CALIFORNIA
FISH

AND

*CON50W«nON OF WILD LIFE THROUGH EDUCATION

Volume 21

San Francisco, July, 1935

23006

25c per copy

$1.00 per year

STATE OF CALIFORNIA
DEPARTMENT OF NATURAL RESOURCES
DIVISION OF FISH AND GAME
San Francisco, California

FRANK F. MBRRIAM GOVERNOR

GEORGE D. NORDENHOLT DIRECTOR OF NATURAL. RESOURCES

FISH AND GAME COMMISSION

DR. E. C. MOORE, President Los Angeles

ELMER HOUCHINS, Commissioner Bakersfield

I. ZELLERBACH, Commissioner San Francisco

HERBERT C. DAVIS, Executive Officer, Division of Fish and Game San Francisco

450 McAllister Street. Phone Underhill 8700.

BUREAU OF FISH CONSERVATION

J. O. SNYDER, Chief San Francisco

J. H. Vogt, Assistant Chief San Francisco

Alan C. Taft, Assistant Chief San Francisco

A. E. Burghduff, Hatchery Inspector San Francisco

L. Phillips, Hatchery Inspector San Francisco

E. V. Cassell, Superintendent of Mt. Shasta Hatchery Mt. Shasta

Geo. McCloud, Superintendent Mt. Whitney Hatchery Independence

D. A. Clanton, Superintendent Forest Home Hatchery Forest Home

Ed Clessen, Foreman Fort Seward Hatchery Alderpoint

Earl Leitritz, Foreman Fall Creek Hatchery Copco

Archie Thompson, Foreman Tosemite Hatchery Yosemite

Wm. Berrian, Foreman Big Creek Hatchery Davenport

Geo. E. West, Foreman Cold Creek Hatchery Ukiah

J. L. Stinnett, Foreman Feather River Hatchery Clio

R. A. McCloud, Foreman Kaweah Hatchery Three Rivers

J. C. Lewis, Superintendent Tahoe Hatchery Tahoe

Donald Evins, Foreman Lake Almanor Hatchery Westwood

H. E. Cole, Foreman Basin Creek Hatchery Tuolumne

Peter Topp, Foreman Burney Creek Hatchery Burney

C. L. Frame, Foreman Kings River Hatchery : Fresno

Allan Pollitt, Foreman Prairie Creek Hatchery Orick

John Marshall, Foreman Brookdale Hatchery Brookdale

Merrill Brown, Foreman Bass Hatchery and Fish Rescue Sacramento

Preston Bills, Superintendent Distribution Car 01 Mt. Shasta

L. Rider, Superintendent Distribution Car 02 Mt. Shasta

BUREAU OF GAME CONSERVATION

J. S. HUNTER, Chief San Francisco

August Bade, Superintendent of Game Farms Yountville

D. D. McLean, Bird and Animal Economist San Francisco

Gordon H. True, Jr., Economic Biologist San Francisco

A. D. McLellan, Game Refuge Supervisor San Francisco

E. D. Piatt, Assistant Superintendent of Game Farms Chino

BUREAU OF COMMERCIAL FISHERIES

N. B. SCOFIELD, Chief San Francisco

S. H. Dado, Assistant Chief San Francisco

H. B. Nidever, Field Inspector San Francisco

L. G. Van Vorhis Terminal Island

BUREAU OF RESEARCH AND ENGINEERING

HERBERT C. DAVIS, Chief San Francisco

W. L. Scofield, Supervisor, State Fisheries Laboratory Terminal Island

Paul A. Shaw, Toxicologist San Francisco

Clarence Elliger, Assistant Hydraulic Engineer San Francisco

BUREAU OF LICENSES
H. R. DUNBAR, Chief Sacramento

California Fish and Game

"conservation of wild life through education"

Volume 21 SACRAMENTO, JULY, 1935 No. 3

TABLE OF CONTENTS

Page
A LIFE HISTORY STUDY OF THE CALIFORNIA QUAIL, WITH

RECOMMENDATIONS FOR ITS CONSERVATION AND MANAGE-
MENT E. Loicell Sumner, Jr. 167

EDITORIALS 257

Reports-
Violations of Fish and Game Laws 265

Statement of Expenditures 267

Statement of Revenue 270

Fresh Fishery Statistics 271

A LIFE HISTORY STUDY OF THE CALIFORNIA
QUAIL, WITH RECOMMENDATIONS FOR ITS
CONSERVATION AND MANAGEMENT

By E. Lowell Sumner, Jr.

PART I

INTRODUCTION

During the last fifty years the California quail (Lophortyx calif or-
nica and its races) has suffered a marked decline in numbers through-
out most of the state, in spite of considerable effort, chiefly in the

2300G

168

CALIFORNIA FISH AND GAME

form of restrictive legislation, directed toward the protection of this
species.

At Ihe present time the Division of Fish and Game is engaged
in a state-wide program of quail refuge establishment, which results in
an acute need for information concerning the habits and requirements
of the birds, so that an outline of management for California quail
is more to be desired than ever before.

Accordingly, in July, 1931, the writer was assigned by the Divi-
sion to a study of the life history of the California quail, with special
reference to its conservation and management. The investigation
was established upon a fellowship basis and has been carried on as a
cooperative project between the Division of Fish and Game and the
University of California, under the supervision of Dr. Joseph Grinnell,

Fig. 1. Portion of the large open-air inclosure used during the present investi-
gation for observation of certain habits of quail which are practically impos-
sible to study when the birds are unconfined. Shows combination laboratory
: i rid observation blind at upper right.

Director of the Museum of Vertebrate Zoology of the University.
Financial support has been given by the Division of Fish and Game,
and the work has also received the general supervision of the division's
Bureau of Education and Research.

ACKNOWLEDGMENTS

The following persons have aided in identifying certain food
items found in quail stomachs : Mr. W. L. McAtee of the United States
Bureau of Biological Survey; Dr. R. V. Chamberlin of the University
of Utah ; Dr. Charles Piper Smith, Dr. Herbert L. Mason, Dr. William

CALIFORNIA PISH AND GAME

169

B. Herms, and Dr. J. F. Lamiman of the University of California.
Mr. Gordon H. True, Jr., has assisted in securing materials and infor-
mation from quail refuges in southern California. Mr. Jack Okell
and Mr. James Rolph III have generously placed their properties at
the disposal of the writer for the quail investigation. The estate of
Mr. Rolph was used as a permanent held headquarters.

To Dr. Joseph Grinnell, Dr. Jean M. Linsdale and Dr. E. Ray-
mond Hall the writer wishes to express appreciation for the helpful
counsel and many suggestions received. Grateful acknowledgment is
also due to Mrs. E. Lowell Sumner, Jr., whose constant help both in
the field and in the laboratory would be difficult to overestimate.

Fig. 2. Part of the interior of the inclosure shown in Pig. 1

were left undisturbed.

all native plants

MATERIALS AND GENERAL METHODS

Most of the information upon which the present report is based
was obtained by the writer as a result of 30 months of field work in a
selected area 50 miles south of San Francisco, in the Santa Cruz Moun-
tains, San Mateo County, California, while trips were also taken to other
parts of the state to obtain data on local conditions. In addition to
the observation and collection of quail in a free condition, it was neces-
sary to build a large open-air inclosure provided with an observation
blind for the study of certain habits of quail which were practically
impossible to observe in a state of nature on account of the secretive-
ness of the birds. The blind was made large enough to serve as a
small field laboratory, and was provided with living quarters enabling
the observer to remain at his post for a week or more at a time
(Figs. 1-2).

170 CALIFORNIA FISH AND GAME

A total of 170 adult quail was collected during every season in
the year for stomach examination and related purposes, and in addi-
tion, nine small young. To facilitate identification of stomach con-
tents, a collection of local plants, with their seeds and fruits, was made.
The relatively few food items which could not be identified by means
of this collection were submitted to various authorities for deter-
mination.

An intensive trapping and banding program was also carried on
uninterruptedly for more than a year, during which 280 individuals
of all ages were captured 1107 times.

The subspecific race of California quail occurring in the Santa
Cruz Mountains and elsewhere along the humid coast region of Cali-
fornia has been designated as the coastal California quail (Lophortyx
calif omica brimnescens) by Grinnell (1931, p. 37). Although there
is some slight difference between this race and the more widely dis-
tributed Valley quail regarding plumage, together with a difference
between their respective habitats as to humidity, the habits of the two
birds are believed to be essentially the same, as are also the principles
involved in their management. Accordingly, in the following report
the two races will be considered together as one species except where
it is specifically stated otherwise.

The original field notes taken by the writer during this investiga-
tion, and also the collection of quail stomachs which furnished the
material for the section on food habits, have been deposited in the
Museum of Vertebrate Zoology, where they may be consulted by persons
interested.

PLAN OF TREATMENT

Part I of this report presents the original information obtained
by the writer on the natural history of the California quail, and pro-
vides most of the foundation for the recommendations given in Part II.

Part II attempts to summarize all that is now known regarding
the management of the California quail. Although largely dependent
upon the preceding section on life history as a basis for recommenda-
tions, Part II also includes reference to certain important findings
and outlines of policy arising from game studies conducted by investi-
gators in the eastern United States.

CALIFORNIA FISH AND GAME 171

PART I -LIFE HISTORY

FOOD HABITS AND REQUIREMENTS

Stomach Collection Represents Each Season of the Year. The

writer attempted to collect a minimum of five quail stomachs from the
Santa Cruz Mountain area for each month of the year, in order to
determine seasonal changes in the diet of the birds. However, in
spite of the most strenuous and prolonged exertions, the impenetrable
nature of the brush land which occupies most of the quail country in
the region studied, together with the wariness of the birds, made the
strict attainment of this ideal impossible. When the number of birds
fell short of the quota, it was necessary to take a compensating num-
ber of extra specimens during the first few days of the following month.
Because of the resulting unevenness of distribution, certain months
have been combined into bi-monthly periods in Table 1, which sum-
marizes the food habits of the birds examined.

Heavy Grazing Results in Inferior Quail Food. In addition to
a total of 74 quail collected from the Santa Cruz Mountain area, where
grazing was light, an additional 28 birds were taken from a rather
heavily grazed region in the foothills of southern Napa County. In
Table 1 the figures for the two lots of stomachs have been combined and
are indistinguishable ; however it is doubtless significant that the stom-
achs of fourteen quail collected from the lightly grazed area during
November and December contained 45 species of food plants, whereas
28 stomachs taken during the same "months from the heavily grazed
region contained only 19 species of food plants.

Of additional significance may be the fact that in November poison
oak (Rhus diver sildba) constituted .027 per cent of the food of quail in
the lightly grazed area, while in the case of the birds from the heavily
grazed region it amounted to 6.11 per cent of the total food eaten.
Errington (1931, pp. 7, 8) has found that fleshy fruits such as those of
the genus Rhus are markedly inferior as food for bobwhite quail and
are only eaten in quantity after the supply of more nutritious foods
approaches exhaustion.

KINDS AND RELATIVE AMOUNTS OF FOODS

FOOD OF ADULTS

Materials. Table 1 summarizes the character and seasonal varia-
tion of the food of 102 adult California quail examined by the writer.
Since the data from 28 stomachs collected during November and
December on a heavily grazed area in Napa County were included
with those from stomachs taken on the main study area, which was
only lightly grazed, the unusually high percentage of bur clover shown
in the table for these two months strikingly reflects the lack of variety
in the food available to the birds living in the overgrazed region.

Methods. Because of the great amount of labor involved in the
"percentage-by-bulk" method of separating the numerous food items

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CALIFORNIA FISH AND GAME 177

in the heterogeneous food mass of a quail stomach (for example the
separating of gravels and small seeds, imbedded in green leaf masses),
it was found most practical to count the individual items without
separating them in every case.

In order that the results of the investigation might be comparable
with the work of others, however, the figures representing the num-
ber of seeds, insects, etc., were mathematically converted into the
percentage-by bulk figures used by the U. S. Biological Survey (McAtee,
1912), and most other authors. The conversion was accomplished by
first determining for each plant species the number of seeds contained
in a cubic centimeter; then by means of these values the amount of
each food item in a given stomach was expressed in terms of cubic
centimeters ; the total number of cubic centimeters of stomach contents
was next found by simple addition ; and finally the percentage of
each food item in a given stomach was computed by dividing the
number of cubic centimeters of each item by the total number of
cubic centimeters of the stomach contents.

The percentage of gravel particles was first computed and this
figure subtracted from the total stomach contents before the per-
centages of actual food materials were determined. The percentages of
green leaf material could not be obtained by the counting method,
and were roughly estimated as to volume in accordance with the cus-
tomary percentage-by -bulk method.

Under "trace" is included percentages of .004 or less.

In the table the various families of plants are listed in order of
decreasing importance as regards their consumption by quail. Of the
green leaf fragments, only the unidentified pieces are listed separately
from the other food items; when identified, which was in the majority
of cases, such fragments are included under the same heading as the
seeds or fruits of the food item in question. For a graphic record
of the seasonal fluctuation in amount of all green leaf fragment mate-
rial, see Fig. 4; statements of the relative proportions of leaf material
to seed material in the case of the individual food items are given in
the detailed treatment of the major food items below.

Instances where the percentages given in the present report differ
from those given by earlier workers (Judd, 1905; Beal, 1910) are
explainable upon the basis of differing local conditions and perhaps
also because of the smaller number of stomachs involved in the present
case. The present list adds 33 new kinds of vegetable food to the
total of approximately 110 kinds previously recorded for the Cali-
fornia quail.

Leguminosae. The pea family provides a larger and more varied
food supply (32.47 per cent of the total food) for the California quail
than does any other, and includes the most important forage plant
of all, namely bur clover. It is doubtless not an accidental circum-
stance that the plants of this family, which play such an important
part in the life of quail, are as nutritious as they are abundant.

Bur Clover. This introduced forage plant is common throughout
California and is highly prized by stockmen because of its exceptionally
high nutritive value at all seasons (Hart, Guilbert and Goss, 1932,
p. 52). It is undoubtedly the most important single item in the diet of

178

CALIFORNIA FISH AND GAME

quail throughout a large part of California. Table 1 and Fig. 3 show
t \\o seasonal peaks of consumption for bur clover, the first in April-
May and the second in November.

The April-May consumption represents the height of the season
during which the birds are forced to depend largely upon a green
leaf diet because of the exhaustion of the seed supply of the previous
fall. The November peak represents the season of maximum seed
abundance. This species of clover is thus available to the birds
throughout the year, especially during the critical winter period of food
shortage.

In the fall and early winter months the burs are swallowed whole,
but about January they commence to open and liberate their seeds
which are thenceforth picked up individually by the birds. The aver-

Fig. 3. Relative proportions of leaves, seeds, and burs of bur clover (Medicago
hispida) consumed by the California Quail through the year.

age number of seeds of this species per stomach (excluding all stomachs
which contained none at all) was 185; maximum 1224.

Other Peas. Other members of the pea family appear to be eaten
roughly in proportion to their abundance. Cultivated trees of the
genus Acacia furnish an important supply of quail food because, like
bur clover, their seeds are available during the midwinter shortage.
This would doubtless have been more clearly indicated in Table 1 but
for the fact that no collections of quail were made in the vicinity
of acacia trees during January.

Numbers Consumed. To give a better idea than is afforded by
the table of percentages, of the quantities of individual seeds eaten
by quail, the average and maximum numbers of seeds of certain com-
mon species is given: Lupinus formosus, 79, 376; Lotus humistratus,
or subpinnatus, 23, 125 ; Trifolium tridentatum, 10, 63.

Gramineae. Cultivated grains make up 14.4 per cent of the food
of quail in the region covered by the present study.

Barley. This item represented 13.30 per cent of the total food in
the case of the present investigation because in the region under con-
sideration hay is the major agricultural crop, with barley second in
importance only to red oats.

CALIFORNIA FISH AND GAME 179

The writer observed that for about three weeks a particular covey
of quail habitually went to a newly sown barley field for its evening
meal. The crops of birds shot from this covey at roosting time were
enormously distended with barley kernels, the number averaging 162
per crop.

Although the barley taken from August to November was waste
grain, this is only 33.3 per cent of all the barley consumed. The large
size of the individual grains is, of course, partly responsible for the
high percentage recorded for this food item.

Red Oats. Red oats constituted only .81 per cent of the food of
quail in the region studied, in spite of the fact that at least twice as
much oats as barley was sown there. The small amount of oats eaten
doubtless results from the difficulty which the birds appear to have
in swallowing the long pointed grains. As noted on page 190, even
barley is difficult for quail to swallow, on account of its length, while
oat grains are still longer. Only 17.7 per cent of the red oats con-
sumed was waste grain.

Wild Grasses. Of the wild grasses, which form 10.07 per cent of
the food of California quail Lolium multiflorum and Bromus hordeaceus
are the most important in the region studied and, while not as con-
sistently nutritious as bur clover (Hart, Guilbert and Goss, 1932,
p. 52), they are nevertheless of great value as forage, and like the
clover they furnish a food supply during the entire year. Average
and maximum numbers of seeds per stomach (excluding all stomachs
not containing any at all) were: Lolium multiflorum, 65, 592; Bromus
hordeaceus, 32, 206.

Geraneaceae. The geranium family is third in importance,
amounting to 7.55 per cent of all the food of quail.

Filaree. The filarees are valuable not only for their seeds, but also
because in late winter they furnish an important source of green leaf
material. As in the case of bur clover, there are two seasonal peaks
of abundance, one in March representing green leaf materials and one
in October representing the height of the seed crop. From the point
of view of nutritive value the filarees are only a little inferior to bur
clover (Hart, Guilbert and Goss, op. cit., p. 52). Average and maxi-
mum numbers of seeds per stomach are 46 and 240.

Geranium. Geranium dissectum occurred in a large number of
stomachs (average and maximum seed numbers per stomach 17 and
243), but the small size of the seeds results in an insignificant per-
centage by volume. The plant is of some importance as a source of
green food.

Compositae. The sunflower family ranks fourth in importance,
and constitutes 5.84 per cent of the total food of quail..

Centaurea melitensis. The Napa thistle, a notorious and wide-
spread agricultural pest, is the the most important of the group ;
however, like all of the Compositae listed, it appears to be largely
unavailable to quail during the period of winter food scarcity.

Average and maximum numbers of seeds of the more important
Compositae as they occur in quail stomachs are given below; an
asterisk indicates that the seeds are not picked up one by one, but are

180 CALIFORNIA FISH AND GAME

.wallowed in quantity while still attached to the flower head : Centaur ea
melitensis, 55, 437; Madia sativa 125, 632; Hemizonia congesta, 368,
1190; *Microseris douglasii, 63, 63 (one stomach only); *Anthemis
ootvla, 248, 548; *Baccharis pihdaris, 1296, 5106.

Ranunculaceae. Although the seeds of the buttercup are eaten
only sparingly, the plant is important because it furnishes a consider-
able amount of green leaf material during the period of winter scarcity.
The blossoms are also habitually eaten by quail.

Rosaceae. All species of Rosaceae together amounted to only
1.5 per cent of the food during the present investigation. This may
be due in part to the fact that the fruits of these plants are largely
out of reach of such ground-foraging birds as quail. There is also
a strong indication that berries are inferior in nutritive quality to
the seeds of legumes, grass, and other weeds, and are of value chiefly
as "salads" or "tonics."

Miscellaneous Families. The following species (listed with figures
I'm- average and maximum numbers per stomach) are eaten during
the major part of the year, but because of the small size of the seeds
they do not bulk large when expressed as percentages : Anagallis
arvensis, 162, 748; Rnmex acetosella, 257, 1819; Brassica sp., 208, 808.

