.) ; San Josef River Valley, Vancouver 
Island, British Columbia, 5 (nos. 140569-140573, skulls only, U. S. 
Nat. Mus., Biol. Surv. coll). 


Comparison WitH Castor canadensis canadensis KunHt, FRoM EASTERN 
CANADA, AND Castor canadensis phaeus HELLER, FROM ADMIRALTY ISLAND 


General external characters—Specimens compared: Castor can- 
adensis leucodonta, nos. 12101-12111, Mus. Vert. Zool., all but the 
last (which is from Great Central Lake, Vancouver Island) are 
from Alberni, Vancouver Island, British Columbia, June 11-27, 
August 25, 1910; Castor canadensis canadensis, nos. 4858 38, U. 8. 
Nat. Mus., Moose River, Ontario, Hudson Bay region, May 21, 1860; 
nos. 174525 3, and 174526 ¢@, U. S. Nat. Mus., Nepisiquit River, 
New Brunswick, Sept. 25, 1911; Castor canadensis phaeus, no. 
128 3, Mole Harbor; nos. 129 6, 185 3, 210 6, 211 36, Hassel- 
borg Lake; no. 209 ¢, Pleasant Bay; all Admiralty Island, Alaska, 
in Mus. Vert. Zool. 

A specimen of leucodonta taken in August (no. 12111) is very 
similar in general coloration to the September examples of cana- 
densis from New Brunswick. The resemblance in coloration of the 
hair dorsally is close. Coloration beneath different, varying about 
hair brown in leucodonta, near bone brown, dark grayish brown, 
dark vinaceous-drab or natal brown in canadensis. The difference 
in ventral coloration is more marked between no. 12111 and no. 
174525, which is darker, than between no. 12111 and no. 174526, 
which is paler. The Vancouver Island series is not comparable 
with the Moose River example of canadensis (no. 4358). The over- 
hair of the latter beneath is darker than in lewcodonta, while the 
underfur beneath is paler. The fact that there has been much 
fading, however, renders comparisons taking account of no. 4358 
of dubious value. 


1916] 


Taylor: Beavers of Western North America 


18.1 
18.5 
20.9 
19.9 


441 


| 
| 


Castor canadensis leucodonta 


Points of Comparison 
(six examples) 


- smount of pelage...-....--..---- Medium. 
’ a 
General coloration -..........- Paler than in canadensis or phaeus. 
Dorsal overhair.........-...------- Varying about cinnamon-buff; no. 12108 
with liberal insprinkling of seal brown 
hairs. 
+ 
Dorsal underfur .............-.--- Varying from fuscous and fuscous-black to 
benzo and hair brown. 
Top of head.............-.-.--------- Not conspicuously different from rest of 


dorsal coloration. 


Ventral overhair................... Lighter than in phaeus; varying about hair 
brown. 
MIG i) rr Paler brown than in canadensis, varying 


from near carob brown to near hazel. 


Base of tail above...............- Varies from near hair brown (no. 12111), 
to a shade between chocolate and bay 
(no. 12107); fringe of hairs near dark 
vinaceous-drab in several specimens 
(notably nos. 12108, 12101, 12102). 


Base of tail beneath.......... Between cinnamon-brown and chestnut or 
bay. 
ithy Tit a Inconspicuous; the few hairs present vary 


about warm buff. 


a Intermediate in width in proportion to 
length, between canadensis and phaeus. 


Castor canadensis pacificus 
(one example) 

Heavier pelage than in lewcodonta. 

Paler than in leucodonta. 

Varying from cinnamon in the middle of 
the back to lighter tints both anteriorly 
and posteriorly, attaining a pinkish buff 
color above the tail and on the cheeks. 


Varying about brownish drab. 


Somewhat lighter than middle of back. 
Difference not conspicuous. 


Cinnamon-drab to light cinnamon-drab. 


Close to benzo brown. 


Varies from the fringe of cinnamon-drab or 
benzo brown hairs to cinnamon-buff or 
even to pinkish buff ten em. anteriorly of 
base of scaled portion of tail. 


Cinnamon-buff to pinkish buff, with a wash 


of cinnamon-drab close to base of scaled 
portion of tail. 


Inconspicuous; lacking as a band. 


Narrower proportionally than in any of the 
others. 


Castor canadensis canadensis 
(two examples) 

Heavier pelage than in leucodonta. 

Darker. 

Similar to phaeus but no specimen as 
dark as no. 209. Hairs a mixture of 
blackish and  ochraceous-tawny (nos. 
174525, 174526); general appearance near 
chocolate or bay. 

Near clove brown (nos. 174525, 174526). 


Similar to rest of upper parts. 


Bone brown, dark grayish brown, or dusky 
drab (no. 174525); a trifle paler, near 
dark vinaceous-drab or natal brown (no. 
174526). 

Near dark vinaceous-drab (no. 174525); 
similar but with tendency to be paler 
(no. 174526). 

Cinnamon-drab to verona brown; hairs at 
immediate base of tail tipped with seal 
brown (no. 174525); chestnut to bay; 
hairs at immediate base of tail tipped 
with seal brown (no. 174526). 

Mixture of chestnut and cinnamon-drab, ap- 
proaching pinkish buff laterally and an- 
teriorly (no. 174525); liver or carob 
brown approaching orange-cinnamon lat- 
erally and anteriorly (no. 174526). 

Definite throat band near cinnamon-buff 
(mo. 174525); nearer ochraceous-bufft (no. 
174526). 


Narrower in proportion to length than in 
either leucodonta or phaeus. 


EXTERNAL CHARACTERS OF Castor canadensis leucodonta GRAY, FROM VAN 
XII. COMPARISON OF ; COUVER ISLAND, BRITISH COLUMBIA, WITH Cast i i 
FROM WASHINGTON STATE, Castor canadensis canadensis KUHL, FROM EASTERN CANADA, AND Castor canadensis phaeus HELLER, FROM ADMIRADED teLee 1 eee 


Castor canadensis phaeus 
(two examples) 
Heavier pelage than in leucodonta. 
Darker. 
No. 210, between cinnamon and _  sayal 
brown; no. 209, seal brown, clove brown 
or blackish. 


Benzo brown to bone brown (no. 210); light 
seal brown to benzo brown (no. 209). 
Conspicuously different from rest of dorsal 
coloration. Orange-cinnamon (no. 210); 

mikado brown (no. 209). 

Darker than in leucodonta. Varying about 
verona brown and warm sepia (nos. 209 
and 210). 


Lighter than in canadensis. Near mars 
brown, burnt umber or Hay’s brown 
(no. 210); deep brownish drab (no. 209). 

Bister immediately at base, becoming paler 
anteriorly (no. 210); near warm sepia 
(no. 209). 


Hazel to chestnut (no. 210); varying about 
verona brown, warm sepia and mars 
brown (no. 209). 


Varying about cinnamon-buff, slightly paler 
dorsally, slightly darker ventrally (no. 
210); few scattered pinkish buff hairs 
(no. 209). 

Broader than in either lewcodonta or cana- 
densis in proportion to length. 


440 University of California Publication 


PASTOR CANADENATN T.RTICONANTA GRA 


ES ip Pt a) 8 Da LATOR 
” rvuert YP MIO 


> rate 36 Bias 


underfur beneath is paler. The fact th 
fading, however, renders comparisons talk 
of dubious value. 


1916] Taylor: Beavers of Western North America 441 


In general, the table of comparisons (see table XII, opposite p. 
440) shows lewcodonta to be paler than phaeus. 

Difference in amount of hair is marked. The pelage is lghter 
weight in lewcodonta than it is in either phaeus or canadensis. 


XIII. MEASUREMENTS AND Ratios or ScALDED Portions or TAILS 


All measurements in millimeters, and taken in dry skins; see fig. H, p. 431 


Ratio 

Museum width to 

Subspecies— number Length Width length 
Gastormc: Leucodomtay <ecrec-ceee------ 12101 237 120 51.0 
Wastorsc lCUCOdOM Lay ce cpcccceceenea-e= 12111 243 108 44.6 
Gastonscaleucodontane..- 12107 270 124 46.0 
@astonie canadensis) y-ce esse 174525 265 116 43.4 
Castonvemcanadensisy coctcc-ccsceene-e 174526 260 108 41.3 
Castor’ cs canadensis” -227--...-.-..- 4358 223 92 41.1 
@astoreephaeus eee eens 209 240 130 54.1 
GAStOT NG DN AGUS cerca ceencnnaae seen ne 210 248 132 53.2 


General cranial characters—Crania compared (being of approx- 
imately same age): nos. 12101 ¢, 12107 ¢, 12111 ¢, Mus. Vert. 
Zool., from Vancouver Island; nos. 174525 ¢, 174526 9, U. 8. Nat. 
Mus., from New Brunswick; nos. 209 ¢, 210 ¢, Mus. Vert. Zool., 
from Admiralty Island. 

Leucondota (see tables of measurements, opp. pp. 426 and 438) is 
separated from canadensis by the following characters: External 
outline of nasals different, tending to be more parallel in leucodonta 
than in canadensis, in which there is a dilation in the outline anteri- 
orly; foramen magnum slightly broader in leucodonta; hamular 
processes (see fig. F, p. 480) definitely broader bladed in lewcodonta 
(no. 12107, 4.4 mm.; 12101, 4.6; 12111, 4.8. No. 174525, 3.5; 174526, 
Sel): 

Distance between most dorsal point on curve of foramen magnum 
and point of junction of lambdoidal ridge and sagittal crest (inion, 
see fig. G, p. 481) greater in the eastern subspecies, as shown by the 
following figures: 


XIV. MEASUREMENTS, FoRAMEN MAcNum To INION 
Measurements in millimeters 


Basilar length Distance foramen 

Specimen number of Hensel magnum to inion 
UPHOiIL Fs Seaodamobosundd SAcocOuUle 111.9 15.4 
MLPA np. fe, Si stevspsretsyeisteterererbeveretel© 110.9 18.1 
IVA Sagonooe SCD Os oO doo DOOooU oOo 122.7 18.5 
IEEPAaY 4 gona focaouer EO oanOMoO 117.6 20.9 


WCPG popomanuboonecoaseNGoD ado 115.2 929 


442 University of California Publications in Zoology (Vor. 12 


C. c. leucodonta is separated from C. c. phaeus by the following 
characters: Nasals less tapering posteriorly and shorter in leuco- 
donta; interorbital constriction broader; foramen magnum nar- 
rower (exception, phaeus no. 210 has a narrower foramen magnum 
than leucodonta no. 12107); maxillary tooth-row shorter in lewco- 
donta; ratio of maxillary tooth-row to basilar length less; palate 
anteriorly (see fig. I) broader (no. 12107, 6.9 mm.; 12101, 6.9; 


Fig. I. Ventral aspect of palate, to show 
method of taking measurements. 
Shorter measurement is “palate 
anteriorly”. Longer dimension is 
that to show the tooth-flare. Ap- 
proximately one-half natural size. 


12111, 6.8. No. 209, 5.6 mm.; 210, 5.1) ; blade of hamulars (see fig. 
F, p. 430) slightly broader (no. 12107, 4.4 mm.; 12101, 4.6; 12111, 
4.8. No. 209, 4.1; 210, 4.1). 


CASTOR CANADENSIS PACIFICUS RHOADS, FROM THE MAINLAND OF 
BRITISH COLUMBIA AND WASHINGTON 


MATERIAL 


Eleven specimens, all from the collections contained in the 
U. 8. National Museum: Puget Sound, 1 (no. 3936, jaws only) ; 
Skagit River [Washington], 1 (no. 36721, skull only); Lake Cush- 
man, Mason County, Washington, 8 (nos. 71276, 71812, 71814, 
78395, 87628, 87629, skulls only, and nos. 71830, 71833, skins only, 
Biol. Surv. coll., in U. 8. Nat. Mus.) ; Keechelus Lake, Washington, 
1 (no. 126190, Biol. Surv. coll.) ; Fisher, Five River Valley, Ore- 
gon, 2 (nos. 136605, 136606, skulls only, Biol. Surv. coll.). 


ComPARISON WitH Castor canadensis leucodonta Gray, FROM VANCOUVER 
ISLAND, AND WITH CERTAIN OTHER SPECIES OF BEAVERS 


General external characters——Material representative of external 
characters is limited. One young topotype skin of Castor cana- 
densis pacificus Rhoads, from Lake Keechelus, Washington, is at 
hand. According to the label it was probably taken in February, 
1903. No example of Castor canadensis leucodonta strictly corre- 
sponding in age and season with this topotype of pacificus is at 


1 This skull was specified by Allen (1877, p. 441) as from Europe, and was 
commented upon as presenting an exceptional character. 


1916] Taylor: Beavers of Western North America 443 


hand. Nearest in these respects are nos. 12103, 12104, from Hall’s 
Ranch, Alberni Valley, Vancouver Island, taken June 15 and 18, 
respectively, 1910. As regards change in external characters with 
age, it should be called to mind that the modifications are not 
great in the material worked over (see p. 420). Respecting seasonal 
change our knowledge is less complete. 

In Castor ec. leucodonta the dorsal overhair appears darker than 
in Castor c. pacificus (see table XII, opposite p. 440, for comparison 
of characters). The general coloration belongs to the cinnamon series 
in both. The darker impression doubtless is the result of the presence 
in leucodonta of more dark hairs sprinkled in with the hght, as, for 
example, in no. 12108. In lewcodonta the dorsal underfur is a darker 
shade of brown than it is in pacificus. 

Ventral coloration shows the same general relation, having a 
darker cast in lewcodonta than in pacificus. The overhair ventrally 
is in leucodonta about as much darker than it is in pacificus as 
benzo brown is darker than cinnamon-drab. No color difference as 
regards underfur ventrally could be distinguished. The difference 
in coloration extends to the feet. The hind feet are hazel to carob 
brown in leucodonta, while in pacificus they are close to benzo 
brown. 

General cranial characters.—Nasals in Castor c. leucodonta give 
the impression of being longer and narrower than in pacificus, 
though the measurements show the reverse to be true in the matter 
of length. Nasals in pacificus have their outlines somewhat more 
invaded laterally by the backward-extending tongues of the pre- 
maxillaries. This lateral invasion of the nasal outline occurs at a 
point farther anteriorly in pacificus than it does in either lewcodonta 
or belugae. Beginning at the anterior end of the skull, and pro- 
ceeding posteriorly, the nasal outlines are at first nearly parallel, 
then the premaxillaries interrupt the outline, and the nasals become 
somewhat narrowed, their lateral outlines again attaiming a nearly 
parallel relation. This posterior parallel portion of the outline is 
longer in pacificus than in leucodonta. looking at the crania in 
their anterior aspect there is manifest a tendency for the nasals to 
fall away in a regular curve from the median line. In lewcodonta, 
on the other hand, the usual condition is a flattish area dorsally on 
the median line. In the mainland examples the nasals extend far- 
ther down on the side of the rostrum anteriorly, showing a condi- 
tion which may be described as a droop. The nasals as a rule do 


444 University of California Publications in Zoology (Vor. 12 


not extend posteriorly of a line connecting antorbital tubercles in 
either leucodonta or pacificus. 

The lateral ridge on rostrum is more prominent in leucodonta 
than in pacificus. 

The hamular blades are broad in lewcodonta (see fig. F, p. 430) ; 
narrow in the Lake Cushman examples, and tending to show an 
intermediate condition in Castor c. belugae from Stuart Lake. 
Broad in two juvenals from Fisher, Oregon (nos. 136605, 136606) 
as well as in a young topotype skull of pacificus from Lake Keeche- 
lus, Washington. Measurements: Castor c. leucodonta, Vancouver 
Island, no. 12107, 4.4 mm.; 12101, 4.6; 12111, 4.8; Castor ec. paci- 
ficus, Lake Cushman, Washington, no. 71814, 2.2 mm.; 87629, 2.8; 
Castor c. belugae, Stuart Lake, British Columbia, no. 77155, 3.1 
mm.; 77150, 3.2; the two juvenals from Fisher, Oregon: no. 136605, 
4.3 mm.; 136606, 3.8; the young topotype cranium of Castor c. 
pacificus: no. 126190, 4.3. In leucodonta this character varies little 
with age. That it may do so in other subspecies, however, is indi- 
cated by the fact that a young specimen of undoubted Castor cana- 
densis frondator (no. 60355), has broad-bladed hamulars, while an 
older specimen of the same subspecies (no. 35946) has narrower 
ones. 

The occipital crest is flattened in leucodonta, upright in all but 
one of the specimens of belugae from Stuart Lake, British Colum- 
bia, and in a majority of those of pacificus from Lake Cushman, 
Washington. The disparity between pacificus and leucodonta in 
this respect is shown by the measurements given in the tables (p. 445, 
most dorsal point on the outline 


ee 


and opp. p. 426) under the heading 
of foramen magnum to occipital crest on the median line’’ (see fig. G, 
p. 431). To make the comparison fair, only the three oldest crania 
of leucodonta (nos. 12111, 12101 and 12107) should be taken into 
account. 

The foramen magnum is similar in general shape. Range of 
the ratio of vertical diameter of foramen magnum to basilar length: 
In belugae, with old and young crania represented, 9.8 percent to 
14 per cent; in pacificus, with adult skulls only represented, 9.0 
per cent to 12.5 per cent; in lewcodonta, with old and very young 
skulls represented, 11.3 per cent to 17.7 per cent. 

The specimens from the mainland of British Columbia and 
Washington are clearly set off from Castor canadensis canadensis 
and its close relative C. c. michiganensis through the characters of 


445 


Beavers of Western North America 


Taylor 


1916] 


XV. CRANIAL MEASUREMENTS* OF Castor canadensis pacificus FROM THE MAINLAND} OF WASHINGTON STATE 


(All measurements in millimeters) 


ee g 
EES E 
ee: St a 
Ric a) ABD) 
Coe 8 og 8 
PD eten ers dae oe eB 
= 2 a F Zig ke 48 33 past ore 
4 S 4 g ‘ c Gi eC ee ee | a eee 
Fi} B = qe 4S E See UB) he bce ae as 
Ee z @ oe. ee & = bag ES OS cn 
gee ee ee ere eee ee ee 
fe 5 = cE he = 5 eER SE 2 Sig oe 38 
M a Eo 4 se ED zc 4 2 es B o S 65 S 
IDE) Sex as N a eg aw E s S88 Se Bs SE ee ge 
EL ae aie olen tem 1167 921 G52) 241 hod B27 "306 207) d4lby glo) | esa roe engin 
(UCTS Wate See (S:7 (94.3 6402) pdlou = 40\s, | e/g) (olay anode dai eg) Om spat oD 
S768 wou ns ee Ws 95:8 646 | 281% | sai <2D1G) 3010) 20s) 13.8) Ave) dd Dyn TORT 
XN Si he ieee ue debian cee 6.5 25.5 50.6 268 312 220 142 204 Iie 248 sao 
ONC oe Mee TIE GRRE) = heyh py ZING) YE ORG) OI Ge I) ala i) Oy 
{ADU eles ee oe 1247 951 68:2 26:9 48.8 87 816 208 dw isin 9 O10) ehisy 00 
S7620: PeAte Pipes (2005 ees CIN Psi TU | RIG LIS SVE IG Slnky PR) DHL 


*For manner of taking measurements see Taylor, 1911, pp. 206, 207, and table opp. p. 426 of the present paper. 
yAll from Lake Cushman, Mason County, except no. 3672, which is from Skagit River. 


446 University of California Publications in Zoology (Vou. 12 


certain cranial dimensions, condition of median process of inter- 
pterygoid fossa, and outline of nasals. 


CASTOR SUBAURATUS SUBAURATUS TAYLOR, FROM THE 
SAN JOAQUIN VALLEY, CALIFORNIA 


MATERIAL 


Eleven specimens, skins with skulls, in collection of Museum of 
Vertebrate Zoology: Westley, near Grayson, Stanislaus County, 
California, 1 (no. 8869); Grayson, Stanislaus County, 2 (nos. 8987, 
8988); Tuolumne River, 5 miles north of Grayson, 2 (nos. 12668, 
12669); San Joaquin River, 5 miles north of Grayson, 1 (no. 
12654); San Joaquin River, 10 miles north of Grayson, 4 (nos. 
16382-16385) ; ‘‘Sespe River, Ventura County, California’’, locality 
possibly erroneous, 1 (no. 4918, skull only). 


CompPartIson WituH Castor canadensis pacificus RHOADS, FROM WASHINGTON 
SraTE, AnD Castor canadensis frondator MEARNS, FROM THE COLORADO AND 
San Prepro Rivers, MEXxIco. 


General external characters 


Specimens compared.—Castor subauratus subawratus, nos. 8869, 
8987, 8988, 12654, 12668, 12669, 16382-16385, Mus. Vert. Zool., all 
from the general vicinity of Grayson, San Joaquin River, Stanis- 
laus County, California, taken November 22 to March 25; Castor 
canadensis pacificus, no. 126190, U. 8. Nat. Mus., Keechelus Lake, 
Washington, February (?), 1903; Castor canadensis frondator, no. 
20751, U. S. Nat. Mus., San Pedro River, Sonora, Mexico, October 
14, 1892. 

Comparative material illustrative of external characters is inade- 
quate, as the list of specimens compared shows. While of swbawra- 
tus there is a good series, one skin only of pacificus is available, 
and this is representative of a young individual. The single speci- 
men of frondator is an adult. Fortunately all the skins were taken 
during the winter months, so seasonal discrepancy is eliminated. 

The California skins have less hair above than has the example 
of pacificus. Several of them have thinner hair below, also. 

Coloration in subauratus is remarkably close to that in pacificus 
as illustrated in the material here compared. The color of the Lake 
Keechelus example has a slightly paler cast than that of the average 
of subauratus, for the reason that the overhair of the example of 


Taylor: Beavers of Western North America 447 


EXTERNAL MEASUREMENTS* OF Castor subauratus subauratus, 
FROM STANISLAUS COUNTY, CALIFORNIA 


XVI. COMPARISON OF EXTERNAL CHARACTERS OF Castor subauratus subauratus TAYLOR, FROM THE 
canadensis pacificus RHOADS, FROM LAKE KEECHELUS, WASHINGTON, AND WITH Cas 


FROM THE SAN PEDRO RIVER, MEXICO 


Castor subauratus subauratus 
Points of Comparison (ten examples) 
eeneral COlOTAtION. <.<..-2<:0<.---2--<-<0----~-- Medium pale. 
MORRRINOVERDOAT 2..25<-ac<-cccceoce-na-n----neenae Cinnamon to light pinkish cinnamon; pos- 
e sessing luster in strong light which is 
not so evident in other forms. 


Varying about dark purple-drab, dark 
vinaceous-drab, dusky brown, and dark 
grayish brown. 


MEM ANOINICETOUE: c2c-.-n.cciesconucsnecnesnnse-o 


SSDS 2 ee re Varying from cream color and light buff 
to somewhat darker shades of the series. 

VOICI eee Ree Varying from vinaceous-buff, avellaneous, 
and wood brown, to fawn color, deep 
olive-buff, and even warm buff. 

ch 0 DG) oe se Varying about benzo brown; no. 12654 near 


chestnut. 


approaching 
Luster in 


Varying about benzo brown; 
light buff about fore limbs. 
evidence. 

Varying about pallid vinaceous-drab. 


VON, OV Gt) 


VOI G15 6 i) 


Base of tail above.................--....--.----- Varying about cinnamon; range of varia- 
tion approximately between pinkish cin- 


namon and cinnamon-rufous. 


Rather narrow zone of hairs varying be- 
tween pinkish cinnamon and chestnut; in 
some specimens there seems to be little 
difference in coloration between rest of 
underparts and base of tail. 

Proportionally broader than in pacificus, 
narrower than in frondator. 


Castor canadensis pacificus 
(one example) 
Similar to subauratus, perhaps slightly 
paler than the average of the series. 
Varying from cinnamon in the middle of 
the back to lighter tints both posteriorly 
and anteriorly, attaining a pinkish buff 
color above the tail and on the cheeks. 
Varying about brownish drab. 


Near deep brownish drab. 


Near avellaneous or wood brown. 


Close to benzo brown. 


Cinnamon-drab to light cinnamon-drab. 


Drab-gray to pale ecru-drab. 


Varying from the fringe of cinnamon-drab 
or benzo brown hairs to cinnamon-buff 
or even to pinkish buff ten cm. anteriorly 
of base of scaled portion of tail. 

Cinnamon-buff to pinkish buff, with a wash 
of cinnamon-drab close to base of scaled 
portion of tail. 


Averages narrower proportionally than in 
either of the other species. 


SAN JOAQUIN VALLEY, CALIFORNIA, WITH Castor 
tor canadensis frondator MEARNS, 


Castor canadensis frondator 
(one example) 
Paler than in subauratus. 


Varying about cinnamon. 


Pale tints of the drab series, near cinna- 
mon-drab, light cinnamon-drab and light 
drab. 

Clay color and darker. 


Mixture of colors which in combination ap- 
proach clay color and hazel. 


Mixture of colors close to clay color and 
chestnut-brown; the chestnut-brown more 
in evidence than the clay color, making 
foot appear darker. 

Close to cinnamon-buff. 


Decidedly lighter than in suwbawratus; ap- 
proaching pale ecru-drab and pale drab- 
gray. 

A fringe of hairs at immediate base of tail 
colored near hazel and chestnut-brown. 


Broad area at immediate base of tail and 
extending somewhat anteriorly from it 
colored near hazel and chestnut-brown. 


Averages broader proportionally than in 
either subauratus or pacificus. 


446 University of California Publications i 


certain cranial dimensions, condition of median 
pterygoid fossa, and outline of nasals. 


OL SWUWUTUWIUS, LUE Ute Leasui una une Uy unas 


1916] Taylor: Beavers of Western North America 447 


XVII. EXTERNAL MEASUREMENTS* OF Castor subauratus subauratus, 
FROM STANISLAUS COUNTY, CALIFORNIA 


Specimens arranged approximately in order of age from the top of table to 
bottom 


(All measurements in millimeters) 


> 3 s es ze 28s 
Museum 3 rl BI B 88 32 $55 
number Sex LOCALITY a eae FH A oe ER aes 
16384 3 10 mi. N. Grayson 834 391 155 27 234 37.6 
12668 é 5 mi. N. Grayson 960 170 265 21 
16385 é 10 mi. N. Grayson 1064 493 185 31 298 124 41.6 
16382 fe) 10 mi. N. Grayson 1108 190 305 137 45.0 
8987 fe) 3 mi. N. Grayson 1038 192 
12669 fe} 5 mi. N. Grayson 1090 185 282 27.5 
16383 é LO%mis N; Grayson) D218 503) Wsi 35; sil! 133 42.8 
8988 fe) GAY SON) sccsesee ee 11385 360 195 310 140 45.2 
12654 fo} 5 mi. N. Grayson 1171 196 31 320 139 39.51 43.4 


*For method of taking measurements see Taylor, 1911, p. 207. 
+From crown of head to tip of ear, measured on medial surface of pinna. 
¢Entrails removed. 


pactficus is thicker than that in the series of subauratus, concealing 
the darker underfur more effectually. 


Illustrative of modification in coloration within the same sub- 
species are the following observations on the coloration of base of 
tail. One specimen of Castor s. subauratus (no. 16383) has the 
coloration dorsally in this region paler than the rest of the series, 
approximately clay color. Ventrally the hairs are chestnut for 
nearly all their length. Im another example (no. 8869) the hairs 
are pale in coloration both above and below, the colored ends being 
hazel instead of chestnut. In no. 12654, the type of subauratus, 
the hairs of this region are darker than the average of the rest of 
the series. Variation of about this amount is noted in all the sub- 
species of which adequate series are available. With all this modi- 
fication, however, the series of ten skins of suwbawratus is remark- 
ably uniform in general coloration. 

The general aspect of the swbawratus series is darker than that 
in frondator, the difference being due to darker underfur. 


448 Unversity of California Publications in Zoology (Vou. 12 


XVIII. MEASUREMENTS AND RATIOS OF SCALED PoRTIONS OF TAILS 


All measurements in millimeters, and taken in dry skins; see fig. H, p. 431 


Ratio 

Museum width to 

Subspecies— number Length Width length 
Castors) subaumatuss eee 16385 298 124 41.6 
Castor s: subauratus -.....-............. 16382 305 137 45.0 
Castor s. subauratus ...................- 16383 311 133 42.8 
Castorsss subauratuse- see 8988 310 140 45.2 
Castors; subauratus: S22. eee 12654 320 139 43.4 
CaStOrnvCmDaCLhCUS pee eee 126190 185 74 38.9 
Gastornie trondaton =e 20751 232 113 48.9 


General cranial characters 


Cran compared.—Castor subauratus subauratus, nos. 16383 2, 
16384 ¢, 8988 2, and 12654 ¢, Mus. Vert. Zool., all from the vicinity 
of Grayson, San Joaquin River, Stanislaus County, California; Castor 
canadensis pacificus, nos. 3672, 71276 ¢, 71812 ¢, 71814 92, 
87628 9, 87629 3, 126190 ¢, U. S. Nat. Mus., all from the main- 
land of Washington State; Castor canadensis frondator, nos. 60354 
6, 8594 38, 35946 ¢°, from the Colorado River, Mexico, 15 miles 
south of Yuma, Arizona, and from the San Pedro River, Mexico, 
respectively. 

Comparable crania of Castor s. subawratus exhibit a tendency 
to be larger than those of Castor c. pacificus. They exceed those 
of pacificus absolutely in width of nasals and ratio of width of 
nasals to basilar length. Vertical diameter of foramen magnum 
less in all comparable crania of swbhawratus than in pacificus or in 
frondator with one exception (pacificus no. 71276 has this diameter 
11.2, which is the same as that in suwbawratus no. 16383); and 
associated with this difference, the ratio of vertical diameter of 
foramen magnum to basilar length is less in the comparable crania 
of subauratus than it is in pacificus (with the single exception men- 
tioned) or in frondator. 

The anterior dilation of the nasal outline in subauratus is 
marked; it clearly separates the California form from pacificus, 
but not from frondator, though absolute measurements in swbawra- 
tus exceed those in the latter. 

In subauratus the extension of the nasals posteriorly of a line 
connecting the points of the antorbital processes is greater than in 
pacificus, but less than in frondator (actual measurements: sub- 
auratus, no. 16383, 1.0 mm.; 8988, 0.8. In pacificus, nasals pos- 


449 


Beavers of Western North America 


Taylor 


1916] 


Museum 
number 


16384 
12668 

8869 
16385 
16382 

8987 
12669 


4918 
16383 
8988 
12654 


XX. CRANI 


Sex 


+0 +0 +0 O3 O3 O3 O35 


+0 +0 O03 O3 


Basilar length of Hensel 


92.5 
107.5 
109.7 
114.2 
112. 
112.6 
116.0 


116.2 
120.1 
119.8 
126.3 


94 
103 


*For manner of tak 
yAll from vicinity o 
Ventura County. 
{Length of fronto-m 
point of meeting of fro) 
point of dividers at firs 


FORNIA+ 


§ 

E 

a 

é 
> cr 
5 bp 
8 5 5 
rt aa sect 

2 js 3 
£ | Ze 
5 25 ee 
& og or 
i= vor ‘'S to 
Ci an 
i a8 == 
8 26.7 14.2 
0 22.2 12.1 
1 27.1 10.2 
19 26.9 9.9 
3 26.4 9.0 
5 26.7 9.4 
2 26.7 10.5 
td 25.7 9.6 
2 26.5 9.3 
5 26.0 8.3 
3 27.3 8.0 

paper. 


ined near Sespe, 


of dividers into 
adjust the other 


XIX. COMPARISON OF CRANIAL CHARACTERS OF Castor subauratus subauratus TAYLOR, FROM THE SAN JOAQUIN VALLEY, CALIFOR 
pacificus RHOADS, FROM WASHINGTON STATE, AND WITH Castor canadensis frondator MEARNS, FROM COLORADO AND ‘SAN PED 


Castor subauratus subauwratus 
(nine examples) 
Not so much constricted on the average as 
in pacificus. 
Thicker and heavier than in either sub- 
species. 
Straightest. 
Heavy and blunt. 


Points of Comparison 
Median constriction of interparietal................ 


Hamular processes of pterygoid bones............ 


Ventral contour of hamulars........................-..- 
Median process in interpterygoid fossa.......... 


co 69 nL) i Little inflated. 


Lateral ridge on auditory bullae...................... Strongly developed. 


Poo oy) More prominent than in the other sub- 


species, extending more posteriorly, flat- 
tened instead of knob-like. 
Relation of condylar foramina to occipital 


oo Condyles more pulled out posteriorly; fora- 
mina in plain view on basi-cranial aspect 


of skull in most specimens. 


Castor canadensis pacificus 
(seven examples) 
More constricted, on the average, than in 
subauratus. 
Lighter than in swbauratus. 


Not so straight as in subauratus. 

Light and sharp (that of no, 71812 ap- 
proaching a blunt condition). 

Comparatively much inflated. 


Weakly developed. 


Not so prominent as in subauratus, extend- 
ing more ventrally, less flattened, more 
knob-like. 


Intermediate in condition between swbau- 
ratus and frondator. 


NIA, WITH Castor canadensis 
RO RIVERS, MEXICO 


Castor canadensis frondator 
(two examples) 
Practically intermediate in degree of con- 
striction. 
Lighter than in subawratus. 


Not so straight as in subauratus. 
Light and sharp. 


Intermediate in condition between subau- 
ratus and pacificus, though nearer the 
latter. 

Intermediate in condition between subau- 
ratus and pacificus, though nearer the 
latter. 

Not so prominent as in swbawratus, extend- 
ing more ventrally, less flattened, more 
knob-like. 


Condyles more folded anteriorly; foramina 
not in plain view on basi-cranial aspect 
of skull, as in most examples of swbau- 
ratus. 


448 University of California Publications i 


XVIII. MEASUREMENTS AND Ratios or SCALED P 


All measurements in millimeters, and taken in dry sk 


Museum 
Subspecies— number Length 
Castors; subauratusy =e 16385 298 
Castor s. subauratus . 305 
Castor s. subauratus .- 311 
Castor s. subauratus 310 


but not from frondator, though absolute meas 
tus exceed those in the latter. 

In subauratus the extension of the nasals 
connecting the points of the antorbital proces 
pacificus, but less than in frondator (actua 
auratus, no. 16383, 1.0 mm.; 8988, 0.8. In 


449 


Beavers of Western North America 


Taylor 


1916] 


XX. CRANIAL MEASUREMENTS* OF Castor subauratus subawratus, FROM WESTERN CALIFORNIA} 
Specimens arranged approximately in order of age from top of table to bottom 
(All measurements in millimeters) 


& § 

2 E 

. Sy, C3 

E aoa = 

f ae a) “a 
: : 23S EP See 
@ g Coe) ae 5 7 o§ 
a 2 ae % Bag FI = s oo 
2 2 > © ep 2 2 G =) Ex 
q E 5 b He Seg Se 3 ¢ Se 2S 
3 4 ee : De ke se es. eke sg sh be <8 
4 3 FE 8 3 ‘s 3 S& Bo jee) eens 2 og as ao 
th z 3 = 5 a s bali a 3 5 2 Es a ao 26 
g S z = ra a % lis ee Abe leh 2S ee gy ae 
re & z 3 Se zr 5 Se rae oa 88 =6 2S ES ale 
2 5 S 2 He a a He OSE ae 258 £6 od od oF 
Ges 3 S E a Bi S Ca ae or er PoC OC See ce Recpte A 
number Sex faa] nN’ = fa yr S = HE PE Ba js} eS mes eS fag 
16384 ¢ Opis ian oR wey BAR WW My fal aii GIR) il 2S OS ON ale 
12668 24 07h 865 61.5 224 465 22/6 23:9 (5.0! 13/0) 1913 942 bbb 210) 22280 lai 
gs69 4 1097 S92 G24 246) 4607 23 | 20% ibs7) Tao) ee igs 974s 57.2) 211 oi el Or2 
16885 ¢g@ 114.2 92.5 62. Os fait) OR BO ee ABYSS) aL) RYT) LY) 9.9 
16382 9 1124 927 65.7 25.6 488 251 29:7 6.0 101 183 99:9 59:3 22.3 26:4 9.0 
9987 © dno6 sod l63is 23h) | 508) 24208 SO | 7515) | MelLGy uuiis 2s so 873) eb 8:4 aon 261 9.4 
12669 9 116.0 94.3 668 255  S11° 25.7 310 56 12.2 188 1048 591 22.2 26.7 10.5 
4918 # 1162 982 66.9 264 49.0 264 299 48 11.2 17.5 104.7 615 22.7 25.7 9.6 
16383 4 1201 96.7 66.2 25.7 49.6 266 321 53 11.2 19.3 105.7 61.7 22.2 26.5 9.3 
8988 9 1198 945 69.4 25.6 513 258 311 46 99 18.6 1045 60.9 21.5 26.0 8.3 
12654 9 126.3 103.4 70.5 28.3 54.6 28.2 34.5 3.8 10.1 182 111.3 65.2 22.3 27.3 8.0 


*For manner of taking measurements see Taylor, 1911, pp. 206, 207, and table opp. p. 426 of the present paper. 

