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consisted of 90% grasses plus other herbaceous stems and leaves (Beeman and
Pelton 1977). Graminoids and forbs bulked 67% of the spring black bear food
in Yosemite National Park (Goldsmith et al. 1980). Black bears in the San
Bernardino Mountains of southern California averaged 53% herbaceous plants
consumed during the spring seasons of 1975 and 1976 (Boyer 1976). Herbage
was the principal food of Yosemite bears during spring and early summer (Gra-
ber 1982). The results of this study parallel the above findings, as well as many
others (Tisch 1961, Hatler 1972, Poelker and Hartwell 1973, Kellyhouse 1975,
Landers et al. 1979).

Graminoid flowers were seldom found in food habit analyses. Protein content
of pre-flowering graminoids is higher than that of post-flowering plants and the
highest protein digestibility also is associated with the pre-flowering phase
(Mealy 1975). Hence, there is an indication that bears were using grass-type
foods when the protein content was greatest.

Soft Mast

The most abundant berry crop in the study area is manzanita. As the nutrition-
al quality of herbaceous plants declined, black bears shifted to this food re-
source. Bears utilized manzanita in all stages: flowers and unripe, ripe, and dried
fruit. Where manzanita occurs at lower elevations and/or on south-facing
slopes, flowers and unripe fruit were eaten as early as May. Dried berries from
the previous year were noted in a few scats collected in early spring. Use of soft
mast continued throughout the year, with peak consumption occurring in late
summer and fall. Manzanita berries, like herbaceous plants, are low in digestible
energy (Goldsmith et al. 1981 ); consequently, large quantities of this food must
be eaten.

Hard Mast

Another dietary shift occurred as the acorn crop matured, but was not as
pronounced as in other California studies. Importance values for acorns in-
creased in September (Figures 2, 3, and 4), but there were indications that acorn
use by bears is greater than these data show. In the fall, scats were difficult to
find because of deciduous leaf cover and general inaccessibility of oak habitats.
Most oak woodlands in the study area are found on steep canyon slopes far from
roads. Helicopter transport to one such area (Royal Gorge) in January 1981
enabled a ground search that revealed, in just a few hours, over 40 bear scats
consisting of manzanita berries and acorns.

The nutritional value of acorns nearly approximates that of corn (Barrett
1971 ), and acorns become available when other foods have lost much of their
nutritional quality (Menke and Fry 1979). Barrett (1971) reports that another
omnivorous, monogastric animal, the wild pig, Sus scrofa, will abandon a diet
of manzanita berries in favor of an adequate acorn supply. In this study area,
bears appear to behave similarly.

During 1979, a failure of the black oak acorn crop occurred, but acorns from


canyon live oak were abundant. When both acorn crops fail, an increase in bear
depredation and nutrition-related mortality can be expected, followed by poor
reproductive success the following winter. Similar occurrences resulting from
mast shortages have been noted in other studies (Jonkel and Cowan 1971,
Rogers 1976). Even though breeding occurs in summer, the blastocyst does not
implant until November or December, but then only if the female is in good
condition. A variety of mature oak species is necessary for optimum black bear
habitat in California.

In 1980 bears did not utilize well-formed, abundant black oak acorns in one
area, but did utilize those from other oak locations. Examination of the unutilized
area revealed that 85% of those acorns had sustained insect damage. Reports
of similar damage to acorns (as high as 80%) have cited the larvae of the filbert
worm, Melissopus latiferreanus, and filbert weevil, Curculio occidentis, as the
responsible insects (Brown 1979). Larvae tunnel throughout the acorn, destroy
the embryo, and deplete its nutritional value.

Bears have been observed climbing oak trees in order to feed on acorns before
they drop (Beeman and Pelton 1977). Arboreal feeding behavior was not ob-
served in this study.

Carpenter ants, Camponotus spp.; termites Zootermopsis spp.; and yellow
jackets, Vespula sp., were consistently represented seasonally during each year
of the study (Figures 2, 3, and 4). Black bears were observed "raking" logs on
occasion, and further evidence of this behavior was often noted throughout the
study area during all seasons. Carpenter ants appeared in the diet all through the
year (Tables 1, 2, and 3). Termites mainly occurred from early spring to mid-
summer. Yellow jackets were noted in the diet from August to October. Insects
were second in importance to herbaceous plants in the spring, to berries in the
summer and to acorns in the fall (Figures 2, 3, and 4). Apparently, black bears
actively seek out social insects for food which may be their only consistent
source of high quality animal protein (Beeman and Pelton 1977). Therefore, the
density, size, and age of dead and down woody material may be critical to
maintaining optimum black bear habitat.