Turkey mullein (Eremocarpus setigerus), elsewhere a favorite food
of quail, was a rare plant in the region where the present study was
carried on, which explains the small percentage recorded for it.

Although one might suppose that various trees and shrubs, espe-
cially those of chaparral, would provide an excellent food supply for
quail, such is not the case. The following species (listed with figures
for average and maximum numbers per stomach) are common in the
region studied, and produce fruit in considerable quantities; never-
theless, as Table 1 shows, they are eaten by quail only sparingly : Poison
oak, 16, 46 ; Snowberry, 9, 168 ; Elderberry, 40, 154 ; Ceanothus sp.,
14, 38; Madrono, 11, 16; Myrica calif ornica, 3, 3 (one stomach only).

Even when the shrubs included under Rosaceae are added to this
list, the total percentage of tree and shrub fruits amounts to only 4.11
per cent of the total food.

Galls. In addition to the two galls listed in Table 1, a third,
minute form occurred in the fleshy walls of nearly all of the poison
oak seeds eaten by quail. This small gall (average long diameter 1.02
mm.) was thought by the writer to be part of the fruit wall until it
was identified by the Division of Food Habits Research of the Bureau
of Biological Survey ; consequently no estimate of its numbers or aggre-
gate bulk was made. However, its size Avas so small that it could not
have rated as more than "trace."

Green Leaf Fragments. Green leaf fragments and similar suc-
culent vegetation constitute an extremely important food for quail
(35.36 per cent of total stomach contents). Although such materials
are eaten during every month of the year, they are especially promi-
nent in the diet during the winter months after most of the seed supply
has become exhausted.

CALIFORNIA FISH AND GAME

181

Relation of Green Leaf Material to Water Consumption. With
the widespread disappearance of green materials following the advent
of the dry season in June, consumption gradually dwindles, until a
minimum of 3.43 per cent is reached in October. This is the season
during which the birds are forced to depend upon springs or pools
for their water requirments.

With the appearance of new vegetation following the first winter
rains, consumption of green leaf material rises abruptly, which in
turn results in a cessation of visits to the water holes.

Relation of Green Leaf Consumption to Seed Supply. As shown
by the curve (Fig. 4), so long as the seed supply holds out it seems
to be preferred to green leaf materials, and doubtless it is much more
nutritious per unit of volume. The writer has seen penned birds
gradually starve to death on a diet of green feed alone, while the
birds showed an extraordinary eagerness for seeds whenever these

Fig. 4. Relative proportions of seeds, green leaf material and animal food consumed
by the California Quail through the year.

were supplied. However, as the seed supply approaches exhaustion
in January and February, the birds are forced more and more to
depend upon green food, until by April and May it constitutes 85.49
per cent of their entire diet.

Possible Relation of Green Leaf Consumption to Reproduction.
Dry winters such as that of 1933-34 (see page 237) work a hard-
ship upon quail by checking the growth of an adequate supply of
green forage, with the result that the birds are short of food and in
poor condition just before as well as during the breeding season. More-
over, green vegetation is recognized by poultrymen as a source of vita-
mins which favor egg laying, so that it is possible that a vitamin
deficiency may be partly responsible for the observed failure of quail
to breed during exceptionally dry years.

The approximate percentages of the various items constituting
the total of all green leaf materials consumed are as follows :

Bur clover, 19.1; unidentified fragments, 38.3; grasses (chiefly
Lolium and Bromus), 20.4; buttercup, 10.8; filaree, 7.4; Madia sativa,

182

CALIFORNIA FISH AND GAME

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184 CALIFORNIA FISH AND GAME

1.0; Agoseris sp., 1.0; Trifolium. sp., .9; Microseris douglasii, .7; gera-
nium, .2; Stellaria media, .2; Polygonum aviculare, .1; Lotus ameri
canus, .1 ; Lactuca scariola, .1.

Other items of green vegetation not found in the stomachs exam-
ined, hut observed in the field to be eaten by quail were as follows (an
asterisk indicates that the plant is an introduced species) :

SisyrincMum helium (blossoms) ; *Avena fatua (leaves) ; blue
grass (Poa sp.) ; *Lolium perenne (florets) ; Bermuda grass (Cynodon
dactylon) ; *Anagallis arvensis (leaves) ; Sanicula menziesii (blossoms) ;
Jt'umex crispus (leaves) ; *hemp (Cannabis sativa) ; *rape (Brassica
napus; leaves, blossoms) ; *scarlet runner bean (Phaseolus multiflorus;
leaves) ; *sour clover [Melilotus indica; leaves) ; Agoseris plebia
(leaves) ; *Picris echiodes (leaves, blossoms) ; Hemizonia congesta
(blossoms) ; * Anthemis cotula (leaves).

Vegetable Debris. In the case of the California quail the amount
of material which could be classified strictly as debris, exclusive of
feather barbules, was infinitesimally small. Some more or less indi-
gestible vegetable fragments were present, but these were taken in as
integral parts of certain more desirable food substances, and accord-
ingly they were not' segregated as debris but included under the
respective food items. Examples of this class of materials are pod
fragments of bur clover, carpels of filaree, bracts of grasses, pods and
placentae of pimpernel.

Feather Barbules. The barbules of feathers were found in about
one-tenth of all stomachs and were doubtless swallowed accidentally
during preening. They occurred more or less throughout the year
and were not more numerous during the period of molt than at other
seasons.

Animal Food. The animal food eaten by adult California quail
in the area studied by the writer amounted to only .38 per cent of
the total food, but, as Table 2 shows, the inclusion of the stomach con-
tents of small young would materially increase the percentage. Ants
comprised 68.4 per cent of the animal food taken by the adult quail,
with Hemiptera (13 per cent) and lepidopterous larvae (10.5 per
cent) next in amount.

FOOD OF YOUNG

Not only does the food of young quail differ from that of the
adults by reason of its higher animal content, but also in that the
individual items are of smaller size. Large seeds such as those of
Lupinus, Yicia, red oats, and the pods of bur clover (see page 190)
are seldom or never attempted at this stage, while in the case of certain
other foods, for example Lolium multiflorum, only the smaller seeds are
eaten. The average size of the grit particles taken by the young is
also less than in the case of the adults.

Age 2-3 Days. The birds of this age (Table 2) were collected

after they had died from overexposure to the heat of the sun, and

, for this reason their stomachs, although moderately full, may not

CALIFORNIA FISH AND GAME 185

indicate the normal ratio between animal and vegetable content. The
percentage of insect matter is unexpectedly low (6.84 per cent) ; the
stomachs show that even at this age the consumption of seeds and
green vegetble material is large.

Age 6-7 Days. The stomach contents of the birds of this group
were unquestionably normal. In addition to the high percentage of
animal food (34.5 per cent), a considerable number of seeds was
consumed.

GENERAL FEEDING HABITS

Times of Feeding. Examination of the crops and stomachs of
California quail collected at various times of the day, together with
field observation, showed that the birds normally leave their roosts in
the morning with crops, and probably stomachs, empty, and feed fairly
steadily for about an hour, or until their crops are about half full.
After this initial meal feeding gradually decreases, and reaches a
minimum at midday; nevertheless it often proceeds intermittently
throughout the whole day up to about 4 p.m., with crops from one-third
to one-half full. After about 4 p.m. feeding activities noticeably
increase and culminate at dusk in a final period of gorging which lasts
for twenty or thirty minutes and fills the crops of the birds until they
bulge.

As in the case of various passerine birds studied by Stevenson
(1933, p. 155), the stomachs of California quail receive a slow stream
of food from the crop throughout the clay (and night also), and are
rarely empty, no matter what the state of engorgement of the crop.

Stormy Weather. Stormy weather shortens the feeding periods,
for at such times the birds often remain in the shelter of a patch <>i'
thick brush from early morning until only fifteen or twenty minutes
before roosting time, when they furtively emerge from cover in com-
pact little bunches, feeding hurriedly, and with only an undertone
of their usual "conversational" clucking and cheeping notes.

Presence of Enemies. If the birds are badly frightened by a
Cooper hawk or other enemy shortly before roosting time, they some-
times refuse to venture forth from cover for the evening meal, and
are obliged to go to roost with nearly empty stomachs. So great
is the urge to feed at this time, however, that the birds usually over-
come their fright much more quickly than at other times of the day,
and are likely to make a concerted last-minute venture into the open
just at dusk, in spite of the presence of the enemy unless the latter is
directly within their sight. Possibly it is for this very reason that the
Cooper hawk is especially active in the pursuit of quail at dusk.

Hours of Feeding During the Breeding Season. At the time of
courtship, the male birds spend a large amount of time driving other
males away and strutting before the females, especially during the
early morning and late afternoon. This leaves them little time for
feeding, so that their stomachs arc often nearly or quite empty during
the day, and are seldom very full even at dusk. The females, however,
still feed in the normal manner.

186 CALIFORNIA FISH AND GAME

With the commencement of incubation, the males not only con-
tinue to drive away other males but in addition they stand guard,
alert and unfed, when their mates leave the nest to secure food; conse-
quently the food intake of the males continues to be intermittent and
limited.

At the time of incubation the females also change their feeding
habits; they leave the nest for the first time at about 8.30 to 9 in the
morning and feed steadily for about twenty minutes. Often upon
leaving they stand upon their tip toes and whir their wings rapidly,
or stretch out the wing and leg of one side simultaneously, like domestic
poultry, thus giving evidence of their relief at having escaped for a
few moments from the cramped confinement of the nest. The females
indulge in subsequent feedings at irregular intervals of about three
hours until about 4.30 to 5.00 in the afternoon, when they return to
their nests for the night.

After the hatching of the young the same general schedule con-
tinues for several weeks, the female and young venturing out at about
9.30 a.m. after the dews have vanished, and feeding under the watchful
protection of the male bird ; other feedings occur at irregular intervals
until late afternoon. The male bird, on the other hand, is still obliged
to obtain his food at odd moments while his family are resting in
thick cover, and it is not until the female has retired with her brood
for the night that he is able to pick up a somewhat more substantial
meal.

As the young grow larger and more independent, and learn to
roost above the ground, the time spent by the male in rigid watchful-
ness lessens, and the feeding hours of the entire family approach the
schedule followed during the remainder of the year. As the dew
hazard vanishes the first morning feeding occurs earlier, the midday
rest period is observed, and the chief meal of the day takes place once
more just before roosting time.

Feeding Mannerisms. Scratching. Quail scratch the ground
vigorously like domestic fowls when searching for food. This habit
is especially useful during the months of late winter and spring when
the birds are searching for the tiny succulent sprouts of new vegeta-
tion. The writer's field notes afford a description of this reaction,
which was observed many times at close range: "June 2, 1932. The
bird scratched four times on each occasion, then took one step back-
ward and looked for food in the little depression made by the scratch-
ing. The four scratchings were always executed two with one foot and
then two with the other, either foot being likely to start first, thus:
right, right, left, left; or left, left, right, right."

Jumping. Quail secure several kinds of food by jumping, notably
the blossoms of the buttercup, yellow sanicle, tarweed and sow thistle.
The wings are not used in jumping, but are held pressed against the
sides, while the neck is stretched to the utmost. When the buttercups
are in bloom it is not uncommon to see the members of an entire
covey bobbing upward by two's and three's to reach the food above
their heads.

In winter, especially when snow covers the ground and shuts off
more accessible supplies, the birds jump vigorously for the flower heads

CALIFORNIA FISH AND GAME 187

of tarweed and any other species which still contain seeds at that time
of year.

The highest that qnail can reach with their bills by jumping is
about sixteen inches, and even then they often miss several times in
succession the object of their efforts. Since many of the blossoms
are above this height, the more lofty plants are thus spared and their
seeds preserved ; hence it is possible that terrestrial feeders like quail
may have contributed to the evolution of the long seed stalks of these
and various other native plants.

Distances Traveled for Food. Under normal conditions the dis-
tance traveled during the day by quail in search of food is slight. In
the early fall, when the annual crop of weed seed is at the height of
its abundance, a covey will often spend one or two whole days at the
edge of a given patch of brush or woodland. Under such conditions
the duration of the birds' stay would be theoretically much longer than
this if adequacy of food supply were the only factor determining their
whereabouts and if the birds hunted for it systematically.

However, a covey gradually moves about over its territory, partly
as a result of harassment by enemies but more because of the innate
tendency of the birds to feed a little here and there and continually
moATe on. regardless of the fact that by so doing they may move from
a plot of ground where food is abundant into one where it is less so.
This instinctive (?) propensity is strikingly illustrated in the case of
quail which have discovered a patch of ground covered with bait. In
spite of the fact that the birds have only to stand still and eat, they
pick up a little here and there, then run a few steps forward and pick
up a dozen more seeds, often stopping to scratch the ground vigorously,
although such scratching is not only unnecessary under these condi-
tions, but actually scatters the seeds far and wide so that many are lost.
Presently, as a result of this aimless feeding, the birds have wandered
entirely away from the site of food abundance, in which case they
usually do not return, at least not for several hours and often for days.

Although such aimless wandering seldom results in a covey travel-
ing more than a few hundred yards in a day, several days carry it
gradually, and erratically, over the whole extent of its territory — which
is largely coincidental with its food supply. The distance covered per
day is not constant, but is governed by chance and by food conditions.
For example, regions with a sparse food supply, such as the semi-
desert portions of southern California, require more daily searching
than do more favorable regions such as the Santa Cruz Mountains.

When the food supply becomes less abundant during midwinter,
the distance covered daily by a covey somewhat increases. However,
except in a few places at high altitudes, it is rare for a covey to be
forced entirely out of its range because the loss by predators auto-
matically increases before this can happen, so that only as many birds
survive as can succeed in maintaining themselves in the territory.

Amount of Food Eaten Daily and Yearly. The weight of food
eaten per day was determined by the writer according to the method
described by Taber (1928, p. 339). This method involves trapping and
weighing the birds just at roosting time, keeping them overnight,
and then weighing them again the first thing in the morning. From

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the loss of weight occurring during the night, the daily (24-hour) loss
may be calculated, and this loss must of course be balanced by the
daily intake. At the time the present experiment was carried on the
average period spent on the roost was eleven hours. Many birds
exhibited an abnormal retention of food in the crop, no doubt as a
result of nervousness induced by confinement; consequently only
the records of birds which had empty or practically empty crops on
the following morning were used. These records comprise night and
morning weighings of 108 birds, as shown in Table 3.

The total weight of food consumed on. the average by an adult
California quail in one year can be estimated, from the above data,
at 8760 grams (19.3 lbs.), or (if calculated upon the basis of the known
value of 60 lbs. of alfalfa seed to the bushel) 11310.7 cc. (.321 bushel;
10.2 qts.).

That this method of estimating the daily food consumption of Cali-
fornia quail is fairly accurate is indicated by the fact that the figures
are close to those obtained by Nice (1910, p. 302), by a different method,
for captive bobwhites, considering the relative sizes of the two species,
which weigh, on the average. 189.5 and 170 grams respectively.

Food Preferences. Quail studied by the writer evidently exer-
cised a certain degree of preference as to what they ate and were by
no means swayed by mere availability. The chosen foods represented
plants of various unrelated groups and seemed to have as their chief
common characteristic a high energy-producing quality.

Although legumes, grasses and composites were abundant and
together made up 62.7 per cent of the food of the birds, such plants
were by no means the only potential source of food. On the other hand,
the writer was surprised to note that certain very common and appar-
ently eminently suitable species of plants were either absent in the
diet or represented by insignificant numbers.

That the scarcity was not due solely to chance, or to the relatively
small number of stomachs examined, is indicated by the fact that no
record of these plants occurs in the reports of Judd (1905) or McAfee
and Bead (1912). Seeds of native plants which were common in the
Santa Cruz Mountain area, but which were not present in quail
stomachs in appreciable quantity, belonged to the following species :

Chlorogalium,. Brodiaea, Calochortus (Liliaceae) ; Collinsia,
Scrophitlaria, MimuUis, Castilleia, Orthocarpus, Cordylanthus
(Scrophulariaceae) ; Trichostema, Micromeria (both very strong
scented), SphaceJe, Stachys, MonardeUa (Labiatae) ; Artemisia, Achil-
lea (Compositae) ; Dodecatheon (Primulaceae) ; Galium (Rubiaceae) ;
Godefia, Baisduvalia, Epilobium (Onagraceae).

Not only are the seeds of these plants abundant,but they are also
within the size limits to which quail are adapted. It will be noted that
the species listed are grouped within a certain few families, and it may
be significant that, with the exception of the Compositae, none of these
families is represented in the list of foods (Table 1) which are regu-
larly eaten by quail. Furthermore, the two members of the Com-
positae which are not eaten are notably strong scented.

The characteristic strong scent of the Labiatae (mints) may
account for their rejection, but why the Liliaceae, Scrophulariaceae,

190 CALIFORNIA FISH AND GAME

and certain others should be refused as food is not clear. Possibly some
of them produce a toxic effect when eaten.

Rejection by Quail of Suitable Foods to Which They Are Unac-
customed. A factor which operates in the case of certain other plant
foods which are rejected by quail is that of novelty, especially if com-
bined with rather large size. The following- excerpts from the field
notes of the writer illustrate this :

"May 15, 1932. The quail inside the large inclosure have difficulty
in eating barley, with which they are unfamiliar. Only about once
out of every six tries, are they successful in swallowing it. It is as
if they couldn't get it down their throats, for the kernels actually
disappear for a moment at the back of the mouth only to fly out again,
as though by the action of the tongue. The birds pick up the barley
with evident eagerness but the swallowing is not so easy."

Three days later they had begun to learn to eat barley, but now they
refused hemp, which was another food strange to them :

"The birds ate a little hemp, but not much. Although hungry,
they would continually walk right over ground where it lay thickly,
ignoring it entirely and searching for other more familiar tidbits, and
for scraps of green vegetation. I remarked, however, that several of
the birds took quite a lot of the remaining barley, picking up six or
seven kernels consecutively without dropping any. Evidently they are
learning to accept this new food."

It is significant that a certain covey of quail three miles away was
accustomed to feed every evening upon barley from a newly sown
field in their territory, the crops of these birds becoming enormously
distended with the kernels.

Sunflower seeds, red oats and large kernels of corn were con-
sistently rejected by the birds within the inclosure, even though they
were brought to the verge of starvation in an attempt to make them
eat these foods. On the other hand, small pieces of cracked corn
were eaten fairly readily, and certain wild individuals were known to
have eaten red oats (see Table 1) ; however, no sunflower seeds were
ever found in the digestive tracts of quail.

Certain native fruits which are larger than sunflower seeds or
corn kernels, but which are nevertheless eaten by quail to an appreci-
able extent, are the hard fleshy fruits of the poison oak and the large
dry berries of the madrono.

That large size and not novelty alone may sometimes be the factor
responsible for the rejection of unaccustomed foods is indicated by
the fact that certain moderate sized or small seeds such as kafir corn
(Andropogon sorghum) and millet {Panicum sp.) were accepted at
once by both penned and wild quail, in spite of being entirely new to
their experience.

Other new foods, however, were refused, in spite of their being
of suitable size. The seeds of Spanish and Scotch brooms, and those of
Fuller's teasel were consistently rejected by the quail hatched inside
the large pen, even when these birds were in a starving condition.
Nevertheless, the quail that lived just outside the pen, where these
plants -were common, ate the seeds in considerable numbers. Stoddard
(1932, p. 125) found that as much as three years might elapse before

CALIFORNIA FISH AND GAME

191

the bobwhite learned to accept ;i new food. That California quail
finally learn to eat the useful new species, however, is shown by the
important place now held in the diet by numerous introduced weeds
and forage plants, notably bur clover.