+All from vicinity of Grayson, Stanislaus County, except no. 4918, which is supposed to have been obtained near Sespe, 
Ventura County. 

{Length of fronto-maxillary suture: taken in region of anterior root of zygomatic arch; fit one point of dividers into 
point of meeting of frontal, premaxilla and maxilla; follow suture between maxilla and frontal laterad; adjust the other 
point of dividers at first point where maxilla touches jugal. 


450 University of California Publications in Zoology (Vor. 12 


Fig. J. Dorsal view crania of Castor canadensis leucodonta and Castor 
subauratus subauratus. Note the different outline of the nasals 
and interparietal; the different conformation of the temporal 
ridges and the external auditory meatus; and the more massive 
character of the skull of the golden beaver. Approximately 
two-thirds natural size. 

a. Castor canadensis leucodonta, no. 12107, 9, Mus. Vert. Zool.; 
Alberni, Vancouver Island, British Columbia. 

b. Castor subauratus subauratus, no, 12654, 9, Mus. Vert. Zool.; 
Grayson, San Joaquin River, Stanislaus County, California. 


Fig. K. Ventral view crania of Castor canadensis leucodonta and Castor 
subauratus subauratus. Note the different breadth of rostrum; 
the difference in squareness anteriorly of the zygomata; the 
difference in hamulars and median process in the interpterygoid 
fossa: in the mastoids, the paroccipitals, the external auditory 
meatus, and the position of the occipital condyles. Approxi- 
mately two-thirds natural size. 

a. Castor canadensis leucodonta, no. 12107, 9, Mus. Vert. Zool.; 
Alberni, Vancouver Island, British Columbia. 

bd. Castor subauratus subauratus, no. 12654, 9, Mus. Vert. Zool.; 
Grayson, San Joaquin River, Stanislaus County, California. 


452 University of California Publications in Zoology (Vo. 12 


biaa ; 
a 


Fig. L. Posterior view crania of Castor canadensis leucodonta and Castor 
subauratus subauratus.2 Note the different outline of the 
foramina magna; the difference in the external auditory meatus, 
and in general massiveness of crania. Approximately natural size. 

a. Castor canadensis leucodonta, no. 12107, 9, Mus. Vert. Zool.; 
Alberni, Vancouver Island, British Columbia. 

b. Castor subauratus subauratus, no. 12654, 9, Mus. Vert. Zool.; 
Grayson, San Joaquin River, Stanislaus County, California. 


teriorly approximate the line in some specimens, in others fail of 
doing so by several millimeters; frondator, no. 60354, 2.6 mm.; 
35946, 3.3 mm.). 


2In figs. J, K, and L comparison has been made between Castor s. 
subauratus and Castor c. leucodonta instead of Castor c. pacificus. For 
textual comparison of cranial characters of leweodonta and pacificus see 
pp. 442 to 446. They differ in nasal outline, breadth of hamular blades, and, 
as a rule, in the condition of occipital crest. 


1916] Taylor: Beavers of Western North America 453 

Rostra attain greatest breadth in subauwratus (measured outside 
of swellings made laterally by ridge involving maxillary and pre- 
maxillary bones [see fig. M]; in swbauratus, no. 16383, 34.8 mm.; 
8988, 33.5; 12654, 37.8. In pacificus, no. 3672, 34.9 mm.; 71276, 
33.6; 71812, 33.4; 71814, 31.7; 78395, 34.5; 87628, 32.7; 87629, 
34.3. In frondator, no. 60354, 35.5 mm.; 35946, 30.2). 

Hamular processes of pterygoid bones (see 
fig. F', p. 480) are broadest in subawratus 
(subauratus, no. 16383, 4.1 mm., 8988, 4.2, 
12654, 4.3; pacificus, no. 3672, 2.1 
71812, 2.5, 71814, 2.2, 78395, 3.4, 87628, 2.0, 
87629, 2.8; frondator, no. 35946, 2.5 mm.). 


The two juvenals of pacificus from Fisher, 


mm., 


Oregon, measure as follows: no. 136605, 4.3 
mm.; no. 136606, 3.8. 
cranium of pacificus from Keechelus Lake, 
Washington (no. 126190), has hamulars 4.8 
The the 
younger erania of pacificus would seem to 


Fig. M. Dorsal view of 
rostrum, to show 
method of taking 
measurement. Ap- 
proximately one-half 
natural size. 


The young topotype 


mm. broad. broader hamulars in 
indicate that this is a character which is different in individuals of 
different ages (see p. 444). 

Mastoid process on the average is closest to auditory bulla in 
subauratus (see fig: N). Auditory bulla is of somewhat different 
shape in swbauwratus than in pacificus or frondator. 

The following comments are in order respecting certain charac- 
teristics of subauratus as shown by the tables of measurements, 
p. 449 and opposite p. 438: 

(1) The golden beaver is not, in most re- 
spects, on the basis of all the material, larger 
Most 


of the dimensions of no. 12654 of subauratus 


than the canadensis series of beavers. 


Fig. N. Ventral view 
of mastoid region, to 
show position of mas- 
toid process relative 
to auditory bulla. 
About one-half nat- 
ural size. 


(2) 


are greater than those of the largest skulls 
of any of the forms tabulated, however, ex- 
cept some of the individuals of its own sub- 
species shastensis, so that with complete ma- 
terial a size difference between subawratus 
and the canadensis series might be proved. 


Small vertical dimension of the foramen magnum would 


serve immediately to distinguish skulls of swbawratus from those of 


any other western form of beaver, except in its own subspecies shas- 


454 University of California Publications in Zoology Vou. 12 


tensis, and in the following additional instances. Nos. 12668 and 
16384, the youngest examples of swbauratus, have foramina magna of 
larger dimensions than the older crania of that subspecies, and are not 
so characteristically marked off from other species of beavers. Fur- 
thermore, no. 4232, from the Cook Inlet region, has as short a vertical 
diameter of foramen magnum as has subauratus, as have also two 
very young specimens of lewcodonta (nos. 12105, 12106), and one 
specimen of pacificus (no. 71276). 

This measurement, the vertical diameter of the foramen magnum, 
is less, with the above exceptions, than in any other specimen of beaver 
measured. The transverse diameter averages less than in phaeus, 
although it averages about the same as in lewcodonta, pacificus and 
belugae, and greater than in frondator or canadensis from eastern 
Canada. 

The ratio of the vertical diameter of the foramen magnum to the 
basilar length brings out coneretely its different shape. This ratio 
is nearly forty-six percent (taken on the basis of comparison of aver- 
ages derived from table X, opposite p. 488) greater in leucodonta and 
fifty-one per cent greater in frondator than it is in subauratus. It 
will be noted that the ratio decreases with age, the foramen magnum 
in the youngest example (no. 16384) being nearest to the canadensis 
series (see table XX, p. 449). 

(3) The ratio of the width of nasals to basilar length is greater 
in every specimen of subawratus, and in four of the five specimens 
of shastensis, than in any other subspecies of western beaver measured 
(except several specimens of belugae, and three examples of leuco- 
donta). 

(4) The ratio of the length of maxillary tooth-row to basilar 
length averages greater in comparable skulls of swhauratus than in 
pacificus, its nearest neighbor on the north, but practically the same as 
in frondator, its nearest neighbor on the south. 


CASTOR CANADENSIS FRONDATOR MEARNS, FROM THE COLORADO 
AND SAN PEDRO RIVERS, MEXICO 


MATERIAL 
Three specimens from United States National Museum: San 
Pedro River, Mexico, 1 (no. $3%¢$, skin and skull) ; Lagoon of Colo- 
rado River, about 15 miles south of Yuma, Arizona, 2 (nos. 60354, 
60355, skulls only). 


1916] Taylor: Beavers of Western North America 455 


Comparison WitH Castor canadensis leucodonta GRAY, FROM VANCOUVER 
IsLAND, Castor canadensis pacificus RHOADS, FROM WASHINGTON STATE, 
Castor canadensis canadensis KUHL, FROM EASTERN CANADA, AND Castor 
canadensis belugae TAyYLor, FROM Cook INLET REGION, ALASKA 

General external characters 

Specimens compared.—Castor canadensis frondator, no. 20751 ¢, 
U.S. Nat. Mus., from San Pedro River, Sonora, Mexico, October 14, 
1892: Castor c. leucodonta, nos. 12101-12111, Mus. Vert. Zool., all but 
the last (which is from Great Central Lake) from Alberni, Vancouver 
Island, British Columbia, June 11 to August 25, 1910; Castor c. 
canadensis, no. 4858 @, U. 8. Nat. Mus., Moose River, Ontario, Hud- 
son Bay region, May 21, 1860; and nos. 174525 ¢, 174526 ¢, both 
U. S. Nat. Mus., Nepisiquit River, New Brunswick, September 25, 
1911; Castor c. belugae, no. 4347 3, Mus. Vert. Zool., Snug Harbor, 
Alaska Peninsula, Cook Inlet region, June 14, 1904. 

Since lewcodonta is, in general, close to pacificus, and since no 
material of pacificus representative of external characters and com- 
parable with frondator is available, comparison has been made between 
frondator and leucodonta. 


XXI. MEASUREMENTS AND RATIOS Or SCALED PorTIONS or TAILS 


All measurements in millimeters, and taken in dry skins; see fig. H, p. 431 


Ratio 

Museum width to 

Subspecies— number Length Width length 
Castor ec. frondator ...... sa 20D 232 113 48.9 
Castor c. leucodonta 12101 237 120 51.0 
Castor c: leucodonta ..................- 12111 243 108 44.6 
Castor ¢; leucodonta 2-222... 12107 270 124 46.0 
Gastornvenicamadensis) 2s... 174525 265 116 43.4 
Gastorscs canadensis) =... = 174526 260 108 41.3 
Castonsc> canadensis) ==. 4358 223 92 41.1 
Castonces beluga creo cceccceeaas 4347 245 115 47.0 


In general, the pelage is not so heavy-weighted, either in frondator 
or in leucodonta as in canadensis. General coloration in canadensis 
dark; in leucodonta paler; in frondator lightest. Frondator has the 
most uniform coloration. The overhair laterally is, however, of a 
brighter tint than that mid-dorsally, the lightest area being the 
cheeks. The single skin of belugae is darker than frondator, but 
paler than the New Brunswick canadensis. In proportions of tail it 
is closer to frondator than are the examples of canadensis. Propor- 
tions of sealed portion of tail are not clearly separative as regards 


456 University of California Publications in Zoology (Vou. 12 


Trondator, leucodonta, and belugae, at least on the evidence here sub- 
mitted, though canadensis would seem to be separated off by its 
narrow tail. 


General cranial characters 


Crania compared.—Castor canadensis frondator, nos. 60354 2, 
39946 9, U.S. Nat. Mus., from the Colorado River, Mexico, 15 miles 
south of Yuma, Arizona, and from the San Pedro River, Mexico, 
respectively ; C. c. pacificus, nos. 3672, 71276 9, 71812 ¢, 71814 9, 
87628 2, 87629 ¢, 126190 ¢, U.S. Nat. Mus., all from the mainland 
of Washington State; C. c. canadensis, nos. 174525 ¢ and 174526 9, 
U.S. Nat. Mus., Nepisiquit River, New Brunswick; C. c. belugae, nos. 
4347 ¢, 4225 36,4224 3, Mus. Vert. Zool., Cook Inlet region, Alaska. 

Nasals of different outline in frondator than in pacificus. Their 
lateral outlines converge more regularly in a posterior direction, there 
being no tendency to be parallel, as is the case in comparable skulls 
of pacificus. The ratio of width of nasals to length tends to be 
greater in frondator than in pacificus, though there is decided over- 
lapping. Measurements of foramen magnum similar, though most 
specimens of pacificus have their foramina broader in proportion to 
vertical diameter than is the case in frondator. Teeth in one specimen 
of frondator flare more laterally than in any specimen of pacificus 
(see fig. I, p. 442, frondator, no. 60354, 34.3 mm., 35946, 30.9; 
pacificus, no. 3672, 33.1 mm., 71276, 31.3, 71812, 30.4, 71814, 34.1, 
78395, 32.5, 87628, 32.6, 87629, 34.3). Ratio of maxillary tooth-row 
to basilar length greater in frondator. Tooth-row longer in frondator 
no. 60354 than in pacificus no. 71812. These specimens have basilar 
length identical. Tooth-row longer in frondator no. 35946, with 
basilar length of 111.7 mm., than in pacificus no. 71812, with basilar 
length of 118.7. 

Median process in interpterygoid fossa longer and sharper in- 
frondator than in pacificus. 

These differences between frondator and pacificus are for the most 
part slight. The outline of nasals is the most dependable single sep- 
arative character. 

Measurements fail to separate frondator either from canadensis or 
the Cook Inlet race. Distance from the inion to dorsal outline of 
foramen magnum less in frondator than in canadensis from New 
Brunswick (frondator, no. 60354, 18.9 mm., 35946, 18.0; canadensis, 
no. 174525, 20.9 mm., 174526, 19.9). Ratio of maxillary tooth-row 
to basilar length slightly greater in frondator than in the New Bruns- 


f. 


XXIL. 


Points of Comparison 
General coloration ...............-.-.-.:.00++- 
Dorsal overhair 


Wrsed| WG e il) 
o 

DTA, 2 

DO) UE 21 ees ae a On 


WENUPEOVEDOAMT w...---.---nnneacnnnn-nnnno F 


Vr Ail tro Gy qb 


DLC C2, [Cs | a a rrr 


COMPARISON OF EXTERNAL CHARACTERS OF Castor canadensis 
leucodonta GRAY, FROM VANCOUVER ISLAND, BRITISH COLU 


Castor canadensis frondator 
(one example) 
Lightest of the three. 
Varying about cinnamon. 


Paler tints of the drab series than are 
found in either leucodonta or canadensis, 
near cinnamon-drab, light cinnamon- 
drab, and light drab. 

Mixture of colors which in combination ap- 
proach clay color and hazel. 


Mixture of colors which approach clay 
color and chestnut-brown; the chestnut- 
brown more in evidence making foot ap- 
pear darker. : 

Close to cinnamon-buff. 


Paler than frondator, approaching pale ecru- 
drab and pale drab-gray. 


Practically undifferentiated, absent, though 
the overhairs on throat and chin are paler 
than those posteriorly, being colored near 
pinkish buff. 

A fringe of hairs at immediate base of tail 
colored near hazel and chestnut-brown. 


Broad area at immediate base of tail near 
hazel and chestnut-brown in coloration. 


Broader proportionally than the average of 
leucodonta; much broader than canadensis. 


frondator MEARNS, FROM THE SAN PEDRO 
MBIA, AND WITH Castor canadensis canadensis 


Castor canadensis leucodonta 
(six examples) 

Darker than in frondator. 

Varying about cinnamon-buff; no. 12108 
with liberal insprinkling of seal brown 
hairs. a 

Darker than in frondator. Varying from 
fuscous and fuscous-black to benzo and 
hair brown. 


Light brownish drab to near russet or mars 
brown. 


Paler brown than in canadensis, varying 
from near carob brown to near hazel. 


Varying about hair brown. 


Darker than in frondator, approaching 
light drab and light cinnamon-drab. 


Inconspicuous; the few hairs present vary 
about warm buff. 


Varies from near hair brown (no. 12111) 
to a shade between chocolate and bay (no. 
12107); fringe of hairs near dark vinace- 
ous-drab in several specimens (notably 
nos. 12108, 12101, 12102). 

Between cinnamon-brown and chestnut or 
bay. 


Broader proportionally than canadensis, 
narrower on the average than frondator. 


RIVER, MEXICO, WITH Castor canadensis 
KUHL, FROM EASTERN CANADA 


Castor canadensis canadensis 
(two examples) 

Darker than in leucodonta. 

Hairs a mixture of blackish and ochraceous- 
tawny (nos. 174525, 174526, former the 
darker); general effect near chocolate or 
bay. 

Near clove brown (nos. 174525, 174526). 


General impression pale brownish drab 
(no. 174525); approaching warm blackish 
brown (no. 174526). 

Near dark vinaceous-drab (no. 174525); 
similar but with tendency to be lighter 
(no. 174526). 


Bone brown, dark grayish brown, or dusky 
drab (no. 174525); a trifle paler, near 
dark vinaceous-drab or natal brown (no. 
174526). 

Shaft of hair pale, nearest pale gull gray, 
tipped with drab-gray (mos. 174525, 
174526). 

Definite throat band present, the scattered 
paler hairs being near cinnamon-buff (no. 
174525); nearer ochraceous-buff (no. 
174526). 

Cinnamon-drab to verona brown, hairs at 
immediate base of tail tipped with seal 
brown (no. 174525); chestnut to bay, 
hairs at immediate base of tail tipped 
with seal brown (no. 174526). 

Mixture of chestnut and cinnamon-drab, ap- 
proaching pinkish buff laterally and an- 
teriorly (no. 174525); liver or carob 
brown approaching orange-cinnamon lat- 
erally and anteriorly (no. 174526). 

Narrowest proportionally of the subspecies 
here compared. 


avd) [endo 


1916] Taylor: Beavers of Western North America 457 


wick examples. Zygomata not quite so square anteriorly in frondator 
as in specimens of belugae from Cook Inlet. Also distance from 
dorsal outline of foramen magnum to inion less (see fig. G, p. 431; 
frondator, no. 35946, 18.0 mm., 60354, 18.9; belugae, no. 4347, 20.5 
mm., 4225, 22.4, 4224, 22.7) ; teeth narrower on average (see table of 
measurements, p. 428). 

Nasal outline, condition of median process in the interpterygoid 
fossa, and certain skull dimensions, show the affinities of frondator 
to be with canadensis rather than with either swbawratus or pacificus. 


OUTLINE OF THE HISTORY OF THE BEAVERS 


The Castoridae and Castoroididae are probably derived from that 
primitive Eocene rodent stock represented in the family Ischy- 
romyidae. Matthew (1910, p. 67) has outlined their possible rela- 
tionships to this family, postulating their descent from the North 
American Paramys, an ancient squirrel-like form which with its con- 
temporary relatives was perhaps ancestral to most, possibly to all, 
the members of the great order Rodentia. It has so far been impos- 
sible definitely to work out whether Plesrarctomys from the French 
Upper Eocene is congeneric with Paramys or distinct from the Ameri- 
can genus. 

According to Matthew’s scheme, Paramys gave rise to Sciuravus 
of the North American Eocene, which in its turn is tentatively placed 
as the ancestor, on the one hand, of the stock which gave rise to 
Eutypomys of North American Middle Oligocene, and on the other, 
of that from which the important genus Steneofiber is derived. These 
two genera are the earliest which are referred by Osborn (1910, 
p. 535) to the Castoridae. 

Concerning Eutypomys very little is known. Osborn refers to it 
as a ground-squirrel or semi-cursorial type, and suggests that it may 
be related to the Pteromyinae. It seems to be of little importance 
to the phylogeny of beavers. On the contrary, Steneofiber assumes 
a place of great importance in this phylogeny, being the supposed 
ancestor of most of the later genera belonging to the beaver family. 

Steneofiber, which was about the size of a marmot, appears first 
in the Middle Oligocene (Stampian) of Europe, also throughout the 
Upper Oligocene of North America, being abundant and characteristic 
in the Upper John Day of Oregon. Evidently it became extinct soon 
after, for it has not been discovered in later deposits. At least by 


458 University of California Publications in Zoology (Vou. 12 


Upper Oligocene, it should be remarked, Steneofiber had attained a 
world-wide distribution. It was replaced, in the Lower and Middle 
Miocene of Europe, by Chalicomys, which died out during the Plio- 
cene. Matthew derived the phylogenetic sideline represented by 
Hystricops of the North American Miocene and Pliocene from Steneo- 
fiber. 

Little is known concerning the genus Huhapsis, a supposed cas- 
torid genus which occurred in North America during Upper Oligo- 
cene. 

Eucastor or Dipoides of Upper Miocene, as well as the genus 
Castor itself, which first appeared in that period, are probably also 
descended from Steneofiber. Hucastor, which was, like Steneofiber, 
about the size of a marmot, is first found in the North American 
Hipparion and Procamelus zones (Upper Miocene) and ranges into 
the Pliocene. Sigmogomphius (Merriam, J. C., 1896, p. 365) from 
the Pliocene near Berkeley, California, and Dipoides, from the Plio- 
cene of Asia, are closely related to if not congeneric with it. Appar- 
ently the Hucastor-Dipoides stock appeared first in North America, 
spreading later into Eurasia. The Hucastor line soon ran out in 
the Old World, while in North America it appears to have given rise 
to the Pleistocene family Castoroididae with its characteristic genus 
Castoroides. 

The earliest species of the European genus T'rogontherium ap- 
peared in the Upper Pliocene, being there represented by teeth much 
smaller than those of its suecessor in the Pleistocene, Trogontherium 
cuviert, the giant beaver of Europe, which was about one-fifth larger 
than the beaver of modern times. This genus coexisted in Europe 
with Castor, but died out at the close of what Osborn calls the First 
Faunal Subzone of the Second Pleistocene Fauna. 

The largest species of the Pleistocene Castorides of North America 
attained the size of a black bear, and was, like the big Huropean 
Trogontherium, preceded by a smaller form. 

The genus Castor, on the basis of evidence now at hand, appeared 
somewhat earlier in Europe than in America. The Pontian Upper 
Miocene deposits are the first in Europe in which Castor has been 
discovered. The recent discovery (Kellogg, 1911, p. 401) of a beaver- 
tooth in the Pliocene of California seems to indicate that it soon 
attained a distribution embracing both continents. Castor is found 
in Pliocene and Pleistocene European formations, and is widespread 
in North American Pleistocene. It is lacking from the Pliocene and 


1916] Taylor: Beavers of Western North America 459 


all earlier formations of the latter continent except for the one 
instance mentioned above, the correlation of the formation in which 
the tooth was found being still somewhat uncertain. 

If there has not been independent origin of Steneofiber, Eucastor- 
Dipoides, and Castor itself on the Eurasian and North American 
continents, it is clear that there have been several intercontinental 
migrations of beavers. Although we are not able to state exactly 
the number of these migrations, the evidence indicates that there 
were at least three. 

While it is realized that negative evidence is likely to prove 
untrustworthy, and that possible errors in correlation introduce a 
further element of uncertainty into general statements as to the 
origin and migration of the castorids, insofar at least as these are 
based on relative time of appearance, it is believed that the following 
tentative propositions merit consideration. 

Steneofiber probably developed first in the Old World, since it is 
found in Europe in Middle Oligocene while it does not appear in 
North America until the Upper Oligocene. By late Oligocene time, 
therefore, its migration had apparently carried it into North America. 

The fact that the John Day epoch of the Oligocene is marked by 
the disappearance of almost all the European migrants which are so 
characteristic of the earlier White River fauna, with other evidence, 
has seemed to show that the land connection with the Old World was 
broken (Scott, 1913, p. 116). The case of Steneofiber might be taken 
to indicate, though such an isolated instance is probably not worth 
much, that the White River land connection was maintained into 
the early part of the John Day phase of the Upper Oligocene. It is 
more probable that Steneofiber crossed the connecting land bridge 
during White River time, but did not attain a widespread distribu- 
tion until the John Day epoch. 

The ELucastor-Dipoides stock would seem to have taken origin in 
North America, since it is first found in the Upper Miocene of that 
continent. By Pliocene time it had migrated at least into Asia. It 
seems certain that Bering Strait was closed during at least a great 
part of that epoch (Scott, 1913, p. 125). 

The genus Castor probably arose in Kurasia from Steneofiber or 
a closely related stock. A wave of migration carried it into North 
America during the Pliocene, probably by way of the North-Pacific 
land-connection, and its remains became widely distributed and fairly 
abundant in the Pleistocene. 


460 University of California Publications in Zoology (Vou. 12 


SumMMaArRyY OF RELATIONSHIPS OF CERTAIN NorTH AMERICAN 


BEAVERS 
SOME DIFFICULTIES TO PRECISE STATEMENT OF RELATIONSHIP 


Gregory (1910, p. 105) has called attention to the stumbling-blocks 
of the phylogenist, namely the two difficulties, first, of distinguishing 
between primitive and specialized characters, and second, between 
resemblances significant of genetic relationship and those indicating 
only convergence or parallelism. 

That parallelism in evolution may be a principle more widespread 
and of greater significance than is ordinarily accorded it by taxono- 
mists has recently been suggested (see Scott, 1913, pp. 649-656, and 
Hopkins, 1914, p. 187). It should be clearly recognized, however, 
that the weight of evidence indicates (Scott, 1913, p. 137) that inde- 
pendent origin of closely similar forms from different stocks in widely 
separated localities is practically unknown. 

Emphasis should also be laid on the importance: (1) of deter- 
mination of the order of appearance of diagnostic characters; and 
(2) of understanding the general adaptive significance of such char- 
acters, so that their broader phylogenetic and systematic value can be 
appraised (Gregory, 1910, p. 112). We are not now in position to 
prove in what order the diagnostic characters in the family here con- 
sidered have appeared, or to state exactly the adaptive significance or 
insignificance of the characters. The fact that it is difficult to ascribe 
any adaptive value whatever to many of the characters which are 
diagnostic between subspecies and species of beavers does, however, 
suggest that these particular characters may be dependable as indi- 
cating relationship. Examples of such characters are: Different out- 
lines of tail; different outlines of nasal bones; different degrees of 
development of median process of interpterygoid fossa; different 
shapes of foramen magnum; different breadths of hamular processes 
of pterygoids; different widths of bony palate anteriorly. 

There are possible further difficulties. Since we do not certainly 
know that the evolution of one form has not taken place somewhat 
more rapidly than that of another, estimates of time of isolation (or 
age of a particular form) and closeness of relationship should only 
cautiously be based on degrees of difference. 


CONSIDERATION OF CERTAIN NORTH AMERICAN FORMS 


Certain problems remain to be considered regarding the relation- 


1916] Taylor: Beavers of Western North America 461 


ships of beavers, some of these being: (1) the relation of the beaver 
of North America to that of Europe and Asia, and (2) the interrela- 
tions of the North American forms. Under the second head arises 
the question whether groupings of the North American forms are 
possible, and since groupings are possible, further problems arise as 
to their interrelationships, their ancestry, and the causes and condi- 
tions in their differentiation. 

The relationships of the American and Eurasian beavers have 
been more or less fully discussed by the following authors: Geo. 
Cuvier (1817, tome 1, p. 191, not seen; and 1825, tome 5, p. 57, not 
seen) ; Fred. Cuvier (1825, liv. 51, not seen) ; Brandt and Ratzeburg 
(1827, pp. 12-30); Brandt (1855, pp. 43-66); Morgan (1868, pp. 
42-45); Ely (in Morgan, 1868, pp. 288-306) ; and Allen (1877, pp. 
437-445). While sufficient material to justify a review of this point 
has not been available, the present writer has been able to compare 
three skulls of Castor fiber from the Elbe River, Germany, with the 
American beavers, and believes that the evidence indicates that they 
are specifically distinct. 

Several points of possible significance arise in connection with 
the problem of the interrelationships of the North American beavers. 
Material illustrative of two species of Nearctic beavers, as well as 
of all their subspecies but two (Castor canadensis carolinensis Rhoads 
and Castor canadensis mexicanus Bailey), has been available in this 
study. Even a cursory examination of this material shows that all 
those beavers listed under canadensis, namely subspecies canadensis, 
michiganensis, belugae, leucodonta, pacificus, frondator and texensis, 
fall into one group, while those listed under subauratus, namely 
subspecies subauratus and shastensis, fall into another. Within these 
groups the subspecies appear to be closely related, their differentia- 
tion apparently having progressed to a slight extent only. From 
their geographical situation and from a consideration of such of their 
characters as may be worked out from the literature, species caecator 
Bangs and subspecies mexicanus Bailey and carolinensis Rhoads 
should be referred to the canadensis group. 

The members of the canadensis group are, on the basis of available 
material, unequally related. For example, Castor canadensis phaeus, 
from Admiralty Island, Alaska, is less closely related to Castor c. 
canadensis of eastern Canada than is Castor c. belugae of the opposing 
mainland. Phaeus is more sharply marked off from the other sub- 
species of canadensis than are Castor c. frondator or belugae. Phaeus 


462 University of California Publications in Zoology (Vou. 12 


cannot be said, on grounds of compared characteristics, to be closer to 
belugae than it is to eastern canadensis. Belugae is more closely 
allied to canadensis of eastern Canada than is either Castor c. leuco- 
donta or Castor c. pacificus. It is impossible to decide to which of 
the two mainland subspecies (belugae or pacificus) leucodonta is most 
closely related. Frondator is more closely allied to canadensis of 
eastern Canada than it is to the California species of beavers or to 
the belugae-leucodonta-pactficus series. Castor c. michiganensis finds 
its closest relative in canadensis. 

The California beavers stand by themselves, having undergone 
considerably more divergence than the other subspecies, and their 
immediate ancestry is decidedly uncertain. Mr. F. H. Holden, of 
the staff of the Museum of Vertebrate Zoology of the University of 
California, who has carefully compared such skeletons of beavers as 
are available, has called the writer’s attention to the fact that the 
majority of skeletal characters of Castor subauratus subauwratus would 
relate it rather to the Eurasiatic Castor fiber than to the North Ameri- 
ean Castor canadensis. This testimony, however, is not borne out by 
the skull characteristics, which would seem definitely to relate subau- 
ratus to the canadensis series. Perhaps the most striking differences 
between the swbauratus series and the canadensis series are the dif- 
ferent shapes of the foramen magnum and of the median process in 
the interpterygoid fossa. Castor c. pacificus and Castor c. belugae 
are somewhat more variable respecting these characters than are any 
other subspecies of canadensis, and individual skulls of these show a 
more decided bent in the direction of swbawratus than do the other 
forms, which suggests the possibility that swbawratus has been derived 
from the form which was ancestral also to the subspecies occupying 
the mainland of Oregon, Washington, British Columbia and Alaska. 
If this should be the ease, the relationship of the California forms with 
canadensis would be close, and the comparatively large amount of 
divergence would be noteworthy as possibly indicating a more rapid 
evolution, or a more complete isolation from the parent stock (possi- 
bly both together) than is exemplified by the other forms. 


REMARKS ON ISOLATION AND ITs RELATION TO SPECIATION 
CONSIDERATION OF SOME OF THE EVIDENCE 
EVIDENCE FROM THE STUDY OF BEAVERS 


We may now profitably undertake a review of some of the facts 


1916] Taylor: Beavers of Western North America 463 


of beaver distribution, with special reference to the problem of specia- 
tion in the group. 

It should be remembered that the beavers of Eurasia and America, 
while not distantly related, are still very clearly marked off spe- 
cifically. It should further be called to mind that there are, on the 
North American continent, two distinct groups of beavers, the cana- 
densis group, including the species canadensis with at least its sub- 
species canadensis, belugae, leucodonta, pacificus, frondator and tex- 
ensis, and the subauratus group, including the species suwbauwratus 
with its subspecies swhauratus and shastensis. 

In the case of each one of these subspecies of beavers, geographic 
range seems to be as characteristic as any physical attribute. It 
should be emphasized that differentiation in beavers has progressed 
only to a slight degree as compared with, for example, the Geomyidae, 
and that all the forms of beavers, even the two groups, are relatively 
closely related. No case of overlapping of subspecific or specific 
ranges is known. Knowledge of the details of the relations of dis- 
tribution of beavers to topography is not sufficient to warrant cate- 
gorical assertions without some qualification, but available evidence 
points to the truth of the following general statements. 

Castor c. michiganensis, a small, dark form, is very closely related 
to @. c. canadensis, its neighbor on the north. The type locality of 
canadensis is Hudson Bay, and the subspecies probably ranges south 
to the chain of Great Lakes, which would seem to be a barrier separat- 
ing it from michiganensis. 

The type locality of Castor c. canadensis (Hudson Bay) is con- 
nected by a nearly continuous series of streams and lakes with extreme 
western Canada. Distance is practically the only deterrence to the 
migration of individuals from eastern Canada to the base of the 
Canadian Rockies and the Alaskan Mountains. Subspecies belugae, 
of canadensis, occupying the Pacific territory from Cook Inlet to 
British Columbia, is closely related to canadensis on the one hand and 
to leucodonta and pacificus on the other. Subspecies phaeus of Ad- 
miralty Island, southeastern Alaska, is comparatively sharply marked 
off. Frondator, type locality San Pedro River, Mexico, which is sup- 
posed to range some distance to the north, is also closely related to 
canadensis. On the other hand, subauratus and its subspecies shas- 
tensis are much less closely related to canadensis than any of the other 
forms, and there do oceur, bounding the range of the subawratus 
group, masses of high mountains on the north, east and south, as 


464 University of California Publications in Zoology (Vou. 12 


well as broad deserts on the east and south. The two subspecies of 
subauratus, namely subawratus and shastensis, are much more closely 
related to each other than is either to any other known beaver in the 
world. Although they are inhabitants of the same hydrographic 
basin, they are efficiently separated by distance in combination with 
the ‘‘narrows’’, by way of which the Pit River cuts through the 
Sierra Nevada Mountains. 

It must be immediately apparent that these facts, so far as they 
go, fulfill the requirements of Wagner and Jordan’s law: ‘‘Given any 
species In any region, the nearest related species is not likely to be 
found in the same region nor in a remote region, but in a neighboring 
district separated from the first by a barrier of some sort.”’ 

It is now quite well established that on continuous land areas 
temperature is the most efficient of all barriers, with humidity of the 
atmosphere a close second. It is a remarkable fact that the genus 
Castor ranges, undergoing at the same time but little change, through 
all the life-zones (based on temperature) from the Hudsonian of the 
northern limit of trees to the Lower Sonoran of the southern deserts, 
and that it is found in faunal areas (based on humidity) as widely 
different as the Sitkan district of southeastern Alaska and the Colo- 
rado Desert of the southwestern United States. Although the semi- 
aquatic environment of the beaver is doubtless more uniform through- 
out this great range of temperature and moisture conditions than is 
the typical terrestrial environment, it must still be conceded that the 
genus Castor is subjected to very different environments in different 
parts of its range. The writer at this moment finds it impossible to 
assign any adaptive significance to the subspecific and specific differ- 
entiatory cranial characters of beavers. It must be confessed that 
the maintenance by the beaver of its chief characteristics through a 
very wide range of environmental conditions, coupled with the further 
fact that it is difficult to attribute any adaptive significance to the 
cranial specific and subspecific characters, invite one to the hypothesis 
that these characteristics of the different races of beaver are due to a 
cumulation of what are for the most part inutile characters through 
the fact of the geographical isolation, alone, of the various beaver 
stocks. 

An alternative view is, of course, that our inability to see the 
adaptive significance of these differentiatory characters, or definitely 
to correlate them with characters which are obviously adaptive, merely 
testifies to the limitations of our own knowledge, and not at all that 


1916] Taylor: Beavers of Western North America 465 


these characters are really not adaptive or not correlated with some 
adaptation. However this may be, it would seem that the possibility 
that geographical isolation alone, with no assistance from natural 
selection, has been a condition in speciation of beavers, is by no means 
excluded. 