Other Animal Food

Mule deer, Odocoileus hemionus, as determined by the presence of deer hair
and one fawn hoof in bear scats, predominated the "other animal food" cate-
gory. One black bear scat was collected on 1 August 1979 close to where a
coyote had killed a doe. On 31 October 1979 several scats containing deer hair
and acorns were located near a deer carcass. A local cattleman (B. Dobbas,
pers. commun.) claims that black bears feed on deer gut piles during the deer
hunting season, which commences in mid-September and terminates in early
November. The study area is part of the summer range of the Blue Canyon deer
herd. Fawn drop begins in early June, peaks in mid-July, and tapers off in August.
In 1 978 and 1 980 the highest occurrence of deer hair in bear scats was associated
with the fawn drop. In 1979 deer hair in scats were most abundant during deer
hunting season; unretrieved or crippled deer surely provide a food source during
this time of year.

Animal foods other than deer occurred sporadically (Tables 1, 2, and 3) and


included Douglas squirrel, Tamiasciurus douglasii; vole, Microtus sp.; shrew,
Sorex sp.; mole, Scapanus sp.; harvest mouse, Reithrodontomys sp.; and \\no
species of ground squirrel, Spermophilus spp. In analysis, carrion v^as so desig-
nated only when animal tissue was associated with substantial amounts of
maggots (Diptera larvae). Most researchers (Beeman and Pelton 1977, Gold-
smith et al. 1981 ) have concluded that mammal and bird food items are proba-
bly taken opportunistically.

Mammalian or avian tissue may be undetected in bear scats unless bone, hair,
or feathers are present, so it is likely that this food source is under-represented
in our data. This would be particularly true with a large animal such as deer with
high body volume (flesh) compared to low body surface (hair).


Garbage had the lowest importance value of any major food type and was
notable only during the summer of 1980 (Figure 4). During that time, bear
disturbances in campgrounds were frequent in spite of the abundance of natural
foods. However, garbage, like forbs and flesh, leaves little evidence in bear scats
and may be underestimated. Black bears should be expected to seek out these
"high quality" human foods, especially if they are available during periods when
natural foods are in short supply (Hatler 1972, Rogers 1976, Goldsmith et al.
1981, Graber 1982).


Black bear feeding patterns in California approximate those cited by other
researchers. In early spring black bears feed primarily on herbaceous plants and
to a lesser extent on over-winter berries and acorns. If, or when, berry crops
become available in summer and fall, a shift to that food source occurs. Bear
foraging strategy again changes when acorns mature in early fall. Social insects
and other animal foods are consumed throughout the year. Artificial food
sources (garbage, camp supplies, orchard crops) are taken locally when oppor-
tunity permits. This information should aid in the prediction of bear food habits
in unstudied areas in California.

This research and other studies demonstrate the omnivorous feeding habits
of black bears (Poelker and Hartwell 1973, Boyer 1976, Beeman and Pelton
1977, Graber 1982). The results emphasize the importance of vegetation in the
bear diet and illustrate cyclic feeding patterns consistent with plant phenology.

The quantity and quality of known bear foods might well be used to determine
relative potential population density. Reproductive failure and/or depredation
should be anticipated if bear foods are eliminated or key mast crop failures
occur. Seasons and bag limits could be adjusted geographically in accordance
with fluctuations in important bear foods. Sufficient quantity and variety of
mast-producing trees and shrubs, particularly oaks, are essential to maintaining
optimum habitat for black bears as well as many other species of wildlife in
California. Timber harvest plans should be designed to provide for the mainte-
nance of mature mast-producing trees and shrubs, dead and down timber, and
grasses and forbs.



Barrett, R.H. 1971. Ecology of the feral hog in Tehama County, California. Dissertation. Univ. of California, Berkeley.

Beeman, L.E., and M.R. Pelton. 1 977. Seasonal foods and feeding ecology of black bears in the Smokey Mountains.

Pages 141-147 inQ. Martinka and K. McArthur, eds. Bears — their biology and managennent. Fourth Int. Conf.