Upper Limit of Size of Food. In the region where the present
study was conducted, the largest food items found in quail stomachs
were as follows:

Species

Average volume of
seed or insect (cc.)

Percent of
yearly diet

Oak gall

Toyon berry

Pentatomid (bug)

Convolvulus sp. seed (morning glory)

Barley

Locustidae (one-third grown grasshopper)

Red oat

Lupinus variicolor

Poison oak

1/8
1/8
1/10
1/14
1/15
1/20
1/26
1/27
1/31

.01

.64

.11

.04

13.43

.03

.81

2.56

1.32

Although Stoddard (op. cit., p. 147) reports that acorns consti-
tuted more than 6 per cent of the yearly food of the bobwhite in the
southeastern United States, and that one bird had eaten fourteen
whole acorns together with fragments of several others, the usual
food of the California quail does not include such large items, and the
results of the present study support the statement of McAfee and
Beal (1912, p. 11) that whereas in some cases "a few bits of acorn and
perhaps nuts were eaten. . . . the quantity is insignificant."

Lower Limit of Size of Food. In general, seeds less than 1.2
mm. in diameter, such as Anagallis arvensis, were not picked up singly,
and probably are either too small to attract the attention of the birds
readily, or represent too small a reward for the amount of energy
involved in picking them up. Many of the seeds which commonly
occur in the stomachs of quail are much smaller than this, but seeds of
this sort are eaten while still enclosed in their relatively large pods,
capsules, or flower heads, and consequently are secured in large num-
bers with each mouthful.

GRIT REQUIREMENTS

Daily Intake and Loss. The number of gravel particles in 102
stomachs varied from 2296 to 10, the average being 508 (14.26 per
cent of total stomach contents). The daily loss of grits through the
intestine can be gauged to some extent by the daily intake of fresh
particles which are found in the crop. Out of a total of 93 crops,
mostly collected in the late afternoon or evening, 47 contained no grits,
while the average for all crops was 5.19 ; the highest number for a
single crop was 163. Since the food content of a quail crop collected
at the end of the day represents roughly one-half of the total food
intake for the day, multiplying 5.19, the average number of gravels per
crop, by 2 gives 10.38, which is probably a fair approximation of the
daily loss of grits. i

192 CALIFORNIA FISH AND GAME

The California quail consumes a much smaller amount of gril
when hard seeds, such as those of poison oak, form a large proportion
of the food. On the other hand, as Table 1 shows, the maximum con-
sumption of grit occurred in late winter and in spring, when seeds of
all kinds were scarce and the principal food Avas green leaf material.

Kinds and Sizes of Grits Eaten. The grit content of stomachs
taken from both the Santa Cruz Mountain area and the foothills of
southern Napa County was similar in being made up of small translu-
cent quartz particles. The particles were irregular in shape, and varied
in adults from about .15 mm. to ,5.7 mm. in diameter, the average
being 1.2 mm. Particles swallowed by two- or three-day old chicks
averaged only about 91.6 per cent as large.

The very minute grit particles which occur in considerable num-
bers are probably not picked up individually by the quail, nor are they
chipped from larger fragments during digestion, since they are also to
be found in the crops. Undoubtedly such particles are swallowed
accidentally as they adhere to seeds, vegetation, or to other larger
pieces of grit. The propensity of quail to pick at nearby objects
while wallowing in their dust baths, and the activities of the
birds in preening their dust-laden feathers, explain the presence of
these microscopic fragments.

WATER REQUIREMENTS

Water Requirements During the Rainy Season. Field records,
supplemented by intensive daily observation of 6 adult quail and their
36 offspring reared within the large inclosure during two successive
years, have proven that after the appearance of the first autumn rains
the birds no longer visit the water holes. The moisture which they
require is then derived from the succulent green vegetation which
carpets the entire country during the rainy season.

Water Requirements During the Dry Season, Including the
Breeding Season. Toward the latter part of April, as the days
become warmer, the birds occasionally take a few sips of water in the
late afternoon, but drinking does not become habitual until the
approach of drought conditions in late June. With the appearance
of the young in July, however, a marked increase in the use of water
is apparent. Often the adults lead their charges to the water holes
three or four times a day, usually taking from three to twenty sips
themselves on these occasions.

With the advent of cooler weather in September, the consumption
of water lessens, and if the sky remains overcast during a large part
of the day, or if there has been a heavy dew the night before, no visits
are made to the water holes. At this time of the year two or three
days may go by without the birds drinkine any fvpp water, and with
the coming of the rains once more total abstinence follows.

'-

Previous Opinions as to Water Requirements. There has been
some discussion among authorities as to whether the quail of the arid
southwestern states must have water to drink. Grinnell (1927)
observed that nests of the California quail were rarely or never located
more than 400 yards from water. He assumed from this circumstance

CALIFORNIA FISH AND GAME 193

that the newly hatched young required water within a few hours after
hatching, and that 400 yards was the maximum that such little quail
could travel for it. From this he concluded that the water factor was a
critical one during this season in the southwestern states for game
birds in general.

Vorhies (1928) brought forward evidence indicating that in
Arizona at least the Gambel quail (Lophortyx gambelii) was not
dependent upon water during the breeding season, often nesting several
miles from water without ever visiting it, and receiving a sufficient
supply from succulent vegetation to satisfy its requirements. Gorsuch
(1931, p. 257) records finding the newly hatched young of the Gambel
quail as far as ten miles from water.

Experiment on Water Requirements. In order to settle the
question as regards the California quail, the writer shut off the water
supply in the large inclosure on February 1, 1933, and left the two
pairs of resident birds without any source of free water until Febru-
ary 1, 1934. As was expected, the birds gave not the slightest evidence
of thirst during the winter and spring months.

With the advent of the summer heat and its accompanying aridity,
the birds began to feed every day upon the succulent leaves of a
small experimental patch of alfalfa within the inclosure. Even when
the maximum daily temperature reached 102° F. during the first two
weeks in June, the quail gave no sign of thirst; however, the alfalfa
patch grew steadily thinner as a result of the attacks of the birds.
That they remained healthy during this period is indicated by the fact
that both pairs produced young during the course of the experiment.

After the hatching of the first brood, the weather remained stead-
ily hot with no dewfall, and the alfalfa patch was entirely eaten up.
However by July 20, when it was gone, the young of the first brood
were able to fly and they, as well as the adults, now commenced to eat
the leaves and floral bracts of a few sunflower plants which had sprung
up inside the inclosure. During a large part of each day the birds
would perch by two's and three's upon the swaying tops of the sun-
flowers, flapping vigorously to maintain their balance and often push-
ing one another off in their eagerness to obtain the green material.

Due to the accidental destruction of a previous nest, the second
pair of adults did not bring off their brood of 12 young until July 21.
By this date all succulent vegetation near the ground, and even within
jumping distance above it, was gone. The newly hatched young of
the second brood were of course unable to climb the sunflowers to
where the larger quail were foraging at a height of three feet above '
the ground, and consequently they perished from thirst during the
ensuing 10 days, although there was an abundance of dry food (seeds)
all about them.

All of the young of the first brood, together with the two pairs of
adults, lived through the remainder of the summer and subsequent
autumn without a drop of free water; however, before the coming of
the rains the birds had obviously become less vigorous. Practically
every shred of green vegetation within the inclosure had been eaten.
The sunflowers were reduced to bare, headless stalks ; every prickly leaf
of the numerous milk thistles present was eaten down to the tough

194 CALIFORNIA FISH AND GAME

midrib; even the rank Spiny Clotburs (Xantium spinoswm), whose
slightest touch is painful to human skin, were reduced to twisted
skeletons.

From this experiment it may be concluded that the California
quail, like various other species, requires no drinking water provided
that there is present either a heavy dewfall, a supply of berries, or
succulent vegetation. However, since most of the native range of
the California quail is subjected to a protracted dry season during
the breeding period (June to August), the amount of succulent vege-
tation approaches zero at that time. The last traces of edible green
vegetation are found at the water holes, and for this reason the birds
are largely restricted to the neighborhoods of water holes and streams
during the nesting season, as pointed out by Grinnell (op. cit.).

The Gambel Quail Nests Under Different Moisture Conditions.

The independence of water which Vorhies has demonstrated for the
Gambel quail is explainable upon the basis of an earlier breeding
season, together with a climate which is different from that of the
region inhabited by the California quail. The Gambel quail nests in
April, at a time when there is an abundance of succulent vegetation
(Vorhies, op cit., p. 450). In addition, the birds are able to use
cactus, which furnish "a very important food during the hottest
summer season," while their diet also includes a considerable amount
of insect material (Gorsuch, 1931. p. 255). Moreover, according: to
Vorhies and Taylor (1933, p. 494), there are two rainy periods in
that part of Arizona under consideration by them, one (December to
March) immediately preceding the breeding season, and the other
(July to September) shortly after it.

GENERAL ECOLOGICAL REQUIREMENTS

The ability of the California quail to exist in a given locality is
largely dependent upon the following three major requirements:

1. Food. Quail food is chiefly supplied by herbaceous weeds and
grasses; not only must it be present in adequate quantity throughout
the year, but it must be close enough to cover that the birds are
not afraid to venture forth to secure it. A food supply which is
more than about fifty feet from cover is largely unavailable for the
birds.

2. Cover. In the ecology of quail the function of chaparral land
and other cover is to supply protection against enemies and weather,
rather than to supply food. Just as food is unavailable to the birds
without cover, so is cover of no use without an adjacent food supply.
Consequently the most satisfactory combination of these two essentials
is represented by an interspersion such as one sees under natural con-
ditions along the edge of woods, at the foot of a chaparral-coverecl
slope, at the margins of grassy valleys, and, under man-made condi-
tions, along weedy fence rows and the sides of roads. The relation
between quail abundance and the extent of "edge" environment will be
more fully discussed in Part II.

In addition to ground cover for the protection of the birds during
the daytime, some sort of roosting cover is also necessary, such as the

CALIFORNIA FISH AND GAME 195

dense foliage of the live oak (Quercus agrifolia) and California laurel
(Umbellidaria californica), or the thick tangles of poison oak (Pig.
11) and sumac {Rhus laurina).

In certain arid portions of the state, notably in western Riverside
County, there are very few dense shrubs; there the quail are forced
to roost within a few feet of the ground in thin stands of brush. The
inadequate protection offered by such scant roosting cover is reflected
in an unusually high mortality due to horned owls, and probably
ground prowlers as well.

3. Water. In addition to the two prime requirements mentioned
above, water in some form is essential.

Topography. The California quail ranges commonly from sea
level to 4000 feet, and occasionally up to 8500 feet or even higher
(Grinnell, Bryant and Storer, 1918, p. 517). Whether the inability of
this species to maintain itself at the higher altitudes, where it is
replaced by the Mountain quail (Oreortyx picta), is due to a lack of
food to which it is adapted (the Mountain quail makes use of mast
and berries), or to the rigors of the winter season, is not indicated
by any of the present findings.

East-West Distribution; Toleration of Humidity and Aridity.

So long as suitable food, cover and water are present the California
quail is able to live with equal facility in all regions from the humid
coastal belt to the margins of the deserts in eastern California.
Although the range of the California quail slightly overlaps that
of the Gambel quail along the desert fringes, the California quail is
unable to compete with the former species throughout the waterless
country lying to the east.

The California quail of the coastal fog belt (Lophortyx californica
brunnescens) is somewhat darker in tone of color than the "Valley
quail" (Lophortyx californica californica) which inhabits the more
arid remainder of the range of the species, and it is possible that the
former is more fog-resistant, although this has not yet been proven.

North-South Distribution; Tolerance of Heat and Cold. The

fact that the natural range of the California quail does not extend
northward beyond the vicinity of Klamath Lake in southern Oregon
is probably due to the inability of the birds to find food when deep
snow covers the ground for long periods. Even in parts of California
the advent of unusually deep snows is accompanied by a heavy quail
mortality. In this respect the species is evidently inferior to the bob-
white of the northern United States, although even this latter species
is by no means immune to the effects of deep snows.

California quail become noticeably sluggish in cold weather, par-
ticularly if there is snow on the ground. The writer's field notes
afford an illustration :

"January 13, 1932. There is about three inches of snow on the
ground and it is cloudy and cold. I saw several quail as they walked
out of the brush onto the road. They moved more slowly than usual,
with an appearance of sluggishness which was enhanced by their
puffing out their feathers and drawing in their heads to an unusual

196 CALIFORNIA FISH AND GAME

degree. Three or four of them stood still on the road in a listless
fashion, only moving away when I had approached to within 50 feet."

Even in rainy, cold weather, with no snow, this same sluggishness
is apparent. For example:

"January 25, 1933. A cold, sleety rain falling. This group of
quail was doing what I have several times seen quail do at roosting
time, namely traversing open, unprotected places on foot on their way
to some distant feeding spot. These birds were about 50 feet from
cover; some were standing still, with feathers fluffed out and heads
hunched in. A Cooper hawk ought to have been able to pick one up
quite easily. I have seen these hawks hunting quail at just this time of
day (early dusk), too."

Southward the range of the California quail extends 1530-odd
miles without a break, from Klamath Lake through a wide gamut of
climatic conditions to Cape San Lucas at the southernmost tip of Lower
California.

Wide Range Results from Great Adaptability as to Food and
Cover Requirements. From an ecological point of view the California
quail may be regarded as a highly successful species because of its
ability to maintain itself in regions which differ widely with respect
to (1) plant species available as food and cover, (2) seasonal rainfall,
and (3) temperature range. It is evident that minor details are
unimportant so long as the environment conforms to a general type,
the specifications of which are :

1. Good interspersion of cover plants (usually of chaparral type,

plus trees for roosting) with food-bearing plants (herbaceous
weeds and grasses) ; and

2. Some available form of water.

EFFECT OF HUMAN ACTIVITIES UPON QUAIL

HUNTING

The Problem. The prominent part which hunting has played in
the decrease of quail is unquestioned. The history of the restrictive
legislation of the last fifty years amounts to a recognition of this fact,
and the early records (Grinnell, Bryant and Storer, op. tit., p. 534;
Welch, 1928, p. 123) of enormous annual kills made by market hunters
furnish additional evidence. In spite of the increasing number of
restrictions on hunting, including the now common practice of posting
private lands, quail populations have been kept at a low point through-
out most of the state by the increase in the number of hunters, the
steady improvement in shooting equipment, the nearly universal use of
the automobile, and the constant extension of roads into regions which
were formerly inaccessible. How to maintain a constant supply of
quail under such adverse conditions constitutes the problem.

The Solution. The only solution for the problem of over-hunting,
aside from eventual complete cessation of the sport, is a system of
game management directed toward the conservation and utilization
of the maximum number of birds which the land is able to produce,
together with efforts to increase the productivity of the land itself.
How this can be done economically is outlined in Part II.

CALIFORNIA FISH AND GAME 197

It would be wrong to imply that hunting is the sole factor respon-
sible for the present relative scarcity of quail. Actually, it is only one
of five major causes of the decrease of the birds, and in many localities
it is over-shadowed in importance by one or more of these other factors,
as indicated below.

AGRICULTURAL PRACTICES

Replacement of Forest, Chaparral and Grass Lands by Cultivated
Crops. Whether a given region has been affected favorably or unfav-
orably, from the standpoint of quail, by the substitution of cultivated
crops for the native vegetation depends upon the original character
of the vegetation. If the region in question was originally composed
of unbroken forest, brush or grass land, with very little "edge" land,
the breaking up of such dominant vegetational types into scattered
farm units will increase the total extent of "edge" environment,
and benefit the quail accordingly. This happened during the early
days of the settlement of forest land in Humboldt County.

In the case of chaparral lands, which occupy a large part of
central and southern California, it is by no means certain that quail
have been benefited by the change to agricultural crops, even where
primitive farming methods prevail. Under prehistoric conditions the
original chaparral was apparently rather open in nature due to fre-
quent local fires (Jepson, 1925, p. 6; Storer, 1932, p. 323), some being
caused by lightning and others being purposely started by the Indians.
This open chaparral permitted a plentiful interspersion of grasses
and herbaceous weeds, thus resulting in extensive tracts of the "edge"
type of environment required by quail. The fact that quail were
"excessively abundant" in the early days in regions quite unchanged
by man (Fisher, 1893, p. 28) supports this idea. On the other hand,
the substitution, under the modern "one crop" system, of unbroken
fields of grain and coverless pasture, orchard and vineyard lands for
the native chaparral has been markedly unfavorable to quail.

Some orange groves may constitute a partial exception to this
rule because they furnish excellent roosting cover; however, because
of the cultivation methods employed, quail food is usually absent
within them; moreover, they can be considered truly beneficial only
where they have not supplanted native trees or chaparral. Eucalyptus
trees sometimes furnish suitable roosting cover, as do certain other
non-native species.

A certain benefit has also come to quail as a result of the introduc-
tion by man of various weeds and forage plants of high food value (see
Table 1 and page 177). Plowing, harrowing and related activities are
also beneficial, if not carried on so as to destroy necessary cover,
because they break up the hardened soil, thereby greatly stimulating
the growth of weeds furnishing quail food.

Primitive Farming versus Clean Farming. Although the Cali-
fornia quail was formerly plentiful in regions where the early "primi-
tive" farming methods favored brushy fence rows, weedy corners
in fields, and similar "edge" conditions, the species has decreased
rapidly in recent times with the advent of the modern practice of
"clean" farming because the latter practice has rendered large areas

198 CALIFORNIA FISH AND GAME

coverless, foodless and utterly unfit for bird life. The adverse effects
of clean farming upon wild life, and other serious disadvantages of
this practice are more fully discussed in Part II.

Exclusive Use by Man of Water Sources. The fatal effect upon
quail in arid regions, of the exclusive use by man of springs and other
water sources is fully discussed in Part II.

Irrigation. Irrigation may benefit quail in some localities not
only by supplying drinking water directly, but also by maintaining
various succulent forage crops, such as alfalfa, which can be used by
the birds in place of free water. However, the total harm clone to
the birds throughout the state by irrigation methods probably exceeds
any local benefits, because the water used for this purpose is usually
obtained at the expense of streams, springs, or accumulated subsurface
water supplies, with a consequent decrease or complete cessation of
lowland stream flow and lowering of the water table — all of which
results in curtailment of the water available to quail over vast areas,
and sometimes even in a marked change in the type of vegetation.

Grazing. As shown in Part II, overgrazing is one of the most
important causes of quail decrease on many range lands, not only
because of the resulting direct damage to nesting cover, but also, even
more significantly, through the wholesale destruction of quail food by
the competing live stock.

Another unfavorable result of grazing, which if not already acting
upon quail and other wild life at least represents a threat, is the
depletion of the mineral supply of upland regions. Thousands of
tons of mineral substances are removed from the soil by forage plants
each year and are then appropriated by live stock for bone formation
and other physiological processes. Instead of being returned to the soil
by the death and decay of these grazing animals, however, such mate-
rials are permanently removed from the regions from which they were
obtained when the live stock is shipped to the markets.

A similar process of mineral depletion also takes place through the
removal of cereal and field crops.

This removal of mineral substances has been going on for about
sixty years, and evidence is already at hand that cattle on certain
California ranges may be suffering from a mineral deficiency during
parts of the year, with a consequent lowering of the reproduction rate
(Hart and Guilbert, 1928, pp. 25, 29, 31). Storer {op cit., p. 324)
thinks that in parts of California the reproduction rate of deer may
also be suffering from a low mineral intake, and, if this is true, it seems
probable that quail may eventually be affected as well.