EVIDENCE FROM CERTAIN OTHER FAMILIES OF MAMMALS 

Its great geographical extent and wide diversity of topographical 
and environmental conditions make California probably as favorable a 
geographic unit as could be found for the study of problems concern- 
ing the origin and maintenance of vertebrate species; and since the 
writer is more familiar with the mammalian fauna of California than 
with that of any other area, this particular field has furnished most 
of the material used. But published facts from mammalian distribu- 
tion in the Great West, and in the continent generally, have been 
Treely drawn upon. 

In cases where the family or genus has recently been monographed, 
relationships as outlined may be regarded as more dependable than 
in those instances in which the group has not undergone adequate 
revision. In the latter the conclusions reached are tentative, and 
are based upon the writer’s familiarity with the mammalian fauna 
in question. 


SorIcIDAE 


Sorex vagrans vagrans, the range of which within California 
(Grinnell, 1913a, p. 270) includes the Upper Sonoran, Transition and 
Boreal zones in the western portion of the state, east to Shasta County 
and south as far as Monterey, finds in Sorex vagrans amoenus its 
nearest relative within the state. The range of the latter form takes 
in the Transition and Boreal zones of the Sierra Nevada Mountains, 
at least from Mono County north to Mount Shasta (Merriam, C. H., 
1895, p. 68, and 1899, p. 87). Sorex halicoetes, of the salt marshes 
bordering the south arm of San Francisco Bay, is more closely allied 
to Sorex vagrans vagrans than it is to any other species of Sorex 
(Grinnell, 1913a, p. 184). It will be remembered that Sorex vagrans 
vagrans is found coastwise in California as far south as Monterey. 

Sorex sinuosus of the brackish marshes of Grizzly Island, Suisun 
Bay, California, is most closely related to Sorex californicus of the 
neighboring uplands. 

Note should be made of the distribution of the tenellus group of 


466 University of California Publications in Zoology (Vor. 12 


subspecies. Sorex tenellus tenellus occurs on the Alabama Hills, 
near Lone Pine, Inyo County, California. Sorex tenellus lyelli is a 
closely related form occurring on Mount Lyell in the Sierra Nevada, 
and Sorex tenellus myops is known only from the White Mountains 
of Inyo County, California, which lie to the eastward of Mount Lyell 
near the California boundary line. Sorex tenellus nanus is an out- 
lying relative found in the Rocky Mountains of Colorado. 

The widely distributed Sorex montereyensis montereyensis, which 
is found in the Transition and Upper Sonoran zones of the northern 
and central coast districts of California from the Oregon line to San 
Luis Obispo County, has as its nearest relative Sorex montereyensis 
martposae, which is also broadly distributed, being a montane form 
found in the Transition zone of the Sierra Nevada and Warner Moun- 
tains, and in the inner coast ranges as far south as Lierly’s Ranch, 
four miles south of Sanhedrin Mountain, Mendocino County. 

Sorex obscurus obscurus occurs in the Transition and Boreal zones 
of the Sierra Nevada Mountains from Mono County north. This is a 
subspecies of very broad distribution, being found in southern British 
Columbia and northern Washington and in the Rocky Mountains, as 
well as in the Sierra Nevada (Merriam, 1895, p. 73). Its closest 
allies are Sorex obscurus longicauda, occupying a narrow strip along 
the Pacific coast from the mouth of the Columbia River to Wrangel, 
Alaska, and Sorex obscurus ventralis, inhabiting the mountains of 
Oaxaca, in southern Mexico. 

Neosorex bendirw bendirvi, occupying in California the Transition 
and Boreal zones of the humid northwest coast belt from Humboldt 
Bay to Gualala, is also found in the Klamath Basin, Oregon, and 
thence northward along the east side of the Cascade Range to Puget 
Sound (Merriam, 1895, p. 96). Its closest allies are Neosorex bendiru 
palmeri, of the coast of Oregon and the Willamette Valley, and Neo- 
sorex bendirvi albiventer, from Lake Cushman, Washington. 

This terminates the list of the California Soricidae, the relation- 
ships of which are best known. Concerning Sorex shastensis from 
Mount Shasta, Sorex pacificus from the north coast district of Cali- 
fornia, and the Sorex ornatus series from Mount Pinos and other 
mountains of southern California, our knowledge is indefinite. 


PROCYONIDAE 


The Pacific raccoon, Procyon psora pacifica, type locality Keechelus 
Lake, Cascade Mountains, Kittitas County, Washington, ranges into 


1916] Taylor: Beavers of Western North America 467 


California on the north, occupying the Transition and Upper Sonoran 
zones along its northern border. The vicinity of Pit River, Shasta 
County, is its southern limit. Occupying both Lower and Upper 
Sonoran and Transition of the rest of the state except the southeastern 
deserts, is Procyon psora psora, the California raccoon, which is very 
closely related to its northern neighbor, and doubtless intergrades 
with it. Favorable situations in the southeastern desert region are 
inhabited by the pallid raccoon, Procyon pallidus, while a fourth 
California form has recently been described from the San Diegan 
region west of the Coast Range Mountains in extreme southwestern 
California (Procyon psora californicus, Mearns, 1914, p. 66). 


MUSTELIDAE 


The closest ally of Martes cawrina caurina, the pine marten, the 
range of which includes the Transition and Boreal of northwestern 
California, is Martes caurina origenes which is found in the Boreal of 
the Rocky Mountains of Colorado (Cary, 1911, p. 189). Martes pen- 
nanti pacifica, the Pacifie fisher, finds its closest living relative in 
Martes pennanti pennanti of eastern Canada. The Sierra Nevada 
and Mount McKinley wolverines, Gulo luscus luteus and Gulo luscus 
hylaeus, are closely related, and both are close to the Gulo luscus 
luscus of Canada. 

Details of distribution and relationships of the weasels are hardly 
complete enough to be satisfactory. However, it is certain that the 
diminutive Mustela muricus of the Sierra Nevada is a member of the 
boreal cicognanti group of weasels. <A close ally of muricus is resi- 
dent on the Pine Forest Mountains of northern Nevada. An unde. 
seribed weasel most closely related to Mustela streatori of the Puget 
Sound district occurs on the north coast of California. The Mustela 
arizonensis is said (Merriam, 1896, p. 22) to be a mountain form of 
Mustela longicauda of the Great Plains from Kansas northward. 
Mustela arizonensis is found in the Transition and Boreal zones of the 
Sierra Nevada from Mount Shasta to Tulare County, as well as on 
the San Jacinto Mountains. Its distribution is stated as follows: 
‘Broadly, the Sierra Nevada and Rocky mountain systems, reaching 
British Columbia in the Rocky Mountain region, but not known north 
of the Siskiyou Mountains in the Sierra-Cascade system’’. North of 
the Siskiyou Mountains in the Sierra-Cascade system occurs its close 
relative, Mustela saturata. The relationships of arizonensis are not 
clear, since Merriam states (op. cit., p. 23) that Sierra specimens 


468 Unversity of California Publications in Zoology (Vou. 12 


show a strong tendency to grade into wanthogenys, the form of the 
California lowlands. 

The closest relatives of Mustela xanthogenys canthogenys of Cali- 
fornia west of the desert divides, from the Mexican line north through 
the San Diegan district, and in west-central California east of north- 
ern humid coast belt, at least to the head of the Sacramento Valley, 
are Mustela xanthogenys munda (doubtfully recognizable). of the 
humid coast Transition north of San Francisco Bay, and Mustela 
zanthogenys oregonensis, of Rogue River Valley, Oregon. 

The Pacific Mink, Mustela vison energumenos, is most closely 
related to its neighbors, Mustela vison nesolestes, of the Alexander 
Archipelago, Alaska, and Mustela vison lacustris of the interior of 
Canada and the northern United States (Hollister, 1913, p. 476). 

Turning to the genus Spilogale, it is noted that two closely related 
forms invade eastern California but do not cross the Sierra Nevada. 
These are Spilogale gracilis saxatilis, found in the northeastern corner 
of the state, and Spilogale gracilis gracilis, inhabiting the Sonoran 
zones of the Inyo region. Spilogale arizonae arizonae of the lower 
Colorado River region is most closely allied to Spilogale arizonae 
martirensis of Lower California. Spilogale phenax phenazx, inhabit- 
ing most of the Pacific slope of California, is apparently replaced by 
Spilogale phenax latifrons in the coast region of extreme northern 
California and Oregon. Another race of the species phenax, Spilo- 
gale phenax olympica, is found on the Olympic Peninsula and the 
shores of Puget Sound (Miller, 1912, p. 106). 

The genus Mephitis is represented by five forms within California, 
three of these belonging to the species occidentalis. This species is 
represented in western North America by five races (Howell, 1901, 
pp. 34-38). Mephitis occidentalis occidentalis is found in northern 
and central California, west of the Sierra-Cascade system, from Mon- 
terey Bay to the Willamette Valley, Oregon. The coast region of 
Washington and northern Oregon, as well as the shores of Puget 
Sound, are occupied by Mephitis occidentalis spissigrada. In the 
eastern part of southern Washington and northern Oregon, on the 
desert side of the Cascade Mountains, is found Mephitis occidentalis 
notata, which intergrades with occidentalis and major. Mephitis 
occidentalis major is found in eastern Oregon, northeastern California 
and Nevada, ranging east to the Wasatch range in Utah. Its closest 
relatives are notata and occidentalis, its neighbors on the north and 
west respectively. In southern California the group is represented by 


1916] Taylor: Beavers of Western North America 469 


Mephitis occidentalis holzneri. The case of Mephitis platyrhina from 
the south fork of the Kern River and Owens Valley, California, is of 
interest and possible significance. Its range (Howell, op. cit., p. 39) 
is overlapped by that of Mephitis occidentalis holznert. If Howell’s 
suggestion that platyrhina may intergrade with major be true, then 
the latter, occupying territory to the north, is the closest relative of 
platyrhina. 

One other Mephitis, namely Mephitis estor, is found in California. 
This is probably an invader from the east, its geographic distribution 
including Arizona, western New Mexico, Sonora, Chihuahua, and 
northern Lower California, as well as the valley of the lower Colorado 
River in California. Although Mephitis estor is a very distinct 
species, its closest relative would seem to be Mephitis mesomelas 
varians, whose range lies to the east. Howell (1901, p. 33) goes so 
far as to suggest that it is possible that intergradation takes place 
between the two. 

The California badger, Taxidea taxus neglecta, unquestionably 
intergrades with and is most closely related to Taxidea taxus taxus of 
eastern America. Specimens of badgers from the Pine Forest Moun- 
tain region of northern Nevada have been referred to the latter 
(Taylor, 1911, pp. 296, 297). Im the arid southwestern United 
States and northern Mexico is found the Mexican badger, Tazxidea 
taxus berlandiert. The fourth form is Taxidea taxus infusca, found 
in Lower California. 

The aquatic genus Latax, which occurs off the west coast of Amer- 
ica and the northeast coast of Asia, finds in the terrestrial and 
fluviatile genus Lutra, which is found both in Asia and North Amer- 
ica, its closest living relative (Taylor, 1914, pp. 498, 495). 


MuriIDAE 


The Muridae are represented in California by so large a number 
of groups and species that details may justifiably be omitted. This 
great family must not be passed over, however, without calling atten- 
tion to the interesting data presented by the Peromyscus maniculatus 
series of mice. This group is represented by forty-three subspecies 
or small species distributed nearly throughout the North American 
continent. 


This was the first to be named and almost the last to be recognized of a 
large group of inosculating forms—the largest and most remarkable of the 
genus and perhaps of American mammals. Its distribution is wider and the 


470 University of California Publications in Zoology  [Vou.12 


number of intergrading forms and of individuals is greater than in any 
similar group of mammals known. From typical maniculatus, development 
may be traced step by step absolutely without break through all the numerous 
subspecies (Osgood, 1909, p. 41, and frontispiece). 

Here we have a spectacular illustration of the principle that the 
closest relative of any form is found, not in a locality far distant nor 
in a locality identical with that occupied by the form, but in a neigh- 
boring region separated from the first by a barrier of some sort 
(see p. 464). 

Respecting the Californian forms of the genus Onychomys it may 
with all assurance be said that no two species are found in one 
locality. This genus has recently been revised by Hollister (1914, 
pp. 427-489). 

The distribution of the Murid genera Reithrodontomys, Sigmodon, 
Neotoma, Evotomys, Microtus, and Fiber, as well as of the genus 
Peromyscus, afford a mass of material similar to that which has 
already been adduced. 


ZAPODIDAE 


Zapus trinotatus trinotatus ranges from the coast region of south- 
ern British Columbia, Washington (including Cascades), Oregon 
(west of western base of Cascades), to northern California, being 
found as far south as Carson’s Camp on Mad River, Humboldt County. 
Its closely related subspecies Zapus trinotatus allent (Preble, 1899, 
pp. 27, 28) oceupies the Boreal zone of the Siskiyou, Trinity, and 
Sierra Nevada mountains, as well as the inner coast range as far 
south as South Yolla Bolly Mountain (between Tehama and Trinity 
counties). 

Zapus major is probably an outlying form with trinetatus affini- 
ties (Preble, 1899, p. 25). It is found on the Warner Mountains of 
Oregon and California. Zapus orarius is found in the humid coast 
belt of California from Marin County to Humboldt Bay. Its rela- 
tionships are not clear, but it is probably nearest to Zapus pacificus, 
which is found in the interior valleys of southwestern Oregon and 
northwestern California. 


APLODONTIIDAE 


Although detailed facts regarding the aplodontias are not so 
numerous as is desirable, it seems obvious that the relationships of the 
California forms are as follows: The recently described Aplodontia 
chryseola (Kellogg. 1914, pp. 295, 296) occupying the Trinity and 


—E 


———eeeE——eEeEeEeEeEeE—EeEeEeE——E—EEEaEE 


1916] Taylor: Beavers of Western North America 471 


Siskiyou mountains is most closely allied to Aplodontia californica. 
of the Sierra Nevada Mountains. On the other hand, the coast forms, 
Aplodontia pacifica, found at Newport, mouth of Yaquina Bay, Lin- 
coln County, Oregon, Aplodontia nigra, Point Arena, Mendocino 
County, California, and Aplodontia phaca, Marin County, California, 
would seem to be most closely related among themselves. 


OCHOTONIDAE 


This small but interesting family is found only upon or near the 
summits of the highest mountains in California. Three species are 
represented within the state, Ochotona taylori, of the high peaks of 
the Warner Mountains of northeastern California; Ochotona schisti- 
ceps, of the central and northern Sierra Nevada; and Ochotona 
albatus, of the Mount Whitney region, southern Sierra Nevada. It is 
not improbable that Ochotona taylori is more closely related to Ocho- 
tona schisticeps of the region intervening than it is to Ochotona 
albatus of the Mount Whitney region of the southern Sierra. 


CERVIDAE 


Odocoileus columbianus columbianus occupies the northwest coast 
region of California and northward at least to the Columbia River. 
It is found as far to the eastward in California as Mount Shasta and 
extends through the inner coast mountains to the Sacramento Valley. 
The Golden Gate, San Francisco Bay, separates it from the closely 
related Odocoileus columbianus scaphiotus, which is found in the 
Transition and Upper Sonoran zones southward through the Santa 
Cruz district into Monterey and San Benito counties. 

Characteristic of the Sierra Nevada and the southern and desert 
ranges within the state is the mule deer. Odocoileus hemionus 
hemionus is found to be generally distributed in eastern California, 
from the southern end of the High Sierra north to the northeastern 
corner of the state. It comes in contact with Odocoileus columbianus 
columbianus in the Shasta region, where there may be some overlap- 
ping of range. From the Tehachapi Mountains (southern end of high 
Sierra Nevada) westward through the Tejon region to the coast moun- 
tains and southward to the Mexican line, west of the desert proper, 
occurs Odocoileus hemionus californicus. There formerly oceurred on 
the deserts of southeastern California the burro deer, Odocoileus 
hemionus eremicus, but the subspecies seems now to be extinct north 
of the Mexican line. These forms of the mule deer are very closely 
related, and it is evident that they occupy adjoining ranges. 


472 University of California Publications in Zoology ‘Vou. 12 


SUMMARIZATION 


Having now adduced evidence from a study of nine families of 
mammals, representative of four orders, we may profitably undertake 
to discover whither it leads us. 

Before proceeding it ought to be stated that this evidence is typi- 
cal of that which would be gathered in course of the consideration of 
any family whatever of California mammals. The evidence might be 
greatly extended, but such extension would imply much unjustified 
repetition, for further evidence would probably not illustrate the 
actual state of things any better than that which we now have at 
hand. 

It should be remembered that our present concern is the gather- 
ing, so far as that may be here practicable, of the evidence on the 
differentiation of species, or polytypic evolution (the splitting up of a 
parent stock into a number of differentiated stocks), as opposed to 
monotypic evolution (continuous change through time and in a given 
direction of the entire stock without divarication). Thus we are not 
discussing the problem of the mode of organic evolution primarily, 
except in so far as this problem is bound up with considerations of 
differentiation. 

Furthermore, we propose to discuss this matter from the stand- 
point of studies, first of mammals, and second, of the higher verte- 
brates in general. It is hardly to be assumed that the same con- 
ditions and factors of organic evolution apply universally, or that, 
where they do apply, they must act in exactly the same way. 

An illustration of the heterogeneity of the organic world is for- 
tunately furnished by the very problem with which we are here deal- 
ing, namely, that of the place of geographic isolation in specific dif- 
ferentiation. We are indebted to Kofoid (1907a, pp. 500-506) for 
calling pointed attention to the possible limitations of isolation in the 
origin of species. After remarking the high degree of cosmopoli- 
tanism of the fresh-water microfauna and flora, Kofoid emphasizes 
the fact that there is not only extensive coincident distribution of 
related species, but that there are numerous cases in which there is 
actual coexistence of the most closely related species in the same 
habitat. 

Similar evidence is adduced by Clark (1911, p. 23), who demon- 
strates the coincident distribution of closely related species among 
the Ophiurans. There seems to be abundant evidence that very 
closely related species of this group, often the most closely related, 


1916] Taylor: Beavers of Western North America 473 


inhabit the same area, and that ‘‘Jordan’s law’’ does not apply to 
this class of Echinoderms; and further, that ‘‘ physiological isolation’’ 
in some form has been a more important factor than geographical or 
bathymetrical isolation in the specific differentiation of Ophiurans. 
This state of things stands, of course, in strong contrast with the 
widespread isolation of related species and subspecies in birds, mam- 
mals, and vertebrates generally (see p. 475). 


RELATION OF EVIDENCE TO SOME CURRENT THEORIES 


Do the facts of distribution which we have before us meet the 
requirements of Darwin’s hypothesis as to the origin of species 


Geographic Aréas 


Species 1 


Species 2 


Species 3 


aan 

a= 

c= 
oibojoog 


SQUIIN 


ce. S| 


Species 4 


Species 5 


Fig. O. Diagram illustrative of species arrangement as we ought to find 
it in nature according to Darwin’s theory of differentiation in 
the same place through the advantage of divergence, 


through natural selection in the same place through the advantage 
of divergence? Do they accord with De Vries’ theory of mutations, 
according to which the parent species, after passing into a mutation 
period, gives rise to one or more elementary species or retrograde varie- 
ties, which may coexist with the parent stock in the same locality? 
Do they fulfill demands of Romanes’ and Gulick’s physiological selee- 


474 University of California Publications in Zoology (Vor. 12 


tion according to which there may be segregation of sections of the 
same species in the same place, and so divergent evolution, through 
preferential mating? 

It will be apparent immediately that the facts here adduced do not 
harmonize with the requirements of any of the hypotheses above 
mentioned; for on any one of them we ought, at least occasionally, 
to find species arranged in nature as illustrated in the accompanying 
diagram (fig. O). According to this diagram each group is repre- 
sented to have undergone divergence in the same place, until its mem- 
bers occupy different ecologic niches, the horizontal lines representing 
physiological, the heavy vertical lines geographical, barriers. What we 
do find in nature is that species are arranged quite otherwise (fig. P). 


Geographic Areas 


Species 1 Species @ Species 3 Species4 Species 5 


Beavers 


m 
Shrews fa) 
e 
fo} 
OQ 
oO 
Martens 
2 
ce) 
= 
a 
w 


Fig. P. Diagram illustrative of species arrangement as we actually do 
find it in nature. 


Each group, speaking now in general terms, for there are some excep- 
tions, occupies the same ecologic niche in different places, rather than 
different ecologic niches in the same place. The two propositions are 
theoretically well balanced, and on @ priori grounds one would per- 
haps be unable to discover any argument in favor of the one arrange- 


1916] Taylor: Beavers of Western North America 475 


ment as over against the other. But the facts of geographical distri- 
bution of higher vertebrates point unequivocally to figure P as a 
more accurate illustration of species arrangement than is figure O. 


RELATION OF EVIDENCE TO WAGNER’S THEORY OF MIGRATION AND GEOGRAPHICAL 
ISOLATION 


There is left Wagner’s theory, that the differentiation of species 
has taken place through migration and geographical isolation, the 
migration and spatial isolation being, according to this view, the con- 
ditions without which specific differentiation does not take place. 
With this proposition the evidence from mammals and higher verte- 
brates in general seems to be, for the most part, in complete harmony. 
The terms of this theory demand that a group made up of closely 
related forms be represented by one species or subspecies, only, in 
each locality, and that the most closely related forms be separated 
by some barrier which shall keep them from interbreeding to an 
extent that would bring about the swamping of incipient characters. 
In other words, on this theory, groups of related species should be 
arranged as illustrated in figure P. And so they are. 

Reference has already been made (p. 472) to Kofoid’s suggestion 
regarding the coincident distribution of some of the most closely 
related species of the fresh-water microfauna and flora. In another 
paper (1907b, pp. 241-251) the same author considers the distribu- 
tion of the Chaetognatha with reference to its bearing on the relation 
of isolation to the origin and preservation of species, coming to the 
following conclusion: ‘‘The apparently wide-spread phenomenon of 
coincident distribution of related species among pelagic organisms 
appears to cast some doubt upon the universality of the operation of 
isolation in the evolution of species as originally maintained by 
Moritz Wagner (1868) and recently revived by President Jordan 
(1905).’’ In the course of this paper Kofoid emphasizes the lack of 
specific distributional data, especially of that pertaining to vertical 
distribution. 

Supplementary in a way to this work of Kofoid is an exceed- 
ingly suggestive paper dealing with the problem of isolation versus 
coincidence in the same group (the Chaetognatha) recently pub- 
lished by Michael (1913, pp. 17-50). The extensive collection of 
specific data with regard not only to latitudinal and longitudinal 
but also to vertical distribution permits the enunciation of the fol- 
lowing principle (Michael, 1913, p. 18) : 


ce 


Pelagic organ- 


476 University of California Publications in Zoology (Vou. 12 


isms may be coincidently distributed as regards latitude and longi- 
tude, and still be completely isolated in their vertical distribution.’’ 

The results of Michael’s work indicate that the inerease of exact 
knowledge in at least one group of oceanic organisms has empha- 
sized the possible significance to these organisms of isolation. 

It is concluded (Michael, 1913, p. 46) that ‘‘Jordan’s Law’’ is 
only partly true, when tested by vertical distribution, for while the 
most closely related species do not inhabit the same environment, 
they do inhabit the most remote environments; and that the more 
closely related species of Chaetognatha are isolated from each other 
either horizontally, vertically, or by virtue of physiological differ- 
ences causing fertilization to take place in different strata of water. 

Attention should be called to the following points in connection 
with the data presented on pages 465 to 472 of the present paper, 
at which the evidence apparently does not clearly harmonize with 
Wagner’s theory. 

Sorex halicoetes of the San Francisco salt marshes is separated 
from its closest relative, Sorex vagrans vagrans of the humid strip 
along the Pacific Ocean, by several miles of upland which is inhab- 
ited by Sorex californicus californicus. According to the strict 
requirements of Wagner’s hypothesis, the ranges of the most closely 
related forms should occupy adjoining territory. If our interpreta- 
tion of the relationship of these forms be correct, Sorex californicus 
californicus must be conceived of as having invaded the territory it 
now occupies at the expense of the vagrans stock, crowding the 
latter to one side or the other. 

Island distribution offers many cases of broad gaps between the 
ranges of species most closely related, but in most cases these gaps 
were formerly nonexistent or else the closely related stocks in ques- 
tion formerly enjoyed some means of transversing them, or are able 
at long intervals still to traverse them, though not to the extent that 
there is a quashing of incipient differential characters through inter- 
breeding. The same principle should be remembered in connection 
with distribution along mountain systems like the Sierra Nevada, 
where, for instance, the family Ochotonidae has a discontinuous dis- 
tribution, being found only in certain ‘“‘islands’’ of high Boreal. 

Two very interesting and instructive cases are furnished by the 
maniculatus series of mice. The relations between Peromyscus 
maniculatus gambelii and Peromyscus maniculatus rubidus, as well 
as those between Peromyscus maniculatus austerus and Peromyscus 


1916] Taylor: Beavers of Western North America 477 


maniculatus oreas, are rather exceptional. Concerning the first 
case Osgood says (1909, pp. 69, 70) : 


The transition from gambeli to rubidus along the line between their ranges 
is rather sudden, suggesting the possibility of hybridizing. From several 
localities specimens fairly typical of both forms are known, from others we 
have both forms and apparent intermediates, and from still others all speci- 
mens thus far obtained are intermediate not typical of either form. This is 
exactly what would be expected upon the theory of hybridism, but of course 
it cannot be considered as conclusive proof. 


Concerning the second he says (pp. 52, 53) : 


The case is very similar to that of gambeli and rubidus in California, the 
complications of which may be due either to hybridization or to intergrada- 
tion. It is already known that oreas and austerus occur together at a number 
of localities and apparently maintain their respective characters. At other 
places only one form has thus far been found, at others extremes of both 
forms and intermediates occur, and at still others intermediates only. There 
is no environmental distinction as in the case of gambeli and rubidus, for 
oreas and austerus live under apparently identical conditions. Although only 
one form has been found at the respective type localities of oreas and 
austerus, both occur together near by, and further collecting may show that 
they do so over a wide area. Specimens which appear to be intermediate 
between oreas and austerus may in reality represent special differentiations 
of the one or the other showing accidental parallelism. 


These cases are very puzzling, as the pairs of species mentioned 
are very closely if not most closely related to each other, and on 
Wagner’s hypothesis should not oceur together. Note should be 
made of at least two possible explanations: It may be that, having 
originated in different geographic areas, and having undergone some 
degree of differentiation, there has been re-invasion, and that, even 
though very closely related, they occupy slightly different ecologic 
niches, and are able to coexist in the same place because of some 
measure of physiological isolation. Or it may be that we have here 
an illustration of Mendelian inheritance in nature, the two sub- 
species crossing freely. The extreme rarity among higher verte- 
brates of this latter type of behavior where ranges overlap at their 
margins, would seem to constitute an argument for the first, rather 
than the second explanation, but the case is not entirely clear. 

It will be remembered that Mustela arizonensis is found in the 
Transition and Boreal zones of the Sierra Nevada and the Rocky 
Mountain systems. On the Cascades and the Trinity Mountains 
its place is taken by Mustela saturata, a closely related form. 
Between the Sierra Nevada and the Rocky Mountains there would 


478 University of California Publications in Zoology (Vou. 12 


seem to be a barrier far greater than that between the Sierra 
Nevada and the Trinity-Cascade mountains, and yet Mustela ari- 
zonensis 18 found on both the Rocky Mountains and the Sierra, but 
is replaced by another species on the Trinity-Cascades! It may be, 
of course, that the real efficacy of the Klamath River gap has not 
been appraised at its true value, and that the effectiveness of the 
barrier between the Rocky Mountains and the Sierra has been 
exaggerated. 

Mephitis platyrhina, as previously stated, is found in the Sono- 
ran valleys around the south end of the Sierra Nevada. This species 
might be conceived to have arisen through mutation, or Darwinian 
divergence, or physiological selection, from Mephitis occidentalis 
holzneri, the range of which partially overlaps the range of 
platyrhina. The following considerations, however, militate some- 
what against these possibilities: (1) The speciation, that is, differ- 
entiation, of the skunk family as a whole seems to have been 
dependent on migration and geographical isolation. (2) The over- 
lapping of the ranges of WM. platyrhina and M. o. holzneri takes 
place over a narrow area only. (3) M. platyrhina seems to be most 
closely related, on the basis of its structure, to MW. 0. major, which 
occurs to the northward, and not to M. o. holzneri (Howell, 1901, 
1 BE) 

One other case: There occur along the Pacific Coast three 
species of the mammalian genus Aplodontia, very intimately related 
one to the other, less intimately related to the members of the genus 
found in the interior mountains (Aplodontia californica and A. 
chryseola). The coast species referred to are Aplodontia pacifica, 
Yaquina Bay, Oregon; Aplodontia nigra, Point Arena, California; 
and Aplodontia phaea, Marin County, California. 

In the case of these forms the Sierran or mountain stock has 
evidently moved westward into the Siskiyou and Trinity mountains, 
where it is represented by the species chryseola, and even to the 
coast itself, where it is represented, in the Humboldt Bay region, by 
an undescribed form nearly identical, cranially, with chryseola. 
Thus the ranges of the closely related A. pacifica of Oregon and 
A. mgra of Point Arena, California, are separated by the chryseola 
stock, which must thus, on the Wagnerian hypothesis, be thought 
to have carried the western boundary of its range to the sea 
since the pacifica-nigra-phaca stock attained its broad coastwise dis: 
tribution. 


1916] Taylor: Beavers of Western North America 479 


It is noteworthy that in all these critical cases there is not a 
sufficient quantity of fossil and Recent material for the adequate 
elucidation of their relationships. This is especially apparent with 
the weasels, skunks, and aplodonts just cited. On the other hand, 
in cases where material is abundant and the status of the forms is 
reasonably well established, the great mass of facts of distribution 
harmonizes with Wagner’s theory. 


How Have Dirrerent Econtocic NicHES BEEN FILLED? 


It should be remembered, of course, that had living forms not 
possessed the power of adapting themselves or becoming adapted to 
different ecologic niches we would not have had the various niches 
filled. But the evidence from mammals and higher vertebrates 
would seem to indicate that geographic isolation is prerequisite to 
any kind of differentiation; physiological isolation, or practical ster- 
ility, being gradually assumed in proportion as the completeness 
and long continuance of geographical isolation, and the association 
of it with diversity of external conditions, permit or condition 
morphological changes extending to the reproductive system. The 
broader the distribution of the species and subspecies of any group 
of mammals or higher vertebrates, the greater the probability that 
some of the forms will find themselves in ecologic niches which, 
while they resemble in a general way the original group niche, will 
still differ from it in some important particulars. In strict terms, 
no two ecologic niches, situated in different geographic areas, can be 
precisely the same. 

Thus it may be noted that while the so-called panamintinus 
eroup of the genus Perognathus occupies the same general ecologic 
niche wherever found, as a matter of fact, the particular ecologic 
niche occupied by each species or subspecies is somewhat different 
from that occupied by any of the other species or subspecies. 
Grinnell and Swarth (1913, pp. 390, 392) have listed in their 
“fourth category’’, Table D, a number of forms of birds and mam- 
mals of which also this may be said to be true, and in the opinion 
of the writer study of group distribution in vertebrates shows this 
to be a rule of practically universal applicability. 

The status of things may be graphically represented by a diagram, 
as in fig. Q. Let the capital letter stand for the stock or species, the 
lower-case letter represent the ecologic niche it occupies, and the 
arabic numeral the geographic area in which the ecologic niche is 


480 Unwwersity of California Publications in Zoology (Vou. 12 


Fig. Q. Diagram illustrating possible explanation of presence of related 
forms in same locality. 


located. The vertical lines represent geographical, the horizontal 
and slanting lines physiological barriers. Thus we have 1, 2, 3, 4 
and so on, contiguous geographic areas, each separated from the 
other by some sort of geographic barrier. If A is the primitive 
mammalian stock, then B may be regarded as a stock which acci- 
dentally or otherwise has become geographically separated from it, 
and finally has become differentiated into a separate species. The 
continuation of this process of migration and differentiation may be 
conceived of as resulting in species G, which has become so far dif- 
ferentiated from the original stock A as not to be included in the 
same subgenus with it. 

The ecologic niche b, occupied by species B, will be shghtly dif- 
ferent from niche a, in which the primitive stock is found. That 
occupied by species C will differ shghtly from that occupied by B. 
This very slight difference in ecologic niche will be apparent up to 
and including g, in which species G, which has attaimed a quite con- 
siderable divergence, is found. 

Thus the series of niches, into which members of the subgenus 
of which species G is representative will be able to enter, will be 
somewhat different from the series to which members of the subgenus 
of which species A is representative will have ready access. Thus it 
may happen, through a further continuation of the process of 
migration and differentiation, that species H to M, members of the 
G subgenus, may arise, and there may even be a re-invasion, on the 
part of some member of the G group, as for instance M, into the 
geographic area occupied by original stock A. But M will have 


1916] Taylor: Beavers of Western North America 481 


become considerably differentiated from A, and probably will be 
physiologically isolated from it, and furthermore, will not be under 
the necessity of competing with A, except in the most general way, 
for it will be fitted to occupy ecologic niche m, which is different from 
ecologic niche a. Two species derived from the same original stock 
will thus be found occupying different ecologic niches in the same 
place. The closest relatives of each will be found, not in the same 
locality but in a neighboring area, the races or species being kept 
separate by a geographic barrier of some sort. Thus B is A’s 
closest relative, and L is M’s closest relative. 

It is to be doubted whether an actual series of cases completely 
illustrative of the steps in such a differentiation and re-invasion can 
ever be demonstrated in nature. The process is an extremely com- 
plicated one, as a moment’s consideration will show. The diagram 
is purposely made very simple; in nature the process must be 
exceedingly complex. The primitive stock and its descendants are 
subject to all the vicissitudes of environmental and topographic 
change, and the process as above outlined would not only require an 
enormous time for its fulfillment, but would depend upon the 
existence of spatial relations of exactly the right sort. Possible 
contingencies in a great many directions are numerous. 

The great weight of evidence from the study of mammals and 
higher vertebrates would seem to indicate that the occupation of 
different ecologic niches in the same place has in every instance 
been the result, not of some process of adaptation of a portion of 
the parent stock to a distinct ecologic niche, and the differentiation 
of this adapted portion while both were living in the same locality, 
but of some comprehensive process similar to the hypothetical one 
above presented, involving migration, differentiation, and re-invasion. 

If, as we believe'the evidence indicates, geographic isolation is, 
in the higher vertebrates, a condition essential to specific differenti- 
ation, then on the assumption that the classes of vertebrates are 
descended from one or a very few original stocks it follows that 
these migrations, differentiations and re-invasions have been very 
extensive. For there are many ecologic niches in almost every 
locality which are now occupied by various mammals, or birds, or 
reptiles. It has been long recognized, of course, that there have 
been numerous migrations and differentiations. But has due recog- 
nition been accorded the principles of invasion and re-invasion ? 
Unless there had been, according to the logical continuation of the 


482 University of California Publications in Zoology (Vou. 12 


two propositions (geographic isolation a condition to divergence in 
higher vertebrates; classes of vertebrates descended from one or a 
very few original stocks) extensive invasions and re-invasions, each 
geographical locality would now be occupied by one mammal, or one 
bird, or one reptile, only. 

Following out this thought it would appear that there have been 
aS many mammalian invasions or re-invasions for each locality as 
there are species or subspecies of mammals within it—unless so 
large a locality is taken as to contain barriers which would serve 
to isolate portions of the stocks from each other. This would seem 
to afford a confirmation from the standpoint of the theoretical 
necessities of isolation, of the axiom, stated by Grinnell (1914, p. 
98) to be a necessary result of the ever-shifting location geograph- 
ically of associational, faunal, and zonal conditions; this axiom is 
that ‘“‘every single element or line of descent, now represented in 
the biota of any locality, must have come either in its present form 
or in some antecedent one from somewhere else.’’ 