Bear Research and Management, Kalispell, Mont.
Boyer, K. 1976. Food habits of black bears (Ursus americanus) in the Banning Canyon area of San Bernardino

National Forest. Unpubl. Thesis. Calif. State Polytechnic University, Pomona, Calif. 63p.
Brower, ).E., and ).H. Zar. 1977. Field and laboratory manual methods for general ecology. Wm. C. Brown Co.,

Dubuque, Iowa. 194p.
Brown, L.R. 1979. Insects feeding on California oak trees. Pages 184-194 in T.R. Plumb, ed. Ecology, management

and utilization of California oaks. USDA Forest Service. Gen. Tech. Rep. PSW-44.
Goldsmith, A., M.E. Walraven, D. Graber, and M. White. 1981. Ecology of the black bear in Sequoia National Park.

Natl. Park Serv. Final report Contract No. CY-8000-4-0022. 64p.
Graber, D., and M. White. 1978. Management of black bears and humans in Yosemite National Park. Cal-Neva

Wildlife, 1978:42-51.
Graber, D. 1982. Ecology and management of black bears in Yosemite National Park. Tech. Rep. #5. Univ. of

Calif. Davis.
Hatler, D.F. 1972. Food habits of black bears in interior Alaska. Can. Field Nat., 86(1):17-31.
Herrero, S. 1978. A comparison of some features of the evolution, ecology, and behavior of black and grizzly/

brown bears. Carnivore, 1 (1):7-17.
Jonkel, C). 1962. Black bear population studies. Montana Dept. of Fish. Wildl. Parks. P-R Completion Rep.

Jonkel, C.)., and I. McT. Cowan. 1971. The black bear in the spruce-fir forest. Wildl. Mon., 27, 57p.
Kelleyhouse, D.G. 1975. Habitat utilization and ecology of the black bear in northern California. Thesis. Calif. State

Univ., Areata. 61 p.
Landers, J.L., R.). Hamilton, A.S. Johnson, and R.L. Marchinton. 1979. Foods and habitat of black bears in southeast-
ern North Carolina. J. Wildl. Manage., 43(1 ):143-153.
Mealey, S.P. 1975. The natural food habits of free-ranging grizzly bears in Yellowstone National Park, 1973-74.

Thesis. Montana State Univ., Bozeman. 158p.
Menke, J.W., and ME. Fry. 1979. Trends in oak utilization — fuelwood, mast production, animal use. Pages 297-305,

in J.R. Plumb, ed. Ecology, management and utilization of California oaks. USDA Forest Service. Gen. Tech.

Rep. PSW-44.
Piekieiek, W., and T.S. Burton. 1975. A black bear population study in northern California. Calif. Fish and Came,

Poelker, R.J., and H.D. Hartwell. 1973. Black bear of Washington. Washington State Game Dept., Biol. Bull. No.

14. 180p.
Rogers, L. 1976. Effects of mast and berry crop failures on survival, growth and reproductive success of black bears.

Trans. N. Am. Wildl. and Nat. Res. Conf., 41:431^38.
Tisch, EL. 1961 . Seasonal food habits of the black bear in the Whitefish Range of northwestern Montana. Thesis.

Montana State Univ., Bozeman 108p.
Verner, J., and A.S. Boss. 1980. California wildlife and their habitats: western Sierra Nevada. USDA Forest Service.

Gen. Tech. Rep. PSW-37. 439p.


Calif. Fish and Came 69(3): 151-171 1983




California Department of Fish and Game

P.O. Box 607, 2440 Main Street

Red Bluff, California 96080

This represents a comprehensive check list of the present status of all known
marine, freshwater, and terrestrial amphibians and reptiles that have been reliably
reported as part of the California fauna. Included is a main list of native and estab-
lished exotic species and four supplementary lists: (i) native species extinct in
California, (ii) distributionally or taxonomically invalid species, (iii) established
exotic species, and (iv) exotic species unsuccessfully introduced or of questionable
status. The main list is composed of 129 full species, comprising 124 native freshwater
and terrestrial species, 5 native marine species, and 5 introduced species. The 129
species comprise 29 families and 66 genera.


Previous listings of the herpetofauna of California include: Cooper (1870),
Van Denburgh (1897, 1922), Grinneil and Camp (1917), Storer (1925), Slevin
(1928, 1934), Smith (1946), and Wright and Wright (1952), with regional lists
by numerous authors. Stebbins (1972) provided the most recent check list of
amphibians and reptiles from the State along with their distributions. Since then,
however, there have been changes in the herpetofauna, its nomenclature, and
the status of many species and subspecies. Also, a current comprehensive check
list of all known amphibians and reptiles (to the subspecies level) in California
has been lacking. This paper is an attempt to enumerate in a single document
the present status of all marine, freshwater, and terrestrial species that have been
reliably reported as part of the California fauna.