Predator Control. In most cases predator control as practiced by
private individuals has resulted in far more harm to quail than good
because of the popular failure to distinguish between harmful and
beneficial predators. For example, such "control" has brought about
little or no reduction in the numbers of Cooper hawks, which are
important enemies of quail, while it has caused the decrease of red-tails
and similar slow-flying hawks, which really are allies of the quail
because such hawks feed upon the rodent competitors of the quail.

CALIFORNIA FISH AND GAME 199

Predator control as carried on by government agencies has been
directed chiefly against the coyote, mountain lion and bobcat, none
of which is an important enemy of quail. On the other hand, coyotes
and bobcats aid in checking the rodent competitors of the birds and to
this extent should be regarded as beneficial.

Rodent Control. Rodent control, particularly as regards ground
squirrels, has been carried on in California by government agencies
upon an enormous scale for many years (Garlough, 1918, p. 786;
Jacobsen, 1918, p. 756; 1931, p. 243 ; Linsdale, 1931, p. 95). Prior to
1928 it may have benefited quail by releasing them from competition
with the squirrels for food. However, with the adoption and wide-
spread use of thallium sulphate in 1928 as a squirrel poison, any benefit
resulting to the birds by release from competition has been far out-
weighed by the destructive effects of this type of poison upon the
birds themselves (Linsdale, op cit., p. 103; Shaw, 1931, p. 33).

LUMBERING

As previously mentioned, where the clearing of forests has led to
the substitution of "edge" types such as slash, chaparral, or primi-
tive farm land, quail have increased in numbers.

ROADWAYS

Linsdale (1929, p. 143) has shown that roadways exert a beneficial
influence upon most bird life far in excess, he believes, of the toll
exacted by speeding automobiles, because the weed patches and tangles
of shrubbery which accompany such roadways afford an abundance
of food and shelter, often in regions otherwise unsuitable for birds.

Although the California quail undoubtedly benefits from the
presence of roadways in this respect, it is the opinion of the writer
that it does so to a lesser extent than do most other roadside inhabit-
ing species. As explained on page 243, this is partly because of the
more than ordinary tendency of quail to collide, while in full flight,
with the fences which so often border roadsides, and partly because of
the reluctance shown by mated pairs during the breeding season to
take wing at the approach of an automobile until too late.

Moreover, if we consider the fact that our present highway system
permits hunters to reach even the most remote strongholds of the
birds, the net effect of roadways upon quail must be regarded as more
harmful than beneficial.

FIRE CONTROL

Relation to Dense Cover. As pointed out in Part II, it is prob-
able that a policy of discouraging the burning of brush areas has had
an unfavorable influence upon quail in some foothill regions by pro-
moting the growth of dense stands of tall brush to such an extent
that food-bearing weeds and grasses have been crowded out.

"protection"

Laws. Efforts to protect quail by regulatory laws have probably
reached their highest development short of a permanent closed season.

3—23006

200 CALIFORNIA FISH AND GAME

Predator Control and Refuge Establishment. Other attempts to
encourage the stock of wild quail have been limited largely to the
killing of predators and the establishment of refuges. How far the
killing of predators has been successful in bringing about an increase
of quail has been indicatel previously.

The establishment of refuges is the first step in the right direction;
however, in the past practically no effort has been made to improve
such refuges from the standpoint of quail aside from the prohibition
of hunting, while such fundamental problems as clean farming, over-
grazing, usurpation of water, and the relations between food supply
and cover, have usually been overlooked. It is in the choice, improve-
ment and maintenance of such refuges that a knowledge of game
management is of vital importance.

CALLS OF QUAIL

Although the calls of the California quail are decidedly varied, the
significance of a given call is rather constant. The difficulty of repro-
ducing in print even a single common note is shown by the wide
diversity of versions listed by Grinnell, Bryant and Storer (1918, p.
518) and by McLean (1930, p. 16) for the wTell-known assembly call.
An additional complication is introduced by the fact that many calls
sound different according to the distance of the birds from the listener.
Due to the large natural inclosure used during the present investi-
gation, the writer was able to make daily observations during two
years upon quail which were calling from a distance of from ten to
thirty feet away. Even in the present instance, however, where the
conditions for the observation and recording of calls were unusually
favorable, emphasis is placed more upon the significance of the calls
than upon an attempt to make accurate typographical representations.
Moreover, there is a decided individual variation among quail as to the
tone, emphasis, and manner of utterance of some of the calls.

The Assembly Call. This is the most familiar and most commonly
uttered call of the California quail, and may be rendered cu-cd-cow
(u as in "luck"; a as in "far"). Although the main accent is usually
on the middle syllable, it is occasionally placed on the last syllable.
Sometimes when the main accent is placed on the last syllable a fairly
strong accent is also given to the first syllable, so that the middle syl-
lable receives the least accent of all, which is the reverse of the usual
condition. Some male birds, especially during the breeding season
when they are more than usually excited, utter several preliminary cu's
in rapid succession before giving the full call, thus : cu, cu, cu, cu-cd-
coiv. The writer once heard a male bird, which was greatly excited
over the presence of a rival male, give voice to a curious combination
of the. assembly call and the alarm note described below, thus: cu-cd-
pwit, cu-cd-pivit, cu-cd-pwit.

The assembly call is usually uttered from one to four times in
succession, with a pause of at least ten seconds before the next series
is given. It is uttered by individuals which are trying to locate other
quail, and is especially in evidence after a covey has recovered from
its fright at being scattered by an enemy and is trying to reassemble.

CALIFORNIA FISH AND GAME 201

There is a strong tendency upon the part of birds within an
assembled covey to answer the calls of strayed or scattered individuals,
although such birds are not themselves lost. This tendency to answer
enables the lost birds to make their way back to the covey, and may
have developed as a result of natural selection. It is so strong that
unless the weather is stormy or very cold, one can nearly always locate
a covey of birds by imitating the assembly call.

During the cold stormy winter months one seldom hears this call,
and it is difficult to get a response from any covey; however, even at
this period, whenever a day or two of warm sunny weather occurs,
the birds commence calling and answering from far and near. By
the last week in February, in normal years, the weather has become
appreciably warmer and the assembly call may be heard at intervals
nearly every day. At this period the instinct to pair is just awakening ;
males call out to females in ringing tones, and the females answer with
similar but somewhat higher pitched, more rapidly delivered notes
which are usually recognizably feminine. Calling females in a cage
or trap very soon became surrounded by unmated males. During the
height of the pairing season such excited calls may be heard all day
long, and sometimes even after dark when the birds are on their
roosts, but with pairing accomplished a comparative quiel reigns
because the two members of a pair seldom become separated by more
than a few dozen yards and do not have to call loudly to learn of
each other 's whereabouts.

The assembly call is used by both parents in response to the
peeping "lost" calls of any of their young which may have strayed
away. It acts like a veritable magnet under such conditions, even
causing chicks which are crouching in fear before the observer to
rise up and scramble in the direction of the calls. Unless the family
group is widely scattered, however, the parents utter the call in very
subdued, muffled tones, often omitting the first syllable entirely, which
is in keeping with the secretive behavior of the birds at this season.

By the first week of September, when the young are well grown
and to a large extent able to take care of themselves, the furtive
watchfulness of the parents relaxes; family groups begin to coalesce
into fall coveys, and with this new gregariousness the assembly call,
so characteristic of the autumn shooting days, once more rings far and
wide over the land, until silenced by the coming of the winter storms.

That the assembly call is primarily used to keep the birds in touch
with one another is shown by the fact that when they are foraging in
grass which is so tall as to hide them completely, they give the call
with increased frequency, as they do also on windy days when hearing
is more than usually difficult.

The Single "Cow!" of the Breeding Season. This note is similar
to the last syllable of the assembly call, but it is louder, is heard only
during the period of courtship, and is uttered by the males alone. Field
notes made by the writer indicate the origin of the call :

"April 1, 1933. Courting time is indeed here, wThat with the warm,
sultry weather which we have had recently, together with the advanced
state of the vegetative season on this (sunny) side of the mountain

202 CALIFORNIA FISH AND GAME

range. A male calling cu-cd-cow twice omitted the first and last syl-
lables of the series, the result being the cow! of the breeding season."

The birds commence to give the cow! call at the first streak of
dawn, while they are still on their roosts, and keep it up from fifteen
minutes to about an hour, after which they descend to the ground to
feed. From that time on, the note may be heard intermittently
throughout the day, although chiefly during the cool hours of the
morning and afternoon. Just at dusk, when the birds are once more
upon their roosts, there is a minor outburst of calls, which lasts for
about ten minutes, or until darkness has definitely fallen.

The cow! note is uttered at regular intervals by the bird as it
perches motionless near the top of a tree or bush, or any other site
which commands a view, yet is not too dangerously revealing. It is
uttered at intervals of from 9 to 127 seconds, the average (of 45 calls)
being 19 seconds. Following the explosive cow!, the bird regularly
utters two or three low, plaintive cheeping notes like "ivliew, ivhew,
whew," but these are only audible if the listener is within fifty feet of
the maker of them.

On the cooler eastern slope of the Santa Cruz Mountains the
first cow! notes of the season were heard between April 15 and May 2
during the three seasons of field observation. On the somewhat
warmer western slope the first calls were heard two weeks earlier
than this. Within two or three clays after the first bird or two com-
mences to call, nearly half of the resident unmated males are in full
swing, and by the end of the week the daily chorus has reached its
height. Toward the end of May, when many of the birds are paired for
the season, calling has diminished; by the middle of June only a few
birds still give the note; and by the last week of June it has ceased
entirely. On rare occasions one may hear some lone bird commence
calling again during the latter part of July, but this lasts only for
about a day. Such sporadic calling must emanate from males which
have either lost their mates or for some other reason are commencing
the reproductive cycle anew.

The number of calling males was never more than a small fraction
of the total number of males known to inhabit the region, and it
was observed that whereas mated birds never gave the call, unmated
birds did so until the time when they, too, were successfully mated.

The cow! call appears to serve as an announcement of territory
and also as notice that the bird is in a condition to pair, as Allen
(1934, p. 183) has shown to be the function of drumming in the
Ruffed Grouse (Bonasa umbellus). The call is also analogous with
the "bobwliite" call from which the quail of the eastern United States
derives its name, and which, as Stoddard (1932, p. 98) has shown "is
largely the call of unmated cocks." If a person answers one of these
calling males with the regular assembly call, the bird will often
travel 200 to 500 yards to the source of the calls, evidently in search
of the female which is supposedly answering him.

The writer has been informed by Mr. E. D. Piatt, in charge of
the State Game Farm at Chino, California, that the cow! note is
only rarely heard from the breeding pens, but that it is sometimes
given by males which are confined by themselves.

CALIFORNIA FISH AND GAME 203

The Explosive " Squill !" of Defiance. This note is the most diffi-
cult of all to describe and has been likened by McLean (op. cit., p. 16)
and Grinnell, Bryant and Storer (1918, p. 519) to an explosive note of
the domestic turkey gobbler. To the present writer it sounded like
a loud, high-pitched, staccato whistle, given here as squill! It is
uttered two or three times in rapid succession, and has something of
the tone of a tightly stretched steel wire which is struck a sharp blow
and instantly muted. The note is uttered by the male alone, is
clearly one of defiance to rival males, and is heard only during the
period of courtship.

Many times the writer has observed a would-be suitor outside
the large inclosure call excitedly cu-cd-cow! to a mated female within,
to be interrupted immediately by a vehement squill!, squill! from her
mate. In other cases, where an unmated female showed interest in
an outside male, while being assiduously courted by a male within
the inclosure, the calling of the outside bird would elicit a perfect
frenzy of squills from his rival inside, which would then be answered
in kind, at which the two males would rush at each other and
exchange lusty, if futile, buffets through the wire.

This note is uttered with all the vehement energy at the birds'
command and fairly explodes from them as they jerk their heads and
necks up to their fullest height, with bills pointed skyward.

The Deep-toned Musical "Wip-Wip" of the Male. This musical
throaty note is uttered by the male alone; it appears to be associated
with sexual activity and perhaps also with the combat of rival males
over possession of a female. It was not heard at any other time. An
extract from the writer's field notes will illustrate:

"June 2, 1932. There was a squabble between the two males while
they and their respective mates were feeding close together. The blue
[refers to color of celluloid leg band] male 'elbowed' the red male
out of the way, pushing against him and pecking the back of his head
and neck, while giving vent to the same deep musical wip-wip which
he uttered a few days ago during coition."

The Alarm Note. This is the plit-plit-plit-plit, or pwit-pwit-pwit-
pwit call of frightened quail which is so well known to the hunter.
There is considerable variation in the number of plits given in a series ;
the minimum number is usually three, while the maximum is leu
or more; usually the second plit of the series receives the strongest
accent, with subsequent syllables trailing off into a faint undertone.
Sometimes, if not greatly startled, a bird will give a few plits one at a
time. In some cases, if the listener is only a few feet away, the note
sounds like whit -whit -whit, wit-wit-wit, what -what -what, or whad-whad-
whad, depending upon the individual bird. It is given by both sexes,
and at all times of the year, and usually indicates a sense of danger on
the part of the birds but not of extreme fright.

When an enemy approaches the nest or young, the alarm note is
uttered by both parents with special vehemence and an almost machine-
like regularity, and is kept up as long as the intruder remains within
sight or earshot. So regularly can a pair with a nest or young be
counted on to give this note that it can be used as a means of taking
censuses at this time of year.

204 CALIFORNIA FISH AND GAME

Often during the nesting season the plit-plit note is uttered by
one or both birds, when no danger, either fancied or real, seems to be
present, especially when the female leaves the nest for a respite from
the duties of incubation. At such times one or both members of the
pair will alight in the open and commence preening while still giving
utterance to the note. Under these circumstances it gradually sub-
sides as the birds go about the business of feeding and dust bathing.

At roosting time one almost invariably hears this note as the birds
clamber about in the dense foliage, settling themselves for the night.
At that time is seems to reflect a general state of nervousness or unde-
eidedness rather than any sense of danger; indeed the whole covey
will usually become silent abruptly if any noise or movement arouses
its alarm.

Young birds which are beginning to acquire adult voices give the
call very frequently, often rising into the air and flying for several feet
at the same time. This is done, evidently, out of sheer animal exuber-
ance and bears no relation to the presence or absence of danger — as
is further testified by the fact that at such times the parents exhibit no
alarm, although at other times of the year they are quick to take fright
at the plit-plit of other quail.

The "Kur-r!" Note Signifying Extreme Fright. If the birds are

startled by the unexpected appearance of an enemy, this low, throaty
note is uttered with a sudden, excited emphasis and repeated three or
four times in rapid succession, sometimes to be immediately followed
by a series of plit-plit^ as the birds dash wildly for cover, or duck
down out of sight in the tall grass. Tf the birds are not very close to
cover, the cry serves as a signal for all members of the covey to freeze
in their tracks, no matter in what position they may be at the time.
An example of this is furnished by the field notes of the writer:

"April 26, 1932. * * * The four quail were feeding busily
when suddenly the yellow [refers to color of celluloid leg hand | male
gave a prolonged low clucking hur-r!, while he held himself perfectly
motionless in a half crouching position. Simultaneously with the first
note of this call the three other birds 'froze' in whatever pose they
happened to be in, the red female looking back over her shoulder, the
red male partly crouched down, and the yellow female with her head
turned to the right. For a full five minutes all four birds stood thus.
None made a sound save the yellow male, who kept up his low note con-
tinuously ; sometimes the note was so low that I could barely hear it at
fifteen feet [location of blind], although the other three quail were
twenty or more feet from the sound. Sometimes I could not hear
it at all, but was sure that it was being given because of the continuing
slight quiver of the bird's body which had marked each audible note.
At last the yellow male commenced to feed, pausing once or twice to
turn into a statue again. Gradually the rest also began to feed, the
danger, whatever it was, being over."

When the field worker hears this call in the Santa Cruz Mountains
he will be correct about 75 per cent of the time in concluding that a
Cooper or sharp-shinned hawk is after the birds.

Both sexes give the call, and it is especially in evidence when the
young are small, at which time the parents become more than ordi-
narily timid, fleeing precipitately at the slightest alarm, real or fancied.

CALIFORNIA FISH AND GAME 205

The hypersensitiveness to danger on the part of the parents at this
time doubtless represents an adaptation which has been developed to
instill caution in the inexperienced but highly imitative young.

The Distress Cry. This call may be written pseu, pseu (as in
"few"). It is uttered loudly and rapidly when the bird is seized, or
is about to be seized, by an enemy such as the Cooper hawk, and some-
times also when the bird is being held for banding, particularly in the
case of immature birds. Although adult birds do not usually cry out
thus if grasped around the body during banding, they nearly always
do so, for some reason, if seized by a leg or wing. When other quail
hear this note they crouch down and remain silent if they are in a
free state; if they are confined in a box or trap, however, they flounder
about wildly in an effort to escape. A single young quail uttering this
note while being banded will often throw a dozen covered cartons full
of banded quail into a pandemonium of scuffling and scrambling.

A similar, though somewhat thinner, note was once uttered by an
adult female which was trying to lure the writer away from its young
by means of the broken wing ruse.

The Peeping Call of Small Young, and Parental Response. This
call may be described as a thin, piping wee-wee-wee-wee-wee, delivered
in a thin, shrill, anxious tone, with a rising inflection on the last
syllables. It is given by the small young as they forage here and there
elose to the parents and serves as a stimulus to the parents to answer
with a low, soft, constantly repeated cluck which prevents the chicks
from straying.

When the young become lost they niter their note more loudly,
plaintively and insistently, and keep this up until reassured by an
answering eu-ca-cow from the parents, toward which they immediately
scramble.

Clucking Notes of Adults. The low soft cluck uttered by the
parents to keep the chicks from straying is more frequently uttered by
the male than by the female, for the former is more assiduous in the
care and protection of the young.

A very low, rapidly uttered clucking note, hardly audible beyond
thirty feet and sounding like mo-mo-mo-mo, is commonly uttered by
quail at all times of the year, often while feeding together. At times
its purpose appears to be to keep the members of the covey or family
group in touch with one another, but at other times its significance is
not so clear; it often grades imperceptibly into the warning kur-r!
note.

"Conversational Notes." These are continually uttered in sub-
dued tones by mated pairs, and by birds of a family group or covey,
while the birds are feeding peaceably together, and consist of a
varied assortment of cheeps and clucks. Their exact significance,
if any, has not been determined, but since they are only made by
individuals when in close proximity to others of their kind, they are
doutbless associated with some feeling of companionship, and possibly
serve to keep the members of the group together.

206 CALIFORNIA FISH AND GAME

SOCIAL ORGANIZATION

Gregariousness. Like other kinds of quail, California quail are
strongly gregarious and, with the exception of a few unmated birds
during the breeding season, spend their entire lives in the company
of others of their kind. Many of the actions of individual members
of the family group or covey are cooperative, even though they may
be largely instinctive and unreasoned. The cooperative nature of quail
behavior is well shown by many of the calls previously described, which
would be meaningless if the birds were solitary in habit.