In almost every locality there are ecologic niches which are 
unoccupied. In many instances there exist animals which could 
occupy these niches if they could get to them; but barriers of one 
sort or another effectually prevent their getting to them. This con- 
stitutes an important line of negative evidence which is comple- 
mentary to the positive line. If an organic stock were capable of 
divarication in a single locality, given geologic time all occupiable 
niches ought to have been filled. 

Of course the species resident in a locality might and probably 
would become modified monotypically through a_ progressively 
changing environment or from some other cause or causes, but there 
would never ensue any differentiation. To increase the number of 
species or subspecies of mammals in the locality further invasions 
or re-Invasions are called for. 


TENTATIVE SUGGESTIONS REGARDING THE MANNER IN WHICH GEOGRAPHIC ISOLA- 
TION ACTS IN THE PROCESS OF SPECIATION 


Although in most cases laboratory work is done with forms low 
in the organic scale, and it is questionable how broadly the conclu- 
sions based upon such work may be applied, some of the results of 
experimental work are exceedingly suggestive as to what may be 
taking place in nature in widely separated classes of organisms. 
As Thomson says (1909, p. 329), “‘The world of organisms is very 


1916] Taylor: Beavers of Western North America 483 


large and heterogeneous, and results that hold good for certain 
forms of life may not be true of others.’’ The variety of the living 
world seems to be no less characteristic than its unity; so it would 
seem appropriate that workers in science should be very chary of 
launching generalizations as universals. 

Feeling, however, that one of the most pressing of present-day 
needs in biology is the correlation of the work of the ‘‘experi- 
mentalist’? and the ‘‘naturalist’’, the writer is led to submit the 
following suggestions, incomplete though they may be. 

When we face the question of the manner in which geographical 
isolation works, some of the results of Tower’s experiments with 
beetles of the genus Leptinotarsa (1906) and of MacDougal’s work 
with plants of the genus Raimannia (1906, p. 422) are of great 
interest. It will be remembered that Tower, by subjecting his 
beetles to varying conditions during the growth-period of the germ- 
cells, effected the production of new species and new characters. 
Although the parents were unaffected, their germ-cells were modi- 
fied, and the offspring grew up different. MacDougal injected salt 
solution into the ovules of Raimannia just previous to fertilization, 
securing potentially new species as a result of the chemical and 
osmotie action exerted on unfertilized ovules. Tower, comment- 
ing on this case (1906, p. 295), says: ‘‘These results of Mac- 
Dougal’s exactly confirm in plants the results that I have obtained 
in these beetles, so that the point is now doubly certain that herit- 
able variations are produced as the direct response to external 
stimuli.’ 

Our studies of the distribution of higher vertebrates invite the 
belief that in the majority of cases differentiation follows migration 
and, clearly in most instances, the exposure to different environ- 
mental conditions. There is thus suggested a possible partial agree- 
ment between these two apparently widely sundered classes of 
observational facts. What has caused the differentiation which we 
observe in the higher vertebrates? The application of the prin- 
ciples of Independent Generation, Independent Variation, or Amixia 
(these three synonymous terms standing for differentiation under 
similar conditions in virtue of geographic isolation alone) to these 
classes of organisms is probably limited. These animals usually 
range so widely that there are obvious environmental differences 
associated with difference in species. The facts of geographical 
distribution of higher vertebrates do not militate against Tower’s 


484 University of California Publications in Zoology (Vou.12 


conclusion that heritable variations are produced as a direct 
response to external stimuli, even if they cannot be shown directly 
to favor it. 

It is noteworthy that Tower insists that although heritable vari- 
ations are produced as a direct response to external stimuli, the 
response itself is absolutely determined within the organism. To 
quote his own words (1906, p. 295) : 


It is true that different intensities of the same stimuli call forth different 
responses, but, as is shown in the chapter on coloration, the response is 
entirely determined within the organism, which is adjusted to different 
intensities of stimuli and reacts according to its own method and on the basis 
of its own constitution, there being no specific reaction called forth by a 
given stimulus. 


The facts from the study of the ‘zoogeography of the higher 
vertebrates appear to harmonize with these conclusions of Tower’s. 
As the evidence becomes more complete it appears more and more 
clear that Darwin was right in assigning to the ‘‘nature of the 
organism’’ a more important place in considerations of speciation 
than that accorded the ‘‘nature of the conditions’’. 

Do we not have here a suggestion as to how isolation works? The 
shaping of the species seems to depend upon two things primarily: 
(1) the nature, rapidity, and intensity of the blows from the 
environmental hammer; and (2) the nature of the organism itself. 
Thus isolation apparently operates, in the higher vertebrates, by 
segregating different sorts of environments and insuring their con- 
tinued moulding of the particular segregated lots of individuals. 
On passing from one faunal area, association, or life-zone to another 
we encounter different, but closely related species. Thus differences 
between geminate species (Jordan, 1908) are apparently dependent 
upon the fact that differential environmental hammers have been 
used on what was originally the same stock. On making a com- 
parison of this sort, we note that there has been great diversity of 
response on the part of the different members of the assemblage to 
the same environmental complex. The entire assemblage in any 
given district has been subject to the same environment, but the 
assumed differences in the stuffs moulded in each case readily 
account for the fact that all have not responded in the same way. 

Tower (1906, pp. 286-296) found that by subjecting a given 
species of potato beetle to unaccustomed environmental stimuli 
there resulted in the offspring a break-up into several forms, some 


ae 


1916] Taylor: Beavers of Western North America 485 


of which resembled the parent stock and others of which exempli- 
fied potentially new species. For example, when Leptinotarsa decem- 
lineata was so treated there was a break-up in the next generation 
into Leptinotarsa immaculothorax, L. pallida, and the unmodified 
L. decemlineata. The evidence from the zoogeography of the 
higher vertebrates clearly indicates, as has been repeatedly stated 
and implied, that isolation of one portion of a parent stock and 
subjection of it to the conditions of one new environment, results in 
the differentiation of but one new form, and that until further 
migration has effected the further isolation of the stock and its 
subjection to some other environment there is no further polytypie 
evolution. 

A further implication contained in the work of Tower should be 
mentioned here. The modifications were definitely brought about in 
his experimental work through the action of the unaccustomed 
environmental stimulus during the period of growth and matura- 
tion of the germ-cells, and at no other time. This invites one to 
the suggestion that the environment may do its work by bringing 
pressure to bear directly or indirectly on the germ-cells, and that 
environmental-somatie impacts may be of no moment in speciation 
if they are not conveyed to the germ-cells. The failure of environ- 
mental-somatic impacts to be translated into environmental-germinal 
impacts may explain the permanence of subspecies amid conditions 
very different from those to which they are accustomed. Ammosper- 
mophilus leucurus leucurus, for example, having its center of dis- 
tribution on the Colorado Desert, ranges notably over into the 
damper San Diegan faunal area, maintaining its desert character- 
istics even under the moister conditions (see Grinnell and Swarth, 
1913, pp. 391-394). Evidently, however, such a translation of 
environmental-somatic impacts into enyironmental-germinal impacts 
has taken place in most cases in the past, and may perhaps be 
anticipated to take place, where it has not yet done so, at some time 
in the future. 

As water, constantly dropping, wears away the stone, so 
the environment, through constant and continuous re-impression, 
seems to find at last some avenue to the germ-cell, until, as believed 
by most students of zoogeography, it becomes true that differentia- 
tion has come to be a part of the connotation of geographical 
isolation. 

That the differentiation of which geographic isolation is a condi- 


486 University of California Publications in Zoology (Vou. 12 


tion is cumulative, is indicated by a great many facts from zoo- 
geography. Species known to have originated recently, as for 
example, those in neighboring areas separated by a barrier not 
absolute, are structurally much more alike than species known to 
have been separated for longer times. Thus Castor subauratus 
shastensis and Castor subauratus subauratus, occupying neighboring 
areas, and only recently separated, are structurally much more 
alike than are Castor fiber and Castor canadensis canadensis oceupy- 
ing areas in different continents, and separated for a much longer 
time. Island forms of mainland stocks are often poorly differenti- 
ated, the degree of divergence seeming in a measure to be asso- 
ciated with the length of time of separation of the island from the 
mainland. This cumulation is, of course, only within limits, for 
there are notable examples of very long separation with compara- 
tively little differentiation, and even a few cases wherein long sep- 
aration has apparently failed to bring about any differentiation 
whatever. 

To summarize the remarks in this section: It may be conceived 
that isolation works with organisms in nature in some such way 
as the experimentalist works with organisms in the laboratory. The 
latter isolates a number of individuals and subjects them to a 
diversity of surroundings known as the conditions of the experi- 
ment. In nature the parent stock becomes separated, by some 
means or other, natural barriers corresponding to the walls of the 
container or of the laboratory, and the different environmental 
conditions in new territory occupied, corresponding to the condi- 
tions of the experiment. Differentiation may ensue in either ease. 
In the higher vertebrates, at least, it has ensued practically in all 
cases. Geographical isolation probably conditions differentiation 
by segregating organism and environment together and maintain- 
ing their mutual interrelations over long periods of time. 

Evidence from geographic distribution of higher vertebrates 
does not oppose, if it does not definitely support, the thesis that 
speciation is dependent on modification in germinal constitution. 
It appears to present unequivocal data that the response to the 
different environments depends upon the nature of the organism. 
It furnishes a mass of evidence against the theory of the possibility 
of the appearance of a new species in territory occupied also by the 
parent, so emphasizing the necessity of migration and geographic 
isolation as conditions of polytypic evolution. It indicates that 


} 
b 
} 
J 
’ 
' 
N 


1916] Taylor: Beavers of Western North America 487 


the differentiation conditioned by isolation is, within limits, eumu- 
lative. 

As remarked previously, these conclusions are drawn up simply 
as a possible means of assisting in understanding the enormous 
and complicated body of facts concerning the speciation of the 
higher vertebrate animals, and may or may not hold for other 
classes of organisms. 


SUMMARY 


1. The ranges of seven recognizably differentiated subspecies 
of beavers touch upon, or lie close to, the Pacific Ocean in western 
North America. 

2. Since beavers grow throughout their life, the consideration 
of changes in form, outline, and dimensions of the various parts due 
to difference in age becomes especially important. There is little 
change in coloration with age. 

3. The beavers possess a highly specialized dentition, as is shown 
by the following facts: (1) the permanent dentition exhibits a high 
degree of hypsodonty, though the milk teeth are brachydont; 
(2) the enamel-fold pattern of the cheek-teeth is much complicated ; 
(3) the dental formula is much reduced. The generalized mam- 
malian formula is 13, Ci, P4, M3 x 2=44, while that of the 
beaver is 11, C9, P1, M3 x 2—20. During ontogeny, the contin- 
uously growing cheek-teeth first increase in size, then decrease 
shghtly. There are no important differences due to age in the pro- 
portional relation of the antero-posterior to the transverse diameter 
of the teeth. The dental armature is a cutting and grinding agency 
of high efficiency. 

4. The similarly rounded instead of plane conformation of the 
palato-maxillary region, and the similar complication of the enamel 
pattern of the cheek-teeth in the Sciuromorph genus Castor and in the 
Hystricomorph genus Hrethizon appear to be parallel adaptations: 
the first condition being an adaptation to the stripping of the bark 
from twigs and branches, the second to the effective mastication of 
this food. 

5. Certain readjustments of current systematic conceptions of 
beaver status are necessary : 

(1) The race of beaver inhabiting the Cook Inlet region of 
Alaska and ranging south into northern and central British Colum- 


488 University of California Publications in Zoology (Vou. 12 


bia is here characterized as a new subspecies, Castor canadensis 
belugae. 

(2) The race of beaver inhabiting the Pit River region of Cali- 
fornia, east of the Sierra Nevada, is here characterized as a new 
subspecies, Castor subauratus shastensis. 

(3) The race of beaver inhabiting the mainland of Washington 
and Oregon is shown to be subspecifically distinct from the form 
on Vancouver Island (Castor canadensis leucodonta Gray) requir- 
ing the rehabilitation of Rhoads’ name pacificus. 

6. The fragmentary palaeontologic history of the Castoridae 

points to at least three intermigrations of beavers between the Old 
World and the New. Steneofiber appears in the middle Oligocene 
of Europe, and in the upper Oligocene of North America. The 
Eucastor-Dipoides stock appears in the upper Miocene of North 
America, and in the Pliocene of Asia. Castor first appears in the 
Pontian upper Miocene of Europe, but in North America it does not 
appear until later, its first known occurrence being in the Pliocene 
of California. Im the case of each of these genera, at least one 
intercontinental migration is indicated. 
7. The beavers of North America (those of which material has 
been available) are separable into two groups: the canadensis 
group, with subspecies canadensis, michiganensis, belugae, lewco- 
donta, pacificus, frondator, and texensis; and the subauratus group, 
with subspecies subauratus and shastensis. The forms making up 
the canadensis group are unequally related. 

8. On the whole, differentiation in the genus Castor is slight. 
The California stock has undergone more divergence than any other. 

9. A consideration of its history and present status shows that 
the widest diversity of opinion as to the isolation concept still pre- 
vails, and emphasizes the necessity for more critical enquiries con- 
cerning it. 

10. The evidence from beavers, and higher vertebrates gener- 
ally, regarding Weismann’s Amixia, Romanes’ Independent Varia- 
bility, and Gulick’s Independent Generation, is inconclusive. In 
the case of beavers it must be conceded, however, that the possi- 
bility is by no means excluded that geographical isolation alone 
has been the chief condition in speciation. 

11. The bulk of the evidence from the zoogeography of beavers 
and other vertebrates seems to indicate that polytypic evolution in 
these groups has been conditioned by the spatial partition of the 


| 


eC 


1916] Taylor: Beavers of Western North America 489 


originally uniform stock as required by Wagner’s isolation theory. If 
any one of the following factors, (1) ‘‘advantage of divergence’’ 
as Darwin used the term, (2) De Vriesian mutation, (3) Men- 
delian segregation, (4) physiological selection, has been operative 
in the differentiation of higher vertebrates, it is not apparent in 
the data of distribution. Im these higher groups differentiation 
becomes, sooner or later, a part of the connotation of geographic 
isolation. 

12. A corollary of the proposition just stated is that in beavers, 
and in higher vertebrates generally, geographic range is exactly as 
characteristic of species as any physical or psychical attribute. 

13. Any definite conclusions regarding certain problematic cases 
would be, premature. Much more testimony must be taken before 
a decision is reached. Such rare occurrences as that of Peromyscus 
maniculatus oreas with its closest relative Peromyscus maniculatus 
austerus in the same territory are susceptible of interpretation 
according to one of two alternatives: (1) the case may be one of 
speciation in accordance with Darwin’s, De Vries’, or Romanes’ 
theories, the two species being preserved intact through Mendelian 
segregation; (2) the speciation may have proceeded according to 
Wagner’s theory of geographic isolation (that is, by migration, 
geographical isolation, and differentiation) ; the physiological isola- 
tion between them may be but partial; and there may have been a 
re-invasion on the part of one into the range of the other. 

14. Certain experimentalists, as Tower and MacDougal, have 
subjected parent generations of insects and plants to unaccustomed 
environmental stimuli, the offspring showing new characters and 
assumed to represent new specifie types. The distribution of the 
higher vertebrates seems to make it certain that in the majority 
of instances differentiation follows migration, and, clearly in most 
cases, subjection to different environmental conditions. It looks 
as if in both cases heritable variations are produced as a direct 
response to external stimuli, the response in each case depending 
on the nature of the organism. 

15. If this be the case, geographic isolation works no less by 
segregating different environments than by separating different lots 
of individuals. Stated differently, geographic isolation conditions 
differentiation through its maintenance of the mutual interrelations 
of organism and environment through long periods of time. 

16. Zoogeography does not furnish evidence whether or not 


490 University of California Publications in Zoology (Vou. 12 


the only effectual environmental impact is that which is received 
by the germ-cells; therefore it is not opposed to such a conclusion, 
already reached experimentally in widely different groups of 
organisms. 

17. There is evidence from vertebrate zoogeography that indi- 
cates the cumulative character of the differentiation associated with 
geographic isolation. 

18. The different ecologic niches in the same locality, so far as 
they are occupied by a given class of organisms, would seem to 
have been filled, not through processes of differentiation and adapta- 
tion in that single locality, but through those of migration, geo- 
graphic isolation, differentiation, concurrent adaptation to different 
niches, and final invasion or re-invasion of the locality in question— 
to occupy different niches there. If geographic isolation is a con- 
dition essential to speciation in the higher vertebrates, and if each 
class of the higher vertebrates is derived from one or a few ances- 
tral stocks, it follows that the tracing of these processes is highly 
important if not indispensable, either to an adequate understanding 
of the course of development of the class, or to a thorough concep- 
tion of any particular facies of the class. Among these processes 
the principles of invasion and re-invasion, which are apparently of 
great significance in explaining the class-assemblage of a given 
locality, have been less emphasized than their importance deserves. 


Transmitted November 18, 1914. 


LITERATURE CITED 


ALLEN, J. A. 
1877. ‘“Castoridae”’ in “Monographs of North American Rodentia” in 
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BAILEY, V. 
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Baker, M. 
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Banes, O. 
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ee 


1916] Taylor: Beavers of Western North America 491 


Branpt, J. F. 

1855. Ueber die Kentnisse, welche die alten Perser, Egypter, Griechen 
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Cuvier, F. 
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Cuvimr, (G. 1 C, EE: D: 
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DaRwIN, C. 
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DAVENPorRT, C. B. 
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1907. Plant breeding comments on the experiments of Nillson and 
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Drxon, C. 
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Ey, W. W. 
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492 University of California Publications in Zoology (Vor. 12 


Gray, J. E. 
1869. On the white-toothed American beaver. Ann. Mag. Nat. Hist., 
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Grecory, W. K. 
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GRINNELL, J. 

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1913b. <A distributional list of the mammals of California. Proc. Calif. 
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GRINNELL, J.. and SwartuH, H. S. 
1918. An account of the birds and mammals of the San Jacinto area 
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GuricKs ence 
1905. Evolution, racial and habitudinal. Carnegie Inst. Wash., publ. 
no. 25, xii + 270. 


HELLER, E. 
1909. “The mammals” in Birds and mammals of the 1907 Alexander 
expedition to southeastern Alaska. Univ. Calif. Publ. Zool., 5, 
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HO.LuIstTer, N. 
1913. A synopsis of the American minks. Proc. U. S. Nat. Mus., 44, 
471-480. 
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Mus., 47, 427-489, with pl. 15. 


HOLMES, S. J. 
1909. The categories of variation. Amer. Nat., 48, 257-285. 


Hopkins, A. C. 
1914. Parallelism in morphological characters and physiological char. 
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(report of paper read before Biological Society of Washing: 
ton). 
HoweEt., A. H. 
1901. Revision of the skunks of the genus Chincha. U. S. Dept. 
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JEFFREY, E. C. 
1914. The mutation myth. Science, n.s., 39, 488-491. 


JORDAN, D. S. 
1905. The origin of species through isolation. Science, n.s., 22, 545-562. 


1908. The law of geminate species. Amer. Nat., 42, 73-80. 


ee 


1916] Taylor: Beavers of Western North America 493 


Kettoce, L. 
1911. A fossil beaver from the Kettleman Hills, California. Univ. 
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KELLoGG, V. L. 
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494 University of California Publications in Zoology ‘Vou. 12 


MICHAEL, E. L. 
1913. The vertical distribution of the Chaetognatha of the San Diego 
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MILLER, G. S., JR. 
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Oscoop, W. H. 

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PREBLE, E. A. 

1899. Revision of the jumping mice of the genus Zapus. U.S. Dept. 
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RuHoaps, S. N. 

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Ripeway, R. 

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ROMANES, G. J. 

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Scorr, W. B. 

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SHELFORD, V. E. 

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SPILLMAN, W. J. 
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2 | 


1916] Taylor: Beavers of Western North America 495 


"TAYLOR, W. P. 
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THOMSON, J. A. 
1909. Heredity. (New York, Putnam’s, London, Murray), xvi + 605, 
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‘Tower, W. L. 
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genus Leptinotarsa. Carnegie Inst. Wash., publ. no. 48, 
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WAGNER, M. 
1868. Die Darwin’sche theorie und das Migrationsgesetz der Organ- 
ismen. (Leipzig, Duncker und Humbolt), viii + 62. 


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335-398, plates 15-18. 

14. An Analysis of the Vertebrate Fauna of the Trinity Region of 
Northern California, by Joseph Grinnell. Pp. 399-410. 

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Ctenophora of the San Diego Region, by Calvin O. Esterly. Pp. 

PS EST Rea '5 rg eae bb Ses Bcae ee tana Cree ere a eer > Sel atl a a5 oe RS 

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UNIVERSITY OF CALIFORNIA PUBLICATIONS 
IN 


ZOOLOGY 
Vol. 12, No. 16, pp. 497-501 May 6, 1916 


TWO NEW APLODONTIAS FROM WESTERN 
NORTH AMERICA 


BY 
WALTER P. TAYLOR 


(Contribution from the Museum of Vertebrate Zoology of the University of California) 


The genus Aplodontia is found west of the Sierra Nevada—Cascade 
mountain system from. southern British Columbia on the north to 
middle California on the south. Study of specimens of the genus from 
the northern part of the range demonstrates the existence there of two 
subspecies hitherto unrecognized. The writer desires to express his 
thanks for the loan of material to the authorities of the Field Museum 
of Natural History, especially to Mr. Wilfred H. Osgood, to those of 
the Museum of Comparative Zoology, particularly to Messrs. Samuel 
Henshaw and Outram Bangs, and to those of the Bureau of Biological 
Survey of the United States Department of Agriculture, especially 
to Messrs. H. W. Henshaw and E. W. Nelson. He is also indebted 
to the authorities in charge of the Museum of Comparative Zoology, 
particularly to the director, Mr. Samuel Henshaw, for the privilege 
of describing a new subspecies of Aplodontia on the basis of material 
loaned. 


Aplodontia rufa grisea, new subspecies 
Puget Sound Mountain Beaver 


Type—FKemale adult, no. 3751, Mus. Vert. Zool.; Renton [near 
Seattle], Washington; October 4, 1907; collected by Frank Stephens; 
orig. no. 294; stuffed skin, with skull and jaws, all in good condition, 
except hamulars broken. 

Diagnosis—Similar to examples of Aplodontia rufa rufa, but 
paler, grayer; separable from Aplodontia rufa olympica through ab- 


498 University of California Publications in Zoology (Vou. 16 


sence of distinct postorbital processes on the jugal; smaller than 
Aplodontia californica columbiana and Aplodontia californica rainieri. 

Comparisons.—In examples of Aplodontia rufa grisea the brown 
coloration ranges from light ochraceous-buff to light buff, while in 
A. r. rufa the range in coloration is from near tawny to light ochra- 
ceous-buff ; grisea for the most part lacks the distinct brown wash so 
often present in rufa. Cranially, grisea tends to have interpterygoid 
fossa narrower than in rufa, audital tube of smaller caliber, post- 
orbital process indicated on the jugal in some specimens, and lesser 
mastoid width. 

Aplodontia rufa grisea may be separated from A. r. olympica, in 
the usual instance, only by the different development of postorbital 
process on the jugal. In grisea these processes are weakly indicated 
in a few examples, in olympica they reach their maximum of develop- 
ment in the genus. Even this character cannot always be relied upon 
as certainly separative, since the postorbital processes are sometimes 
weakly developed in olympica. Available skins of grisea and olympica 
are not strictly comparable, for the majority of the specimens of 
grisea were collected in winter, and the series of olympica was taken 
in summer. Seasonal variation in the genus is usually exceeded by 
individual variation, however, so cross-comparisons may perhaps legit- 
imately be made. A summer skin of grisea from Sumas, British 
Columbia (no. 88008, Biol. Surv. Coll.), is identical in coloration with 
certain summer specimens of olympica. Comparison of this skin with 
the entire series of examples of olympica, however, demonstrates the 
presence of more blackish dorsally in the latter and more of a tendency 
toward a brown wash ventrally. 

Aplodontia rufa grisea is smaller than Aplodontia californica 
columbiana in both external and cranial characters, with slighter tend- 
ency to whitish ventrally ; tendeney for zygomatic and mastoid widths 
to be more nearly equal, lesser tendeney to accentuation and approxi- 
mation of temporal ridges, and incisive foramina tending to be shorter. 
From A. c. rainieri, grisea is separated by smaller size and by having 
basilar length averaging less, maximum length of nasals in grisea 
about equal to minimum in rainiert, nasals tending to be narrower 
anteriorly and posteriorly, interpterygoid fossa averaging narrower, 
mastoid width of cranium averaging less, conformation of rostrum 
more plane. 

Material—Fourteen specimens, as follows: one (no. 88008, Biol. 
Sury. Coll., taken by A. C. Brooks) from Sumas, British Columbia; 


1916 | Taylor: Two New Aplodontias from North America 499 


four (no. 94348, Biol. Surv. Coll., and nos. 6822, 6824, and 6825, Mus. 
Comp. Zool., all taken by A. C. Brooks) from. Chilliwack, British 
Columbia; one (no. 6823, Mus. Comp. Zool., taken by A. C. Brooks) 
from Mount Baker Range, British Columbia; one (no. 7388, Field 
Mus. Nat. Hist., taken by L. M. Turner) from Ravenna, Washington ; 
three (nos. 3749-3751, Mus. Vert. Zool., taken by Frank Stephens) 
from Renton, near Seattle, Washington; four (nos. 7385-7387, Field 
Mus. Nat. Hist., taken by E. C. Starks, and no. 3748, Mus. Vert. Zool., 
taken by Frank Stephens) from Seattle, Washington. 
Measurements.—Of type (adult female): total length, 330 mm.; 
tail, 25; hind foot, 55; basilar length, 59.1; leneth of nasals, 26.7; 
width of nasals, 11.7; length of audital tube, 19.6; length of incisive 
foramen, 7 
fossa, 5.3; mastoid width, 52.5; alveolar length of superior cheek teeth, 
19.6; distance between infraorbital foramina, 15.2; mandible, trans- 


.2; zygomatic width, 57.5; greatest width of interpterygoid 


versely across angular process, 22.3; greatest length of mandible, 49.5. 

Remarks—A plodontia rufa grisea of the Puget Sound and Sumas 
districts is apparently nearest to A. r. olympica of the Olympie 
Mountain region. The characters of the Puget Sound form intergrade 
not only with those of A. r. olympica but also with those of A. r. rufa 
of the lower Columbia River. 

Specimens referred to Aplodontia rufa grisea from Sumas, Chilli- 
wack, and the Mount Baker Range, British Columbia, show tendencies 
toward greater dimensions in some respects, and are otherwise un- 
typieal, indicating a slight geographic variant in that region. 

Although Aplodontia rufa grisea is not strongly marked at best, 
its relationships would seem to be indicated more accurately by its 
recognition as a subspecies of rufa than by its direct reference to that 
form. 


Aplodontia californica columbiana, new subspecies 
British Columbia Mountain Beaver 


Type.—Male adult ; no. 1899, Coll. KE. A. and O. Bangs, Mus. Comp. 
Zool.; Roab’s Ranch, Hope, British Columbia, June 14, 1894; col- 
lected by W. C. Colt; stuffed skin, with skull and jaws, in good con- 
dition, except skin with foreleg injured in trap, skull with left audital 
tube, region of foramen magnum, and hamulars somewhat injured. 

Diagnosis —Similar to Aplodontia californica rainieri, but larger ; 
males having white patches beneath; nasals tending to be longer and 


500 University of California Publications in Zoology | Vou. 16 


broader, zygomatic arches heavier and more expanded at posterior 
root, caliber of audital tubes tending to be less, a more pronounced 
tendeney apparent toward approximation of temporal ridges, less of 
a hollow in skull outline dorsally (looking at skull in side view). 

Comparisons.—In coloration dorsally Aplodontia californica colum- 
biana is not conspicuously or appreciably different from A. c. rainieri. 
The tendency observable in males of columbiana to have irregular 
patches of white hair beneath is not expressed in available examples 
of rainieri. Total length externally is nearly 14 per cent greater in 
columbiana, figured on the basis of the total length in the type of 
rainieri and that in nine specimens of columbiana. The following 
cranial measurements average greater In columbiana than in rainieri: 
length and width of nasals, length of incisive foramina, zygomatic 
width, mastoid width, and greatest length of mandible. Available 
material shows the length of nasals in columbiana to be absolutely 
greater than in rainiert. 

From Aplodontia californica californica the British Columbian 
form is distinguished, among other characters, by its larger general 
size, and by having, in the usual instance, zygomatic arches less square 
anteriorly, lighter in weight and more expanded in the region of the 
posterior root, caliber of audital tubes less, and external auditory 
meatus of different shape. 

Comparison with Aplodontia rufa grisea is perhaps not strictly 
necessary. From this race A. californica columbiana is separated by 
the more pronounced tendency observable in columbiana to have irreg- 
ular white patches beneath, by larger size in general, different outline 
of nasals, heavier zygomatie arches and their greater expansion pos- 
teriorly, lesser caliber of audital tubes, and different outline of ex- 
ternal auditory meatus. The last-mentioned character is a conspic- 
uous one, the meatus in columbiana being pinched up anteroposteri- 
orly, making the dorsoventral diameter of the meatus greater than 
the anteroposterior. In grisea the outline of the meatus approximates 
a circle. 

Material—Nine specimens, all from British Columbia: four (nos. 
1892-1895, Mus. Comp. Zool., taken by W. C. Colt) from Lake House, 
Hope; five (nos. 1896-1900, Mus. Comp. Zool., taken by W. C. Colt) 
from Roab’s Ranch, Hope. 

Measurements—Of type (adult male): total length, 470 mm.; 
tail, 22: hind foot, 20; width of nasals, 13.0; length of incisive fora- 
men, 7.8; zygomatic width, 66.0; mastoid width, 61.2; alveolar length 


— 


OOO eee 


oe 


1916] Taylor: Two New Aplodontias from North America 501 


superior cheek teeth, 19.5; distance between infraorbital foramina, 
16.6; mandible, transversely across angular process, 25.2; greatest 
length of mandible, 50.2. 

Remarks.—Aplodontia californica columbiana is a strongly marked 
form, being in general the largest species of mountain beaver de- 
seribed up to the present time. It averages above the maximum in 
any other species or subspecies in length of nasals, zygomatie width, 
and, with only two exceptions, in mastoid width. In total length 
externally it averages decidedly above the maximum in any other race. 

Although it is true that so far as at present known there are 
broad gaps between the geographic ranges of the three subspecies of 
Aplodontia californica, and that no intergradation between them has 
been demonstrated, nevertheless their mutual relationships as well 
as their status with reference to other forms of the genus would seem 
to be best shown by referring them all to this species. 


Transmitted February 28, 1916. 


UNIVERSITY OF CALIFORNIA PUBLICATIONS— (Continued) 


12. Batrachoseps major and Bufo cognatus californicus, New Amphibia 
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‘ Vol. 12, No. 17, pp. 503-544, pls. 19-22 August 12, 1916 


rk . 4, 
NOTES ON THE LOCAL DISTRIBUTION AND 
HABITS OF THE AMPHIBIANS AND 
REPTILES OF SOUTHEASTERN 
“CALIFORNIA IN THE VICINITY 

OF THE TURTLE MOUNTAINS 


tional Mus 


BY 


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UNIVERSITY OF CALIFORNIA PUBLICATIONS 
IN 


ZOOLOGY 
Vol..12, No. 17, pp. 503-544, pls. 19-22 August 12, 1916 


NOTES ON THE LOCAL DISTRIBUTION ANI 
HABITS OF THE AMPHIBIANS AND 
REPTILES OF SOUTHEASTERN 
CALIFORNIA IN THE VICINITY 
OF THE TURTLE MOUNTAINS 


BY 
CHARLES LEWIS CAMP 


(Contribution from the Museum of Vertebrate Zoology of the University of California) 


CONTENTS 

PAGE 
Introduction: geography, climate, plant and animal associations .................... 503 

Check-list of the species of amphibians and reptiles of the Turtle Mountain 
TCP OTD oS Ee So eS oes A oe ee eter 508 
General notes on the amphibians and reptiles of the Colorado Desert .- 509 
Annotated! list of amphibians and! reptiles)... -s-.c.scsceeeecnceccnsevecensnseeeeceeesveesee 512 

INTRODUCTION 


A prominent feature in the topography of southeastern California 
is a chain of rugged hills known as the Turtle Mountains. These he 
thirty miles west of Parker, Arizona (on the Colorado River), and 
extend north from the San Bernardino-Riverside county line for about 
forty miles. The author’s survey of this particular part of the desert 
extended over a period of nineteen days, in May and June of 1914. 
In July, 1909, a day was spent at Goffs, a station on the Santa Fe 
Railroad thirty miles west of Needles, California, and about sixty 
miles north of Blythe Junction. At the latter point, on the Parker 
branch of the Santa Fe, six days were spent in June, 1914. At all 
these places especial attention was paid to the vertebrates. Altogether 
one hundred and thirteen specimens of reptiles and amphibians were 


Mohave, California ............ 2 


Los Angeles, California ...... 293 15.86 1878-1909 60.3 


504 University of Califorma Publications in Zoology — [Vou. 12 


collected ; these comprise twenty species. Habit notes were recorded at 
time of observation. The Turtle Mountain work was undertaken for 
the purpose of studying a definite fauna in an arid locality, where 
animal habitats are reduced nearly to simple topographical terms. 
(See map, pl. 19.) 

All the specimens obtained have been added to the collection of the 
California Museum of Vertebrate Zoology, and it is under the numbers 
of this Museum that they are individually referred to in this paper. 
To the Director of this institution, Dr. Joseph Grinnell, the writer 
wishes here to express appreciation for criticism and general super- 
vision during the preparation of the manuscript. 

The color names used in this paper are taken from Ridgway’s 
(1912) Color Standards and Color Nomenclature. 

The desert bordering the lower Colorado River, its tributaries and 
distributaries, is not subject to the low winter temperatures prevalent 
in other continental parts of the United States, and it seems reasonable 
to suppose that this high winter (as well as summer) temperature 
might account for the greater number of species of lizards in the 
Colorado Desert than in the higher and colder Mohave Desert, the 
more enclosed and colder Death Valley region, and the plateaus of 
Nevada and Utah, farther north. Certainly the climate of the regions 
mentioned appears to differ in no other decided respect than in winter 
temperature, and this makes it seem likely that the latter must be 
reckoned as an important control in the distribution of saurians. 
Climatic characteristics of the Colorado Valley are: moderate winter 
and very high summer temperatures, during both night and day, 
scanty and sporadic rainfall, very low relative humidity and much air 


CLIMATOLOGICAL TABLE 
(Compiled from Bulletins L and W, U. S. Weather Bureau) 


Altitude Mean annual Mean Mean 
(feet) rainfall annual Mean July 


above = temperature relative temperature 
igh Bee 


Locality sea level inches from humidity 


(57% 8 a.m.) 


aYiam as PAS OT ay eee 141 3.31 1870-1907 72.1 136% 8 p.m.( 90.9 
Ranker Arizona 35) 4.12 1894-1907 WO! . (25 eee 92.5 
Needles, California —.......... 477 3.49 1892-1908* W330) jee 94.0 


751 A079) NSiTS1900" * Se eee 
Mammoth Tank, California 257 1.99 1878-1908 16.04) oe 98.5 
(79% 8 a.m.) 
164% 8 p.m. 


* There are at Needles only eleven days in the year on the average with .01 
inches or more of rain. 


1916] Camp: Amphibians and Reptiles 505 


movement, the last two resulting in a high rate of evaporation. No- 
where in the United States are lizards so numerous, both in species and 
individuals, as along the lower Colorado River. 

The Turtle Mountains have never been surveyed. They rise to 
certainly not more than three thousand feet above the lowest surround- 
ing depressions, the latter being about nine hundred feet above sea- 
level. At no points are their summits high enough to support the 
pifion and golden-oak associations such as occur on nearby desert 
mountains. Detrital materials washed from the slopes, and lying up 
against the hill-sides in great fans, nearly bury the lowest passes. In 
many places beds of brown scoriaceous rock cover the alluvial slopes 
and mountain sides. Farther out on the plains are stretches of 
gravelly ash peppered with lapillae. The scoriae are glossy and re- 
fleet much heat on bright days. Igneous rocks make up a large part of 
the mountain mass and there are a few eroded voleanic plugs, up to 
four or five hundred feet in diameter. Other formations occurring in 
some abundance are granites, quartz ledges, and, in the canon bottoms, 
conglomerates; the latter, judging from their position, are of recent 
formation. 