Like other check lists of the various natural faunas of California, the purpose
of this list is to establish the basis for compilation of a detailed handbook of these
animals, and to promote stability and uniformity in both common and scientific
names. Since Stebbins (1951, 1954, 1966, 1972) has largely achieved these two
goals, this list is mainly an update based on new information. It is hoped that
the list will become the basis for future editions as major revisions become
necessary. This list also complements the comprehensive check lists of other
vertebrate groups found in the State ( most recently: Hubbs, Follett, and Dempst-
er 1979; Shapovalov, Cordone, and Dill 1981).


The main list covers both native and established exotic species. The supple-
mentary lists include: (i) native species believed to be extinct in California, (ii)
distributionally or taxonomically invalid species, (iii) established exotic species,
and (iv) exotic species unsuccessfully introduced or of questionable status.

' Accepted for publication March 1982.

' Present address: School of Renewable Natural Resources, 210 Biological Sciences East Building, The University
of Arizona, Tucson, AZ 85721.


An attempt has been made to include all native forms whose occurrence has
been reported and not disproved in the literature. The existence of most of these
species and subspecies will not be questioned as they are known to have
breeding populations in the State; however, there are a few exceptions. None
of the marine reptiles reproduce in California, but since they occasionally enter
state waters, they must be included in the main list. Such criteria allow the
inclusion of the sea snake, Pelamis platurus, which has been reported several
times in southern California waters (Kropach 1975).

There is the distinct possibility that some native species are no longer part of
the California fauna. One such native form that now appears to be extinct is the
Sonoran mud turtle, Kinosternon sonoriense. Since it is virtually impossible to
prove or disprove its absence, however, this turtle is still included in the main
list. On the other hand, only those exotic species which are known to have
successfully reproducing populations in the State are included.

For the purposes of this paper, the definition of the State includes the entire
Colorado River where it forms the California boundary, and the Pacific Ocean
within 805 km of any point of land in the State between the seaward projections
of its northern and southern boundaries. The westward limit coincides roughly
with the outer edge of the California Current (Hubbs et al. 1979).

The status of endangered, threatened, and rare species, along with species of
special concern, is based on information presented by Stewart (1971), Bury
( 1 972 ) , Ashton ( 1 976) , the California Administrative Code ( 1 980) , Mallette and
Nicola (1980), the Federal Register, and the Department of Fish and Games files.
The status of exotic species is determined by information presented by Stebbins
(1966, 1972), Bury and Luckenbach (1976), Smith and Kohler (1977), personal
communications with various authorities, and other papers cited in the text.

Hybrids have been omitted. Intraspecific, interspecific, and intergeneric hy-
brids of a number of subspecies listed have been recorded from California. The
most striking examples are salamanders of the genus Ensatina (Brown 1974).


Uniformity in the usage of scientific and common names continues to be a
never-to-be attained goal (Shapovalov et al 1981 ). This is due, in part, to our
ever increasing knowledge of the relationships of amphibians and reptiles at the
family, generic, species, and subspecies levels. New techniques and discoveries,
along with painstaking research, have contributed to the problems of nomen-
clatural confusion. This is especially true with such variable groups as Batra-
choseps, Crotaphytus, Gambelia, and Thamnophis, which seem to defy
taxonomic classification at times.

In preparing this list, care has been taken to follow a standard set of rules and
not add to the nomenclatural confusion. Classification schemes follow those
employed by Coin, Coin, and Zug (1978) with few exceptions. Scientific names
used conform to the provisions of the International Code of Zoological Nomen-
clature, 1964, and the amendments adopted by the Monaco (1972) Congress.
Common names, however, have proven to be a never ending source of contro-
versy. The previous authorative list of common names for amphibians and
reptiles (Conant et al. 1956) was an outgrowth of Schmidt's (1953) check list.
Accepted by most herpetologists, this list served as the basis for the common
names utilized by Conant (1958, 1975) and Stebbins (1966, in prep.) in their


field guides. Unfortunately the list remained unrevised over the years except for
a quick update by Dowling (1974) who did not include subspecies. Collins et
al. (1978) was the first group to revise this list in depth, but their disregard of
the criteria they proposed in adopting common and scientific names, along with
many other inconsistencies in taxonomy and spellings, has resulted in this list
being unacceptable as the definitive work for North American amphibian and
reptile nomenclature. Because of these problems, I decided to utilize the criteria
adopted by Collins et al. (1978) and not the list itself. Thus, common names are
based on Conant era/. (1956) and Stebbins (1966), except for those species and
subspecies that were established after these publications were compiled or that
are of uncertain status. In these cases, the original published descriptions (or
revisions) or the accounts contained in the Catalogue of American Amphibians
and Reptiles (American Society of Ichthyologists and Herpetologists 1963-1970;
Society for the Study of Amphibians and Reptiles 1970-present) are utilized as
the final authority. These exceptions are mentioned in the text.