Imitativeness as a Basis for Mass Action. The reactions of each
member of a covey are also indicative of a fairly well disciplined social
unit. For example, with the exception of a few inexperienced young
of the year, when one quail gives the warning ku-rr! note and
"freezes," they all freeze, and if one bird bursts into the air and sails
away from an enemy the others usually follow. That this disciplined
obedience has its beginning in the strong tendency of young birds to
imitate their parents, which in turn are more than usually furtive
and wary during the very time that the young are growing up, has been
pointed out previously. The fact that late winter coveys tend to be
more alert and disciplined in their reactions to danger is explainable
partly on the ground of gradual learning by the young, and also partly
from the fact that the more sluggish and least cooperative tend to be
weeded out by enemies. '

Birds which have become separated from their fellows, and indi-
viduals which have been liberated alone in new localities, show marked
uneasiness and run about distractedly, giving the assembly call in loud,
far-carrying tones. The anxiety of a flock to reassemble at roosting
time is so great that it will often do so even in the face of a danger
which still threatens.

The imitative tendency asserts itself throughout life, as is shown by
the manner in which the adult birds will follow one another as they
move from place to place when feeding ; inside the large inclosure it
was common to observe that if one pair of birds left off dozing and
resting in the thick brush and came out to feed, another pair imme-
diately would follow the first pair to the feeding grounds. On the
other hand, whatever may be the nature of ' ' thought transference ' ' in
certain kinds of birds which form highly compact social groups, the
members of which appear to act in complete unison (Selous, 1931), in
quail such direct thought transference is slightly or not at all developed ;
quail follow one another only as they see or hear what the others are
doing, and even then their reaction time is sometimes slow, as the
following field notes show:

"October 15, 1932. At 5.47 p.m. one of the birds crouched for an
instant and then launched into the air toward the roosting tree. At
5.48 the remainder of the flock, except one, sprang into the air and
followed the first bird. They did not start exactly all at once, for I
noticed a slight delay in the taking off of one or two of the birds. One
bird remained behind, standing motionless where the rest had been.
Half a minute later he, too, left for the roosting: tree."

CALIFORNIA FISH AND GAME 207

Among adult quail, leadership seems to be determined by chance,
with neither sex preponderating. Of course when the group consists
of parents and their small young the parents are the acknowledged
leaders. Like the bush-tits, described by Miller (1921, p. 125), one or
more "leaders" may move off nearly simultaneously in different direc-
tions, in which case it is merely a matter of chance, together with the
degree of proximity of the potential followers, as to which direction
the main body of the flock may take. Occasionally the covey may be
temporarily, or even more or less permanently, split through the follow-
ing of more than one leader.

If any one of the "leaders" is not soon followed by others, it
abandons its exploring, even though it may have discovered an excep-
tionally good food supply, and hurries back to rejoin the main covey.
Unlike bush-tits, however, such loiterers, except in the case of very
small young, rarely become lost, perhaps because of the relatively slower
method of progress of quail and the greater carrying power of their
calls.

Cooperative Action. Some of the social habits of quail are
decidedly cooperative, one might almost say altruistic. Thus when one
bird finds an unusually good supply of food it often calls the rest to
it, for example :

"May 8, 1932. All at once the blue [refers to color of celluloid
leg band] male discovered a concentrated area of millet seed, and gave
a low rapid clucking while he scratched and fed. It was just the
reaction one sees in a domestic hen which has found something good to
eat and is calling her chicks to come and get some. The blue female,
two feet distant, came running to his side and began picking up the
grains. The red male, which was ten feet away, ran in a beeline to
where the blue male was feeding, and all three fed side by side for
nearly five minutes."

One bird will also warn the others of danger, thus :

"October 20, 1932. At 3.35 p.m. I went hunting for the covey at
McLellan's and soon located them; however, at my coming they flew
in scattered detachments up into the live oak trees. Here it was
impossible to see any of the birds, even though I was within sixty feet
of some of them. For one and one-quarter hours I stood stock still on
the steep, leaf-strewn canyon side, beneath the rapidly darkening canopy
of trees, waiting for the birds to "unfreeze." I used my quail caller
frequently and would often get a response from some birds in the
farther trees ; however, one particular bird which was nearer to me
and which evidently saw me plainly, kept giving a low but far-carrying
mo-mo-mo whenever the other birds began to answer me, at which they
would become silent and remain so for five or ten minutes thereafter.
Night fell without my getting a shot."

Sentry Duty. One of the most striking and highly evolved of the
social habits of the California ouail is that of the sentry duty that is
regularly performed bv the males during the breeding season and for
several months thereafter (Fig 5).

Prior to the development of this habit both birds of the pair regu-
larly feed together ; however, the male gradually tends more and more
to cease feeding after a few minutes and stand motionless and alert,

L'OS

CALIFORNIA FISH AND GAME

ready to give warning to his mate, of any danger that threatens. At
such times the female feeds busily, and contrary to the custom at other
seasons, spends little or no time looking around for possible danger. If
all is well, the male utters at short intervals a low, soft cluck ; in case of
danger he utters the kur-r! note and plit-plits loudly, whereupon the
female vanishes.

During the incubation period the male still accompanies the female
to the feeding grounds, often flying to meet her as she Leaves the nesl ;

Pig. 5. Male California Quail doing "sentry duty" while his mate and young
(hidden in the grass below) forage without the necessity of keeping a lookout
for danger.

but now, except for an occasional disinterested peck at some object,
he spends the entire time doing sentry duty until the female is ready
to return to the nest. What nourishment he gets is picked up at odd
moments during the day, especially just before roosting time, since the
female returns to the nest some little time before the coolness of dusk
has settled.

CALIFORNIA FISH AND GAME 209

At hatching time the duties of the male as guardian are enormously
increased, for he must now maintain a constant vigilance in behalf
of the young birds while they forage throughout the long summer day.
Food is eaten by him only intermittently, and he loses weight accord-
ingly. At the least sign of alarm he gives the sharp, warning kur-r!
note which sends the family scurrying for shelter ; and when the coast
seems clear, it is he who unobtrusively creeps out of cover before the
others and remains motionless for many minutes, on the lookout for
any return of danger. Tf he commences again to utter at intervals
his soft clucking nolo, the others venture out; otherwise they do not.

After the young have grown to nearly adult size and have attained
a measure of alertness to danger, the impulse of the male to do sentry
duty gradually wanes. Some time elapses, however, before the impulse
completely disappears, and the writer has observed it in force as late
as December 3, although this was exceptional. By the last week of
July (in the Santa Cruz Mountains) the family groups have begun to
combine into larger units, with the result that there are usually several
males available for sentry duty, which relieves any one of them from
full-time responsibility in the matter.

Williams (1903, p. 146) has reported that a given sentry is
relieved of his duty after a few minutes by some other male, and that
a rough system of rotation is practiced as the feeding flock moves
slowly along. The writer has not had an opportunity personally to
observe this rotation system, but, in view of the demonstrably limited
mental capacity of the birds (page 233), it seems likely that any appear-
ance of system must be due to chance. It must he conceded, however,
that one seldom, if ever, observes more than one sentry at a time on duty,
which indicates that the remaining birds feel a sense of security and a
freedom from any necessity personally to watch for danger so long as
another of their number is doing so. On the other hand, the bird which
is acting as sentry often jumps down into the tall grass and mingles
with the rest without being replaced, which is a further indication of the
haphazard nature of the system, if such it can be called.

COVEY FORMATION AND RANGE

Method of Investigation. The size, permanency, interrelationships
and range of coveys was studied in part by daily field observations and
mapping of the locations of birds, but chiefly by means of an intensive
banding campaign involving the frequent recapture of every living
quail inhabiting a selected portion of quail range. The area chosen
for this wrork comprised about 60 acres (Fig. 13), and since it was
almost entirely surrounded by natural boundaries, the frequent capture
of every resident bird proved possible.

Eight traps, located at strategic positions in the territory, were
operated practically continuously for a year, effecting a total of 1042
captures representing 239 individual birds. Previous to this concen-
trated drive, 41 additional birds had been trapped 65 times, but these
earlier records, although used for other purposes, have not been included
in the present calculations.

210 CALIFORNIA FISH AND GAME

In addition to being banded with Biological Survey bands, the
birds were marked with colored celluloid bands used in various com-
binations so that individual birds could be recognized in the field by
means of binoculars without having to be trapped. Although occasion-
ally productive of worthwhile information, in actual practice the method
of marking with celluloid bands proved less useful than was antici-
pated. Not only did the density of the cover combine with the wari-
ness of the birds and the quickness of their motions to make observation
of band colors difficult, but in addition the celluloid proved too pliable,
so that many of the birds soon lost one or more bands while running
about or scratching for food.

Covey Formation. The first stage in covey formation is initiated
at the hatching of the young, at which time they and their parents
constitute a compact social unit. This unit endures until the young
have acquired their juvenal plumage and are from one-third to one-half
grown, or much longer if the family remains isolated. About the last
week in July, however, neighboring family groups commence to merge,
and this process continues through late summer, to the daily accom-
paniment of the assembly call, until by the middle of September the
full-sized coveys have been formed. At all times, even when first
hatched, the young will follow any adults which they may happen to
find, regardless of blood relationship.

Size. Since the average number of young in a successful brood
is about nine (page 235), five or six such family groups will constitute
an early fall covey of the size usually encountered in the Santa Cruz
Mountains. In addition to the parents which have successfully reared
their young, such coveys usually also include two or three unsuccessful
adult birds.

As the season advances and the covey is harassed by natural
enemies, notably Cooper hawks, or by man, it gradually shrinks in size
unless reinforced by birds from other remnant coveys. In the Santa
Cruz Mountains, coveys which have been reduced to less than ten birds
usually join with neighboring groups, even if this means partial
abandonment of their original territory, and this tendency results in
the maintenance of a fairly uniform size in many coveys, even though
the total number of coveys is gradually decreasing. In December,
1933, the average size of four more or less distinct coveys was 34.75
birds, which is perhaps smaller than in the days of quail abundance
thirty to fifty years ago.

On the other hand, there may simply have been more coveys in the
early days ; indeed, it is the opinion of the writer that the very nature
of covey organization is such that the permanent coherence of large
groups is impossible. Since, as previously pointed out, there is no
acknowledged leader, the number of potential leaders increases with
the size of the covey until a point is reached where it becomes impossible
for all the members to follow one bird, which results in splitting.

On the well stocked area chosen by the writer for the banding
study, the ranges of the various coveys intersected, and it was when
these coveys happened to meet on the edges of overlap that the large
coveys were formed. Under such conditions, splitting and recombining
occurred repeatedly throughout the season, the groupings being largely

CALIFORNIA FISH AND GAME

211

a matter of chance and often resulting in the more or less permanent
transference of some individuals from the range of one covey to that of
another. The movements are briefly summarized as follows (see
Fig. 13)

Covey

Number of
Individuals*

Nu

nber of captures in territory
of another covey

Number of

captures in

own territory

1
2
3
4
5

82
59
69
51

25 +

Covey No.
2-122
1- 44
1- 33
1- 2
1- 2

Covey No. Covey No. Covey No.
3-14 4- 7 5-0
3-41 4-7 5-0
2-18 4-12 5-1
2-18 3-17 5-3
2- 3 3-11 4-4

130
55

166
65

4t

* Number of individuals at hatching of young.

t But little trapping was done in the territory of covey No. 5.

Daily Range. The banding study supplemented by field observa-
tions showed that, in the Santa Cruz Mountains, the territory covered
daily by a flock of quail is often less than 200 feet in diameter, while
if food is abundant, or if the weather is stormy so that the birds
remain all day in dense cover, the distance traveled in three days may
be only 100 feet. Of course the total distance traveled within this
territory will be considerably greater than the figures given, due to the
irregular, more or less aimless wandering of the birds which involves
a considerable retracing of steps.

When food became hard to find during midwinter, the birds were
obliged to cover larger areas in order to satisfy their needs.

During the reproductive season the individual pairs have a sharply
circumscribed foraging range, due to the inability of the young to
travel far, together with the abundance of food, and this explains the
paucity of trapping records at this time of the year.

Seasonal Range. Although the daily range of a quail covey is
usually small, the total year-round range of many of its individual
members may be fairly extensive, in some cases exceeding two miles.
This is because during the breeding season, when the covey breaks up,
many of the birds scatter widely, invading regions where quail are
seldom or never seen at other times of the year ; then, with the advent
of the fall season, such strayed individuals may join some group which
is nearer to their breeding grounds than was the original covey.

Hunting also causes local emigrations of birds from the area of
harassment, as was shown during the present study when two outside
birds suddenly joined a covey of banded birds following a shooting
foray on some neighboring property one-eighth of a mile distant.

With the appearance of great quantities of green vegetation about
the first week of May, some coveys have been observed to shift from
their winter headquarters (Fig. 6) to areas of particular abundance
such as the margins of grassy slopes on the sunny side of a hill ; such
shifts rarely amount to more than a few hundred yards, however.

212 CALIFORNIA FISH AND GAME

Similar small-scale migrations often take place during the latter
part of October when the birds temporarily move into the woods to feed
upon the fruits of the madrono, snowberry, wild honeysuckle, etc.

Permanency of Coveys. Although certain individuals of a given
covey may fail to return in the following autumn, the covey usually
receives a sufficient number of new birds, most of which are young of
the year, that its size remains fairly constant. In reality the numbers
of a covey and its location are governed largely by the available food
and shelter rather than by the fortuitous presence of its individual
members ; and in this respect the covey is analogous to a river, which
constantly maintains itself although the individual drops of which it is
composed continually pass onward.

Fig. 6. An ideal winter quail range. The brushy slope shown harbored a flock
of about 35 quail at all times other than the nesting season. Dominant shrubs
are poison oak, of which dense tangles used for roosting are visible in the
foreground at left, and Baccharis pilularis, represented by the extensive
dark areas of brush. Numerous live oaks (lower right hand portion of the
-slope) and California laurels (conical trees at upper left of slope below
horizon line) furnished other roosting sites. Food (including newly sown
barley) was gleaned from the adjacent field. The birds spent a whole winter
on this area of approximately seven acres.

Homing Instinct in Wild and Artificially Propagated Birds. The
behavior of quail which have been raised in captivity and then liber-
ated in a strange environment is often abnormal, as is also that of
wild quail which have been trapped and subsequently released in a
different locality. Even under such conditions the tendency of the
birds is to remain where liberated, provided the environmental condi-
tions suit them, so that quail may be said to have only a very slightly
developed homing instinct. Of three wild quail trapped in Oakland,
California, and released in the Santa Cruz Mountain study area, all
remained where released, and raised young ; of four wild birds trapped
three miles away and on the other side of the ridge from the study area,
and released at the study area, two remained, while the other two,
which were never seen again, may possibly have died.

In the case of 67 artificially propagated birds, which were
released (with 9933 others) by the Division of Fish and Game in
suitable localities in southern California, and which were subse-
quently shot by hunters or otherwise retaken during the ensuing 16

CALIFORNIA FISH AND GAME 213

months, the average distance between point of release and point of
capture was 2.6 miles; the maximum was 10.5 miles; 17 per cent of
the birds remained in the locality where liberated. Some of the
birds that traveled the farthest ceased their wanderings only when
they readied the outskirts of the nearest town and found themselves
in the proximity of farm buildings and other evidences of human
occupation with which they had previously been associated all their
lives.

COURTSHIP BEHAVIOR

The courtship behavior of quail is exceedingly difficult to
observe under ordinary conditions because of the secretive nature of
the birds at such times, together with the effective concealment
afforded by the dense vegetation. In the case of the present study
an unusually favorable opportunity to observe courtship activities
was afforded by the large inclosure frequently referred to above.
However, even with the aid of the inclosure and adjacent blind, many
details in the mating procedure were missed because of the dense
cover which, like other features of the interior, had been left in a
natural condition.

The dates given here for the various stages of the reproductive
cycle are averages of three years' observation in the Santa Cruz
Mountain region, at 1700 to 2000 feet altitude. For other regions
they will be slightly earlier, or later, according to the climate. Thus
m San Diego County the whole cycle occurs about three weeks earlier
than in the region under discussion. In Oakland, Berkeley, and Palo
Alto, California, whose respective climates are warmer and the vegeta-
tional cycles earlier than in the Santa Cruz Mountains (although these
cities are all within 50 miles of the mountains in question), the repro-
ductive season commences about seven to ten days earlier.

Pugnacious Stage. About the last week in February, if the
\vea.ther lias not been unusually cold, the first signs of pairing com-
mence In he in evidence. Certain males which have been peaceable
enough until now will occasionally lower their heads, with crest
feathers laid hack, and make little runs at other males in the covey,
al which the latter usually dodge away without offering resistance.

With the coming of warm weather, however, as such minor
encounters become numerous, the males grow more truculent. The
males also give the assembly call with increasing frequence, apparently
to attract the females, and these, in turn, can often be heard calling in
answer. Unfortunately at this time the birds become more secretive
and as they carry on their courtship activities in dense cover, the
observer can only obtain brief glimpses of the drama which is being
enacted.

Apparently each male tends to attach himself to a particular
female which he follows about, while constantly endeavoring to drive
away any other male which comes near. The fighting between males
occurs almost any time that two males get within two or three feet of
each other. At such times one male runs at the other with lowered head
and ruffled feathers, but now, instead of running away as formerly,

214 CALIFORNIA FISH AND GAME

the other will often stand his ground with lowered head, returning
peck for peck amid a chorus of excited clucks and loud squill! notes.
The birds rush at each other and dodge with great rapidity, sometimes
even fighting for a moment breast to breast, when they peck at each
other's faces and leap simultaneously into the air like fighting roosters.
Quoting from the writer's notes: "Sometimes while thus engaged, one
would come down squarely upon the back of the other, which resembled
the position taken during coition; perhaps the birds momentarily got
their instincts mixed. ' '

This seems significant in the light of Allen's observation (1934, p.
183) that sex as such is not recognized in birds and that a dominant
male will attempt to copulate with a bird of the same sex if the latter
has been reduced to submission. The writer has also observed a male
drive away a female on one occasion, which is similar to the behavior
of some of Allen's grouse. Usually, however, one of the contestants
turns and bolts before hostilities have become violent, and in any case
the fighting never lasts more than a minute.

The vanquished male is not usually chased beyond ten or fifteen
feet, but occasionally the pursuit lasts longer as the winner follows
every twist and turn of his foe with furious determination. On one
rare occasion the loser was actually pursued through the air for fifty
feet before the chase was abandoned. Immediately upon the conclusion
of a battle the victorious one gives the squill! note with great energy
and defiance.

In a state of nature, where the vanquished bird has an opportunity
to escape, such fights must seldom prove harmful, but if the birds are
confined in a trap, one bird will often kill the other by picking the
base of the skull and adjoining neck vertebrae clear to the bone.

During this period the covey still preserves its identity but it is
now split into discordant groups and torn by the excitements accom-
panying the gradual development of sexual awareness, so that its final
dissolution is imminent. Although the writer was not able to observe
the details of the process, it is clear that as a result of several strenuous
weeks of fighting and calling, each successful male wins a place by the
side of a particular female and prevents all other males from approach-
ing her. As in the case of the bobwhite, female California quail take
no part in the scuffling of the males, nor do they fight among them-
selves, but that they are interested bystanders is indicated by the fact
that they frequently give the assembly note in response to that of the
males and often come running in answer to the calls of the latter.

Like certain pigeons observed by Whitman (1919, pp. 37, 154), the
females seem to exercise some choice regarding their future mates,
responding to the attentions of one suitor while ignoring those of
another which is equally assiduous in courtship. The writer trapped a
female, together with a male which was paying court to her, and
released them inside the large inclosure. In spite of the constant
attendance of this male, she would have nothing to do with him for
several weeks, but spent nearly all of each day answering the calls of
various males on the outside, and trying to get out to them. At length
his perseverance was rewarded, but only after most of the unmated
cocks outside had procured mates and left the vicinity.