The hillsides are uniformly steep. They are covered with loose 
rocky material and retain but little soil. The canon beds are boulder- 
strewn and often very narrow. At their mouths they are encroached 
upon by alluvial fans. These mesa-like benches are covered with 
scoriae, or with smooth pavements of flat pebbles, and show little soil 
on the surface. The arroyos dissecting the mesas are uniformly broad 
and sandy, giving evidence of copious, even though rare, floods. Under- 
cut caves in the solidified gravels of the wash banks are frequently 
present at various levels above the wash floor. These holes furnish homes 
for the rough-scaled lizards and the desert wood rats (Neotoma), the 
latter industrious rodents being responsible for loosely piled accumu- 
lations of stones and dead twigs in and about the caves (see pl. 21, 
fig. 5). 

The typical desert plains vegetation is not supported on the leached 
sand of the washes, which, probably because it is poor in nitrogen, has 
a distinct flora of leguminous plants such as palo verde (Cercidium 
torreyanum), smoke tree (Dalea spinosa), and ironwood (Olneya 
tesota). On-the gravelly surface of the plains grows that most 
abundant of all desert plants, the creosote bush or greasewood (Larrea 
tridentata). Wash deltas and gentle slopes lying above 2000 feet in 
elevation usually bear, in addition to the creosote, stands of the long- 


506 University of California Publications in Zoology [ Vou. 12 


leafed tree yucca (Yucca mohavensis), a smaller species than the 
twisted form so common on the western stretches of the Mohave Desert. 

Long belts of drifting sand traverse the California deserts at vari- 
ous places. These obviously owe their geographic position to the pre- 
vailing direction of wind, in connection with the place of occurrence 
of original stream deposits. Running east and west two miles south of 
Blythe Junction is a wind-drifted strip of fine sand blown into small 
dunes that are being carried eastward across the desert. In places 
this eolhan material has been driven up over mountains and forms 
white easeades down their eastward slopes. The flora of the dunes 
south of the Turtle Mountains is dominated by a large species of 
Ephedra, or desert tea, and bunches of the wiry ‘‘galleta grass.”’ 

The plant and animal associations (see Ruthven, 1907, and Grin- 
nell, 1914) in the region under discussion can be definitely correlated 
with the terrain, and the two together may conveniently be classed as 
environments, of which the following eight, at least, may be recognized 
in this region (see pl. 19). 

(1) The rocky hillside environment, characterized by the low, pale, 
rounded shrub Encelia, and a small species of Ephedra. 

(2) The eafion-bed environment, with the green catelaw (Acacia) 
as its most typical plant. 

(3) The rocky mesa environment, possessing few species of either 
plants or vertebrate animals. 

(4) The wash-bed environment, which is a direct continuation out- 
side the mountains of the canon bed and is in general slightly below 
the level of the rocky mesa. The characteristic association of legumin- 
ous desert trees lives in this environment. 

(5) The low plain environment, distinguished by the predom- 
inance of the creosote bush (see pl. 20, fig. 3). 

(6) The high plain environment, with the tree yueca, in addition 
to the low plain flora (see pl. 20, fig. 2). 

(7) The drifting-sand environment, supporting a large Ephedra 
and patches of galleta grass (pl. 21, fig. 4). 

(8) The canon spring environment, represented in this region only 
by the three or four permanent springs of the Turtle Mountains, and 
characterized by the hydrophilous arrowweed association. 

The water-supply, except after the rare showers, is extremely 
scanty. Behind dykes in three or four of the larger canons feeble but 
lasting springs issue from the wash gravel. At some times of the 
year water can be found under the sand, or exposed in ‘‘tanks,’’ pot- 


1916] Camp: Amphibians and Reptiles 507 


holes hollowed out of the rock by stream action, and shaded by sur- 
rounding cliffs. 

The desert terrestrial animals are for the most part independent 
of water ; those apparently needing it, during the summer at least, are: 
coyotes, foxes, badgers, bob-cats, bighorn, and quail. The animals 
which fly to the water holes are perhaps more numerous, and include 
the bats, most birds, bees and wasps. 

The following table indicating the environmental distribution of 
reptiles in the Turtle Mountain vicinity shows how sharply marked in 


TABLE TO SHOw HasiTatT LIMITATIONS OF AMPHIBIANS AND REPTILES IN THE 
TURTLE MOUNTAIN REGION 


Environments with characteristic plants 


2 ar a 
o s a3 = = 
Boy. g eeegecs nee ee 
Ss 25 8 Sek Sa S95 wh 
ae gS a Sono (om bie 
Species 28 oe 2 = 2 3 3 Bae a8 
1. Bufo punctatus -............. 
2. Testudo agassizii -........ x xt x 
3. Dipsosaurus dorsalis... -... xx* xe xt xe fee 
aU AOL AL AN enenenaseansen-cn= Ere ah aa aS oo aS Rw eee 
5. Callisaurus ventralis 
ventralis ? xX x Xx x xt 
6. Crotaphytus collaris 
Joye WKS yal) eee x xX Xe snes = otis an 
7. Crotaphytus wislizenii....  -... sn x xx xx xx xx 
8. Sauromalus ater -.........-- x xx xe es hb as oe zaze 
9. Uta stansburiana elegans x x x x xx ee 
10. Uta graciosa a ees as — Ne xe OG 
11. Sceloporus magister........ x Xx = x? 3 ? = 
12. Phrynosoma platyrhinos — -... ee xt x xx? q Pe ae 
13. Xantusia vigilis -.......... Ess see sats = fase ne 
14. Cnemidophorus tigris 
(HIGAINE) cetecceseeeereeseenoes x x sec x x x x 
15. Sonora episcopa -......-.----- x : 
16. Sonora occipitalis —....-.  -.-- aes ee sass ME) teres q : 
17. Lampropeltis boylii -.... 2 xe Sei =i Be ae se 2 
18. Bascanion flagellum 
POW AG WM ease -neeaceesesnen= Xe 2 q 2 ? ? 
19. Crotalus cerastes pat Sx eS 9 2 2 x! 
20. Crotalus mitchellii__...... = xe ? x 


Nore.—Large crosses mean abundant; two small crosses, fairly common; one 
small cross, rare. Small numbers opposite crosses refer to number of individuals 
upon which the generalization is based. A question mark (?) means that these 
species oceur elsewhere on the desert in this environment and may do so in 
the Turtle Mountain region. 


508 University of California Publications in Zoology [ Vou. 12 


most cases are the habitat preferences of these easily observed animals. 
An occasional species, like Uma notata and Xantusia vigilis, is re- 
stricted entirely to one kind of habitat, and no species, even though 
abundant, is known to range into every type of environment, although 
there are no apparent physical reasons why it should not do so. Theo- 
retically, therefore, it may be possible for isolation to occur in a very 
limited region of well differentiated habitats and associations. This 
suggests interesting problems concerned with habitat limitations (see 
Grinnell, 1914, p. 102), and the separation of species and genera 
within a faunal area. Of the eight genera of iguanine lizards here 
represented only three range much beyond the limits of the arid 
Southwest. All are closely related and belong to a single group within 
the sub-family. Many of the differences between these genera seem to 
be adaptive and to be correlated with the differences in the nature of 
their habitats. 


CHECK-LIST OF THE SPECIES OF AMPHIBIANS AND 
REPTILES OF THE TURTLE MOUNTAIN REGION 


Little uniformity prevails in the use of vernacular names for most, 
reptiles. Nearly all our species have several book names, and others 
are burdened with appellations belonging to more than one species. 
The matter might well be put into the hands of a committee appointed 
to edit the names that have been used, and to decide upon suitable 
ones, if vernaculars are not to be dropped from the literature alto- 
gether. The common names employed in the following list have been 
selected for descriptive appropriateness or because of long use. The 
authorities here followed in the adoption of vernacular names are 
Ditmars (1907), Grinnell (1908), Grinnell and Grinnell (1907), Meek 
(1906), Merriam in Stejneger (1893), Stone (1911), Van Denburgh 
(1897), and Yarrow (1882). Whenever a descriptive name could be 
found it has been employed ; in a few cases vernaculars in the follow- 
ing list have not been used before. 

. Bufo punctatus Baird and Girard. Spotted Toad 

. Testudo agassizii (Cooper). Desert Tortoise. 

Dipsosaurus dorsalis (Baird and Girard). Desert Iguana. 

Uma notata Baird. Ocellated Sand Lizard. 

Callisaurus ventralis ventralis (Hallowell). Gridiron-tailed Lizard. 


Crotaphytus collaris baileyi (Stejneger). Bailey Collared Lizard. 
Crotaphytus wislizenii Baird and Girard. Leopard Lizard. 


NO ew poe 


1916] Camp: Amphibians and Reptiles 509 


8. Sauromalus ater Duméril. Chuckwalla. 

9. Uta stansburiana elegans Yarrow. Desert Brown-shouldered Lizard. 
10. Uta graciosa (Hallowell). Long-tailed Swift. 

11. Sceloporus magister Hallowell. Rough-sealed Lizard. 

12. Phrynosoma platyrhinos Girard. Desert Horned-toad. 

13. Xantusia vigilis Baird. Desert Night Lizard. 

14. Cnemidophorus tigris tigris Baird and Girard. Desert Whip-tailed Lizard. 
15. Sonora episcopa (Kennicott). Texas Ground Snake. 

16. Sonora occipitalis (Hallowell). Desert Burrowing Snake. 

17. Lampropeltis boylii (Baird and Girard). Boyle King Snake. 

18. Bascanion flagellum frenatum Stejneger. Red Racer. 

19. Crotalus mitchellii (Cope). Pallid Rattlesnake. 

20. Crotalus cerastes Hallowell. Sidewinder. 


GENERAL NOTES ON THE AMPHIBIANS AND REPTILES 
OF THE COLORADO DESERT 


The most common amphibians of the Colorado Desert are toads. 
Three hardy species of Salientia (Bufo cognatus cognatus, Bufo 
alvarius, and Bufo lentiginosus woodhousti) inhabit the Colorado 
River valley, and one other species, Bufo punctatus, though not known 
to be found along the river (south of the Grand Canon), occurs in 
some of the desert springs, mountain cafions, and stream-beds (see 
(Stone and Rehn, 1903, p. 34, and Stejneger, 1893, p. 219). The 
Great Basin leopard frog, Rana pipiens brachycephala, also lives in 
and along the Colorado River. 

Recent reptiles are represented on the Colorado Desert in Cali- 
fornia and Arizona by one species of tortoise, one gecko (Coleonyx), 
one night lizard (Yantusia), the ‘‘Gila monster’’ (Heloderma), one 
teiid (Cnemidophorus) and thirteen iguanine lizards, one blind snake 
(Rena), six colubrine and two crotaline snakes. A mud turtle (Kinos- 
ternon) occurs along the lower Colorado River. It might be noted 
here that Stephens (1914, p. 134) denies reports of the presence of 
“Gila monsters’’ in southeastern California. Neither is there any 
authentic record of Holbrookia or Elaps from California. The igua- 
nine lizards include nine genera, the ecolubrine snakes five, and the 
crotaline snakes two. Among the genera of iguanine lizards are found 
herbivorous, carnivorous, omnivorous, insectivorous and myrmeco- 
phagous forms. Most species of desert lizards live on the ground 
among rocks or on sand and some spend their time in bushes and 
desert trees. In comparison with the lizards, the snakes of the Colo- 
rado Desert are few in number of individuals. 


510 University of Califorma Publications in Zoology [ Vou. 12 


Glanecing over the characters which distinguish desert reptiles we 
find that great power of lecomotion is possessed by all but a few of the 
forms. The latter imelude the rattlesnakes and ‘‘Gila monsters,’’ 
which have special means of protection, and the chuckwalla, gecko and 
night lizard, which seem to be restricted to habitats where they find 
immediately accessible retreats. The desert whip-tailed lizard, the 
gridiron-tailed lizard, the desert iguana, the ocellated sand lizard and 
the red racer all live in the open and forage at great distances from 
cover. The leopard lizard, probably the greatest runner of all, preys 
on other swift saurians, and the Bailey collared lizard does likewise. 
The latter, however, lives among the rocks where cover is always 
quickly available, and its agility is doubtless associated with its ear- 
nivorous habits. In many of the swifter lizards the tail is held above 
the ground and used as a counter-balanece while running. 

Loose sand characterizes one of the typical desert habitats. Many 
remarkable adaptations in the desert reptiles are developed or pre- 
served by this kind of an environment. In the burrowing snake, 
Sonora occipitalis, in the gridiron-tailed lizard, in the ocellated sand 
lizard, and in the desert horned-toad, the rostrum is specially devel- 
oped, protruding beyond the mouth. These reptiles swim into the sand 
by lateral movements of the head, and the lizards mentioned do not 
employ the fore feet in digging as does the Cnemidophorus, or whip- 
tail group. The eyelids of the burrowing lizards are fringed and meet 
tightly along thickened edges and the nostrils are collapsible. In the 
sand-loving desert horned-toad the ear opening is usually covered by 
the granular integument, while in other species of the same genus 
it is not. 

The toes on both front and hind feet of the most characteristic 
American sand lizard, Uma notata, are broadened by a fringe of 
elongate scales, and in this respeet parallel certain species native to 
desert regions in other parts of the world. 

Keen vision and alertness are attributes of most desert lizards. In 
Dipsosaurus, Callisaurus, and Uma, alert lizards inhabiting open 
plains with sparse vegetation, the head is held aloft and the body is 
propped up on the fore limbs when the creature gazes about (see pl. 22, 
mee Te 

The tail-dropping faculty of the species of lizards here discussed 
varies from almost perfect autotomy in the night lizard to entire 
absence of this function in the chuckwalla. The latter uses its tail as 
an organ of defense. Cnemidophorus, Uma, Callisaurus, Uta, Scelo- 


1916] Camp: Amphibians and Reptiles 511 


porus, and Dipsosaurus are known to part with their tails, the first 
easily, the last with difficulty. In Sauromalus, Crotaphytus and 
Phrynosoma autotomy does not oceur. 

Notable in the case of many species, for example the desert horned- 
toad, the gridiron-tailed lizard, the ocellated sand lizard, and the side- 
winder, is the close resemblance between the color of the reptile as 
viewed at a little distance and the color of its surroundings (see Atsatt, 
19138, p. 49; Stejneger, 1890, pp. 114-115; and Yarrow, 1875, p. 512). 
Color change in lizards may be seasonal, as in the orange-red nuptial 
colors of female Uma, Callisaurus, and Crotaphytus wislizenii, tran- 
sitory, as the colors exhibited by the long-tailed swift when under ex- 
citement, or adaptive, as the dusky shades of gridiron-tailed lizards 
and horned-toads when found on dark soil (see in this connection 
Franklin, 1913, and Stejneger, 1890). 

The femoral pores in lizards secrete a substance which seems to be 
of use during the breeding season. In many desert reptiles the period 
of greatest sexual activity, from April to July, is accompanied in the 
male by an abundance of the waxy secretion of these glands. Lizards 
in which a decided difference was found at this time of year between 
the size of the femoral pores in males and females were the chuckwalla, 
the rough-sealed lizard, the gridiron-tailed lizard, the long-tailed swift, 
the Bailey collared lizard, the ocellated sand lizard, and the desert 
whip-tail. The pores function most actively therefore in adult males 
during the early summer. 

The daily and seasonal habits of desert reptiles seem to show some 
correlation with the distribution of the species. Those forms lke 
Dipsosaurus dorsalis, Uma notata, Sauromalus ater, Uta graciosa, and 
Crotalus cerastes, which have ranges restricted to the hottest parts of 
the desert, may be seen abroad at all hours of the day during the sum- 
mer and seareely ever appear in the winter. Uta stansburiana, which 
is the only desert lizard ranging into the cooler Pacific coast district, is 
the last to be driven into cover by the cool of winter and the first to 
emerge on warm days. 

It is reasonably certain that most, if not all, of the American desert 
reptiles do not require water. 


512 University of California Publications in Zoology [ Vou. 12 


ANNOTATED LIST OF THE AMPHIBIANS AND REPTILES 


Bufo punctatus Baird and Girard 
Spotted Toad 


Forty-three larvae and recently metamorphosed young of a species 
of toad (no. 5539) are at hand from North Mountain Spring. I have 
available no comparable young of punctatus and so cannot be quite 
certain of the above identification of these young specimens. They are 
about half the length of Bufo halophilus of corresponding stage of de- 
velopment; they are broad-headed and dotted on the back with many 
small tubercles (red, in life) surrounded with indistinet black circles. 
The underparts are white and gray in alcohol (bronze-colored beneath, 
in life). The lower labial teeth in the tadpoles are in two long rows 
and one short one. The first of these (anterior), in the specimens 
examined, contained 126 teeth, the second 120, and the third 31 teeth. 
The comb-like upper lip is made up of from 55 to 65 solidly united 
teeth, and the lower lip of from 78 to 95 teeth (in three specimens 
examined). Total lengths, in millimeters, of nearly grown tadpoles: 
25.4, 24.5, 23.0, 25.0, 25.0. Total lengths of recently metamorphosed 
young: 10.3, 9.7, 10.2, 10.5, 9.4. 

The tadpoles were found on May 28 in a water-hole at the south end 
of the Turtle Mountains, five miles from any other spring and thirty 
miles from the nearest permanent stream. A search revealed some 
young toads huddled together in the crevices of planks about the pool, 
and some in wet sand nearby. The toads were active and apparently 
well fed, having an abundance of small flies to prey upon about the 
foul water. The sluggish tadpoles, swimming slowly to the surface 
of the murky, red pool were easily taken in the hand. 

The sporadic occurrence of this amphibian in the driest of North 
American deserts is noteworthy. 


Testudo agassizii (Cooper) 


Desert Tortoise 


Four tortoises were collected on July 20, 1909, at Goffs. None was 
seen at the south end of the Turtle Mountains, though their dens and 
broken skeletons were found there. 

A large living specimen was obtained in 1908 at Mecca, Riverside 
County, California. It was said to have been taken in the Cottonwood 


1916] Camp: Amphibians and Reptiles 513 
Mountains twelve miles northeast of the locality mentioned. This 
specimen was measured on February 23, 1909, and again on May 21, 
1912. It was thirteen inches in length and had grown scarcely at all 
during these three years. It differed from the examples taken at Goffs 
in the color of the iris, which was yellow rather than brown as in the 
more northern examples. A desert tortoise was found near Victor- 
ville in April, 1906, and another at Barstow on March 16, 1914. In 
the Museum are additional specimens (nos. 3550 and 3609) from 
Kramer, San Bernardino County, and one-half mile east of Mohave, 
Kern County, California. A dead tortoise was seen three miles south 
of Palmdale, Los Angeles County, California, on July 24, 1914. Tor- 
toises, though widely distributed, appear to be common at few places 
in the desert. 

From what has been published concerning Testudo polyphemus of 
the southern states, a close relative of the desert tortoise, it appears 
that the most western North American species and the gopher tortoise 
of the East do not differ much in their general habits. Both live for 
the greater part of their lives in holes, both inhabit sandy or other 
loose soil suitable for the burrowing of their dens, and both dig their 
own refuges by the crude methods at their command. Like all known 
members of the genus, both are herbivorous. 

About a mile north of Goffs station a number of desert tortoise 
burrows were found, some of which were occupied. The holes were 
dug slantingly into the rather firmly packed sand and gravel, usually 
at a grade of about forty-five degrees. Often the places chosen were 
under creosote bushes and in the banks of small dry washes. The 
tunnels were from two to eight feet long, with a slight widening at 
the bottom. They varied in diameter with the size of the tortoise that 
inhabited them, being in every case about the shape of a longitudinal, 
vertical section of the animal’s shell. Sometimes the tortoise could be 
seen lying at the inner end of its burrow. In the deeper holes a stick 
thrust in would reveal the presence of the creature which, lying partly 
outstretched, would draw up its feet and head when it felt the touch ; 
and this diminution of respiratory space beneath the shell would be 
accompanied by a noisy expiration like the rapid blowing of a bellows. 
When seized by the back of the carapace to be drawn out the tortoises 
would sometimes stick fast in the holes, hooking their crooked front 
legs into the sand. One deep burrow, otherwise empty, contained the 
broken halves of two white, hard-shelled eggs which appeared to have 
been spherical and about an inch in diameter. No brush or food of 


514 University of California Publications in Zoology [ Vou. 12 


any kind was found in any of the dens. Late in the afternoon of a 
hot July day a large tortoise was surprised in the act of coming out 
of its burrow. When it saw me it turned immediately and ambled 
back to safety. 

Desert tortoises are said to come out in great numbers after thun- 
dershowers. But this is by no means the only time of their activity, 
for they appear to wander abroad at all seasons, frequenting rocky 
and uneven as well as level ground. One meets with them plodding 
steadily across-country, occasionally stretching their short necks down 
over the pointed extremities of the plastron and testing the ground 
with the sensitive tip of the snout or stepping aside to crop some small 
annual plant growing in the shade of a boulder. Their usual gait 
does not carry them along at a rate of more than four or five miles a 
day (twenty feet a minute by test), and they live at such great dis- 
tances from water that in places it would seem impracticable for them 
to get a drink from one year’s end to another. Sometimes when 
roughly handled a tortoise will void the contents of its cloaca and blad- 
der. The foeces are black and about the size of those of the fox. 

The author kept a number of desert tortoises at his home for a 
time in an enclosure out-of-doors. The captive tortoises paced their 
pens and frequently attempted to dig out under the fence. They 
were not able to burrow into hard ground, but in a gravel bank they 
would scrape away alternately with the fore limbs and when the hole 
became deep enough would turn around and push the dirt out with 
their shoulders. They grazed contentedly upon the lawn or in patches 
of green weeds, and also ate lettuce or cabbage leaves thrown to them. 
They were never active unless warm and seldom so even then, coming 
out in bright sunshine to lie outstretched or seeking the warmth of a 
stove or radiator in a room. They were handled a great deal and were 
never known to open their mouths to bite. They soon became used to 
human surroundings and would pursue their accustomed activities 
with people about. Some of their habits are worthy of notice. 

When one tortoise meets another in the course of its journeying 
each, whatever the sex, nods its head rapidly up and down as if in 
salutation, and sometimes noses are touched before passing along. If 
two males happen to meet, a fight is likely to ensue. After the pre- 
liminary nodding the tortoises separate a little distance and then rush 
toward one another with the heads drawn part way into the shell. The 
combatants meet head on and the curved horns projecting from the 
anterior end of the plastron are butted rather violently against the 


1916] Camp: Amphibians and Reptiles 515 


adversary, but do him no damage except sometimes to turn him upon 
his back ; he may then struggle for some time with one fore leg vibrat- 
ing vigorously in the air and the other pawing for a foothold in the 
ground before he ean right himself. 

The males court their mates by biting them gently around the edges 
of the shell. During copulation the male stamps his hind feet and 
utters a mechanical grunt with the head hooked over the end of the 
plastron and the mouth half open. 

Stephens (1914, p. 135) writes that teeth marks are sometimes seen 
on shells of living tortoises and believes that the shells ‘‘generally 
prove too hard for the coyotes.’’ The younger tortoises are soft-shelled 
and delicate. They probably fall prey in numbers to raptorial mam- 
mals and birds. The old ones are a favorite delicacy among Indian 
and Mexican section-hands who live with their families along the rail- 
road lines. Some tortoises kept as curiosities at Needles on a grass plot 
in front of the Santa Fe hotel are thought to have been gradually de- 
pleted by the inroads of the Indians, many of whom lounge about the 
place. 


Dipsosaurus dorsalis Baird and Girard 


Desert Iguana 


Four desert iguanas (nos. 5499-5502) were taken in the vicinity of 
Blythe Junction. In two specimens the rostral plate is separated from 
the nasals by one row of scales, and in the other two individuals by 
two rows. The femoral pores are 18 in two thighs, 21 in four, and 22 
in one; being 18 right: 18 left once, 21:21 twice, and —:22 once. 

The ground color varies slightly from light grayish to yellowish. 
One specimen has wide, brown reticulations enclosing lighter spots on 
the sides and back. A smaller example has narrow brown dots and 
dashes in place of the broad reticulations. 

The total length of the largest individual is 360 millimeters; the 
tail is partly regenerated and measures 242 millimeters. 

These round-nosed, large-tailed lizards are fairly common in the 
low plain environment in the sandy tracts south of Blythe Junction, 
and in the washes traversing the alluvial slopes about the Turtle Moun- 
tains. They appear to avoid rocky ground, being absent from the hill- 
sides and mesas. They are shy when approached and run swiftly, with 
tail slightly raised, to the shelter of a bush, or into a chipmunk’s or 
kangaroo-rat’s burrow. When wounded they puff themselves up till 


516 University of California Publications in Zoology [ Vou. 12 
their sides become taut, and may then be pulled from a small hole only 
with difficulty. With curiosity aroused they prop themselves high on 
their fore limbs, attentively viewing the passer-by and seldom ‘‘show- 
ing off’’ with up-and-down movements of the body. 

These lizards are phytophagous and may be sometimes surprised 
in the act of raiding the young leaves of low bushes, in the upper 
foliage of which they forage during the hottest part of the day. The 


stomach of a medium-sized individual contained two grams of the 


leaves and fruit of a malvaceous annual, Sphaeralcea ambiqua. 

Remains of a desert iguana were found below the cliff-side nest of a 
prairie falcon where, on June 6, 1914, two nearly grown young falcons 
set up a squawking chorus at my approach. 


Uma notata Baird 
Ocellated Sand Lizard 


In the drifting sand two miles south of Blythe Junction, thirteen 
specimens of this extraordinary species were obtained (nos. 5444— 
5456). Some have the identical characters ascribed only to Uma rufo- 
punctata by Cope (1900, pp. 279-281), while the smallest specimens 
are referable to Uma notata as described by Cope (1900, pp. 277-279) 
and Baird. It seems almost certain that the separation indicated is 
based on nothing more than individual and age differences. It ap- 
pears also likely that the other two described species of this genus are 
but extreme variations of notata. Both scoparia and inornata were 
described from localities which lie within the probable range of notata, 
and in each case the type is the only known example. Three specimens 
at hand from Imperial County, California (doubtless near the type 
locality of inornata), have distinct spots on the belly and are not dif- 
ferent from individuals in the Blythe Junction series. Most of the 
characters supposedly distinctive of scoparia, including the diamond- 
shaped dorsal seales and black dorsal spots, are expressed in indi- 
viduals of the present series. The increased number of femoral pores 
in the type specimen of scoparia finds a parallel in individuals of other 
species of lizards and is probably also in this ease not to be considered 
of systematic importance. 

The femoral pores (including the accessory row) in the series col- 
lected near Blythe Junetion number 25 in one thigh, 27 in one, 28 in 
one, 29 in three, 30 in four, 31 in four, 32 in four, 33 in one, 34 in 
four, and 35 in two; being ¢ 35 right: 35 left once, 2 34:34, 9 34:29, 


1916] Camp: Amphibians and Reptiles 517 


J 33:31, J 32:32, J, 2 31:32 twice, 2 30:34, $ 30:30, 2 29:31, 2 29:30, 
2 27 :28, and 9 —:25. They are small in the eight females and medium- 
sized in the five males of the lot. In six eases they are in one and in 
seven instances in two rows. An accessory row is not present in the 
thighs with 25, 27, 28 and 35 pores, but is represented in most of the 
intermediate thighs and contains from one to three pores. 

The supra-ocular rows of scales are 8 in one individual, 9 in five, 
10 in five, 11 in one, and 12 in one. The loreal rows, counted where 
they join the line of the orbit, are 7 in six and 8 in seven specimens. 


The labials are strongly keeled ; they number 7 in one, > in two, 


1 14 
8 10; : 
Ta i; in two, and 4° in one. The 


in one, » G6 in two, - 7: 
keeled suborbitals are 6-6 in one, 6-5 in two, 5-5 in four, 5-4 in 
one, 4-4 in two, 3-4 in one and 3-3 in one. The occipitals are 
separated from the small supra-oculars by 3 to 4 rows of scales. The 
scales on the outer edge of the gular fold are smaller than those in 
the middle, which in turn are larger than those on the throat. Ear 
lappets are 4 to 6. The points on the fringes of the lower eyelids are 
longer than those on the upper. 

In all the specimens of the present lot a black spot from four to 
twelve millimeters in diameter is present on each side of the abdomen 


. 9 . 
in two, |; In two 


(see pl. 22) ; in some of the smaller specimens these spots are narrowly 
margined with green. In three the black markings on the throat are 
indistinct or reduced; in others two or three crescents and one to 
three V-shaped marks are present on the throat (see pl. 22, fig. 6). The 
black spots on the tail number 0 to 7; they are apparently not present 
on regenerated tail-tips and are never continued on the dorsal surface 
as in Callisaurus. Four of the smaller specimens are dorsally of the 
greenish notata type of coloration, with the ground color of the back of 
a pale greenish blue, near pale glaucous blue. Five of the intermediate 
examples are spotted with cinnamon on a background of greenish 
yellow, with the ocellations and brown crescents ascribed to rufo- 
punctata; and the four largest individuals (all males) are of the 
scoparia type of coloration, being covered with rich black ocellations 
each bordered with a fine line, one scale wide, of orange-rufous. Each 
spot of light color is centered with a black dot, itself encircled by a 
narrow orange-rufous ring. The ground color is maize yellow. The 
eyelids and sometimes the sides of the head and tail in females are 
tinged with orange. One so colored contained three eggs, each 9 
millimeters in diameter. Another contained one egg 18 millimeters 
long. Black spots are present behind both femurs in only two indi- 


518 University of California Publications in Zoology [ Vor. 12 


viduals and behind one femur in three; they are in all cases very small 
and obscure. 


MEASUREMENTS IN MILLIMETERS OF Uma notata FROM NEAR BLYTHE JUNCTION, 
RIVERSIDE COUNTY, CALIFORNIA 


Sex and number .... 5445 54529 5450¢ 54499 54569 54549 54519 54539 


Total length .... 215 181 193 172 201 180 202 139 
Tail length ... 112 94 96 90 108 94 98 69 
Body length .......... 103 76 97 82 93 86 104 70 
Hand toot ee 31 27 30 27 32 27 32 26 
Base of 5th to end 

of 4th toe. 27 23 26 22 27 24 27 20 
Snout to ear = 19 18 19 18 19 18 18 15 
Head width .......... 16 15 17 15 als 15 18 12 


The ocellated sand lizard has so far been taken only at a few locali- 
ties within a cireumscribed area on the deserts of the southwest. It 
is strictly confined to belts of wind-blown sand, and in the vicinity of 
Blythe Junction was never seen beyond the borders of a narrow zone 
of sand dunes two miles south of the railroad. Many of these shy 
lizards were abroad in the hotter part of the day, scurrying over the 
fine sand, with a cloud of dust in their wake, or foraging beneath 
squaw-tea bushes on the dunes. Seldom was an individual taken un- 
awares, and it was found difficult in most cases to approach an alert 
animal close enough for a successful shot with the .82 caliber auxilliary. 
Plate 22, figure 7, from an animal in captivity, shows the posture when 
fully alert. 

The speed attained by these heavy lizards on loose sand is not so 
great as that of Crotaphytus, Cnemidophorus and Callisaurus in the 
same situations, despite the broadening fringe of scales on the toes. 
When alarmed they make for the nearest dune and turn behind it to 
enter a Dipodomys or Citellus hole or to bury themselves quickly in 
sand as the gridiron-tails do, the broad nose of which lizards their own 
shovel-like snout resembles. The tracks of the hind feet of a Uma 
running at top speed are five to six inches apart; and the deep imprints 
of these members indicate that most of the work is done by the hind 
legs, the fore limbs being merely used to balance the creature. The 
tail is curled upward while running, as in Callisaurus. 

Their curious color patterns, though they may seem unduly strik- 
ing when viewed in the specimen in hand, really harmonize in strong 
light with the buff tint of the sands, and the lizards are seldom de- 
tected until they begin to move. 


1916] Camp: Amphibians and Reptiles 519 


One stomach was filled with a great number of ants. Another con- 
tained two grasshoppers entire, one large hemipter, eight red ants, 
two brown ants, two beetles, a pebble, and several pieces of vegetation. 
Another held seven brown and seven red ants, one beetle, several para- 
sitic nematodes, and two fresh leaves and the terminal bud of a plant. 
One specimen when shot had a plant stem in its mouth. 


Callisaurus ventralis ventralis (Hallowell) 
Gridiron-tailed Lizard 

Highteen specimens of this abundant lizard were shot (nos. 5457— 
5474). All have four dark patches on the belly, two on each side, and 
are in other ways typical. The femoral pores are 12 in one thigh, 13 
in one, 14 in four, 15 in six, 16 in six, 17 in eight, 18 in five, and 19 
in three; being J, 9 19 right 18 left twice, 9 18:18 once, ¢ 17:19, ¢ 
17:18, § 17:16, ¢ 17:15, g 16:17 twice, ¢ 16:16, 9 16:14, # 15:17, 
3, 2 15:14 twice, ¢ 14:15, 9 13:15, J 12:15 and ¢—:17. Among the 
thirteen males five have large femoral pores, six have medium-sized 
ones, and two have small ones. All the five females have small pores. 
The males have large postanal scales, the females small ones. 

In the examples before me the color above grades from pale smoke 
eray, with white on top of the head and white in spots down the back, 
to neutral gray with the usual dark dorsal markings. A specimen 
taken in the zone of drifting sand below Blythe Junction is the hght- 
est of the lot, and some taken upon a mesa covered with brown lava 
are among the darkest of the specimens represented. It would appear 
that in this lizard, as in Phrynosoma, the tone of color is changeable 
in the individual to suit the surroundings. The throat is dusky in 
some specimens and light in others. There is a reddish spot behind 
the arm in the females. The females also all have pink saes beneath 
the throat which are not ‘‘inflated,’’ but are sometimes drawn down 
by inuscles connected with the hyoid apparatus. The pink throat saes 
are present in only three of the males in the series at hand. The 
underparts (with the exception of the two black wedges and the blue 
and green patches of the males) are white and not yellow as in speci- 
mens taken at Barstow, farther west. The tail bands number 4 in one, 
6 in three, 7 in three, and 8 in eleven specimens. The anterior three 
or four of the ventral tail spots are sometimes entirely blue, and the 
posterior bands are often margined with blue below. Several of the 
largest males are nearly equal in length, but do not approach in size, 
individuals collected at Barstow in March, 1914. In the largest speci- 


520 University of California Publications in Zoology [ Von. 12 


men from Blythe Junction (no. 5472), the total length is 194 mm., 
tail length 111 mm. A specimen from Barstow, San Bernardino 
County (no. 5385), is 223 mm. in total length, and 130 mm. in length 
of tail. 

This lizard is abundant on the open desert around the Turtle 
Mountains. It does not occur on the rocky hillsides, and even in the 
sandy canon bottoms is found but sparingly. It is on the open 
stretches of desert dotted with creosote bushes that this species is 
typically at home; and here it outnumbers all the other diurnal verte- 
brates combined. Individuals may be observed bobbing up and down, 
switching the tail from side to side, walking jerkily along with the 
tail curled over the back, or running with such speed that the eye can 
barely follow. The writer estimated that one of these swift lizards 
covered a distance of 90 feet in four seconds, which would be travel- 
ling at a rate of about fifteen miles an hour. The lizards can stop and 
start with the most confusing abruptness, and rarely run straight 
away but describe a circle when pursued. When tired out they may 
crouch close to the ground and will then permit themselves to be 
caught; or they may burrow into loose sand by wriggling the head 
from side to side and pushing with the hind feet while the front feet 
remain pressed close to the side. Sometimes when closely pursued 
they enter holes. 