In regard to the spelling of scientific patronyms emended to represent a man's
name (such as Ensatina eschscholtzii, Rana boylii, Copherus agassizii,
Phrynosoma douglassii, Thamnophis couchii, etc.), I chose to follow the lead
of Schmidt (1953), Stebbins (1954, 1966, 1972), Conant et al. (1956), Wright
and Wright (1957), Conant (1958, 1975), Leviton (1972), Cochran and Coin
(1978), Collins et al. (1978), Smith (1978), and Behler and King (1979), and
use single -/ endings for the sake of clarity. This approach is also taken by
ichthyologists (Robins etal. 1980). This nomenclatural enigma and its surround-
ing controversy is amply treated by Jennings (1982).


Native Species and Established Exotic Species

This list consists of 129 full species, which may be subdivided as follows: 124
native species (5 marine) and 5 established introduced species.

Species which have been introduced into California are denoted by an asterisk
(*), marine reptiles by an (O), and extinct species by a dagger {]). The
following symbols are used to denote current status:
SE: State-listed endangered species.
SR: State-listed rare species.
SP: State-listed protected species.
FE: Federally listed endangered species.
FT: Federally listed threatened species.
FP: Species protected by other federal laws (principally those relating to national parks and

C: Species common in the State.

S: Species of special concern in the State. (Those species which may become listed as rare,
threatened, endangered, or protected in the near future due to habitat modification or
destruction, excessive collecting, disease, or impact of exotic species.)
r: Species of very limited distribution only in California. Common in adjoining states or

Native and Established Exotic Species

Order Caudata — Salamanders.

Family AMBYSTOMATIDAE— Mole Salamanders and Relatives'

^ Edwards (1976) places Dicamptodon and Rhyacotriton in a separate family— DICAMPTODONTIDAE.


1. Ambystoma gracile (Baird). northwestern salamander.

la. Ambystoma gracile gracile (Baird). brown salamander C

2. Ambystoma macrodactylum (Baird). long-toed salamander.

2a. Ambystoma macrodactylum croceum Russell and Anderson. Santa Cruz

long-toed salamander SE, FE

2b. Ambystoma macrodactylum sigillatum Ferguson, southern long-toed salamander C

3. Ambystoma tigrinum (Green), tiger salamander^

3a. Ambystoma tigrinum californiense (Gray). California tiger salamander' S

4. Dicamptodon ensatus (Eschscholtz). Pacific giant salamander C

5. Rhyacotriton olympicus (Caige). Olympic salamander.

5a. Rhyacotriton olympicus variegatus Stebbins and Lowe, southern Olympic

salamander C


6. Taricha granulosa (Skilton). rough-skinned newt.

6a. Taricha granulosa granulosa (Skilton). northern rough-skinned newt C

7. Taricha rivularis (Twitty). red-bellied newt C

8. Taricha torosa (Rathke). California newt.

8a. Taricha torosa torosa (Rathke). Coast Range newt C

8b. Taricha torosa sierrae (Twitty). Sierra newt C

Family PLETHODONTIDAE— Lungless Salamanders.

9. Aneides ferreus Cope, clouded salamander C

10. Aneides flavipunctatus (Strauch). black salamander* C

11. Aneides lugubris (hiallowell). arboreal salamander C

12. Batrachoseps aridus Brame. desert slender salamander SE, FE

13. Batrachoseps attenuatus (Eschscholtz). California slender salamander C

14. Batrachoseps camp/ Marlow, Brode, and Wake. Inyo Mountains salamander* S

15. Batrachoseps nigriventris Cope, black-bellied slender salamander C

16. Batrachoseps pacificus (Cope). Pacific slender salamander*.

16a. Batrachoseps pacificus pacificus (Cope). Channel Islands slender salamander S, FP

16b. Batrachoseps pacificus major CAmp. garden slender salamander C

16c. Batrachoseps pacificus relictus Brame and Murray, relictual slender salamander S

17. Batrachoseps simatus Brame and Murray. Kern Canyon slender salamander SR

18. Batrachoseps stebbinsi Brame and Murray. Tehachapi slender salamander SR

19. Ensatina eschscholtzi Gray, ensatina.

19a. Ensatina eschscholtzi eschscholtzi Gray. Monterey salamander C

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