CALIFORNIA FISH AND GAME 215

In ordinary years it is common to see the birds of a covey feeding
or running along the roadsides in pairs by the middle of March; at
night the birds still roost together in flocks, the birds of a mated pair
probably perched close together, but this is the final stage prior to the
breaking up of the covey.

Dissolution of the Covey. Coincident with the pairing of the birds
within the covey there develops a strong tendency on the part of the
various pairs to stray away in various directions without heed as to
the direction taken by others. By the middle or latter part of April,
as a result of this tendency the flock has almost entirely vanished as
such, and one now encounters single pairs of quail far from the normal
range of the covey. Such mated pairs seem to be looking for a nesting
site, and in their travels they are often seen in the most unexpected
and apparently unsuitable places, such as the scanty shrubbery around
public buildings, or along trails in the thick woods.

Behavior of Unmated Males, and Its Relation to the "Cow"!
Note. The significance of the cow! note as an announcement that the
male bird has appropriated territory and is in a condition to mate ha.s
been discussed on page 201. The majority of the males in a given area
are able to secure mates at the commencement of the courtship period
and, like Allen's grouse which did not drum, are not impelled to give
this note; instead, they pass into the next phase of the reproductive
cycle.

In marked contrast to their furtive, timid habits after the hatch-
ing of the young, is the boldness, or, more correctly, obliviousness to
danger, shown by the males during the period of courtship. While
intent upon reaching a female which they have espied they will often
allow an observer to approach them in full view to within forty feet, at
which they will merely run forward a few feet and then stop, even
though still far from shelter.

i&j

The Completion of Mating ; Loss of Pugnacity by the Male. After
the middle or later part of April, when the fighting and courting is
largely over, the two birds of a pair remain closely associated ; wherever
one goes the other follows, and if they become separated by the harass-
ment of an enemy they call anxiously to one another as soon as the
danger has vanished, and quickly find each other again. Neither indi-
vidual of a given pair has exclusive leadership during their daily travels,
but the female seems to strike out the route more often than the male.
At the roosting hour the two birds fly up at the same time into the
foliage and spend the night close beside each other. If they happen to
find themselves separated by a few feet upon arriving, one or the other
may be seen to leave its perch and fly over to that of its mate.

The male has now become less noisy and has lost most of his
combativeness, only driving other males away when they approach to
within five or six feet of his mate or himself, and not always doing so
even then. More and more he displays the impulse to stand guard
while the female feeds, alert for danger and silent except for a low,
soft clucking.

After an undetermined interval following pairing, coition takes
place, the delay apparently resulting from the time required by the

4 — 2 300G

216 CALIFORNIA FISH AND GAME

i'emale (or perhaps the male) to reach the appropriate stage of the cycle.
Readiness to copulate is shown in the male by his walking slowly and
somewhat hesitatingly up behind the female as she moves leisurely
along feeding, and placing one foot upon her back. If she squats down,
with partly extended wings, the act takes place.

Although the thesis that birds do not recognize sex has been
demonstrated in a number of cases by Allen (op. cit.), and although
it has been increasingly in vogue since the publications of Herrick
(1910) and Howard (1920, 1929) to interpret the reproductive activi-
ties of birds in terms of instincts and blind physiological cycles, this
does not mean that birds are without powers of keen discrimination as
to the personalities of others of their kind, or that they fail to act
consistently toward these personalities, whether with motives of anger
or of affection. An example in illustration of this latter idea is fur-
nished by the reactions of two mated pairs of quail within the large
inclosure, as observed from a distance of fifteen feet. The reproductive
cycles of the blue (refers to leg bands) pair coincided, whereas those
of the red pair evidently did not as shown by the fact that the red
female refused for about a week to allow her mate to copulate (although
they later reared a brood of young). Balked in this direction, the red
male tried repeatedly during the week to copulate with the blue female
whenever both pairs came out of the brush to feed together at twilight.
In every instance observed, the blue female refused to accept his
attentions, although she received the advances of her own mate, while
the blue male was evidently roused to anger (the word is used after
due consideration) by these attempts upon his mate. One instance out
of many recorded in the writer's notes may be quoted as an illustration :

"June 2, 1932, 7.16 p.m. . . . Just at this time when the
blue male had his back turned on the other three and all of the birds
were standing idly about, I saw an act which looked highly intelligent
and purposive for a bird. The red male suddenly walked rapidly, and
half crouching, in an unequivocally stealthy manner (none of his usual
hesitation) over to the blue female three and one-half feet away — the
blue male remaining oblivious to it all. The red male mounted upon her
back, but she, as always, refused to requite his amour. Instead she
walked toward her own mate, carrying the would-be adulterer, perched
standing, upon her back — a most amazing sight. As the blue female
drew near, her mate suddenly caught sight of the two; instantly he
charged at the red male, knocked him off his mate's back, pecked him
once and then gave him a distinct bite — a. hard, determined, savage
nip on the back of the neck. The blue female ran away, while the red
female, as usual, continued feeding unconcernedly."

NESTING HABITS

Unfortunately the extreme density of the cover, which has been
commented upon previously, rendered the location of quail nests more
than usually difficult in the region covered by this study.

Four successful nests were available for observation at unknown
points inside the large inclosure, but the likelihood that a search for
them would result in desertion, or other abnormal behavior on the part
of the adults, appeared so strong that no effort was made to locate

CALIFORNIA FISH AND GAME 217

them. Several nests were found by accident, and these, together with
field observations on the behavior of the adults and the subsequent
appearance of the young, have resulted in the accumulation of a certain
amount of information. An additional season devoted solely to the
study of nesting habits would have yielded highly worthwhile results.

Season of Nesting. In the Santa Cruz Mountains, altitude 1700 to
2000 feet, the earliest date for the completion of a set of eggs was
(approximately) April 18; the latest date was June 27; while the
average for twenty-one observed instances was about June 9.

Nest Site. Authorities are agreed that California quail are "far
from particular" as to the choice of nest sites, which is equivalent to
saying that they are highly adaptive. All nests found by the writer
were in slight depressions beneath but at the edge of clumps of over-
hanging brush or artificial shrubbery; likewise, the descriptions of
nest sites given by other authors indicate that, whatever their diversity,
all of the nest sites have as a common characteristic their location in
an "edge" type of environment.

The vicinity of Strathmore, Tulare County, is flat valley land, tree-
less except for cultivated varieties, and is largely devoid of cover other
than that provided by grasses and forage crops. The quail have never-
theless been able to invade this region, finding food in the rich farming
zone, and nesting sites in the vineyards, where they place their nests
against, and, when possible, partially beneath, the straggling trunks of
the vines.

Incubation. None of the male California quail within the inclosure
was ever observed to incubate the eggs ; however, Mr. John B. Price has
reported that, on the Stanford University Campus in 1933, when a
female quail deserted her nest the male took over the incubation and
successfully brought off the young.

The male California quail observed by the writer spent their time
dozing, feeding at odd intervals, or doing sentry duty, while the females
were on the nests. When the latter left their nests to feed, their mates
flew to meet them, and remained on guard over them.

The incubating female first leaves the nest at about 8.30 to 9.00 in
the morning, flying to the nearest feeding ground, where she stretches
her legs and wings, fluffs out her feathers, and relaxes generally, return-
ing to her duties in about twenty minutes. Other recesses occur at
irregular intervals — of about three hours each on bright, warm days —
until 4.30 to 5.00 p.m., when she settles upon the nest for the night,
leaving the male to feed and do sentry duty until dusk, at which time
he flies up to roost alone.

Female California quail, because of their protracted stay upon the
nest, have less opportunity to defecate than under ordinary circum-
stances, with the result that the droppings which they pass during the
brief intervals away from the nest are of the large size characteristic at
this period.

Number of Broods per Year. No indication of the raising of two
broods of young by a given pair of birds was encountered during this
study, in spite of the fact that the intensive banding program of 1933
should have revealed such a condition had it existed. Although many

218 CALIFORNIA FISH AND GAME

groups containing young of two distinct ages were encountered, calcu-
lation of their probable ages showed a difference of less than thirty days,
which would be the shortest lapse of time possible between the hatching
of the first and second broods (the minimum figure for incubation period
given by Grinnell, Bryant and Storer [1918, p. 529], is twenty-three
days, plus the time required for deposition of second set).

A few records exist which indicate the possibility of a female pro-
ducing two broods in one season (McLean, 1930, p. 19), but if this
does take place the instances must unquestionably be rare.

CARE OF YOUNG

Brooding. During the first few days after hatching, the female
broods the young during most of the time that they are not actively
foraging. "Whether or not the male also broods was not learned. The
female and young retire for the night at about 4.30 to 5.00 o'clock, or
as soon as the air becomes cool, which is several hours earlier than is
customary for full-grown birds, this early habit representing an adap-
tation for the protection of the young, from the chilling to which they
are exceedingly susceptible. For the same reason, the quail family does
not stir abroad in the morning until about 9.00 o'clock, or until the sun
has thoroughly warmed the grass. On mornings following a heavy dew,
the young quail are often brooded until 11.00 o'clock, while on the other
hand, if the night has been exceptionally warm and dewless, they may
appear as early as 5.00 a.m.

Susceptibility of Young to Chilling. The susceptibility of young
quail to fatal chilling, when they are exposed to dampness or even cool
air, is so great as to be a matter of common knowledge. An illustration
is afforded by an experience of the writer as follows :

"July 5, 1932. I kept the day-old young in a closed carton for
about ten minutes before weighing them. It was a hot morning, and
thinking that they might become overheated I put a bunch of damp
clover stems into the box in the shade.

The result was almost fatal. When I returned, all but one of the
youngsters were in a huddle at one corner of the box. The one
remaining baby had got its under surface quite soaked, probably as a
result of having been pressed against the clover by the others. It was
very feeble, and when laid upon its back it had not even enough
vitality to turn over. I thought that it would die shortly; however, I
placed it in the warm sun (only its head being shaded), and in fifteen
minutes it had entirely regained its strength."

Resistance to Wetting. On the other hand, if the weather be dry
and warm, young quail easily endure an accidental wetting. Thus,
the writer saw a day-old youngster inside the large inclosure instantly
paddle out of a pool of water one and one-quarter inches deep into
which it had fallen, without being harmed by the experience. Undoubt-
edly the breeding of quail has become timed, through a process of
evolution, to coincide with the driest portion of the year commensurate
with the maturing of the young before the rigors of winter set in.

CALIFORNIA FISH AND GAME

219

That California quail, at least of the coastal subspecies, are
accustomed to withstand the heavy ground fogs which regularly inun-
date their territory, is self-evident. On the Pacific slope of the moun-
tain crest upon which the present study was carried on, cold dense fogs
blanketed the quail country practically every summer night from
about 6 p.m. until about 9 a.m. of the following day (Figs. 7-8), yet
produced no visible ill effect upon the birds; moreover, this is the very
region which not only supports a considerable quail population at the
present time but which is described by Welch (1928) as a center of
quail abundance in the days of market hunting.

Undoubtedly the ability of the young to survive in this fog belt is
due to the fact that they are brooded until after the sun has driven

Fig. 7. Area of quail abundance on humid ocean-facing slope of San Mateo
County, showing the blanket of cold fog which inundates this country each
night in summer without detriment to young quail, because they are brooded
until it disappears at about 9 a.m.

Fig. 8. Same area as above, a few hours after the disappearance of the fog.
One of the reasons that this region has always supported a numerous quail
population is that a large amount of "edge" land is present. Unfortunately,
over-grazing has greatly reduced its value to quail in most places.

away the fog in the morning. From this it will be seen that the
activities of an enemy which forced the family to move during the
period of dampness would be injurious even if such an enemy failed
actually to capture any of the birds. So, too, it would be impossible
for quail to breed in a region where the fog did not disappear for a
long enough period each clay to allow adequate foraging on the part of
the young.

Although brooding by the female might also be necessary under
certain conditions to protect small young from the intense rays of the
sun,- such protection is ordinarily supplied by the dense cover into
which the family retreats during the midday period. As a matter of

220 CALIFORNIA FISH AND GAME

fact the young withstand high temperatures much more readily than
low ones.

Feeding. The parents do not feed their young directly, but merely
serve as a focal point for the foraging of the baby quail ; yet they are
markedly solicitous for the welfare of their offspring, especially in the
ease of the male. If one of the parents discovers some unusually
attractive or abundant food supply it gives a low, excited series of soft
clucking sounds, like those of domestic chickens under the same condi-
tions, upon which the young come running to the feast.

Relative Solicitude Displayed by the Two Sexes. The greater
solicitude of the male is well shown when one surprises a family of
small young and their parents. If there is the slightest chance of
escaping detection, the male will remain with the young rather than
slip away as the female frequently does. An illustration of this is
afforded by the writer 's notes :

"July 5, 1932. Mention should be made of the extreme tenacity
with which the parents (especially the male) held to the dense cover
right at my feet while I was catching the baby quail. The cover was
dense enough to completely conceal the old quail, and since they still
had a fevr of the uncaught young with them, they were extremely loth
to leave. I could wTalk up to within two and one-half feet of them
without knowing they were there, and would only become apprised of
llieir whereabouts when they burst out of the clover with a whir of
wings and sailed off, to alight thirty feet away and call plit-plit-plit
frantically. No wonder I couldn't locate any of the breeding quail up
at the pasture recently ! ... As usual the male was more solicitous,
or at least more bold. He always returned first, called to the young
more frequently, and remained longer."

Defense of Young by Parents. Adult quail are by no means as
helpless in the defense of their young as one might suppose. Even
large animals are sometimes driven away from the vicinity of a brood,
the male usually being the more aggressive, as in other activities per-
taining to the care of the young.

The writer saw a male quail chase meadow mice (Microtus calif or-
nicus) away from the vicinity of the brood several times, and has
observed the routing of thrashers (Toxostoma redivivum) on many
occasions ; in addition he has received reports from other observers, as
follows :

Mr. A. B. McLellan, a rancher living on part of the area covered by
the present study, saw a male quail fly at his dog and chase it away
from a brood of young. Mr. Donald D. McLean reported another
instance of a dog being driven away; also an attack upon a spotted
owl (Strix occidentalis) which he had flushed, by a male quail, the
latter hurtling with considerable force at the owl as it flew by. Mr.
McPherson Lough, deputy game warden at Palo Alto, watched a male
California quail as it successfully defended a brood of small young
from the attacks of a California jay (Aphelocoma californica) ; he
also saw a male quail repeatedly rush at a king (?) snake in an open
field, but was unable to find any of the young. Mr. Allan McKean,
formerly bird and animal economist for the Division of Fish and Game,

CALIFORNIA FISH AND GAME 221

reported that a male quail made repeated dashes, on foot, at a road-
runner (Geococcyx calif or nianus) which was near a brood of young
quail until the road-runner was driven away.

There is a published record of a California quail chasing away a
jay (Post, 1932, p. 96) ; while Jaeger (1926, p. 271) records that a
female quail "in close cooperation" with an English sparrow {Passer
domesticus) drove a weasel (Mustela xanthogenys) across a vacant lot
;ind an adjoining street, until they were frightened by a workman.

Feigning Injury. On one occasion when the writer suddenly came
upon a brood of young accompanied by their parents, the female
toppled over upon her side, and uttering a thin, peeping distress cry,
struggled and fluttered in the dnst as though mortally hurt. In the
meantime, of course, the chicks scrambled, cheeping, into the dense
brush. Although this ruse has been recorded once before, by Proud
(in Bendire. 1892. p. 29). it does not seem to be common, for none of
the parents of numerous other broods discovered by the writer acted
this way.

Timidity of Adults in Relation to Imita'tiveness of Young. As

mentioned previously, the adults arc far more timid during the time
that the young are small than at any other season, and this doubtless
has an adaptive significance. As an example of this hvper-sensitivity
to danger, the writer observed many times the seemingly panic-stricken
state of parent birds inside the large inclosure, caused by a glimpse
of some one walking along a certain, not far distant, path. The birds
would cry Jcur-r! and instantly dive into the tall grass, their imitative
young scrambling pell mell after them, and all remaining motionless as
stones for long minutes thereafter. Yet these same adults, both before
and after the breeding season, were accustomed to feed in the open
with perfect unconcern while numerous persons passed back and forth
along this path.

Limitations of Instinct as Displayed in the Care of Young. That
the behavior of adult quail in caring for the young rests upon no more
exalted an intellectual level than do the other instinctive activities of
birds is well shown by the rather common instances of maladjustment
betwreen instinct and need. For example, no direct effort is made by
the parents to help their chicks out of difficulties, no matter how simple
the rerpiired act of deliverance may be. If one or more of the young-
sters happen to tumble into a ditch or become causrht in thick weeds the
adults will remain in the vicinity for a time, lending encouragement by
answering with frequent calls of cu-ca-cow the disconsolate cheepings
of their offspring, but that is all. If the youngsters are able to extri-
cate themselves unaided and catch up with the gradually receding
family group they are saved ; otherwise they are not. Tt is for this
reason that one finds abandoned stragglers fairly often during the
summer months.

Of course this rough-and-ready system of culling tends to insure
that in the long run only the most nimble and vigorous individuals
shall survive. A compensating circumstance is that the adults show no
aversion to adopting the lost young of broods other than their own.
This readiness to adopt other young1 explains many if not all of the
cases where young of two distinct sizes have been observed following
a single pair of adults.

222 CALIFORNIA FISH AND GAME

GROWTH AND BEHAVIOR OF THE YOUNG

Since the present investigation was conducted primarily in order
to learn the environmental requirements of quail, little time was avail-
able for a detailed study of the growth and behavior of the young.
For this reason the data presented on this subject are not as complete
as desired.

HATCHING

In the Santa Cruz Mountains the first brood of small young was
seen on May 11 ; the latest was seen July 19 ; while the average date
was about July 2.

FIRST FIVE DAYS

Frailty. Some of the characteristics of the behavior of small
young have been noted above under previous headings. For the first
three or four days after hatching, baby quail are distinctly weak and
frail, in spite of the agility which they display when following their
parents. During the first day, especially, they repeatedly fall headlong
over small obstructions, such as clods and grass stems, and it is evident
that their precocious activity is possible chiefly because of the small size
of their bodies, which requires relatively slight muscular strength to
move. The frailty of the youngsters at this time is also shown by the
ease with which they succumb to chilling, as mentioned above.

Initial Fear Reactions. Like the young of other species of birds,
newly hatched quail are equipped with only a generalized fear
mechanism ; they probably give the fear reaction to any large moving
object, irrespective of whether it may be harmful or not, and only
learn what to fear as the result of an extensive period of tutelage
under their parents.

When a human intruder approaches a family group, both adults
and young usually crouch down in the long grass and remain perfectly
still, thus being likely to escape observation. However, if escape by
crouching proves impossible, and the adults are forced to leave in the
explosive manner previously described, the young instantly scramble in
all directions, uttering in chorus their fine cheeping wee-wee-wee.
When they come in contact with any object such as a clod, clump of
weeds, or a dry leaf, into or under which they can squeeze themselves,
they become silent and motionless unless actually seized by the enemy.
They abandon their crouching tactics only after the frenzied plit-plit
of the parents has subsided and the low clucking notes and subdued
cu-ca-cow of parental encouragement are heard. Even when the young-
sters have been captured and are huddling silently in a closed carton,
they will come to life with loud peeps and frantic scrambling if they
hear this call.