Of eight stomachs examined not one contained plant remains, the 
contents being insects, small pebbles, part of a shed lizard skin, and 
parasitic nematode worms. Perhaps, like some of the geckos, these 
lizards eat their own shed integument. The insects represented in- 
eluded eight Orthoptera, eight ants, and several small Coleoptera. 
Some of the grasshoppers and crickets were of large size (40 mm. long) 
and had been swallowed entire. These lizards sometimes spring a foot 
or more to seize a tempting bait; and I saw one, probably by mistake, 
leap over the edge of an eight-foot wash-bank while jumping for a 
grasshopper in a bush. At Blythe Junction a gridiron-tailed lizard 
was seen regularly at a certain doorstep picking up dead erane-flies 
and other night-flying insects thrown there by the housewife. The 
lizard apparently became so absorbed in picking up, shaking and 
swallowing the gauzy-winged flies that it many times permitted the 
observers to touch it lightly upon the back. 

After sundown the gridiron-tail buries itself in sand, and when 
alarmed as by an approaching team or pedestrian will start up sud- 


denly and dash away. 


1916] Camp: Amphibians and Reptiles 521 


i ‘ : 

Some of the females taken in July contained eggs. Two egos, 18 
by 9 millimeters in the two diameters, were taken from one lizard: 
these had coriaceous coverings and were apparently ready to be laid. 


Crotaphytus collaris baileyi Stejneger 
Bailey Collared Lizard 


Eight specimens (nos. 5480-5487) represent this form in the col- 
lection from the Turtle Mountain region. There are five males and 
three females, the former being readily distinguished by the large 
postanal plates. All have the interorbital scales in two distinct rows. 

The femoral pores are 16 in three thighs, 17 in five, 19 in four, 20 
in two and 21 in two; being 9 21 right: 21 left once, 9 20:20, 4, 9 19:19 
twice, gj 17:17 twice, ¢ 16:17, and ¢ 16:16. They are very small in 
the three females, medium-sized in three males, and large in two of 
the males. Stejneger (1890, p. 105) describes the color of a living 
Bailey collared lizard from the San Francisco Mountain plateau, Ari- 
zona. None of our specimens show strongly marked reticulations as 
do examples from farther east and north. All the females have faint 
indications of whitish cross-bars, and in them the black collar is not 
joined below the neck. The males have the collar connected ventrally, 
and the largest males show not a sign of dorsal cross-bars. The total 
length of the largest specimen, a male, is 310 millimeters, the tail 
length 214 millimeters. 

These grotesque lizards inhabit the rocky slopes of the Turtle 
Mountains in numbers and live also among the rocks about the bases 
of the hills, but they were never seen on the open desert. This agrees 
with observations by Taylor (1912, p. 326) in northern Nevada. Like 
the chuckwallas, the Bailey lizards mount rocky eminences and lie for 
hours in the sun during the hottest part of the day. When approached 
they slip down into crevices or run with alacrity over the roughest 
ground, clearing obstacles up to two feet in height with great leaps. 
The males distend their dark throats when ‘‘showing off.’’ They seem 
hard to kill, and when thought dead will sometimes ‘‘come to life’’ in 
the collecting sack, blinking their yellow eyes and looking ferocious. 
When under excitement the brilliantly colored throat is distended and 
the huge mouth is sometimes opened in anger. Coues (1875, pp. 598— 
599) has recorded interesting observations upon the habits of this 
species. 

One stomach contained two chewed grasshoppers, and another 
three orthopterous insects, more or less chewed, and four small beetles. 


522 University of California Publications in Zoology [ Vor. 12 


Crotaphytus wislizenii Baird and Girard 
Leopard Lizard 


Six specimens of this species were secured (nos. 5488-5493) of 
which two are females and four males, the latter with large postanal 
scales. The femoral pores number 20 in one thigh, 22 in one, 23 in 
three, 24 in four, and 25 in three; being ¢ 25 right: 25 left onee, ¢ 
25 :23, 9 24:24, J, 2 23:24 twice, J 20:22. One of the females taken in 
July still displays the red nuptial coloration: the bars on sides of 
neck, back and hind legs (in ordinary coloration white or yellow) are 
peach red to scarlet ; base and tip of tail beneath are shrimp pink. The 
ground color of this individual is light neutral gray on the lighter 
parts of the back; fuscous spots oceur on the back and sides, with 
bands of the same color on the tail. A female (no. 5489) not exhibit- 
ing red coloration contained one large egg. A large male has scarcely 
a trace of the ordinary reticulation on the back, and the brown dorsal 
spots are reduced to small dots on the body and tail. The longest 
specimen, a male, has the following measurements: total length 364 
millimeters, tail length 260 millimeters. 

The leopard lizard, probably the swiftest of North American desert 
reptiles, was fairly common in the Turtle Mountain district at the 
time I was there. This species does not inhabit the rocky hillsides in 
that vicinity; unlike the Bailey collared lizard, it appears to haunt 
the more level plains and sandy places. Individuals are wary and 
take to retreats, often before it 1s possible to get a shot at them. The 
tracks of the hind feet of leopard lizards running swiftly in sand were 
found to be ten inches apart. 

A grown gridiron-tailed lizard swallowed whole and head first was 
found in one stomach. The flabby sides of the leopard lizard are often 
distended with the remains of smaller lizards which they have run 
down and swallowed. Taylor (1912, p. 348) and Franklin (1914) have 
seen this species eat cicadas, leaping into the air to catch them. 


Sauromalus ater Duméril 


Chuekwalla 


Chuckwallas were common on the rocky sides of gulches at the 
Horn Mine. Seven specimens were taken there, and one specimen in 
a level field of scoriae at Blythe Junction. These specimens (nos. 
5518-5525) show considerable individual variation in width of head, 


1916] Camp: Amphibians and Reptiles 523 


size of scales on side of neck, and coloration, but fall easily within the 
general range of characters given for this species. The femoral pores 
are much enlarged in the males and almost indistinguishable in the 
female, a condition holding in many other species of lizards of the 
locality in the early summer season when collecting was done. ‘The 
femoral pores number 15 in one thigh, 16 in three, 17 in one, 18 in 
three, 20 in two, 23 in one, and 24 in one of the thighs, where counts 
could be made; being ¢123 right: 24 left, 7 20:20, (18:19, (18:16, 
bo —:18, J 16:17, J 15:16, 9 —:—. One individual exhibits an ac- 
cessory row of pores on each side. 

Some of the specimens show a great amount of red on both dorsal 
and ventral surfaces and a few do not; some are banded on the tail 
and some exhibit scarcely a trace of this characteristic. In a young 
specimen there are around the tail four broad, encircling bands of 
brown alternating with three circles of yellow. An adult male (no. 
5520) is colored as follows: top of head dark brown, nearly black, 
with many yellow scales scattered over the oeciput and head and small 
patches of orange in the ear just behind the tympanum; back speckled 
with black seales in lichen-like pattern; about an equal number of 
orange and of yellow scales, evenly dispersed, covering most of back; 
shoulder patches large, dark brown, and dorsal surfaces of limbs dark 
brown; feet spotted with yellow; head and limbs beneath, black; belly 
almost uniform dark moroceo to brick red; tail abruptly lighter than 
rest of body, deep colonial buff, faintly banded with three broad rings 
of deep olive buff. The measurements of the largest male are: total 
length 371 millimeters, tail length 198 millimeters. 

This clumsy, vegetarian lizard is common on the rocky slopes of 
the Turtle Mountains and may occasionally be seen in the patches of 
scoriae out from the base of the range. It was never noticed else- 
where, and having rather feeble powers of locomotion, doubtless de- 
pends upon the security of the rocks to a greater extent than do swifter 
lizards. It lacks the curiosity of smaller species and loses no time in 
slipping to safety at the approach of danger. 

Chuckwallas were seen perching on rocks so hot as to be unbear- 
able to the hand, and big, gorgeously colored males were noted in pur- 
suit of each other over hillsides in the middle of the hottest July days. 
In the latter part of June one pair, male and female, were seen near 
one another. 

The chuckwalla has a curious habit of sticking out the fleshy 
tongue at every few steps when walking along. Like toads and 


524 University of California Publications in Zoology [ Vou. 12 


Phrynosoma, Sauromalus can be made to assume a rigid hypnotic 
posture by gentle rubbing on the belly. In this condition an individual 
may remain half an hour without moving. 

The half-eaten body of a large female was picked up near a nest con- 
taining two young prairie falcons. When attacked in its retreats, the 
chuckwalla inflates itself and lashes the heavy stub-tail about vigor- 
ously. Aside from this it appears to be utterly innocuous, and the 
writer has never known one to attempt to bite even when handled 
roughly. Mr. Dane Coolidge states that the desert Indians, to whom 
the chuckwallas are a delicacy, puncture the lizards’ sides with sharp- 
ened wire in order to deflate them and then draw them from their 
retreats among the rocks. 

Old chuckwallas often have scars on the back caused perhaps by 
crawling about in crevices. One individual secured had lost the front 
foot on one side and the hind foot on the other, and in spite of its mis- 
fortune was lively and had a stomach full of food. One chuckwalla 
was seen up in a small creosote bush from which most of the leaves 
had been stripped. The three stomachs examined contained plant re- 
mains. In two cases the leaves were swallowed entire and belonged to 
a composite (Franseria dumosa) and a spurge (Euphorbia poly- 
carpa) ; the other stomach contained many chewed leaves and stems. 


Uta stansburiana elegans Yarrow 


Desert Brown-shouldered Lizard 


The present writer follows Richardson (1915, p. 473) in the use of 
the above name. The characters ascribed to elegans are exemplified 
fairly well in the nine specimens (nos. 1099-1100, 5526-5532) from the 
vicinity of the Turtle Mountains. The dorsal horizontal scale rows 
number 76 in two specimens, 82 in two, 84 in one, 85 in two, 86 in one, 
and 100 in one. The average number of rows is 84, and the error of 
numbering, as ascertained by repeated counts, is certainly not greater 
than 7 per cent. The average number of dorsal scale rows in six 
specimens of hesperis at hand is 100.6. These averages agree quite 
well with determinations by Richardson of 86.5 and 102 for the two 
subspecies elegans and hesperis respectively. Individuals of the two 
subspecies cannot always be separated by the number of dorsal scale 
rows alone. The present series of elegans is much bluer in dorsal col- 
oration and smaller in size than in the large series of hesperis at hand. 
The femoral pores number 13 in three thighs, 14 in ten, and 15 in 


1916] Camp: Amphibians and Reptiles 525 


four; being 15 right: 15 left once, 14:15 twice, 14:14 three times, 
14:13, —:14, and 13:13. All have scattered bright blue seales over the 
back, and some are green along the sides. In one female the indigo 
patches in the axilla are almost entirely lacking. Both striped and 
spotted types of coloration are exhibited in the present series. The 
total length of the largest example, a female, is 139 millimeters, and 
the tail length is 89 millimeters. 

The tiny desert brown-shouldered lizard oceurs throughout all the 
environments of the Turtle Mountains district except in the tracts of 
eolian sand near Blythe Junction. It seems to be most common in the 
more rocky localities, particularly on the lava fields. Those taken on 
brown scoriae were noticeably bluer than those found elsewhere. The 
stomach of one individual contained several small ants and beetles, 
and one spider. 


Uta graciosa (Hallowell) 
Long-tailed Swift 


Seven specimens of this arboreal lizard were colleeted (nos. 1102, 
5533-5538), one at Goffs and six near Blythe Junction. These in- 
clude five males and two females. All the males have large postanal 
plates and blue patches on the belly. The females do not possess 
either of these characters. The seutellation of the back in the present 
series is typical. The femoral pores number 10 in three thighs, and 
11 in nine; being 9, 4, J, 11 right: 11 left three times, 3, 2 11:— twiee, 
J 11:10, 10:10. The pores are large in the males only. The colors 
are rapidly changeable in life, as deseribed below. A male in alcohol 
has the patches on the belly olympic blue in color, thickly flecked with 
white, and divided by a light line. Each white dot involves one scale. 
The sides are yellowish, and the back grayish with reticulations of dark 
gull gray. The alcoholic females are yellowish beneath. The largest 
specimen, a male, measures 181 millimeters in total length and 127 
millimeters in tail length. A female measures in total length 168 milli- 
meters and in tail length 114 millimeters. 

A number of long-tailed swifts were seen in the vicinity of Blythe 
Junction. Some were in creosote bushes on the open desert, some in 
squaw-tea on the sand dunes, and some on the branches of smoke trees 
in the washes. They like to sun themselves on the topmost twig of a 
bush, hanging motionless and head downwards as though pinned there 
by a shrike. If disturbed they drop to the middle of the bush and 


526 University of California Publications in Zoology [ Vou. 12 


flatten themselves against a limb lengthwise, keeping on the side away 
from the intruder, their wiry tails stretched out stiffly in line with the 
body. When alarmed while on the ground they make for the nearest 
bush and jump up into it, there to dodge actively about among the 
branches, quite unlike their brown-shouldered relatives which usually 
retreat beneath stones or into holes when pursued. The species under 
discussion appears to be active at least till dark in the evening, and 
early in the morning, as well as in the middle of the day. 

A pair was seen copulating on July 13, in the hottest time of the 
day. The two lizards were clinging to the inclined branch of a ereo- 
sote bush and the female was colored for the occasion, being light 
orange with two longitudinal black stripes down the sides and a row of 
black lozenges down the center of the back. The male was grayish 
over the back and yellowish on the sides. The power of color change 
in these lizards is greater and more rapid than in any other Cali- 
fornian reptile. A nearly white male held in my hand changed rapid- 
ly in two or three minutes to yellowish with black cross bands on the 
back, the originally hight greenish ventral patches became blue, and 
a yellow spot appeared under the throat. 

I saw a female of this species swallow a large-winged insect it had 
picked up from the sand. The stomach of a male contained chewed 
plant stems and what appeared to be the broken shells of insect eggs. 
An elongate, white, tick-like parasite was seen affixed head downwards 
in the axilla of a long-tailed swift. 


Sceloporus magister Hallowell 
Rough-sealed Lizard 


Five specimens (nos. 5475-5479) of this brilliantly colored lizard 
were taken. Four are males and one is a female. The femoral pores 
number 13 in five thighs, 14 in two, and 15 in three; being 9 15 right: 
15 left once, J 15:14, J 14:13, ¢ 13:13 twice. The pores are small in 
the female and greatly enlarged in the males. The anterior auricular 
denticulations are long and tapering. 

The coloration of the adult males varies a good deal, and this varia- 
tion is especially noticeable in the vivid ventral colors. One male has 
the neck band pure black, the throat patch olympic blue of the sheen 
of porcelain, the darkest belly scales urania blue of a porcelain cast, 
and the seales laterally on the ventral patch variscite green to Blane’s 
blue. Many of the seales along the sides of the body are edged with 


1916] Camp: Amphibians and Reptiles 527 


rufous and have brown centers. Scales on the sides of the tail are opa- 
line green. The general color of the upper parts and the top of the head 
is deep olive buff to buffy brown. The scales of the dorsal surface are 
edged with dark brown. The ground color of the ventral surface is 
whitish. 

The ventral patches are in three specimens divided and in one 
united. There are no indications of dorsal cross bands or spotting in 
any of the males. The female is marked dorsally with sixteen brown 
patches, about a scale in width. The lower surface is creamy white 
lightly tinted on scattered scales with pale greenish, pale orange and, 
beneath the throat, pale blue. The collar of the female is brown. 

The largest male measures 266 millimeters and the tail length in 
the same specimen is 149 millimeters. 

The rough-sealed lizard was only occasionally seen in the Turtle 
Mountain region. It lives in and beneath catclaw bushes, on boulders 
in the canon bottoms, and in eaves in the undereut wash-banks. In 
only one instance were any of these lizards noticed far from safe re- 
treats. During the forenoon of June 2, while the ground was still 
damp from a recent thunderstorm, two large male rough-sealed lizards 
appeared at intervals on a bare hillside. An explanation of their un- 
wonted fearlessness may have been that they were in an active sexual 
state. 

A large orthopterous insect, somewhat chewed, a fly, a beetle, and 
several other insects were found in one stomach. Another stomach 
contained a grasshopper, a beetle, a lepidopterous insect, several small 
red ants, and some pebbles. A third contained a caterpillar, five 
Coleoptera, one hemipter, three small red ants, the fruit and green 
leaves of a small plant (identity uncertain) and a few dry leaves (per- 
haps taken accidentally). 


Phrynosoma platyrhinos Girard 


Desert Horned-toad 


Five specimens of this species (nos. 5494-5498) were collected, in- 
cluding two females and three males; and these were all the horned- 
toads seen in the Turtle Mountain vicinity. The males can be dis- 
tinguished at once by the large postanal plates. The ear opening is 
covered in all the above specimens. The femoral pores number 7 in 
three thighs, 8 in five and 9 in two; being ¢ 9 right: 8 left once, 2 8:9, 
Jo 8:8, $7:8,97:7. The red of the dorsal parts varies much among in- 


528 University of Califormia Publications in Zoology [ Vou. 12 


dividuals, being light coral red in some and brick red in one, which 
latter also has the sides of the head, the horns, and the tail sprinkled 
with light red. The underparts are pure white, or else spotted with 
from 60 to 70 black dots. Red in the dorsal coloration occurs in both 
males and females. The total length of the longest specimen, a male, 
is 137 millimeters, and the tail length is 52 millimeters. Another male 
measures 122 millimeters in total length and has a tail length of 48 
millimeters. 

Desert horned-toads about the Turtle Mountains are occasionally 
seen in the sandy wash-bed and low-plain environments. They seem 
to be less frequent on the rocky mesas, and are wholly absent on the 
hillsides. Their activity in the hot season appears to be restricted to 
the morning and afternoon hours. When alarmed they often retreat 
to the shelter of an Atriplex or other low-growing bush, dodging about, 
when pursued, on the ground beneath the thickly matted lower 
branches (see Richardson, 1915, p. 423). One was found on an open 
mesa after sunset, appressed to a small brown piece of lava and ap- 
parently asleep. This individual was dark gray when discovered, but 
became very light in color the next day. 

Phrynosoma platyrhinos seems to be a more agile species than P. 
blainvillit of the Pacifie coast district in southern California. Bryant 
(1911, p. 16) interestingly describes the burrowing habits of Phryno- 
soma. 

The examination of the stomach of a preserved specimen revealed 
four parasitic nematodes, six beetles, one orthopter, many black ants, 
a leaf, a seed, five pebbles and some gravel and earth. Contents of an- 
other stomach were: fifteen parasitic nematodes, six Coleoptera, one 
orthopter, 145 red-headed ants, all apparently of the same species 
and swallowed whole, and one pebble. 


Xantusia vigilis Baird 
Desert Night Lizard 


The only specimen found (no. 1101) was taken near Goffs. It has 
116 transverse and 38 longitudinal scale rows on the back. The fronto- 
nasals are joined on the median line, the frontal is entire, and the 
specimen seems identical with examples of vigils from the western 
Mohave Desert. There are seven femoral pores on the right thigh. 
The color above is cream buff to chamois; slightly lighter below; faint 


1916] Camp: Amphibians and Reptiles 529 


white lines on back of neck; back speckled with brown scales. Total 
length 86 millimeters, tail length 52 millimeters. 

The specimen secured was found in the usual habitat of the species, 
under a prostrate tree-yuecca branch in a small grove of Yucca mo- 
havensis. The species is rare in this locality, which appears to be the 
eastern limit of its range. Unsuccessful search was made for night 
lizards among the rather scattering tree yuccas along the east base of 
the Turtle Mountains, five miles north of the Horn Mine. 


Cnemidophorus tigris tigris Baird and Girard 
Desert Whip-tailed Lizard 


Fifteen individuals (nos. 5503-5517) of this forked-tongued lizard 
were secured in the vicinity of the Horn Mine and Blythe Junction. 
This series illustrates some phases of variation as pointed out by 
Gadow (1906) for this remarkably unstable genus. The scales along 
the edge of the gular fold are all smaller than those under the chin, 
and are of equal size throughout. The number of large, transverse 
seales in front of the forearm run from 6 to 9. The femoral pores are 
19 in one thigh, 20 in five, 21 in six, 22 in nine, and 23 in seven of the 
thighs in which counts could be made; being 23 right: 23 left twice, 
23:22, g 22:23, ¢ 22:22 three times, 22:—, 21:22, 21:21, 20:21 three 
times, 20:20, and 19:—. The femoral pores are large in four, medium 
in six (at least two of which are males), and small in five specimens. 

Light, almost wholly unspotted, specimens were taken on the glar- 
ing sand south of Blythe Junction. The throats in these are cream- 
colored and much lighter than in other specimens; and the sides of the 
head are yellowish and show no dark markings. Other specimens col- 
lected among dark rocks, in washes, and on rocky hillsides exhibit the 
bluish gray throats, dusky shoulders, and yellowish hind quarters typ1- 
cal of the species. The areas between the darker spots on the neck, 
and the spot in front of the ear (light yellow in C. stejnegeri and in 
C. tigris undulatus), are in the present series dusky, and in many speci- 
mens this dusky suffusion obscures the darker markings on the sides 
of the neck. The dorsal spotting, striping and cross-banding seems to 
occur rather indiscriminately among both large and small individuals. 
The examples from sandy areas, as before noted, have the dorsal pat- 
tern almost obliterated. The dorsal black dashes in some of the others 
are in six to eight longitudinal series joined obliquely to form zigzag 
stripes, with the yellow ground color showing as longitudinal lines be- 


530 University of California Publications in Zoology [ Vou. 12 


tween. In other examples the black longitudinal lines are broken up 
into squarish patches, and in a few these patches are joined trans- 
versely in the posterior region giving a tiger-like banding. 


Museum snim|b ery eee ee 5513 Jo 5516 Q 
Total length (in millimeters) - 335 309 
Tail length (in millimeters) 22. .-2.--0.-0.s.eseeseeseeee--e 246 * 220 
Length of fourth toe (in millimeters) -.................. 25 23 


The whip-tailed lzard seems to occur abundantly in the Turtle 
Mountain vicinity in every phase of environment, except the rocky 
mesa, from rocky hillside to sand dune (see table, p. 507). It was espe- 
cially well represented over the rocky hillsides, where individuals 
ceaselessly forage, sticking their sharp noses into little piles of leaves 
and debris or picking up small bits of food with their active tongues. 
They slink about hesitatingly on the sand, with their tails dragging 
behind them, thus leaving a characteristic track. When running 
swiftly this lizard elevates its tail, so that the ground is just cleared; 
and the tip lashes about as the lizard runs. 

Though usually timid, the whip-tails, hke Callisaurus, seem to be 
almost devoid of fear when feeding. I saw two come into a room and 
gather crumbs from the floor while several people were about. They 
sometimes rest with their hind feet raised clear of the hot sand. They 
exhibit a tendeney to burrow with their forefeet when annoyed. 

The stomach of one whip-tail contained a large grasshopper, slight- 
ly chewed. Another had eaten a small beetle, a spider, and a quantity 
of tiny yellow ants. 


Sonora episcopa (Kennicott) 
Texas Ground Snake 


The one specimen (no. 5549) was taken in the rocky hills four 
miles northwest of Blythe Junction. So far as known to the writer, 
this is the first reeord of this snake from California. The scales are 
in 15 longitudinal rows, the loreals are 1-1, the gastrosteges 185, the 
urosteges 50, the total length 405 millimeters, and the tail length 73 
millimeters. 

The coloration differs slightly from that given by Van Denburgh 
(1912, pp. 153-154) for two specimens collected at Yuma, Arizona. 
The head is orange (a variation also recorded by Brown, 1901) instead 
of the usual yellowish brown color, and is identical in tone with the 
‘‘vinaeeous rufous’’ dorsal band. The darker patches on the head 


1916] Camp: Amphibians and Reptiles 531 


are only barely distinguishable. The dorsal stripe is three whole and 
two half seales wide on the body, and two whole and two half scales 
wide on the tail. The orange colors have turned to light pink after 
nine months immersion in alcohol. In the adoption of the scientific 
name of this and the next following species the present writer follows 
Van Denburgh and Slevin (1913, p. 411). 

The specimen captured was found on June 8 at six p.m., coiled be- 
side a stone in front of a hole, into which it abruptly disappeared when 
approached. The red colors were conspicuous in the living snake from 
the moment it was discovered. The stomach was apparently empty. 


Sonora occipitalis (Hallowell) 
Desert Burrowing Snake 
The two specimens secured (nos. 5547, 5548) were the only ex- 
amples of this species noted. In each the body scale rows are 15 and 
the loreals 1-1; other features as follows: 


Black bands Total Tail 
Gastro- Uro- —————_>7. length in length in 
Nos. steges steges on body on tail millimeters millimeters 
5547 161 46 31 9 104 36 
5548 163 41 32 10 318 53 


The life-colors of this species (see Richardson, 1910, p. 383) have 
faded more quickly and completely in alcohol in the present specimens 
than in any of the other reptiles in the collection. The yellow and red 
bands are now, after nine months, entirely white, although the speci- 
mens have been kept in the dark. 

Two of these docile little snakes were found on the gravelly, 
ereosote-dotted plains south of Blythe Junction. One was taken late 
in the afternoon, the other early in the morning, and neither was 
active. One was caked with clay as though it had just emerged from 
the soil. Mr. H. A. Smith of Blythe Junction, to whom I showed 
one of the above examples, said he once found one of these snakes 
in the hard soil of his yard and some distance below the surface. 


Lampropeltis boylii (Baird and Girard) 
Boyle King Snake 
The only specimen taken (no. 5543) shows no trace of longitudinal 
striping as in ‘‘californiae’’ from San Diego, Riverside, San Bernar- 
dino (Waterman Canon), and Fresno counties. A very few of the 


532 University of California Publications in Zoology [ Vou. 12 


white scales on the sides are narrowly bordered posteriorly with brown; 
this may indicate a leaning toward the conjwncta type, described from 
Cape San Lucas and Yuma, Arizona (Van Denburgh, 1895, pp. 142, 
143). 

In the Blythe Junction specimen the loreals are distinet, there are 
two postoculars, the anterior temporals are three on each side, and 
there are nine inferior labials, of which the fifth is the largest. The 
scale rows are 23-21, the gastrosteges 256, and the urosteges 54 (all 
divided). The total length is 912 millimeters, and the tail length is 
115 millimeters. 

The rostral plate is yellowish, margined above with brown. All 
the other parts are dark brown and white. The color pattern is typi- 
eal. There is a small white patch on the middle of the nape one scale 
behind the parietals. The body is encireled with thirty-five white 
rings and the tail with eight. 

Only the one ‘‘milk snake’’ was seen in the Turtle Mountains. It 
was found on May 380, crawling over the rocks in a canon bottom at 
about nine o’clock in the morning. An example of this species, taken 
in the river bottom (arrowweed association) at Needles, on July 15, 
1909, was trying to swallow head-first a harvest mouse (Reithrodon- 
mouse trap. 


’ 


be] 


tomys) caught in a ‘‘gee-whiz 


Bascanion flagellum frenatum Stejneger 
Red Racer 


Two specimens of this variable form are at hand (nos. 5545, 5546). 
The scales are in 17 rows in both, and in each the anal plate is divided. 
Loreal fused with posterior nasal on each side in no. 5546; partly 
fused on left, distinet on right side, in no. 5545. Other characters as 
follows: 


No. 5545 5546 
Supertorpl alba) sess eee 7 right, 9 left 8 
Inferior labials -... = i 11 
GASUROSLE RCS teense crane eens eee ees 201 207 
NODOSA SYS EES| rr eecner eee tec E eae 110 111 
Total length (millimeters) - =. 1291 1237 
Tail length (millimeters) 22--2s.-2-2 2 s---n 341 327 


Color descriptions of this subspecies by Cope (1898, p. 801), Stej- 
neger (1893, pp. 208-209), and Van Denburgh (1897, p. 187) taken 
probably from preserved material, do not mention the vivid red color 
of this snake. In both of the present examples there are traces of at 


1916] Camp: Amphibians and Reptiles 533 


least three black cross-bars on the nape, and no. 5545 has a series of 
lighter scales in regular transverse intervals down the back on the 
anterior half of the body. Both specimens are heavily marked with 
brown and red spots about the face, neck, and throat. 

The red racer seems to be the most generally distributed snake on 
the Colorado Desert. It occurs on mountain and plain alike, and is 
far swifter in movement than any other desert snake. The two present 
examples were taken at the Horn Mine near a tent floor under which 
they had apparently been living. 


Crotalus mitchellii (Cope) 
Pallid Rattlesnake 


Three specimens (nos. 5540, 5541, 5544) of this desert species were 
captured near the Horn Mine. In scutellation and coloration these 
examples show departure from some of the typical characteristics of 
mitchell and seem to approach to a certain extent those of tigris. In 
no. 5540 the rostral is in contact with the anterior nasals on both sides, 
but on the left a small scale has started to split off from the nasal be- 
tween the latter and the rostral. In the other two specimens the ros- 
tral is separated from the anterior nasals by one row of three scales on 
each side. Rostral higher than wide in no. 5540, equilateral in no. 
5544, and wider than high in no. 5541. Seales on body all keeled; in 
23 rows in no. 5541, and in 25 rows on the other two specimens. Supra- 
oculars striate and rugose. Three rows of scales between suborbital 
chain and labials. Other characters as follows: 


Vins emia Nui Greene sacs ore ener eee eseneeeerenacess OOO. 5541 5544 
Rows of scales between supra-oculars —-.-..........--- 7 6 5 
FS iryaxeveniae TE NES) ee ce eee 17 15 16 
Tiransivageone Te oy IS} eos aee eee eee ee 16 7, ily 
(CiDST AROS GER) ceeceseeeees Steere ere pee eee See 179 183 179 
LUIS PADS See eee oe eceer ee SE ee eee Cee 18+ 21+ 19+ 
Total length (in millimeters) -................--------------- 653 773 616 
MavlVeneth (Gn millimeters) 2-2 ---cecencne-e---e 41 55 47 


7 Last one divided. 
¢ First and last three divided. 


The color varies widely. In no. 5540 the transverse bands of 
ground color are light pink, the muzzle greenish, the top of the head 
pinkish yellow speckled with yellow, and the sides of the head gray 
over the temporal region and corner of the mouth; dark patches on 


534 University of California Publications in Zoology [ Vou. 12 


back indistinct, brown; sides gray; a few scales surrounding darker 
patches on back, bluish gray ; underparts whitish; a broad light sub- 
ocular stripe including six of the upper labials. 

No. 5541 has more red in the coloration of the sides, and the lighter 
dorsal bands are flesh pink. Gray predominates in the dorsal colora- 
tion, and the pink ground color becomes tawny on the posterior end 
of the body. Distinet reddish brown bands on posterior third of 
the body; anteriorly each of these bands splits into three transverse 
blotches, the outer two being small, and the middle one large as in 
tigris. Belly white; lateral edges of gastrosteges in median abdominal 
region speckled with red and gray. Light area below eye, covering 
five upper labials. No. 5544 is in the red phase. The coral red of the 
top of the head and back almost obscures every other marking, but the 
dorsal bands can be made out because of their darker shade, near rus- 
set vinaceous. The pink color becomes yellowish near the tail; the 
latter is marked by seven fairly distinct blackish bands which do not 
meet below. The sides and top of the head are obscurely stippled with 
gray. No light markings below or behind eye. In all three specimens 
the sides of the head do not exhibit the postocular stripe common to 
so many species of rattlesnakes (see Stejneger, 1895, pp. 423-424). 

Only the three pallid rattlesnakes taken were seen; two of these 
were in a rocky wash at the Horn Mine, and one (no. 5541) was beneath 
a dead palo verde in a wash about a mile from the foot of the moun- 
tains. The stomach of no. 5544 contained a Stephens canon mouse 
(Peromyscus crinitus stephensi). Since this rodent is wholly noc- 
turnal, the instance might be taken as showing nocturnal habits on the 
part of the snake. 


Crotalus cerastes Hallowell 
Sidewinder 


One horned rattlesnake (no. 5542) was captured in the drifting 
sand near Blythe Junction. This specimen has the anterior and pos- 
terior nasal plates divided on the right side and united on the left. 
One internasal is present on each side. There are 13 superior and 13 
inferior labials, 143 gastrosteges, and 24 urosteges of which the pos- 
terior four are divided. The total length to the base of the rattle is 
504 millimeters, and the tail length 39 millimeters. There is a wart- 
like growth on one side near the neck, consisting of eight elongated 
scales arranged in rosette fashion. 


1916] Camp: Amphibians and Reptiles 53! 


The writer is by no means convinced that this rattlesnake is ex- 
clusively nocturnal in habits as suggested by Meek (1905, p. 18). 
Both at Needles and near Blythe Junction individuals were traced by 
the characteristic tracks in the sand. Each was found closely coiled in 
a symmetrical pad and partly buried flush with the surface in the 
hot sand right out in the noonday sunshine of midsummer. In neither 
case were the snakes easily seen, as they were of the exact color of 
their sandy surroundings. Both, though alert, allowed themselves to 
be noosed without moving away or doing more than rattle feebly. 
That they eat the diurnal lizards Uta and Cnemidophorus (see Van 
Denburgh and Slevin, 1914, p. 429) is an evidence of daytime activity. 


Transmitted August 20, 1915. 


536 University of California Publications in Zoology [ Vou. 12 


LITERATURE CITED 
AtsatT, S. R. 

1913. The reptiles of the San Jacinto area of southern California. Univ. 

Calif. Publ. Zool., 12, 31-50, 2 tables. 
Brown, A. E. 

1901. A review of the genera and species of American snakes north of 

Mexico. Proc. Acad. Nat. Sci. Phila., 53, 10-110. 
BryANntT, H. C. 

1911. The horned lizards of California and Nevada of the genera Phryno- 
soma and Anota. Univ. Calif. Publ. Zool., 9, 1-84, pls. 1-9, 6 figs. 
in text. 

Corr, E. D. 

1898. The crocodilians, lizards and snakes of North America. Rep. U. 8. 

Nat. Mus., 1898, 151-1294, 36 pls., 347 figs. in text. 
Cougs, E. 

1875. Synopsis of the reptiles and batrachians of Arizona with critical and 
field notes and an extensive synonomy. Jn Wheeler, G. M., Report 
Geogr. Geol. Expl. and Surveys West of 100th Meridian, 5, pp. 
585-633, 10 pls. 

Dirmars, R. L. 

1907. The reptile book (New York, Doubleday, Page & Co.), xxxii + 472, 

136 pls. 
FRANKLIN, D 

1913. Color changes in collared lizards. Copeia (New York), 1, no. 1, Dec. 
27, 1913. 

1914. Notes on leopard lizards. Tbid., 1, no. 5, April 15, 1914. 

Gapow, H. 

1906. A contribution to the study of evolution based upon the Mexican 
species of Cnemidophorus. Proc. Zool. Soc. London, 1906, 277-375, 
1 pl., 22 figs. in text. 

GRINNELL, J. 

1908. The biota of the San Bernardino mountains. Univ. Calif. Publ. Zool., 
5, 1-170); pls. 1-24. 

1914. An aeeount of the mammals and birds of the lower Colorado valley 
with especial reference to the distributional problems presented. 
Ibid., 12, 51-294, pls. 3-13, 9 figs. in text. 

GRINNELL, J., and GRINNELL, H. W. 

1907. Reptiles of Los Angeles County, California. Throop Inst. Bull. (Pasa- 

dena, Calif.), no. 35, 1-64, 23 figs. in text. 
MEEK, 8. E. 

1906. An annotated list of a collection of reptiles from southern California 
and northern Lower California. Field Mus. Publ., Zool. Series, 7, 
1-19, 3 pls., 1 table. 

RICHARDSON, C. H. 

1910. Notes on a little-known species of snake, Chionactis occipitalis. Science 
(n. s.), 32, 383-584. 

1915. Reptiles of northwestern Nevada and adjacent territory. Proce. U.S. 
Nat. Mus., 48, 403-435. 


1916] Camp: Amphibians and Reptiles 537 


RUTHVEN, A. G 
1907. A collection of reptiles and amphibians from southern New Mexico 
and Arizona. Bull. Amer. Mus. Nat. Hist., 23, 483-603, 22 figs. 
STEJNEGER, L. 
1890. Annotated list of reptiles and batrachians collected by Dr. C. Hart 
Merriam and Vernon Bailey on the San Franciseo Mountain 
Plateau and Desert of the Little Colorado, Arizona, with descrip- 
tions of new species. U. 8. Dept. Agric., Div. Ornithology and 
Mammalogy, N. Amer. Fauna, no. 3, 103-118. 
1893. Annotated list of the reptiles and batrachians collected by the Death 
Valley expedition in 1891, with descriptions of new species. Ibid., 
7, 159-228, 394-398, 4 pls. 
1895. The poisonous snakes of North America. Rep. U. 8. Nat. Mus., 1893, 
337-487, pls. 1-19, 70 figs. in text. 
STEPHENS, F. 
1914. Arid California and its animal life. Biennial Rep. Calif. Fish and 
Game Comm., 3, 1912-1914, 127-135, 2 figs. in text. 
STONE, W. 
1911. On some collections of reptiles and batrachians from the western 
United States. Proc. Acad. Nat. Sei. Phila., 43, 222-232. 
STONE, W., and REHN, J. A. G. 
1903. On the terrestrial vertebrates of portions of southern New Mexico 
and western Texas. Ibid., 55, 16-34. 
Taytor, W. P. 
1912. Field notes on amphibians, reptiles and birds of northern Humboldt 
County, Nevada, with a discussion of some of the faunal features 
of the region. Univ. Calif. Publ. Zool., 7, 319-436. 
VAN DENBURGH, J. 
1895. A review of the herpetology of Lower California. Part I, Reptiles. 
Proce. Calif. Acad. Sei. (2), 5, 77-162, 11 pls. 
1897. The reptiles of the Pacific Coast and Great Basin. Occasional papers, 
Calif. Acad. Sci., 5, 1-236, many figs. in text. 
1912. Notes on a collection of reptiles from southern California and Ari- 
zona. Proce. Calif. Acad. Sci. (4), 3, 147-154. 
VAN DENBURGH, J., and SLEVIN, J. R. 
1913. A list of the amphibians and reptiles of Arizona, with notes on the 
species in the collection of the Academy. Jbid., (4), 3, 391-454, 
pls. 17-28. 
Yarrow, H. C 
1875. Report upon the collections of batrachians and reptiles made in por- 
tions of Nevada, Utah, California, New Mexico, and Arizona dur- 
ing the years 1871, 1872, 1873, and 1874. In Wheeler, G. M., Re- 
port Geogr. Geol. Expl. and Surveys West of the 100th Meridian, 5, 
pp- 509-584. 
1882. Check list of North American reptilia and batrachia, with catalogue 
of specimens in U. S. National Museum. Bull. U. 8. Nat. Mus., 24, 
3-249. 


PLATE 19 


Map of Turtle Mountain region, southeastern California, showing animal | 
environments. 


e 


[538] 


UNIV. CALIF. PUBL. ZOOL. VOL. 12 [CAMP] PLATE 19 


PLATE 20 
Fig. 2. High plain environment near volcanic plug at south end of Turtle 
Mountains, San Bernardino County, California. Yucca mohavensis in foreground 
and middle distance. 
Fig. 3. Low plain environment near Blythe Junction, Riverside County, 
California. Dipodomys deserti burrows in foreground, sand dunes in distance. 


[540] 


UNIV, CALIF. PUBL. ZOOL. VOL. 12 


Vt 
Wee 


TO) Pare 
eae 
uv y u 


PLATE 21 


Fig. 4. Drifting sand environment near Blythe Junction, Riverside County, 
California. Home of Uma notata; kit fox scratchings, and burrows of Citellus 
tereticaudus, in foreground. 


Fig. 5. Small wash in rocky mesa environment at south end of Turtle 
Mountains. Caves in foreground inhabited by Neotoma, and by Sceloporus 
magister. 


[542] 


UNIV, CALIF. PUBL. ZOOL. VOL. 


12 


: Ce 
ies 


ae: ry 


PLATE 22 


Fig. 6. Uma notata, from specimens. Dorsal view from no. 1285, Mus. Vert. 
Zool.; ventral view from no. 1286, Mus. Vert. Zool. : 


Fig. 7. Uma notata, from life, showing posture when alert 


[544] 


UNIV. CALIF. PUBL. ZOOL. VOL, 12 CAMP] PLATE 


“eT ee 
r % j 


UNIVERSITY OF CALIFORNIA PUBLICATIONS— (Continued) 


* 12. Batrachoseps major and Bufo cognatus californicus, New Amphibia 
from Southern California, by Charles Lewis Camp. Pp, 327-334. 
Nf ag EE Ls 65 RRA oe AR 5 ral a Sec Re ie ay RR Bes Nee a 

18. Report upon Mammals and Birds found in Portions of Trinity, Siskiyou, 
and Shasta Counties, California, by Louise Kellogg. Pp. 335-398, 

plates 15-18. 

14, An Analysis of the Vertebrate Fauna of the Trinity Region of Northern 
California, by Joseph Grinnell, Pp. 401-412. 
Nos, 13 and 14 inone cover. January, 1916 22. cece 

15, The Status of the Beavers of Western North America, with a Con- 
sideration of the Factors in their Speciation, by Walter P. Taylor, 
Pp. 413-495, 22 text-figures.= March, 1916 cs. ne 

16. Two New Aplodontias from Western North America, by Walter P, 
Taylor. Pp. 497-501. May, 1916 

17. Notes on the Local Distribution and Habits of the Amphibians and 

Reptiles of Southeastern California in the Vicinity of the Turtle 

Mountains, by Charles Lewis Camp. Pp. 503-544, plates 19-22, 

PUT VIS ye BO dh coe as hk ans cee wa ct ccree op kta nace poaatnn Sancho soe Sucewt docs chaps coed occn 


Vol. 13. 1. The Schizopoda of the San Diego Region, by Calvin O. Esterly. Pp. 
1220; “platesi-2.5 Aprile LOi4 oe See ee Oe eee 
-2. A Study of the Occurrence and Manner of Distribution of the Cteno- 
phora of the San Diego Region, by Calvin O, Esterly. Pp. 21-38. 
April, 1914 
3. A New Self-Regr ating Paraffin Bath, by C. W. Woodworth. Pp. 39- 
Zi we COS TeUres..- April, 19 Uae asa catec anda eee a 
4, Diplodinium ecaudatum, with an Account of Its Neuromotor Apparatus, 
by Robert G. Sharp. Pp, 43-122, piates 3-7, 4 text figures. May, 
EB SR INI RS RG NL rT ORNS Cah n9, sR TN ons ARRAS, Pe SrA Ri ar EAE 
5. The Vertical Distribution and Movements of the Schizopoda of the 
San Diego Region, by Calvin O. Esterly. Pp. 123-145,. May, 1914... 
6. The Anatomy of Heterodontus francisct. I. The Exoskeleton, by. J. 
Frank Daniel, Pp. 147-166, plates 8-9, 4 text figures, May 23, 


se Sac oe Pact an cc a eg NX Se ea 
7. The Movements and Reactions of the Isolated Melanophores of the 
Frog, by-S. J. Holmes. Pp. 167-174, plate 10. August, 1914 _.u....... ra 


8. Polychaetous Annelids of the Pacific Coast in the Collections cf the 
Zoological Museum of the University of paltoras, by Aaron L. 
Treadwell. Pp. 175-234, plates 11-12. 


9. New Syllidae from San Francisco Bay (collected by the U. 8. 8. 
‘* Albatross’’), by Aaron L. Treadwell. Pp. 235-238, 7 text figures. 
Nos, 8 and 9 in one cover. October, 1914 222 ci cee eee 

10. Note on the Medusan Genus Stomolophus, from San Diego, by Henry 
B. Bigelow... Pp. 239-241. September, 1914 2... 

11. A Study of the Structure of Feathers, with Reference to their Taxo- 

nomic Significance, by Asa C, Chandler, Pp, 243-446, plates 13-17, 

of aR Seo ET bre Aiea = 9 2p 3 0 a A LY SA RGcs et See pe Aang ln = Rank Se eRe 

- 12, Anatomical Adaptations in the Thoracic Limb of the California Pocket 

Gopher and Other Rodents, by Charles Daniel Holliger. Pp. 447- 

494, plates 38-39, 20 text-figures. March, 1916 —...W00002 2c 


Vol. 14, 1. A Report upon the Physical Conditions in San Francisco Bay, Based 
upon the Operations of the United States Fisheries Steamer ‘‘Alba- 
tross’’ during the Years 1912 and 1913, by. F. B. Sumner, G. D, 
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* Vol.-15, Introduction. Dependence of Marine Biology upon Fodaachs and 
yi Necessity of Quantitative Biological Research. Pp. i-xxiii, June, 
+ UE Eas Sc St te ee ee ee Sm eS 
: 1. Hydrographic, Plankton, and Dredging Records of the Scripps Institu- 

tion for Biological Research of the University of California, 1901 to 
1912, compiled and arranged under the supervision of W. EH. Ritter 
by Ellis L. Michael and George F. McEwen. Pp. 1-206, 4 text figures 
pos IS 0G Paategs 18 A: Aas oh ea a dace eo Se ee nS OP NO RR anc Tes er es 


WO VoL 16. 1. An Outline of the Morphology and Life History of Crithidia lepto- 
RE as coridis, sp. nov., by Irene McCulloch. Pp. 1-22, plates 1-4, 1 text 
fipirel Sapsem ber, 1915 ya rae ascakdetamste cuecosrndinane cca sence es 


10 


+75 


85 


10 


65 


05 


2.00 


45 


2.25 


225 


25 


UNIVERSITY OF CALIFORNIA PUBLICATIONS— (Continued) 


2. On Giardia microti sp. nov., from the Meadow Mouse, by Charles 
Atwood Kofoid and Elizabeth Bohn Christiansen. Pp. 23-29, 1 figure 
in text. 


8, On Binary and Multiple Fission in Giardia muris (Grassi), by Charles 
Atwood Kofoid and Elizabeth Bohn Christiansen. Pp. 30-54, plates 
5-8, 1 figure in text. 
Nos. 2 and $8 in one cower, November, 1915 2..0...2.--.cccccecccseetereceseee 
4. The Cultivation of Tissues from Amphibians, by John C. Johnson. 
Pp. 55-62, 2 figures in text. November, 1915 co...o.cc cc ceee spe eeteenteeeseeee 
5. Notes on the Tintinnoina. 1. On the Probable Origin of Dictyocysta 
tiara Haeckel. 2. On Petalotricha entzi sp. nov. by Charles Atwood 
Kofoid. Pp. 63-69, 8 figures in text. December, 1915-00. -22...2202..-.0-- 


6. Binary and Multiple Fission in Hexamitus, by Olive Swezy. Pp. 71- 88, 
plates 9-11. 


7. On a New Trichomonad Flagellate, Trichomitus parvus, from the Intes- 


tine of Amphibians, by Olive Swezy. -Pp. 89-94, plate 12. 
Nos. 6 and 7 in one cover. December, 1915 oo... cccceccccecesnceeceeneee am 


8, On Blepharocorys oat Sp. Nov., 2 New Ciliate from the Caecum of the 


Horse, by Irwin C. Schumacher. Pp. 95-106, plate 13. December, 
NES SS eS PS SS ONE Ce ae ag ee ne ee eR 


9. Three New Helices from California, by S.. Stillman Berry. _ Pp. 107- 
TE VS anuary, 1910 ee a ee ee ree ae ened a 


10. On Trypanosoma triatomae, a New Flagellate from a Hemipteran Bug 
from the Nests of the Wood Rat Neotoma fuscipes, by Charles Atwood 
Kofoid and Irene McCulloch. Pp, 113-126, plates 14-15, February, 
DOL As aa a sae Mice ee ete ee Gua sek eet ceaeae 


11. The Genera Monocercomonas and Polymastiz, by Olive Swezy. Pp. 127- 
188; plates:56-17~; February, ‘19160. Ss Sh a ee ee 


12. Notes on the Spiny Lobster (Panulirus interruptus) of the California 
Coast, by Bennet M. Allen. Pp. 139-152, 2 figs. in text. March, 1916 


13. Notes on the Marine Fishes of California, by Carl L. Hubbs. Pp. 153- 
169;-plates: 18-20.3¢ March,: 1916 <2 a ee ee 


14, The Feeding Habits and Food of Pelagic Copepods and the Question 
of Nutrition by Organic Substances in Solution in the Water, by 
Calvin O. Esterly. Pp. 171-184, 2 figs. in text. March, 1916 -............. 

15. The Kinetonucleus of Flagellates and the Binuclear Theory of Hart- 
mann, by Olive Swezy. Pp. 185-240, 58 figs. in text. March, 1916... 

16. On the Life-History of a Soil Amoeba, by Charlie Woodruff Wilson. 
Pp. 241-292, plates 18-23. Durky, 1916 oan ccc cece acne antec ecccesenconetes 

17. Distribution of the Land Vertebrates of Southeastern Washington, by 
Lee Raymond Dice. Pp. 293-348, plates 24-26. June, 1916 


10. 


AQ re 


15 


10- 


15 


Bae 


Cr tae 


50 


+60 


60 


aie) Cis Mier ork eee bake 


Abpadetin Sh iho % 


we 


Pe 


INDEX* 


Titles of papers in this volume, and names of new species, are printed in a 
bold-faced type 


Abronia villosa, 78. 

Acacia greggii, 83, 282. 

Accipiter cooperi, 69, 124, 380, 404. 

velox, 69, 75, 124, 380, 404. 

Achyronychia cooperi, 241. 

Actitis macularius, 68, 121, 379, 404. 

Aéronautes melanoleucus, 86, 96, 143. 

Age, change due to, in a single spe- 
cies, 418. 

Agelaius phoeniceus neutralis, 162. 
nevadensis, 383, 403. 
sonoriensis, 65, 69, 72, 97, 161, 

162. 

Alexander, Annie M., 1, 52, 336, 405; 
gifts by, 306, 416. 

Aluco pratincola, 127. 

Ameiurus nebulosus, 62. 

Amixia, 485. 

Ammospermophilus harrisi, 288, 292. 
harrisi, 80, 86, 101, 102, 219, 225; 

measurements of, 220, 222. 
saxicola, 221. 
leucurus leucurus, 80, 86, 101, 102, 
221, 225; measurements of, 222, 
223. 

Amphibia, new, from Southern Cali- 
fornia, 327. 

Amphibians, of Southeastern Cali- 
fornia, 503; habitat limitations 
of, in Turtle Mountains region, 
507. 

Amphibians and Reptiles of South- 
eastern California in the Vicin- 
ity of the Turtle Mountains, 
Notes on the Local Distribution 
and Habits of, 503. 

Amphispiza bilineata deserticola, 78, 
80, 81, 86, 173. 

nevadensis nevadensis, 78, 80, 173. 

Analysis, An, of the Vertebrate 
Fauna of the Trinity Region of 
Northern California, 399; litera- 
ture cited, 410. 

Anas platyrhynchos, 67, 115. 

Anser albifrons gambeli, 11. 

Anthus rubescens, 68, 206. 

Antrozous pallidus pallidus, 70, 263. 

Aphelocoma californica californica, 
382, 404. 

Aplodontia, A Previously Unde- 
scribed, from the Middle North 
Coast of California, 297. 

Aplodontia californica, 295, 296, 370, 
401; figure of skull, opp. 398. 


* Univ. Calif. Publ. Zool., vol. 12. 


[545] 


columbiana, 499; comparison 
with A. e. rainieri, with A. e. 
californica, and with A. rufa 
grisea, 500. 
chryseola, 295, 296, 297, 369, 370, 
401, 402, 407; measurements, 
371; pictures of burrows, opp. 
394; picture of habitat, opp. 
596; figures of skulls, opp. 398. 
major, 295. 
nigra, 297, 298, 299; 
ments, 300. 
phaea, 296, 297, 298, 299, 370; 
measurements, 300. 
rufa, 299. 
grisea, 497; comparison with 
A. r. rufa, with A. r. olym- 
pica, 497, 498; with A. eali- 
fornica eolumbiana, and with 
A. ¢. rainieri, 498. 
Aplodontia chryseola, a New Moun- 
tain Beaver from the Trinity 
Region of Northern California, 
295. 
Aplodontias from Western North 
America, Two New, 497. 
Archilochus alexandri, 69, 71, 82, 143, 
144, 146. 
Ardea herodias treganzai, 67, 96, 116, 
118; measurements of, 118. 
Arizona elegans, 43. 
Aselepias subulata, 87, 146. 
Associational areas of the Colorado 
Valley, 66; river, 67, 68; willow- 
cottonwood, 69, 70, pictures of, 
opp. 278; tule, 72; arrowweed, 
73, picture of, opp. 278; quail- 
brush, 74, 75, picture of, opp. 
280; mesquite, 75, 76, pictures 
of, opp. 280; saltbush, 77, 78, 
pictures of, opp. 282, 290; creo- 
sote (mesa), 80; catelaw (or 
wash), 82, picture of, opp. 282; 
Saguaro, 85; Encelia (rocky 
hills), 86, picture of, opp. 288; 
three factors in restriction of, 
96. 
Associations, plant and animal, of 
southeastern California, 506. 
Astragalinus lawrencei, 76, 166. 
psaltria hesperophilus, 73, 76, 82, 
86, 165, 383, 403. 
tristis salicamans, 883, 403. 


measure- 


. Astur atricapillus striatulus, 380, 404. 


Index 


Asyndesmus lewisi, 381, 400. 
Atriplex, confertifolia, 87. 
lentiformis, 75, 94, 280. 
polyearpa, 78, 84, 241; picture of, 
282. 
Atsatt, S. R., 31. 
Auriparus flaviceps flaviceps, 76, 82, 
211. 
Avocet, 120. 
Baccharis glutinosa, 70, 189. 
Badger, 507. 
Mexican, 259. 
Baeolophus inornatus inornatus, 387, 


403. 
Bailey, Vernon, acknowledgment to, 
Si), 321. 


Barn owls, 26. 

Barriers, problem of, with regard to 
birds and mammals, 107; elassi- 
fication of, 109; as agents in 
multiplication of species, 110; 
faunal, 407, 408, 410. 

Bascanion flagellum frenatum, 42, 46, 
48, 507, 509, 532. 

laterale, 43, 46, 48. 
Bassariscus astutus raptor, 354, 402. 
Bat, brown, large, 268, 352. 
eanon, 267. 
cave, 266. 
foliage, oak, 317. 
free-tailed, Mexican, 268. 
Hollister, 263. 
leaf-nosed, California, 269. 
long-legged, 352. 
lump-nosed, intermediate, 320. 
pale, 263. 
pallid, desert, 263. 
little, Stephens, 265. 
silver-haired, 352. 
Tejon, 318. 

Bats, vespertilionid, 
Races of, 317. 
Batrachoseps attenuatus, 328, 329; 

measurements of, 330. 
caudatus, measurements of, 330. 
major, 327; measurements of, 330. 
pacificus, 328; measurements of, 


Three New 


330. 
Batrachoseps major and Bufo cog- 
natus californicus, New Am- 


phibia from Southern California, 
327; literature cited, 334. 
Bear, black, 353. 
Beaver, mountain, British Columbia, 
499. 
Point Arena, 297. 
Puget Sound, 497. 
Trinity, 295, 296, 369; picture of 
burrows of, opp. 394. 
Beavers, nomenclature of, 417; 
teeth of, as cutting and grind- 
ing instruments, 426, measure- 
ments of, 428; description of 


[546] 


new subspecies, 429, 433; com- 
parisons of, 437; comparative 
measurements of adult skulls, 
inset opp. 438; history of, 457; 
relationships of, 460; of Califor- 
nia, 462; of Eurasia and Amer- 
ica, 463; semi-aquatie environ- 
ment of, 464. See also Castor. 

Beavers, The Status of the, of West- 
ern North America, with a Con- 
sideration of the Factors in 
their Speciation, 413; summary, 
487; literature cited, 490. 

Beck, R. H., 1. 

Bernicla occidentalis, 6, 10. 

Bighorn, desert, 218, 507. 

Birds of the Colorado Valley, check- 
list of, 110-113; general aec- 
counts of, 113. 

Birds and mammals of the Lower 
Colorado Valley, 51. 

Blackbird, Brewer, 164, 383. 

red-winged, Nevada, 383. 
Sonora, 161. 
yellow-headed, 160. 
Bluebird, mountain, 389. 
western, 216, 280, 389. 

Blythe Junction, California, pictures 
of environments near, opp. 540, 
542. 

Boa, California, 41. 

Bob-eat, 507. 

Bombycilla garrula, 187, 385, 405. 

Branta canadensis, 10, 11. 
canadensis, 1, 2, 8, 13, 14, 15, 19; 

measurements, 16, 17, 18. 
hutchinsi; 62) 3) ds a Lol 
measurements, 16, 17, 18. 
minima, 4, 13, 15, 19, 22, 24; 
measurements, 16, 17, 18. 
occidentalis, 1, 5, 8, 9, 13, 14, 19. 
occidentalis, 9. 

Branta canadensis group, study of a 
collection of geese of the, 1. 

Bryant, Harold C., 25. 

Bubo virginianus pallescens, 65, 69, 
82, 86, 129. 

Bufo alvarius, 509. 

Bufo cognatus californicus, 331; 

measurements of, 333. 
cognatus, 331, 509; measure- 
ments of, 333. 
halophilus, 331, 512. 
lentiginosus woodhousii, 332, 509. 
punctatus, 507, 508, 509, 512. 
Bunting, lark, 181. 
lazuli, 181, 385. 

Bush-tit, coast, 387. 

Buteo borealis calurus, 69, 86, 125. 

Butorides virescens anthonyi, 67, 119. 

Calamospiza melanocorys, 181. 

Cactus, giant, picture of, opp. 286. 


— —  ———— 


Index 


California. See Climatological Table 
of Southern California; Environ- 
ments of southeastern Califor- 
nia; Reptiles of the San Jacinto 
Area. 

California pocket gopher. See 
Gopher, Pocket, California. 
Callisaurus ventralis, 33, 46, 47, 48, 

49, 507, 508, 519. 

Callospermophilus chrysodeirus, 374. 
chrysodeirus, 400. 
trinitatis, 874, 400, 402, 407; 

picture of habitat, opp. 392. 

Calypte costae, 82, 86, 145. 

Camp, Charles Lewis, 827, 503. 

Canis lestes, 353, 354, 400. 

ochropus estor, 65, 80, 254. 
Carp, 62. 
Carpodacus cassini, 383, 404. 
mexicanus frontalis, 82, 85, 86, 164, 
383, 404. 
purpureus californicus, 383, 404. 
Castle Lake, Siskiyou County, 349. 
Castor, history of, 458, 459; ranges 
of, 464; and Erethizon, parallel- 
ism in, 426. See also Beaver. 
Castor, canadensis belugae, 418, 429, 
461, 463, measurements of super- 
ior cheek teeth, 428; description 
of, 429; comparisons with: C. ec. 
leucodonta, 430, 455; with C. e. 
canadensis, 432, external char- 
acters, inset opp. 482, 455; with 
C. ¢. phaeus, 432, external char- 
acters, insert opp. 482, 438; with 
C. e. frondator, 455; with C. ec. 
pacificus, 455; cranial measure- 
ments, insert opp. 430; measure- 
ments of tails, 455. 
eanadensis, 461, 463; measure- 
ments of superior cheek teeth, 
428; drawing of tail of, 431; 
comparisons with: C. ¢. be- 
lugae, 432, external characters, 
inset opp. 432, 455; with C. e¢. 
phaeus, 432; external char- 
acters, inset opp. 440, 455; 
with C. ce. leueodonta, 440, 
external characters, inset opp. 
440, 455; with C. e. pacificus, 
external characters, inset opp. 
440, 455; with C. ec. frondator, 
455, external characters, 456; 
measurements of tails, 441, 
455. 

frondator, 68, 97, 98, 101, 225, 
461; measurements of superior 
cheek teeth, 428; drawing of 
tail, 431; comparisons with: C. 
subauratus subauratus, 446, 
external characters, inset opp. 
446, cranial characters, inset 
opp. 448; with C. ¢. pacificus, 
446, cranial characters, inset 


[547] 


opp. 448; with C. ¢. pacificus, 
446, 455, external characters, 
inset opp. 446, cranial char- 
acters, inset opp. 448; with C. 
ce. leucodonta, 455, external 
characters, inset opp. 456; 
with C. ¢. canadensis, 455, ex- 
ternal characters, inset opp. 
456; with C. ce. belugae, 455. 

leucodonta, 461; change in, due 
to age, 418, 424 fig. B; external 
measurements, 419; coloration, 
420; pelage, 420; cranial char- 
acters, #22, measurements of, 
inset opp. 426; measurements 
of teeth, 423, 428; comparison 
with: C. ¢. belugae, 430, 455; 
with C. ce. canadensis, 440, ex- 
ternal characters, insets opp. 
440 and 456, 455; with C. e. 
phaeus, 440; with C. e. paci- 
fieus, 442, 455; with C. e. fron- 
dator, 455, external characters, 
456; drawings of tail, 431; 
measurements of tails, 441, 
445; figure of: dorsal view 
erania, 450; ventral view cra- 
nia, 451; posterior view crania, 
452. 

michiganensis, 461, 463. 

pacificus, 461; measurements of 
superior cheek teeth, 428; com- 
parison with C. ¢. leucodonta, 
440, 442, 455, external char- 
acters, inset opp. 440; with 
C. e. canadensis, 455, external 
characters, inset opp. 440; 
with C. subauratus subauratus, 
446, external characters, inset 
opp. 446; with C. e. frondator, 
446, 455, external characters, 
inset opp. 446, cranial char- 
acters, inset opp. 448; with C. 
c. belugae, 455; cranial meas- 
urements, 445; measurements 
of tails, 448. 

phaeus, 463; measurements of 
superior cheek teeth, 428, 
drawing of tail of, 431; com- 
parison with C. e. belugae, 432, 
438, external characters, inset 
opp. 482; with C. ¢. canaden- 
sis, 432, 437, external char- 
acters, inset opp. 432; meas- 
urements of tails, 441. 

texensis, 461. 

subauratus shastensis, 418, 433, 

461, 463; type locality, 435; 
eranial measurements, 436. 

subauratus, 461; measurements 
of superior cheek teeth, 428; 
drawing of tail, 431; compari- 
son with: C. ¢. pacificus, 446, 
external characters, inset opp. 


Index 


446, cranial characters, inset 
opp. 448; with C. c. frondator, 


446, external measurements, 
inset opp. 446; external meas- 
urements, 447; measurements 


of tails, 448; cranial measure- 
ments, 449; figure of: dorsal 
view crania, 450; ventral view 
crania, 451; posterior view 
erania, 452. 

Castoroididae, 458. 

Cat, ring-tailed, 354. 

Catfish, 62. 

Cathartes aura septentrionalis, 123. 

Catherpes mexicanus conspersus, 86, 
90, 209. 

Centurus uropygialis, 65, 69, 75, 85, 
86, 133. 

Cercideum torreyanum, 505. 

Cereus giganteus, 85; picture of, opp. 
286. 

Certhia familiaris zelotes, 387, 400; 
picture of habitat, opp. 396. 

Ceryle alcyon, 68, 131. 

Chaetura vauxi, 143. 

Chalicomys, 458. 

Chamaea fasciata henshawi, 387, 403. 

Chat, long-tailed, 204, 386. 

Chen hyperboreus, 11. 
hyperboreus, 67, 116. 

rossi, 11. 
Chickadee, chestnut-sided, 387. 
mountain, 387. 
Oregon, 587. 
Chickaree, Sierra, 375. 
Chipmunk, Allen, 373. 
Klamath, 373. 
Sonoma, 321. 

Chipmunk, A New, Eutamias sono- 
mae, from the Inner Northern 
Coast Belt of California, 321; 
distribution of, in northwestern 
California, 323. 

Chondestes grammacus strigatus, 167, 
383, 403. 

Chordeiles acutipennis texensis, 69, 
77, 80, 81, 142. 

Chuck-a-walla, 34. 

Chueckwalla, 509, 510, 511, 

Cinclus mexicanus unicolor, 386, 405. 

Cireus hudsonius, 68, 72, 123. 

Citellus douglasii, 349, 372, 404. 

tereticaudus, 290, 542. 
tereticaudus, 65, 78, 80, 101 

Climatological table for southern 
California, 504. 

Cnemidophorus stejnegeri, 39, 40, 46, 
47, 48, 49, 529. 

tigris, 39, 47, 48. 
tigris, 507, 509, 529. 
undulatus, 529. 

Coffee Creek, Trinity County, 342. 

Colaptes cafer collaris, 69, 134, 381, 
404; measurements of, 138. 


599 


osa, 


994. 


> oe 


[548] 


chrysoides mearnsi, 65, 69, 85, 135; 
measurements of, 138. 
Colorado Desert, fauna of, 65, 97; 
climatie features of, 504; am- 
phibians and reptiles of, 509. 


Colorado River, influence of: on 
associational areas, 90; on dis- 
persal, and differentiation, of 


species, 97, 100; as an absolute 
barrier, 101, 102; map of, opp. 
274; view of, from Mellen, Ari- 
zona, opp. 276, and at The 
Needles, 288. 

Colorado River expedition, plants se- 
cured by, 53. 

Colorado salmon, 62. 

Colorado Valley, zonal and faunal 
position, 62; sectional profile of, 
88, 89; influences of Coloradu 
River on, 90; isolation of, geo- 
graphic, physiographic, and asso- 
ciational, 99; degree of isolation, 
102; birds, check-list of, 110-113, 
general accounts of, 113; mam- 
mals, check-list of, 217, general 
accounts of, 218. See also Asso- 
ciational areas. 

Colorado Valley, Lower, An Account 
of the Birds and Mammals of, 
with Especial Reference to the 
Distributional Problems Pre- 
sented, 51. 

Coon, Pacifie, 355. 

pallid, 260. 

Coot, 121. 

Cormorant, Farallon, 114. 

Corvus corax sinuatus, 68, 86, 155; 
measurements of, 156. 

eryptoleucus, 155. 

Corynorhinus macrotis intermedius, 
320. 
pallescens, 87, 263, 320. 
townsendi, 320. 

Cottontail, Arizona, 250. 

Cougar, northwestern, 360. 

Yuma, 251. 

Cowbird, dwarf, 156. 

Coyote, desert, 254, 507; cannot eat 
tortoises, 515. 

mountain, 353. 

Crane, little brown, 120. 

Creeper, Sierra, 387. 

Creosote bush, 505. 

Crotalus cerastes, 44, 46, 48, 49, 507, 
509, 511, 534. 

lucifer, 44, 46, 48, 49. 
mitchelli, 44, 46, 48, 507, 509, 533. 
ruber, 44, 46, 48, 49. 
Crotaphytus collaris baileyi, 33, 46, 
48, 507, 508, 521. 
wislizenii, 34, 46, 47, 48, 507, 508, 
522. 
Cucurbita palmata, 79. 


Index 


Cyanocitta stelleri frontalis, 382, 
401; picture of habitat, opp. 396. 

Cyprinus carpio, 62. 

Dafila acuta, 67, 115. 

Dalea spinosa, 83, 505. 

Deer, black-tailed, Columbian, 378. 

burro, 219. 

Dendragapus obscurus fuliginosus, 
380, 402. 
sierrae, 380, 401; picture of 

habitat, opp. 396. 

Dendroica aestiva aestiva, 197, 386. 
brewsteri, 69, 196, 197, 200, 405. 
rubiginosa, 69, 200. 
sonorana, 65, 69, 71, 97, 195; 

measurements of, 198, 199. 
auduboni auduboni, 69, 76, 200, 
386, 400; picture of habitat, opp. 
396. 
nigrescens, 69, 76, 200, 386, 400. 
occidentalis, 69, 201, 386, 400; pie- 
ture of habitat, opp. 396. 
townsendi, 69, 73, 201, 386, 405. 

De Vries’ theory of mutations, 473. 

Diadophis amabilis, 41, 46, 48. 

Dipodomys ecalifornicus, 367. 
ealifornicus, 366, 402. 
pallidulus, 367. 
trinitatis, 366, 367, 402, 403, 407. 

deserti, 79, 80, 224, 241, 243, 290; 
picture of burrows of, opp. 
540. 

deserti, 65, 78, 80, 92, 93, 101, 
240. 
merriami, 243, 282. 
merriami, 78, 79, 80, 82, 101, 128, 
241. 
simiolus, 242. 

Dipoides, 458, 459. 

Dipper, American, 386. 

Dipsosaurus dorsalis, 507, 508, 511, 
515. 

Distribution of River Otters in Cali- 
fornia, with Description of a 
New Subspecies, 305; literature 
cited, 309, 

Distribution of amphibians and rep- 
tiles of southeastern California, 
503. 

Dixon, Joseph, 52. 

Dove, western mourning, 122, 380. 

white-winged, 123. 

Dryobates pubescens gairdneri, 381, 
403. 

sealaris cactophilus, 65, 69, 75, 82, 

132. 

villosus orius, 381, 400. 

Duck, lesser scaup, 115. 

ruddy, 116. 

Ecologic niches, how filled, 479. 

Egret, American, 119. 

Empidonax difficilis difficilis, 69, 150. 

griseus, 69, 76, 153; measurements 

of, 152. 


hammondi, 69, 151, 382, 404; meas- 
urements of, 152. 

trailli trailli, 69, 151, 382, 404. 

wrighti, 69, 152, 382, 400; meas- 
urements of, 152. 

Eneelia farinosa, 87, 248, 288. 

Environments of southeastern Cali- 
fornia, named and characterized, 
506; canon bed, 506; rocky mesa, 
506; picture of, opp. 542; wash- 
bed, 506; low plain, and high 
plain, 506; pictures of, opp. 540; 
drifting sand, picture of, opp. 
542; canon spring, 506; animals 
of Turtle Mountain region, map 
of, opp. 538. 

Eptesicus fuscus, 80, 268. 
fuseus, 352, 404. 

Erethizon, and Castor, parallelism in, 
426. 

Eriogonum inflatum, 79, 81. 

Erismatura jamaicensis, 67, 116. 

Eucastor, 458, 459. 

Euhapsis, 458. 

Eumeeces (sp.?), 49. 
gilberti, 49. 
skiltonianus, 40, 46, 48. 

Euphagus cyanocephalus, 164, 383, 

404. 

Eutamias amoenus amoenus, 373, 
400; pictures of habitat of, opp. 
392, 396. 

hindsi, 321, 323, 329. 

merriami pricei, 324, 

quadrimaculatus, 324, 373. 

senex, 373, 400; picture of habitat, 
opp. 396. 

sonomae, 321, 323, 324. 

townsendi ochrogenys, 323, 324. 

Eutamias sonomae, A New Chipmunk 
from the Inner Northern Coast 
Belt of California, 321; litera- 
ture cited, 325. 

Eutypomys, 457. 

Evolution, considerations of, 414, 
460; polytypic, 415, 472; current 
theories of, 473. 

FEvotomys obscurus, 562, 401. 

Fagonia californica, 87. 

Falco columbarius richardsoni, 126. 
mexicanus, 86, 125. 
sparverius deserticola, 126. 

peninsularis, 127. 
phalaena, 69, 85, 126. 
sparverius, 126, 380, 404. 

Faleon, prairie, 125. 

Fauna of the Colorado Desert, 65, 97. 

Faunal gaps, 407. 

Faunas, nomenclature of, 408. 

Felis aztecus browni, 2/ 
hippolestes aztecus, 253. 
oregonensis browni, 65, 70, 76, 251; 

measurements of, 2538, 
oregonensis, 360, 404. 


Index 


Field Museum of Natural History, 
acknowledgment of material 
loaned by, 416, 497. 

Finch, purple, California, 383. 

Cassin, 383. 

Fisher, W. K., discovery of Microdi- 
podops in California, 301. 

Fisher, Pacific, 356. 

Flicker, Mearns gilded, 129, 135, 286. 

red-shafted, 134, 381. 

hybrid, discussion of, 136. 
Flycatcher, ash-throated, 147, 286. 

gray, 153. 

Hammond, 151, 382. 

olive-sided, 150, 382. 

Traill, 151, 382. 

vermilion, 153. 

western, 150. 

Wright, 152, 382. 

Four New Pocket Gophers from Cali- 
fornia, 311. 