The crouching of young quail in crevices when under the stimulus
of fear resembles the thigmotactic reactions of young terns under the
same stimulus (Watson, 1908, p. 242). Like the latter, the quail chicks
are often satisfied to sit quietly with their heads in a cranny, even
though their bodies can be seen by the enemy.

CALIFORNIA FISH AND GAME 223

Feeding. The instinct to scratch in surface litter and peck at
small objects is present from the first, but it is doubtless necessary for
quail chicks to learn to distinguish edible from inedible objects by a
process of trial and error, as is the case in the domestic chick. The
propensity to eat insects must also be instinctive, since it can hardly
be learned by imitating the 99.62 per cent vegetarian parents.

Iris. The iris color at this age is light brown instead of dark
brown as in the adults.

SIXTH TO FOURTEENTH DAY

By the sixth day the young quail are noticeably stronger and no
longer tumble over every little obstacle. They often jump into the air
to a height of three or four inches, thereby showing the first signs of
xhe instinct to play. Small clods or other low objects are hurdled or
used as lookout stands by the youngsters as they scamper about in
response to the urge to exercise their developing muscles.

A slight increase in efficiency of the fear reactions was shown by
the youngsters in the large inclosure on this day, in that when actually
pursued they ran far into the tall grass, threading their way among
the tangled stems, instead of merely crouching down next to the nearest
clod. On the other hand, they had as yet acquired little or no sense
of potential danger, for while feeding they ran here and there around
the old birds, cheeping loudly and showing none of the cautious watch-
fulness of their parents.

FIFTEENTH TO TWENTIETH DAY

By the fifteenth clay the young are just learning to use their
wings. They run and jump more vigorously than ever, often leaping
into the air when there is no obstacle to hurdle, and fluttering along
at a height of two and one-half to three feet above ground for a dis-
tance of seven or eight feet. When suddenly confronted by danger,
they spring into the air and fly toward the nearest cover, and once
they have gained it, they can be forced to leave only with difficulty.

During this period their thin peeping note grows stronger and
deeper in tone, and in addition the birds now commence to give the
plit-plii note of the adults. The latter note is as yet decidedly squeaky
in tone, as well as being less loud than that of the adults; however, it
is recognizable as the same vocal process.

Although the youngsters are thus manifestly more able-bodied,
they have as yet learned very little caution from their parents, and
are noticeably self-willed. An example from the writer's notes will
illustrate :

"July 19, 1932. . . . After they had drunk, the male tun km I
and walked back toward the grass. He clucked several times, but
only a few of the chicks followed him, the rest being scattered about
in all directions. It is evident that they are becoming more self-willed
and independent. I approached so close to the window of the blind
that the male saw me. He clucked more emphatically and retreated
farther into the grass, but still the chicks paid little heed. A few
more came to him, and once, when he clucked a little more insistently,
in fact giving the danger call, one of the chicks jumped away from
him and into a clump of weeds — but it reappeared a moment later

224 CALIFORNIA FISH AND GAME

and continued feeding. After several minutes they all gradually
joined the parent."

Such laxity in obedience to parental warnings doubtless results in
the destruction of many young birds of this age.

TWENTIETH TO FORTY-FIFTH DAY

Between the twentieth and the forty-fifth day the initiative instinct
of the youngsters commences to assert itself, and they readily ' ' freeze
at the danger note of the male, although they still show a marked
tendency to relax and commence feeding again before their more
cautious parents are ready to do so.

During this period the young birds continue to exercise then-
developing wings, and also their vocal powers. Often at this time
several individuals spring into the air at one time with excited,
squeaky plit-plit notes, and nutter into the nearby cover, as if threat-
ened by danger. Actually no danger exists, and such play activities
serve merely as a drill in preparation for future necessity.

At this stage of development the youngsters are in full juvenal
plumage, with a sufficient coating of feathers that brooding by the
female is no longer necessary. Accordingly, the whole family now
roosts together in the trees, retiring at the regular roosting hour
observed by the adult birds.

Dust bathing is indulged in by the young from this time forth,
and the diet changes gradually from the markedly insectivorous type
of previous weeks to the nearly 100 per cent vegetarian fare of the
adults.

LATER STAGES

As the birds begin to acquire their first fall plumage, about the

eighth week, their flight becomes considerably stronger, taking on
the powerful whirring wing motion of the adults. Accordingly, the
young birds come to trust more to their wings when escaping enemies,
and less to their earlier crouching concealment tactics. Indeed, at this
time they flush more readily than the adults, if one beats the cover
where they are hiding.

At this stage they have commenced to absorb some of the wariness
of the old birds, responding with celerity to the signal to "freeze," and
diving quickly into cover when alarmed. However, many of them still
have the perilous habit of perching in plain sight on the tops of bushes
after having been flushed, and it is doubtless significant that more quail
were observed to be captured by Cooper hawks at this season than at
any other time of the year. The gregarious habit of quail provides an
efficient if somewhat rough and ready school of training from which the
young birds continue to learn, even after attaining adult stature.

On the fifty-second day the young quail in the large enclosure
were heard giving the assembly call for the first time. The notes were
faint and shrill, while the amount and placement of the accent given
the various syllables varied greatly, thus: cu-ci't-cu, cu-cu-cn ; cu-cu-cu,
vu-cu-cu.

The "conversational" cheeping notes uttered by feeding quail
gradually developed from the peeping of the young as their voices
became deeper and less shrill; however, the clucking mo-mo-mo-mo

CALIFORNIA FISH AND GAME

225

note was not heard until the birds inside the inclosure were sixty
days of age.

Weight Increase. Young quail grow slowly during the first
week after hatching, as the following figures, from one brood, show:

Age (days)
o

6

20

21

76

77

78

adult__.

No. of birds
weighed

ir>

10

6

3

4

1

1

550

A verage weight

(grams)

6.1

8.4

19.3

27.2

128.5

121.3

128.8

189.5

Fig. 9. Wing of adult quail (left), and of immature quail, less than one year oM
(right). The latter shows the buffy-white spots on the upper primary coverts
which serve to distinguish birds of less than one year from the adults.

Use of Juvenal Plumage Characters for Age Diagnosis. Upon
the acquisition of the first fall or adult plumage, commencing about
the eighth week, all of the grayish-buff, brown spotted juvenal plumage
is lost except the primary coverts and outer pairs of primaries.
The innermost six primary coverts of the juvenal plumage are "clay
color," barred and striped with buffy white, in which respect they
differ from the fully adult plumage, as shown in Fig. 9.

Because these mottled coverts are not lost until the second autumn
after hatching, they furnish a reliable indication that birds possessing
them are not more than one year old. Birds with mottled coverts
breed during their second summer (that is, when one year old) just.

226 CALIFORNIA FISH AND GAME

as freely as do the older birds ; however, since in their second fall molt
they finally lose these feathers, early in August before the ensuing
crop of young (which are markedly smaller in size) has begun to
take on this plumage, there is never any chance of confusing the two
generations upon the basis of this character.

The fact that the young of a given season may be distinguished
from their parents for a whole year thereafter is of considerable
importance to hunters and game managers, as pointed out hereinafter.

GENERAL BEHAVIOR AND MISCELLANEOUS HABITS

OF QUAIL

ROOSTING HABITS

Behavior at Roosting1 Time. Unless they have been disturbed
a short time earlier, the members of a covey come out into the open
just at dusk and feed rather close together, uttering their low cluck-
ing and cheeping notes. Then, as the proper degree of twilight arrives,
one or two birds will stop feeding, take several hesitant steps in the
direction of cover, and suddenly launch into the air with a whir of
wings, heading straight for the roosting site.

If the covey has been feeding some distance from cover, as for
instance in the middle of a barley field, so that the birds are a little
nervous, the entire flock rises at once, with a thunderous roar, and
accompanies the leaders. On the other hand, if the site of the even-
ing meal is close to cover, so that the birds feel at ease, they may leave
in straggling groups of three or four. Even in such a case the entire
covey is at the roost within about K- minutes following the arrival of
the first individuals.

Having reached the roosting tree, or trees, the birds take from two
to about ten minutes to settle themselves for the night. Some indi-
viduals hop from twig to twig, calling plit-plit excitedly, and causing
the branches to bounce and sway with their motions; others come sail-
ing in on set wings and plump themselves down into the foliage with-
out any further noise. As the minutes pass and the twilight deepens,
while the horned owls commence to boom, the commotion in the roost-
ing tree subsides ; a single bird may call cu-cd-cow once or twice as if
to reassure itself of the near presence of its companions, and then all
becomes silent.

Ground Roosting. It is commonly believed that California
quail always roost in trees, or at least off the ground. This is not
strictly true, although the proportion of ground to arboreal roostings
is probably less than 1 to 1000. The one instance of ground roosting
personally observed by the writer was recorded in the field note book
as follows :

"Sept. 5, 1933. . . . Incredible though it may seem, about
twelve quail were roosting in a compact bunch on the ground in an open
field. The field is covered with a dense, unbroken stand of tarweed
(Hemizonia congesta [Fig. 10]). I first saw the birds while driving
over the field at about 9 p.m. ; the birds did not fly, but their running
could be traced by the waving of the dense herbage in front of the
head lights. Jumping out of the car, and pursuing them with a
flashlight, I caught four by hand, the dense viscid tarweed hampering

CALIFORNIA FISH AND GAME

227

them in their efforts to take wing; the others eventually succeeded in
launching into the air. I cannot be sure of the cause of this extraor-
dinary choice of a roosting site. Of course the Hemizonia was seed-
ing abundantly, providing a fine evening meal, as well as dense cover.
A Cooper hawk was in the vicinity this evening and may have fright-
ened the birds so that they did not dare to fly to the trees."

Types of Roosting Cover. The preferred type of roosting cover
is furnished by a considerable variety of trees having characteristically
dense foliage, and in addition to the species listed in this paper (Part
II) there are doubtless others in use locally.

Fig. 10. Ground roosting site in field as described in the text. Solid area of
low, light colored plants represents tarweed (Hemizonia congesta) ; tall,
dark weeds in left foreground are curly dock (Rumex crispus).

The height above the ground chosen by the birds seems to be
determined largely by the location of the densest shelter in the tree
concerned. Thus, in small dense live oaks, it is sometimes no more
than 11 or 12 feet, while in the case of very large, densely foliaged
California laurels it is occasionally as great as 35 feet. In the majority
of cases, however, the roosts of the birds are to be found between 20
and 25 feet.

In the arid parts of southern California, trees dense enough for
roosting are scarce or even absent, and in such cases the birds are
obliged to use the low, open brush for this purpose. Under such con-
ditions they may roost anywhere from a few inches to two or three
feet above the ground, or possibly on it in some cases; but the inade-
quacy of the protection offered by such cover is reflected in the abnor-
mally heavy mortality caused by owls.

L'L'S

CALIFORNIA FISH AND GAME

Occasionally, quail roost in dense tangles of brush when these offer
unusual security, even where tree roosting- sites are abundant. For
example, a covey of quail in the Santa Cruz Mountains regularly
roosted at a height of two to ten feet above ground in various impene-
trable tangles of poison oak which covered large areas in their
territory (Figs. 6, 11).

4i

Fig. 11. Dense tangle of poison oak (Rhus diversiloba) about
seven feet high and covering many hundred square feet, in
which a covey of quail habitually roosted. This tangle is one
of several shown in Fig. 6.

Position During Roosting. Numerous observations made at
night with the aid of a flashlight showed that, although some birds might
roost within a few inches of each other, occasionally even touching,
the covey was usually scattered rather widely through the roosting tree,
provided perching facilities were adequate. If two or more suitable
roosting trees were close together, the birds would spread out into these,
while certain lone birds might spend the night fully twenty feet from
their nearest neighbors. Sometimes at roosting time a covey would
divide into two groups which would fly to trees as much as 200 feet
apart.

CALIFORNIA FISH AND GAME

229

Fig. 12
ing posi

Roosting birds showed a marked tendency to perch in the crotches
of small branches, and in places where small lateral branches diverged
from an upright trunk (Fig. 12). The birds would snuggle as close as
possible to the upright branches, which reduced the chances of their

outlines forming a silhouette from any
angle and rendered them almost invisible.
On more than one occasion the writer lias
crouched for hours beneath a low, dense
tree which was known to contain ten or
more roosting quail, without being able to
locate more than one or two of them,
despite the fact that the belly patches of
the birds are fairly conspicuous when seen
from below by the glare of a powerful flash-
light.

Whether or not quail roost at night
with their heads beneath their scapulars

Diagram showing roost- was no+ flotprnn'ned bnt a miail which w;is
edition commonly assumed " °>s nur ue lei mmeu, out a quail \wuui was

by California quail in forks of asleep during the daytime was observed in
S, branches in dense this position.'

Power of Vision at Night. Like the majority of day birds,
quail display poor vision at night, and once darkness has set in are
loth to leave their roosts unless driven by the most extreme urgency.
Birds which are frightened from their roosts fly hesitantly and act as
though lost, even on moonlight nights in territory where they have
spent their entire lives. Although they seem to know enough to steer
toward trees or brush under such circumstances, they alight blunder-
ingly on top of the foliage and remain there for several minutes
before making their way gropingly down into the sheltered interior.

Undoubtedly the slow flight and bewildered manner shown by the
birds at night makes them easy prey for owls and other night enemies,
once they have been forced from their roosts. Their inability
to see in the dark also explains their tendency to remain on their
perches in the face of a brilliant flashlight beam — often to the point
where it is possible, by careful maneuvering, to catch them by hand,
as is the case of various other day birds.

Time of Roosting. The time of roosting is determined by the
intensity of the light, rather than by the hour as recorded by the
clock, and consequently it varies according to the season of the year,
as shown by Table 4. The state of the light at roosting time is
what may be described as "deep dusk," in the absence of foot-candle
ratings, and it undoubtedly represents the minimum illumination
at which the birds are able to perceive the approach of enemies or
pick out the proper sites for roosting. At such times there was
always enough light to enable the writer to distinguish nearby
objects, but coveys of quail feeding on plowed ground at a distance
of 100 yards were invisible; it was the time of day when the small
passerine birds gave their evening chorus and flitted restlessly
through the shrubbery as they, too, prepared to go to roost.

That quail react with remarkable uniformity to changing light
intensity is shown by the fact that all the birds in a given section go

230

CALIFORNIA FISH AND GAME

TABLE 4

RELATION OF HOURS OF ROOSTING AND AWAKENING TO THE

SEASON OF THE YEAR

. * = 2nd Year of Observation

Month

Day

Roosting

hour

(P.M.-)

Awakening

hour

(A.M.)

Time spent

on roost
(hrs., min.)

December-
January

January

January

February..
February-
February..

March

March

March

March

March

April

April

April

April

April

April

April

April

April

April

April

April

April

May

May

May

May

May

May

May

May

May

May

May

May

May

May

May

May

May

May

May

June

June

June

June

June

June

June

June

July

July

July

5

8

14

6

9

18

4

18

21

27

29

7

8

13

14

16

17

18

22

23

24

26

27

28

3

4

8

10

11

12

15

16

17

18

19

20

22

23

24

25

27

28

31

1

2

6

S

14

15

19

20

3

5

28

5.00

5.10*

5.25*

5.34*

6.00*

6.00*

6.08*

6.22*

6.43*

6.45
6.49*

6.58

7.00*

7.03*

7.00

7.03

7.03

7.04

7.07
7.03
7.07

7.10

7.17
7.23*

7.25
7.23

7.25*

l .£.

99*

7.19 (clou
7.19 (clou

dy).
dy).

7.35

7.34

7.31

7.14 (clou

7.20 (clou

7.35*

7.30

7.36

7.30

7.45*

7.47

7.50*

7.52

5.56*
5.51*

5.31*

5.10

4.50
5.10

4.35

4.05

3.56

4.08

3 55

dy)

dy) 3.15*

11.34
11.07

10.37

10.06

9.43

8.42

8.37

8.21

4 . 10 (clou
3.59

4.05*

dy)'-

8.21

8.19

4.10*
5.00*

8.18

CALIFORNIA FISH AND GAME

231

TABLE 4— Continued

RELATION OF HOURS OF ROOSTING AND AWAKENING TO THE

SEASON OF THE YEAR

* = 2nd Year of Observation

Month

August

August

August

September.
September.
September.
September

October

October

November.
November.
November.
November.

Day

22
25
31

5
19
20
25
15
16

5
23
24
26

Roosting
hour

(P.M.)

7.05

7.09

7.00

6.55*

6.29

5.47*

5.26*
5. 17*
5.11* (clo
5.15*

Awakening
hour

(A.M.)

5.35
5.30*
5.30*
6.53*

udy) .

6.48

Time spent

on roost
(hrs., min.)

11.06

13 . 33

to roost within a period of about ten minutes. Coveys on the east
side of a ridge go to roost slightly earlier than those on the wesl
side of the same ridge because darkness comes a tritle earlier for
the former. On cloudy evenings, when the illumination is less than
usual, the birds fly up to their perches at a correspondingly earlier
time, as Table 4 shows. A still more remarkable illustration of the
uniformity of the roosting hour under given light conditions is
provided by the fact that, although the figures given in the table
represent observations during two different years, yet when com-
bined into one seasonal cycle they dovetail perfectly; moreover, the
covey upon wihich the observations were made was not entirely
composed of the same individuals during the two years. As a matter
of fact one can predict within two or three minutes, often less, just
when a given covey will go to roost, when once the daily rhythm for
that particular locality has been ascertained.

As shown in the table, due to the seasonal fluctuation in the
roosting hour, from 5.00 p.m. to approximately. 8.00 p.m. together with
a corresponding but reverse fluctuation in the hour of awakening, the
length of time spent by quail on their roosts varies from 8 hours,
18 minutes, in July, to 13 hours, 33 minutes, in November, and pos-
sibly still more in December and January. These figures indicate
that possibly the northward limit of range of the California quail
may be determined in part by a less than optimum daily period of
time available for foraging during the winter months, especially
when the presence of snow increases the difficulty of obtaining food.

Frequency of Use of a Given Roost. The frequency with which
a given roost is used depends in part upon local circumstances sucn
as the availability of other roosts, and the distance of the covey from
the roost at the end of the day. In the Santa Cruz Mountains, where
roosting cover is extensive, the birds frequently fly to the nearest

5—23006

232 CALIFORNIA FISH AND GAME

available tree, and since the whereabouts of a covey within its terri-
tory is largely dependent upon chance, the choice of a roosting site
depends upon the same factor. Sometimes a given roosting tree will
not be revisited for more than a week; and because of this irregu-
larity, it has not proved possible in the region mentioned to use
droppings as indicators of quail abundance or for any other purpose
involving accurate estimates.

Awakening Habits. In the morning, quail frequently start call-
ing from their roosts at an hour when the degree of illumination
corresponds roughly to that of roosting time. Low, "conversational"
clucking notes can be heard from the dark canopy of foliage, and
occasionally a male bird will burst out with a ringing cu-ca-cow. Dur-
ing the breeding season this period of early dawn is enlivened by the
chorus of cow! notes uttered by the unpaired males. This is the time
of day when the song-birds are also chirping and calling, but whereas
such small birds are already up and about, the quail have not yet
moved from their roosts.

Contrary to prevalent opinion, California quail are late risers as
compared with the passerine birds. Usually quail will remain on their
roosts from six to fifteen minutes after the smaller birds are astir,
while on especially cold winter mornings they sometimes do not fly
down for nearly an hour. Like the bobwhite, which also dislikes to
move about on cold or wet mornings (Stoddard, op. cit., p. 55), Cali-
fornia quail even after having left the roost will often stand about
listlessly in groups, with their feathers fluffed out, or run stiffly for a
few steps to peck at some morsel, and then stop and stand all hunched
up. It is because of this dilatoriness that the hours of awakening
recorded in Table 4 show less uniformity than do the hours of roosting.