Fox, gray, Arizona, 255. 

Townsend, 354. 
kit, desert, 255, 542. 

Franseria dumosa, 79, 81. 

Frog, leopard, Great Basin, 509. 

Fulica americana, 67, 72, 121. 

Gaps, faunal, 407; zonal, 407. 

Gavia immer, 67, 113. 

Gecko, 509. 

Geese, A Study of a Collection of, 
of the Branta canadensis Group 
from the San Joaquin Valley, 
California, 1; summary, 19; lit- 
erature cited, 20. 

Geococcyx californianus, 73, 77, 82, 
100, 130. 

Geothlypis trichas occidentalis, 72, 
74, 204, 386, 405. 
scirpicola, 69, 72, 202, 276; meas- 

urements of, 203. 

Gerrhonotus scincicauda ignavus, 38, 
43, 46, 48. 

Gila elegans, 62. 

‘*Gila monster,’’ 509, 510. 

Gilbert, C. H., 53. 

Glaucomys sabrinus flaviventris, 375, 
401, 402, 407; picture of locality, 
opp. 396. 

Gnateatcher, plumbeous, 214, 282. 

western, 213. 

Goldfinch, green-backed, 165, 383. 

Lawrence, 166. 
willow, 383. 

Goose, lesser snow, 116. 

Gopher, Diablo, 313. 

La Puerta, 315. 
Carrizo Plain, 314. 
Gopher, pocket, 25, 26. 

California, 25. 
Ehrenberg, 239. 
Imperial Valley, 259. 
Red Bluff, 364. 
Trinity, 365. 

Yolla Bolly, 312. 


[550] 


Gopher, California Pocket, Nocturnal 
Wanderings of the, 25. 

Gophers, trapping, 26, 27, 28. 

Gophers, Four New Pocket, from 
California, 311. 

Goshawk, western, 380. 

Grass, galleta, 506. 

Greasewood, 505. 

Grinnell, H. W., 317. 

Grinnell, Joseph, 51, 301, 305, 311, 
321, 399, 504. 

Grizzly Creek, Trinity County, 345; 
picture of locality, opp. 396. 

Grosbeak, black-headed, 179. 
Pacific, 385. 

blue, Arizona, 180. 
evening, western, 383. 

Grouse, Sierra, 380. 
sooty, 380. 

Grus canadensis, 67, 120. 
mexicana, 120. 

Guiraca caerulea lazula, 69, 74, 97, 

98, 180. 

Gulo luseus hylaeus, 467. 
luseus, 467. 
luteus, 467. 

Habitat limitations, 508; of am- 
phibians and reptiles in the 
Turtle Mountain region, 507. 

Hall ees Mee: 

Hasselborg, Allen E., 10. 

Hawk, Cooper, 124, 380. 
marsh, 123. 
pigeon, Richardson, 126. 
red-tailed, western, 125. 
sharp-shinned, 124, 380. 
sparrow, 380. 
desert, 126. 

Texas nighthawk, 142. 

Helena, Trinity County, 339. 

Heleodytes brunneicapillus couesi, 76, 
82, 208. 

Henshaw, H. W., acknowledgment to, 
317, 321. 

Hermit thrush, 7. 

Herodias egretta, 119. 

Heron, night, black-crowned, 119. 
blue, pallid, 116. 
green, Anthony, 119. 

Hesperiphona vespertina 
383, 404. 

Hypsiglena ochrorhynehus, 42. 

Hirundo erythrogastra, 185. 

Holliger, C. D., collector of mammals, 
301. 

Horned toad, desert, 509, 511, 527. 

Hummingbird, black-chinned, 143. 
ealliope, 382. 

Costa, 145. 
rufous, 382. 

Hunters’ Camp, Trinity County, 345. 

Hylocichla guttata, 7. 
guttata, 70, 215. 
nanus, 215, 388, 405. 


montana, 


— 


Index 


slevini, 388, 402; picture of lo- 
eality, opp. 596. 
ustulata swainsoni, 388, 405; pic- 
ture of locality, opp. 396. 
ustulata, 70, 215, 388, 405; pic- 
ture of locality, opp. 396. 

Hyptis emoryi, 87, 146. 

Hystricops, 458. 

Ibis, glossy, white-faced, 116. 

wood, 116. 

Ieteria virens longicauda, 70, 204, 
386, 405. 

Teterus bullocki, 69, 82, 164, 3883, 404. 

eucullatus nelsoni, 69, 163. 

Iguana, desert, 508, 515. 

Tridoproene bicolor, 185. 

Ironwood, 505. 

Ischyromyidae, 457. 

Isolation, associational, 99; geo- 
graphic, 99, 481, manner of oper- 
ation, 482; geographical, and mi- 
gration, Wagner’s theory of, 
475; physiographic, 99; in poly- 
typic evolution, 415; its relation 
to speciation, 462. 

Ixoreus naevius naevius, 388, 405. 

Jack rabbit, California, 376. 

Jolorado Desert, 250. 
Jackson Lake, Siskiyou County, 341. 
Jay, blue-fronted, 382. 

California, 382. 

Jones, L. Hollister, 52. 

Junco, Sierra, 172, 384. 

Juneco oreganus thurberi, 172,. 384, 
401; picture of locality, opp. 396. 

Kangaroo Creek, Siskiyou County, 
347. 

Kangaroo Mouse, Mono, 302. 

Kangaroo rat, big desert, 240, 290. 

Merriam, 241. 
Kellogg, Louise, 295, 335, 405. 
Killdeer, 121, 379. 
Kingbird, western, 146. 
Kingfisher, belted, 151. 
Texas green, 131. 

Kinglet, ashy, 212. 
golden-crowned, western, 388. 
ruby-crowned, 388. 

Kofoid, C. A., 53, 416, 472, 475. 

Lampropeltis boylei, 41, 46, 48, 49, 
507, 509, 531. 

pyrrhomelaena multicincta, 41, 45, 
46, 

Lanius ludovicianus excubitorides, 
7SuSones: 

Lanivireo solitarius ecassini, 69, 82, 
189, 385, 405; picture of locality, 
opp. 396. 

Larrea, 315. 

divaricata, 79, 80, 87, 282. 
tridentata, 505. 
Lasionycteris noctivagans, 352, 404. 
Latax, 469. 


Lepus californicus ealifornicus, 377, 


402. 
deserticola, 65, 78, 80, 82, 101, 
250. 

washingtonii klamathensis, 376, 
401. 


Lichanura roseofusea, 41, 46, 48. 

Light, J. E., 25. 

Linnet, California, 164, 383. 

Literature cited, 20, 29, 50, 269, 304, 
309, 325, 334, 389, 410, 490, 536. 

Lizard, color change in, 511; tail- 
dropping faculty of, 510; fe- 
moral pores of, 511. 

Lizard, alligator, San Diegan, 38. 

Bailey, 33. 
brown-shouldered, 35. 
collared, Bailey, 508, 511, 521. 
desert, 509, 524. 
night, 509, 528. 
whip-tailed, 509, 511, 529. 
dusky, sealed, 37. 
fence, 35. 
gridiron-tailed, 33, 508, 511, 519. 
horned, Blainville, 38. 
desert, 38. 
leopard, 34, 508, 522. 
Mearns, 34. 
mountain, 36. 
orange-throat, Belding, 40. 
rough-sealed, 87, 505, 509, 511, 526. 
sand, ocellated, 508, 511, 516. 
whip-tailed, Stejneger, 39. 
Loon, common, 113. 
Lophortyx californica californica, 
403. 
vallicola, 379. 
gambeli, 65, 73, 74, 75, 77, 82, 122. 
Lutra californica, 309. 
canadensis brevipilosus, 307; meas- 
urements of skulls of, 308; 
plates of, opp. 310. 
pacifica, 307. 
periclyzomae, 307. 
sonora, 305. 
Lycium andersoni, 79, 84, 143, 144, 
145. 
parishi, 84. 

Lynx eremicus californicus, 254. 

eremicus, 76, 82, 253. 
fasciatus, 360, 402. 
pallescens, 360. 

Macrotus californicus, 80, 269. 

Mallard, 115. 

Mammalian Genus Microdipodops 
from California, A Second 
Species of the, 301; literature 
eited, 304. 

Mammals and Birds of the Lower 
Colorado Valley, An Account of 
the, with Especial Reference to 
the Distributional Problems Pre- 
sented, 51; literature cited, 269. 


Index 


Mammals of Colorado Valley, check- 
list of, 217; general accounts of, 
218. 

Mammals and Birds Found in Por- 
tions of Trinity, Siskiyou and 
Shasta Counties, California, Re- 
port upon, 335; literature cited, 
389. 

Mammals of the Trinity region, 
check-list of, 350; general ac- 
counts of, 351-379. 

Marila affinis, 67, 115. 

Marten, pine, northwestern, 555. 

Martes caurina, 356. 
caurina, 355, 400, 467; picture 

of habitat, opp. 396. 
origenes, 467. 
pennanti pacifica, 356, 400, 467; 
measurements of, 357. 
pennanti, 467. 

Martynia proboscidea, 79. 

Mayten, Siskiyou County, 339. 

Meadowlark, 163. 

western, 383. 
Meadow mouse, 27. 
Melanerpes formicivorus bairdi, 381, 
404. 
Melopelia asiatica trudeaui, 69, 123. 
Melopiza lincolni, gracilis, 176. 
lincolni, 69, °73, 176, 384, 400; 
picture of habitat, opp. 396. 

striata, 176. 

melodia fallax, 69, 73, 74, 173. 

fisherella, 384. 

merrilli, 384. 

montana, 174. 

rufina, 384, 402. 

saltonis, 65, 69, 72, 73, 74, 91, 
97, 174, 276. 

Mendelian inheritance, 477. 

Mephitis estor, 65, 70, 73, 76, 97, 
257, 258, 294, 469; measurements 
of, 258. 

mesomelas varians, 469. 
occidentalis, 257, 258, 402, 468. 
holzneri, 469. 
major, 468. 
notata, 468. 
occidentalis, 359. 
spissigrada, 468. 
platyrhina, 469. 

Merganser, red-breasted, 114. 

Mergus serrator, 67, 114. 

Merriam, J. C., cited, 105. 

Michael, E. L., 475, 476. 

Microdipodops, the Mammalian 
Genus, a Second Species of, 
from California, 301. 

Microdipodops californicus, 301, 302, 
304. 

megacephalus, 302, 304. 
oregonus, 302, 304. 

pallidus, 802, 304. 

polionotus, 302, 3038. 


| 
Micropallas whitneyi, 85, 129. | 
Microtus californicus, 27. | 
ealifornicus, 363, 402. 
montanus montanus, 363, 401. 
mordax mordax, 363, 400; picture 
of locality, opp. 396. 
Mimus polyglottos leucopterus, 76, 
82, 206. 
Mink, Pacific, 359. 
Mockingbird, western, 206, 280. 
Mojave Desert, climatic features of, 

504. 

Mole, Central California, 351. 
shrew, large, 351. 

Moles, trapping, 26. 

Molothrus ater artemisiae, 158. 

obseurus, 69, 71, 76, 156, 157, 

158; measurements of, 159, 
160. 
Mount Eddy, Siskiyou County, 348. 
Mountain sheep, 218. 
Mouse, cafion, Stephens, 229, 534. 
harvest, desert, 233. 

Klamath, 360. 
jumping, Allen, 359. 
kangaroo, Mono, 302. 
meadow, 27. 

California, 363. | 

cantankerous, 363. 

Peale, 363. 
pocket, Colorado Desert, 245. 

intermediate, 248. 

long-tailed, 244. 

spiny, 249. 

Yuma, 243. { 
red-backed, dusky, 362. 
white-footed, Boyle, 361. 

desert, 229. 

Gambel, 361. 

Gilbert, 361. 

Sonora, 227. 

Miihlenbergia debilis, 87. 
Muridae, 469. 
Muskrat, pallid, 237. 
Mustela arizonensis, 467. 
longicauda, 467. 
muricus, 358, 401, 467. 
saturata, 358, 401, 467. 
streatori, 467. 
vison energumenos, 349, 404, 468. 

lacustris, 468. 

nesolestes, 468. 
xanthogenys muna, 468. 

oregonensis, 468. 

xanthogenys, 468. 

Mustelidae, 467. 

Mutations, De Vries’ theory of, 473. 
Myadestes townsendi, 388, 400. 
Mycteria americana, 67, 116. 
Myiarchus cinerascens cinerascens, 

69, 75, 82, 85, 147, 148; measure- 

ments of, 148. 

Myiochanes richardsoni richardsoni, 

69, 75, 150, 382, 404. 


Index 


Myotis californicus californicus, 266, 
317, 318. 
pallidus, 65, 76, 78, 265; meas- 
urements of, 266. 
quercinus, 317, 318. 
longicrus, 319. 
longicrus, 352, 404. 
occultus, 65, 70, 263; measurements 
of, 264; figures of dental 
series, 265. 
velifer, 80, 266. 
yumanensis saturatus, 318, 319. 
sociabilis, 318, 319. 
yumanensis, 318, 319. 
Natural selection, 473. 
Neosorex bendirii albiventer, 466. 
bendirii, 466. 
palmeri, 466. 
Neotoma, picture of caves inhabited 
by, 542. 


Neotoma albigula venusta, 65, 73, 75, 
76, 97, 98, 101, 128, 233, 234, 
280. 

cinerea occidentalis, 362, 400. 
eumulator, 234. 
fuscipes fuscipes, 361, 403. 
intermedia desertorum, 87, 100, 
101, 102, 104, 233, 235, 288, 505. 
lepida, 236. 
stephensi, 336. 
Nettion carolinense, 67, 115. 
Neurotrichus gibbsi major, 351, 402. 


Nighthawk, Texas, 142. 

Nocturnal Wanderings of the Cali- 
fornia Pocket Gopher, 25; liter- 
ature cited, 29. 

Nomenclature of beavers, 417. 

Notes on the Local Distribution and 
Habits of the Amphibians and 
Reptiles of Southeastern Califor- 
nia in the Vicinity of the Turtle 
Mountains, 503; literature cited, 
536. 

Nucifraga columbiana, 382, 400. 

Nuteracker, Clarke, 382. 

Nuthateh, red-breasted, 387. 

slender-billed, 387. 

Nuttallornis borealis, 69, 150, 382, 
404; picture of locality, opp. 396. 

Nycticorax nycticorax naevius, 67, 
119. 

Nyctinomus mexicanus, 70, 268. 

Odocoileus columbianus columbianus, 
378, 402. 

hemionus eremicus, 65, 70, 76, 82, 
219. 

Olneya tesota, 83, 505; figure of, opp. 
284. 

Ondatra zibethica pallida, 68, 72, 97, 
98, 101, 237, 276. 

Oporornis tolmiei, 73, 74, 201. 

Oreortyx picta picta, 379, 402; pic- 
ture of locality, opp. 396. 

Oreoscoptes montanus, 76, 82, 206. 


[553] 


Oreospiza chlorura, 69, 73, 74, 178, 
385, 400. 

Oriole, Bullock, 164, 383. 

hooded, Arizona, 163. 

Osprey, 127. 

Otters, River, in California, Distribu- 
tion of, with Description of a 
New Subspecies, 305; literature 
cited, 309. 

Otus asio gilmani, 65, 69, 85, 128. 

Ovis canadensis gaillardi, 219. 
nelsoni, 86, 218. 

Owl, barn, 26, 127. 

elf, 129. 

horned, western, 129. 

sereech, Saguaro, 128, 286. 
Gxyechus vociferus, 379, 404. 


vociferus, 68, 72, 121. 

Pandion haliaétus carolinensis, 68, 
127. 

Parallelism in Castor and Erethizon, 
426. 


Paramys, 457. 

Parkinsonia torreyana, 83; figure of, 
opp. 284. 

Passer domesticus, 166, 383, 404. 

Passerculus sandwichensis alaudinus, 
78, 167. 
nevadensis, 78, 167. 

Passerella iliaca, 7. 
megarhyncha, 384, 401. 
meruloides, 384, 404. 
unalascheensis, 384, 404. 

Passerina amoena, 69, 181, 385, 404. 

Pelecanus erythrorhynchos, 67, 114. 

Pelican, white, 114. 

Penthestes atricapillus occidentalis, 
387, 402. 

gambeli gambeli, 387, 400; picture 
of locality, opp. 3 
rufescens rufescens, 
Perityle emoryi, 87. 


96. 
387, 402. 


Perognathus bombycinus, 80, 101, 
243. 
formosus, 80, 82, 87, 92, 101, 104, 
244, 246, 
intermedius, 65, 80, 87, 101, 246, 
248, 288. 
panamintinus bangsi, 243. 
penicillatus, 224, 243, 246, 282, 290. 


angustirostris, 247. 
penicillatus, 65, 73, 76, 78, 80, 
82, 84, 93, 101, 128, 245. 
spinatus, 245, 246. 
spinatus, 65, 80, 82, 87, 95, 101, 
249. 
Peromyscus boylii boylii, 361, 402. 
erinitus stephensi, 86, 90, 101, 104, 
229, 230, 534. 
eremicus, 224, 230, 243, 290. 
eremicus, 65, 78, 80, 82, 90, 101, 
229. 
maniculatus, 469. 
gambeli, 361, 404; picture of lo- 
eality, opp. 396. 


rubidus, 361. 
sonoriensis, 70, 71, 73, 75, 76, 97, 


truei gilberti, 3861, 402. 
Petrochelidon lunifrons lunifrons, 68, 
183. 
tachina, 185. 
Pewee, wood, western, 150, 382. 
Phainopepla, 187, 280. 
nitens, 76, 77, 82, 187. 


Phalacroecorax auritus albociliatus, 
67, 114. 

Phalaenoptilus nuttalli californicus, 
141. 


nuttalli, 69, 77, 82, 139; measure- 
ments of, 141. 
nitidus, 69, 77, 82, 159; measure- 
ments of, 142. 
Phloeotomus pileatus abieticola, 581, 
404. 
Phoebe, black, 149, 382. 
Say, 149. 
Phoradendron ecalifornicum, 77, 84. 
Phragmites communis, 71; figure of, 
opp. 276. 
Phrynosoma~ blainvillei 
38, 46, 48, 49, 528. 
platyrhinos, 38, 46, 48, 49, 507, 509, 
527, 528. 
Physiological selection, 473. 
Pintail, 115. 
Pipilo aberti, 65, 69, 74, 75, 76, 77, 91, 
97, 98, 100, 177. 
erissalis earolae, 385, 403. 
maculatus eurtatus, 73, 74, 177. 
faleineilus, 385, 403. 
megalonyx, 177. 
montanus, 177. 
Pipistrellus hesperus, 266. 
hesperus, 78, 82, 87, 267. 
merriami, 268. 
Pipit, 206. 
Piranga Iudoviciana, 69, 76, 80, 182, 
380, 404. 
rubra cooperi, 65, 69, 71, 97, 98, 
182. 
Pisobia minutilla, 68, 121. 
Pituophis catenifer, 43, 46, 48. 
Planesticus migratorius propinquus, 
70, 76, 216, 388, 405. 
Plegadis guarauna, 67, 116. 
Plesiarctomys, 457. 
Pluchea, 315. 
sericea, 73; figure of, opp. 278. 
Pocket gopher. See Gopher. 
Polioptila caerulea obscura, 70, 76, 
82, 213, 214. 
plumbea, 65, 74, 76, 82, 214. 
Poecetes gramineus confinis, 77. 
Poor-will, frosted, 139. 
Nuttall, 139. 
Populus fremonti, 70. 
Procyon pallidus, 65, 68, 70, 72, 97, 
260, 467; measurements of, 262. 


blainvillei, 


[554] 


psora, 260. 
californica, 467. 
pacifica, 355, 404, 466. 
psora, 467. 

Procyonidae, 466. 

Prosopis juliflora, 76; figure of, opp. 
280. 

pubescens, 70. 

Psaltriparus minimus minimus, 387, 
403. 

Ptychocheilus lucius, 62. 

Pyrocephalus rubinus mexicanus, 69, 
76, 153. 

Quail, desert, 122, 507. 

mountain, 379. 
valley, 379. 
Querquedula cyanoptera, 67, 115. 
Rabbit, brush, redwood, 378. 
snowshoe, Oregon, 276. 

Rabbit. See also Jack rabbit. 

Racer, California, 43. 
red, 42, 509, 532. 

Rana pipiens brachycephala, 509. 

Rat, cotton, desert, western, 230. 

kangaroo, big desert, 240, 290. 
Merriam, 241. 
Trinity, 366. 

wood, bushy-tailed, western, 362. 
Colorado Valley, 233. 
desert, 235, 505. 
dusky-footed, 361. 

Rattlesnake, bleached or pallid, 44, 
509, 533. 

Paecifie, 44. 
red, 44. 

Raven, western, 155. 

Recurvirostra americana, 120. 

Regulus calendula, 388, 405. 
eineraceus, 70, 212, 213. 

satrapa olivaceus, 388, 405; pic- 
ture of locality of, opp. 396. 

Reithrodontomys megalotis deserti, 
65, 70, 72, 97, 101, 233. 
klamathensis, 340, 360, 403. 
longicaudus, 360. 

Report upon Mammals and Birds 
Found in Portions of Trinity, 
Siskiyou and Shasta Counties, 
California, with Description of a 
New Dipodomys, 335; literature 
cited, 389. 

Reptiles of Southeastern California, 
503; habitat limitations of, in 
Turtle Mountains region, 507; 
vernacular names for, 508; des- 
ert, characters of, 510. 

Reptiles, The, of the San Jacinto 
Area of Southern California, 31; 
literature cited, 50. 

Reptiles and Amphibians of South- 
eastern California in the Vicin- 
ity of Turtle Mountains, Notes 
on the Local Distribution and 
Habits of the, 503. 


Index 


Rhinocheilus lecontei, 42, 46, 48, 49. 

Richmond, C. W., 1. 

River Otter. See Otter. 

River Otters in California, Distribu- 
tion of, with Description of a 
New Subspecies, 305. 

Roadrunner, 130, 131. 

Robin, western, 216, 280, 388. 

Rowley, John, acknowledgment to, 
SLT. 

Rush Creek, Siskiyou County, 346. 

Salamander, garden, 327. 

Salix fluviatilis, 70. 

nigra, 70. 

Salmon, Colorado, 62. 

Saloon Creek Divide, Trinity County, 
343; illustration showing nature 
of, opp. 3892. 

Salpinctes obsoletus obsoletus, 80, 86, 
90, 208. 

Salvadora grahamiae, 42, 46, 48, 49. 

San Jacinto area of Southern Cali- 
fornia, reptiles of, 31. 

Sandpiper, least, 121. 

spotted, 121, 379. 
Sapsucker, red-breasted, Sierra, 381. 
red-naped, 132. 

Sauromalus ater, 34, 46, 48, 49, 507, 
509, 511, 522. 

Sayornis nigricans, 68, 149, 382, 404. 

sayus sayus, 80, 86, 149. 


Scapanus latimanus latimanus, 351, 
401. 

Sceloporus biseriatus, 35, 36, 46, 48, 
49. 


graciosus, 36, 45, 46, 49. 

magister, 37, 46, 47, 48, 507, 509, 
526; picture of caves inhabited 
by, opp. 542. 

oreutti, 37, 38, 46, 47, 48, 49. 

Seiuravus, 457. 

Seiurus douglasii albolimbatus, 375, 
401. 

griseus griseus, 375, 404. 

Scott River Valley, Siskiyou County, 
340; illustration showing nature 
of, opp. 392. 

Second Species, A, of the Mammalian 
Genus Microdipodops from Cali- 
fornia, 301; literature cited, 304. 

Selasphorus rufus, 382, 404. 

Shasta County, map of portion of 
traversed in zoological explora- 
tions, 337. 

Shasta, Mount, included 
fauna, 406, 408. 

Shoveller, 115. 

Shrew, Monterey, 352. 

Sierra Nevada, 351. 

Shrew-mole, large, 351. 

Shrike, white-rumped, 188. 

Sialia currucoides, 389, 400. 

mexicana anabelae, 216. 
bairdi, 216. 


in Sierra 


occidentalis, 76, 216, 389, 405. 
Sidewinder, 44, 509, 511, 534. 
Sigmodon hispidus arizonae, 233. 

eremicus, 70, 72, 97, 98, 101, 230; 

measurements of, 232. 

Sigmogomphius, 458. 

Siskin, pine, 383. 

Siskiyou County, map of portion of 
traversed in zoological explora- 
tions, 337. 

Sitkan district, 7. 

Sitta canadensis, 387, 405. 

carolinensis aculeata, 387, 405. 

Skink, western, 40. 

Skunk, spotted, Arizona, 259. 
California, 359. 

striped, Arizona, 257. 
Northern California, 359. 
Snake, burrowing, desert, 509, 510, 
531. 
faded, 45. 
garter, California, 43. 
gopher, western, 43. 
ground, Texas, 509, 530. 
king, coral, 41. 

Boyle, 509, 531. 
long-nosed, 42. 
milk, Boyle, 41. 
night, spotted, 42. 
patch-nosed, 42. 
ring-necked, western, 41. 

Snake. See also Rattlesnake; Side- 
winder. 

Solitaire, Townsend, 388. 

Sonora episcopa, 507, 509, 530. 
occipitalis, 507, 509, 510, 531. 
Sonoran, Lower, zonal diagnosis, 65. 

Sorex californicus, 465. 

halicoetes, 465. 

montereyensis mariposae, 402, 466. 
montereyensis, 352, 402, 466. 

obseurus longicauda, 466. 
obseurus, 466. 
ventralis, 466. 

sinuosus, 465. 

tenellus lyelli, 466. 

myops, 466. 

nanus, 466. 

tenellus, 466. 

vagrans amoenus, 351, 401, 465; 

picture of locality, opp. 396. 

vagrans, 401, 465. 

Soricidae, 465. 

Sparrow, black-chinned, 172. 

Brewer, 171, 282. 
chipping, western, 170, 384. 
desert, 173. 

English, 283. 

fox; 7. 

Shumagin, 354. 

thick-billed, 384. 

Yakutat, 384. 

golden-crowned, 384. 
intermediate, 168. 


Index 


lark, western, 167, 383. 
Lincoln, 176, 384. 
sage, Nevada, 173, 282. 
Savannah, Nevada, 167. 
western, 167. 
song, Modoe, 384. 
Rocky Mountain, 173. 
rusty, 384. 
Salton Sink, 174. 
vesper, western, 166. 
white-crowned, 167. 

Spatula clypeata, 67, 115. 

Speciation, of beavers of western 
North America, 413, 495; rela- 
tion of isolation to, 462; action 
of geographic isolation in process 
ot, 482. 

Sphaeralcea ambigua, 516. 

Sphyrapicus varius daggetti, 381, 
401, 

varius nuchalis, 69, 75, 132. 

Spilogale arizonae arizonae, 73, 259, 

468. 
martirensis, 468. 
gracilis gracilis, 468. 
saxatilis, 468. 
phenax latifrons, 359, 468. 
olympica, 468. 
phenax, 359, 402, 468. 

Spinus pinus pinus, 383, 404. 

Spirostachys occidentalis, 79. 

Spizella atrogularis, 86, 172. 

breweri, 78, 80, 86, 171, 172. 

pallida, 172. 

passerina arizonae, 9, 76, 86, 170, 
384, 404. 

Squirrel, flying, Trinity, 375. 

gray, California, 375. 

ground, antelope, 221. 
Douglas, 372. 
golden-mantled, Trinity, 374. 
Harris, 219, 292. 
round-tailed, 224. 

Stelgidopteryx serripennis, 68, 86, 
186. 

Stellula calliope, 382, 400. 

Steneofiber, 457, 459. 

Stephens, Frank, 52, 219. 

Stern, N., collector of mammals, 301. 

Sterna forsteri, 67, 114. 

Study, A, of a Collection of Geese 
of the Branta canadensis Group 
from the San Joaquin Valley, 
California, 1; literature cited, 20. 

Sturnella neglecta, 77, 163, 383, 404. 

Suaeda suffrutescens, 79. 

Summerville, Siskiyou County, 544. 

Swallow, barn, 185. 

cliff, 183. 
northern violet-green, 185. 
rough-winged, 186. 
tree, 185. 
Swarth, Harry S., 1, 53. 


Swift, long-tailed, 509, 511, 525. 
Vaux, 143. 
white-throated, 143. 
Sylvilagus auduboni arizonae, 70, 73, 
75, 76, 78, 101, 250. 
bachmani ubericolor, 378, 403. 
Tachycineta thalassina lepida, 185. 
Tanager, Cooper, 182. 
western, 182, 380. 
Taxidea taxus berlandieri, 78, 259, 
469. 
infusea, 469. 
neglecta, 469. 
Taylor, W. P., 53, 297, 413, 497. 
Tea, desert, 506. 
Teal, cinnamon, 115. 
green-winged, 115. 
Telmatodytes palustris plesius, 72, 
100, 211. 
Tern, Forster, 114. 
Testudo agassizii, 507, 508, 512. 
polyphemus, 513. 
Thamnophis elegans, 49. 
Hammondi, 43, 46, 48. 
Thomomys albatus, 65, 78, 101, 239. 
angularis angularis, 313, 314. 
pasealis, 314, 315. 
bottae bottae, 25, 313, 314. 
pasealis, 314, 315. 
cabezonae, 316. 
chrysonotus, 80, 101, 222, 239. 
diaboli, 313. 
infrapallidus, 314, 315. 
leucodon navus, 313, 340, 364, 402. 
monticola monticola, 312, 365. 
pinetorum, 312, 365, 401, 402, 
407; picture of locality of, opp. 
392, 396. 
premaxillaris, $12. 
nigricans nigricans, 313, 315. 
puertae, 315. 
perpallidus, 239. 
Thrasher, crissal, 207, 280. 
sage, 206. 
Three New Races of Vespertilionid 
Bats from California, 317. 
Thrush, hermit, 7. 
Alaska, 215. 
dwarf, 215, 388. 
Monterey, 388. 
olive-backed, 388. 
russet-backed, 215, 388. 
varied, 388. 
Thryomanes bewicki drymoecus, 387, 
403. 
bewicki eremophilus, 78, 82, 209, 
210. 
Titmouse, plain, 387. 
Toad, Arroyo, 331. 
horned, desert, 509, 511, 527. 
spotted, 508, 512. 
Tortoise, desert, 508, 512, 518, 514. 
Tower House, Shasta County, 339. 


Index 


Towhee, Abert, 177, 280. 
brown, northern, 385. 
green-tailed, 178, 385. 

Nevada, 177. 
Sacramento, 385. 

Toxostoma crissale, 65, 74, 75, 76, 77, 
97, 98, 100, 207. 

Tree, smoke, 505. 

Trinity County, report upon mam- 
mals and birds found in portions 
of, 335; map showing portions 
traversed in zoological explora- 
tions, 337; description of locali- 
ties in region of, 338-350. 

Trinity region, description of mam- 
mals of, 3850-379, of birds of, 
379-389; analysis of vertebrate 
fauna of, 399, 405, 406, 409, 410; 
fauna of, allied with Sacramento 
fauna, 407. 

Troglodytes aédon parkmani, 70, 210, 
387, 405, 

Trogontherium, 458. 

Tropical zone, lower Colorado River 
valley referred to, 66. 

Turtle Mountains, California, deserip- 
tion of, 505; amphibians and rep- 
tiles of, 503, their habitat limi- 
tations, 507, check-list of, 508- 
509; canon spring environment 
represented in, 506; map show- 
ing animal environments opp. 
538; picture of high plain en- 
vironment at south end of, opp. 
540, of rocky mesa environment, 
opp. 542. 

Two New Aplodontias from Western 
North America, 497. 

Tyrannus verticalis, 69, 146. 

Uma notata, 507, 508, 510, 511, 516; 
measurements of, 578; picture of 
home of, opp. 542; pictures of, 
opp. 544. 

rufopunctata, 516, 517. 
scoparia, 516, 517. 

United States Department of Agri- 
culture, Bureau of Biological 
Survey, acknowledgment of loan 
of material, 416, 497. 

United States National Museum, 
acknowledgment of loan of ma- 
terial, 1, 416. 

Urocyon cinereoargenteus californi- 
eus, 255, 404. 
scotti, 70, 82, 255; measurements 

of, 256. 
sequoiensis, 404. 
townsendi, 354, 404. 

Ursus americanus, 353, 404. 

Uta graciosa, 507, 509, 511, 525. 
mearnsi, 54, 46, 48, 49. 
stansburiana, 35, 46, 48, 507, 509, 

511. 
elegans, 524. 


Verdin, 211, 282. 

Vermivora celata celata, 69, 73, 194. 
lutescens, 69, 194, 195, 386, 405; 

picture of locality of, opp. 396. 
luciae, 65, 76, 77, 97, 98, 191, 292, 
294. 
ruficapilla gutturalis, 69, 193, 194, 
385, 400; picture of locality of, 
opp. 396. 
ruficapilla, 194. 

Vertebrate Fauna of the Trinity 
Region of Northern California, 
An Analysis of the, 399; litera- 
ture cited, 410. 

Verticaria hyperythra beldingi, 40, 
46, 48, 49. 

Vespertilionid Bats from California, 
Three New Races of, 317. 

Vesperugo merriami, 267. 

Vireo belli arizonae, 65, 69, 71, 73, 76, 
97, 98, 189, 190. 

huttoni huttoni, 285, 405. 

Vireo, Cassin, 189, 385. 
Hutton, 385. 
least, Arizona, 189. 
warbling, western, 189, 385. 

Vireosylva gilva swainsoni, 69, 76, 
189, 380, 405. 

Vulpes macrotis arsipus, 65, 80, 255; 
measurements of, 255. 

Vulture, turkey, 123. 

Wagner and Jordan’s law, 464. 
Wagner’s theory of migration and 
geographical isolation, 475. 

Warbler, Audubon, 200, 386. 
Calaveras, 193, 385. 
gray, black-throated, 200, 386. 
hermit, 201, 386. 

Lucy, 191, 292, 294. 
lutescent, 195, 386. 
orange-crowned, 194. 
pileolated, Alaska, 205. 
golden, 205, 386. 
Tolmie, 201. 
Townsend, 201, 386. 
yellow, Alaska, 200. 
California, 200, 386. 
Sonora, 195. 

Waxwing, Bohemian, 187, 385. 

Weasel, least, Sierra, 358. 
Siskiyou, 358. 

Wildeat, barred, 360. 
desert, 253. 

Wildeat Peak, Siskiyou County, 

California, 342. 

Wilsonia pusilla chryseola, 70, 76, 
205, 386, 405. 
pileolata, 70, 76, 205. 

Wood rat, Colorado Valley, 233. 

desert, 235. 

Woodpecker, cactus, 132. 
California, 381. 
Gairdner, 381. 

Gila, 133, 286. 


Index 


hairy, Modoe, 381. 
Lewis, 381. 
pileated, northern, 381. 
white-headed, northern, 381. 
Wren, bewick, desert, 209, 282. 
cactus, 208, 282. 
canon, 209. 
house, western, 210, 387. 
marsh, western, 211. 
rock, 208. 
San Joaquin, 387. 
Wren-tit, pallid, 387. 
Xanthocephalus xanthocephalus, 69, 
72, 160. 
Xantusia vigilis, 507, 508, 509, 528. 
Xenopicus albolarvatus albolarvatus, 
381, 400. 
Xyrauchen cypho, 62. 


Yellowthroat, tule, 202. 
western, 204, 386. 
Yucea mohavensis, 506, 526; picture 
of, opp. 540. 
Yueea, tree, 506. 
Zamelodia melanocephala capitalis, 
180, 385, 404. 
melanocephala, 69, 82, 179, 180. 
Zapus trinotatus alleni, 369, 401; pic- 
ture of locality, opp. 396. 
Zenaidura macroura marginella, 77, 
82, 122, 380, 404. 
Zonal gaps, 407. 
Zonotrichia coronata, 384, 404. 
leucophrys gambeli, 69, 73, 74, 76, 
78, 82, 168. 
leucophrys, 73, 74, 76, 82, 167. 


ERRATA 


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For scalded read sealed. 


Page 476, line 11 from bottom. For transversing read traversing. 


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