Of course, during summer, when the weather is warm even in
the early morning and there is little or no dew, the birds are more
lively, especially the unmated males that rise with the song-birds and
commence their matutinal calling.

DUST BATHING AND RESTING

With the exception of the breeding season, when courtship activ-
ities and the care of the young occupy most of the day, there is little
left for quail to do after they have breakfasted and visited water.
Some wander about picking up a morsel here and there in a leisurely
fashion ; others stand for long intervals dozing, with feathers fluffed
out and head depressed, their eyes slowly closing and then blinking
open again almost immediately as the birds glance around for danger
before venturing upon another momentary nap. Still other individuals
scratch and wallow in the earth, dust bathing like chickens; they kick
out with first one foot and then the other while lying over on the
opposite side; and then, returning to a sitting position, with fluffed
up, vibrated plumage, they work down into the loose earth which they
have scratched up. Damp earth, close to a spring or when caused by a
light shower, seems to stimulate the birds to unusual bathing activity.
It is also common to see two or three individuals use a ' ' bathing hole ' '
one after another, which of course may facilitate the transmission of
lice and other organisms capable to bearing disease.

CALIFORNIA FISH AND GAME 233

The period of inactivity just described is especially characteristic
of the warm hours of the forenoon and early afternoon; however, local
incidents may modify the schedule at any time. The sudden appear-
ance of a Cooper hawk, for example, may force the covey to drop its
indolent repose at an instant's notice and scramble wildly for cover,
where the birds may crouch for an hour before they dare venture
into the open again.

With the passing of the hours the urge to drink may assert itself
again, or the birds may all get up and wander off for no discernible
reason. Then, as the afternoon becomes cooler, a certain amount of
feeding commences, together with more dozing, dust bathing, and
occasional sudden frights, until at length it is evening. Appetites
now begin to assert themselves and feeding becomes the main business,
with the additional necessity of another drink if it is midsummer,
after which it, is time to go to roost, and another day is done.

PSYCHOLOGICAL TRAITS

Although quail are noted for their wariness, numerous observations
made during this study have shown that this wariness results from a
natural timidity and nervousness, rather than from any unusual mental
powers. However, they undoubtedly possess a retentive memory as
regards fear experiences. Certain coveys which were hunted regularly
in order to obtain specimens soon became so wild that they would
flush at a distance of 150 feet or more and immediately fly into the
depths of the woods, where they would perch in scattered formation
in the tree tops and utter no sound for more than an hour thereafter.
In another case wherein a covey of birds inside the large inclosure
was caught bj^ hand at night and greatly frightened, the birds did
not once venture out into the open to feed during the following forty-
nine days!

The natural timidity of the birds is further shown by the fact
that several days after small passerine birds have learned to visit
the traps freely, the quail are still reluctant to enter such structures.

The rather low degree of intelligence which quail display, disre-
garding their natural timidity, is shown by their inability to cope with
situations requiring more than the stereotyped reactions provided by
habit and instinct. For example, when a family of young quail was
confined by itself in a small, open-topped pen whose wire sides were
only twenty-one inches high, the parents, instead of hopping or flying
over this insignificant barrier to reach the young, ran back and forth
all day long at the base of the wire, futilely trying to thrust their
heads through the meshes. Even when a slanting wire screen was laid
from the top of the pen walls to the ground outside, so that the parents
could easily walk up the ramp thus provided, they took no advantage
of this aid but continued their endless pacing up and down. In
another case a female, confined with her brood inside this small pen
tramped to death by her blind, interminable running along the base
of the walls, three of the thirteen young which trailed along, cheeping
and tired, behind her. To other kinds of birds, especially jays and
thrashers, which are not so circumscribed by inborn terrestrial ten-
dencies as quail, this twenty-one inch strip of netting constituted no
obstacle at all, either to egress or ingress.

234 CALIFORNIA FISH AND GAME

The lack of modifiability shown by quail, which sometimes require
two or three years to learn to eat a new food, even though it may be
abundant, is a further indication of the intellectual limits of the birds.

In contrast to the lack of mental adaptability just cited, may be
mentioned the unique incident described on page 216 wherein a male
quail seemed to know that he could force his attentions upon a certain
female with impunity at a time when the true mate of this female
was not on guard. That quail may learn to modify their innate habits
after a time is also demonstrated by the reactions of certain birds
inside the large inclosure, that at first used to bump their heads severely
against the low wire top at roosting time, but after the first ten days
learned to check their flight before striking it.

In short, the California quail appears to be a creature possessing
no more than average avian intelligence, whose everyday activities
are largely dominated by instinct reinforced by habit. Certain funda-
mental behavior patterns, together with a marked natural timidity,
suffice it for solving most of the emergencies which it encounters,
and where this equipment fails and the individual perishes, the species
is preserved by reason of its high fecundity.

RELATIVE UNIMPORTANCE OF HEARING IN DETECTION OF DANGER

Like various other species of birds observed by the writer, quail
place little reliance upon sound (other than their own alarm notes) as a
warning of danger, unless they have learned through experience to
associate a definite sound with some definite danger. Even hearing
the whirring noise of some other covey in full flight seems to make
little impression upon them, provided no alarm notes are audible ;
likewise the commonly uttered kek-kek-kek note of their arch enemy,
the Cooper hawk, produces no observable effect upon quail which are
nearby, perhaps because the hawk makes this noise when it is perched
in the thick foliage where the birds can not see it.

Similarly the birds inside the large inclosure, although they fled
precipitately whenever they perceived the least motion through a hole
in the blind, yet showed no concern when loud hammering sounds were
made inside. The most striking example of the indifference of quail
to such sounds was shown by a male quail which walked along the top
of a fence with perfect composure while eight shots were fired toward
him from a concealed point thirty feet distant with a .22 rifle.

The warning cry of the California jay, which is sometimes heeded
by quail, has probably become associated with danger as a result of
actual experience. The utility of this note, from the standpoint of
quail, is discussed on page 240.

MORTALITY

NEST FAILURES

The percentages of successful and unsuccessful nestings of quail
were not determined directly in the case of the present study, due to
the; difficulty -of locating enough nests; however, an approximation of
the percentage of nest failures may be obtained by another method.

CALIFORNIA FISH AND GAME

235

Various observers have recorded the percentage of nest failures for
different species of birds as follows:

Species

Nest location

Percentage
of failure

Authority

Bobwhite. ._ ...

Ground

64 (nests) __
57 (nests) __

58 . 7 (eggs) .
59 (eggs)

52 (eggs)...

40 (eggs) —
25 (eggs)...

Stoddard (1932, p. 184)
Bump (1933, p. 389)

Pickweil (1931 p. 110)

Ruffed grouse.

Ground

Horned lark (Otocoris

alpest7'is)

Ground

Song sparrow (Melospiza
melodia) .

Open nest, above
ground . .

Nice (1933, p. 130)

Miscellaneous passerine
species. _

Mostly open nests,
above ground

Tree swallow (Iridoprocne
bicolor)

Clabaugh (1925, p. 115)
Low (1933, p. 87)

Blue bird (Sialia sialis) . .

Bird boxes. .

Low (1934, p. 41)

From these data it might be judged that the percentage of nest fail-
ures for California quail amounts to near 59 per cent ; on the other hand,
it seems likely that it was not quite as high as that of the bobwhite
quail, in the unusually favorable region covered by the present study,
where the grazing factor was insignificant, farming activities unim-
portant, rains absent, and cover ideal. Accordingly, we may arbitrarily
assume that the percentage of nest failures was about 60, and then
check this figure with the known resident population of adults and
young, as revealed by banding studies.

As recorded below (page 246), 175 young were produced during
the breeding season of 1933 in an area which supported an adult
population definitely known to total 31 potential pairs. If 40 per cent
of the 31 pairs, that is to say 12.40 pairs, successfully hatched out 9
young each (average number according to field observation) at the
first attempt, there would be produced a total of 111.6 young, lint of
the 18.60 unsuccessful pairs, most or all (for the sake of simplicity we
will here assume all) probably made a second attempt, which, btH&g
also 40 per cent successful, resulted in 7.44 additional pairs, producing
9 young each, that is to say 66.96 young at the second attempt. Tlic
total number of young hatched out would, according to the above calcu-
lations, amount to 178.56, which is as close to the number observed
in the field — 175 individuals — as one could reasonably expect. Third
attempts at nesting, as also the possibility that because of sterility or
similar reason the observed number of 34 males and 31 females failed
to make 31 pairs, have not been considered in the above calculations
because such factors would tend to counterbalance one another, and
because their inclusion would over-complicate the presentation of the
matter. Another inaccuracy is introduced by the fact that the average
figure of 9 young, used here, pertains to families seen in the field
from one to seven days after hatching, during which time, of course,
such family groups have already suffered a varying, undetermined
mortality. Furthermore, some young undoubtedly perished before
the census of the total quail population could be taken by trapping.

236 CALIFORNIA FISH AND GAME

However, these two factors also tend to cancel each other. Of course
the mortality rate will vary from year to year, even in the same locality,
due to climatic conditions, while in localities whose respective environ-
ments differ widely the discrepancy bet ween the mortality rates may
be still greater. Nevertheless, according to the figures given above,
the figure 60 per cent, which was assumed to represent nest failures
in this particular locality, seems to be substantiated.

This heavy loss of nests is compensated for under normal circum-
stances by the large number of eggs laid by quail, and is therefore not
in itself serious from the point of view of the game manager.

MORTALITY AFTER HATCHING

It is exceedingly difficult to follow the fortunes of any consid-
erable number of broods of small young in the field on account of the
extreme secretiveness of the parents during this period. For this
reason no exact figures are available to show the actual extent of
mortality of such small young. However, since the average number
of eggs laid by the California quail is about fourteen (Grinnell, Bryant
and Storer, 1918, p. 522), and since the average number of young per
family was observed to be only nine at the end of the first week, as
previously mentioned, the loss during this first week (disregarding the
possible failure to hatch, of any of the eggs of a set, which appears
to be comparatively rare) is seen to be %4 or 35.7 per cent. After
the first week, by which time the least fit individuals will have been
weeded out, the mortality rate doubtless decreases, so that the loss for
the first month may be slightly more than the figure of 39.6 per cent
estimated by Errington (1933b, p. 132) for the bobwhite at the end
of this period.

By late fall the now well disciplined coveys, made up of the rem-
nants of several family groups, are able to pass through the winter with
relatively small losses. The subject of mortality rate of quail after the
first month of life is treated more fully on page 246.

Elements. As regards the small young, the poor protection
against rain and fog offered by the natal down and the fatal effects of
heavy fogs and late rains upon such birds have already been pointed
out (page 218), as has also the fact that the breeding season of quail is
so timed as to escape such adverse conditions during normal years.
Accordingly, the mortality from this source must ordinarily be slight.

In the case of the adults in the Santa Cruz Mountain area, where
the winters are mild, with snowfall light and often wholly absent, only
insignificant losses are directly attributable to the elements; however,
in the northern parts of their range, and in the foothills of the Sierra
Nevada, considerable mortality is sometimes caused by heavy snows.

Of more serious consequence to the California quail than stormy
weather is a period of drought of a degree sufficient to check the
growth of green vegetation prior to the breeding season. If the winter
rains fail to produce an adequate supply of green forage, the resulting
food shortage (together with a possible vitamin deficiency) is reflected
in the poor physical condition of the birds, which may entirely prevent
them from breeding in those parts of the State most seriously affected.

CALIFORNIA FISH AND GAME 237

Thus, during the winters of 1931-1932 and 1932-1933, when the
rainfall was fairly adequate, quail increased noticeably throughout
the State. During the winter of 1933-1934, which was exceptionally
dry, with green food reduced to a fraction of the normal amount,
many flocks of quail in the arid parts of Los Angeles and San Diego
counties failed to break up into pairs, so that no young at all were
produced, while in other large areas also affected by drougth the crop
of young birds was far below the average.

Natural Enemies. Although the natural enemies of quail are
responsible for a slow, steady drain upon quail populations, this
mortality does not reach serious proportions as long as the food supply
of the birds remains adequate for maintenance of vigor. When the
food supply dwindles, however, especially during the winter season,
the birds become weakened and are easily captured by even the
slower predators ; moreover, the strong, vigorous birds are compelled
to forage farther from protective cover, so that they, too, fall prey to
enemies in greater numbers than normally. "When the surplus has
been removed, and the birds reduced once more to the carrying capacity
of the range, losses fall back to normal.

COOPER AND SHARP-SHINNED HAWKS

Cooper and sharp-shinned hawks, especially the former, are the
most destructive of all enemies of quail. If unsuccessful at the first
attempt to catch a quail, these hawks frequently perch motionless in
a nearby tree or bush, waiting for the covey to venture out into the
open again after it has recovered from its fright.

According to field observations, Cooper hawks can fly about as
fast as quail, but not much, if any, faster, so that if the latter have a
few feet head-start their chances of reaching safety are good. A hawk
relies upon surprising its quarry, as a rule, rather than in overtaking
it by sheer speed ; however, if cover is adequate, the wariness of the
covey is usually sufficient to enable it to escape, since it has the
advantage of its many pairs of eyes to keep watch for danger.

Both Cooper and sharp-shinned hawks are far more adept at
flying through thick cover than other hawks; indeed, the writer once
saw a sharp-shin fly faster and with more agility than any quail
through a dense stretch of woods which was almost impenetrable to a
human being, the bird holding its course only three feet above the
ground, where it encountered a minimum of obstructing branches.
Hawks of these species readily enter quail traps and are thus a
serious nuisance when banding activities are being carried on.

Continued harassment by enemies may have an adverse effect
upon game by preventing it from feeding, thereby weakening it. In
this connection it is worthy of note that the stomach of a quail, shot
by the writer from a covey which had been prevented by a Cooper
hawk from feeding just before going to roost, was practically empty.

On the other hand, the damage done to quail by these hawks in
localities where there is plenty of food and cover is not serious. In
the region covered by the present study, Cooper hawks were unsuc-
cessful in every attempt to catch adult quail that was observed by
the writer ; two three-quarters grown young were seen to be taken.

Male sharp-shinned hawks, because of their small size, appear
not to be able to capture adult quail; furthermore, they are on their
breeding grounds, which are for the most part north of the range of

238

CALIFORNIA FISH AND GAME
TABLE 5

STOMACH CONTENTS OF COOPER HAWKS (ACCIPITER COOPERI) TAKEN ON

QUAIL REFUGES

Date

Jan.

3,

1932

Nov.

30,

1932

Nov

30,

1932

Mar

26,

1933

Sept

20,

1933

Oct.

1.

1933

Oct.

7,

1933

Dec.

21,

1933

Dec.

28,

1933

Jan.

5,

1934

Jan.

12,

1934

Locality

Santa Cruz Mountain
study area

Santa Cruz Mountain
study area

Santa Cruz Mountain
study area

Santa Cruz Mountain
study area

Santa Cruz Mountain

study area

Near Lancaster, Los

Angeles County

Santa Cruz Mountain

study area

Near San Bernardino,

San Bernardino Co

Near San Bernardino,

San Bernardino Co

Near San Bernardino,
San Bernardino Co

Near San Bernardino,
San Bernardino Co.- .

Contents

Feathers of domestic
pigeon

Golden-crowned sparrow
(Zonotrichia coronata) . .

Meadowmouse (Microtus
californicus)

Steller jay (Cyanocitta
stelleri)

California quail, juv. male
2 lizards (Sceloporus sp.)_.

California quail

California quail

California quail

California shrike (Lanius
ludovicianus gambeli)

California quail

Remarks

Many killed by these
birds

Game-farm-raised
bird

Game-farm-raised
bird

STOMACH CONTENTS OF SHARP-SHINNED HAWKS (ACCIPITER VELOX)

TAKEN ON QUAIL REFUGES

Oct. 10, 1933

Near San Bernardino,
San Bernardino Co

Near San Bernardino,
San Bernardino Co

Near San Bernardino.
San Bernardino Co...

California quaiL . . _.

Game-farm-raised

Nov. 14, 1933

California quail --

bird
Game-f arm-raised

Nov. 27, 1933

California quail .

bird
Game-farm-raised

bird

the California quail, during the season when the young of the latter
would be vulnerable to their attacks.

RED-TAILED HAWK

The unimportance of the red-tailed hawk as an enemy of quail,
as well as the benefits resulting from its presence, will be pointed out
in Part II. In the Santa Cruz Mountain area, where red-tails were
abundant, none was ever seen trying to catch quail, nor would they
have been able to, under ordinary circumstances, on account of the
density of the cover. In parts of southern California, where cover

CALIFORNIA FISH AND GAME

239

was markedly inadequate, red-tailed hawks were occasionally observed
to catch inexperienced, game-farm-raised birds, as indicated in Table 6 ;
but such occurrences were obviously not representative of the normal
relations between the two species.

TABLE 6
STOMACH CONTENTS OF RED-TAILED HAWKS (BUTEO BOREALIS CALURUS)

TAKEN ON QUAIL REFUGES

Date

March, 1933

April, 1933
Oct. 24. 1933

Nov. 14, 1933

Dec. 10, 1933
Dec. 15, 1933

Locality

Near San Bernardino,
San Bernardino Co..

Near Lancaster, Los
Angeles Co

Near Lancaster, Los
Angeles Co

Near San Bernardino,
San Bernardino Co..

Near San Bernardino,
San Bernardino Co._

Near San Bernardino,
San Bernardino Co._

Content -

California quail

G round squirrel

Pocket gopher (Thomomys
s\>.) '

Cottontail rabbit (Sylvil-
agus auduboni sancti-
diegi)

Meadow mouse

Sparrow (Zonotrichia sp.?)
Meadow mouse

Remarks

Game-farm-raised
bird

OTHER HAWKS

For a general consideration of the relation of other hawks to quail,
see Part II. In addition, the following records of stomach contents
were obtained from quail refuges :

Marsh hawk (Circus hudsonius) , collected October, 1933 near Lan-
caster, California — one meadow mouse; Ferruginous roughleg (Buteo
regcdis), collected December 26, 1933 near San Bernardino, Califor-
nia— one ground squirrel.

HORNED OWL

Horned owls appear to do no serious damage to quail in areas
where roosting cover is abundant. In regions where roosting cover is
scanty, horned owls are sometimes important enemies of quail, but this
circumstance is primarily due to a faulty environment. The, writer
has many times observed coveys of quail feeding at dusk in open places,
such as barley fields, while several horned owls were booming near-by
and some were even visible in the tops of the trees; nevertheless, no
attempt was made by these owls to catch the quail, probably because,
like red-tailed hawks, horned owls are no match for quail as regards
speed of flight during the hours of daylight. Probably mosl of the
quail which fall prey to horned owls do so as a result of being flushed
from their roosts, in some cases, perhaps, by another enemy.

OTHER OWLS

For a general consideration of the relations of other owls to quail,
see Part II. In addition, the following records of stomach contents
were obtained from quail refuges :

Burrowing owl (Speotyto cunicalaria kypogaca) , collected Novem-
ber, 1933, near San Bernardino, California — finely triturated insect

240

CALIFORNIA FISH AND GAME
TABLE 7

STOMACH CONTENTS OF HORNED OWLS (BUBO VIRGINIANUS PACIFICUS)

TAKEN ON QUAIL REFUGES

Date

Locality

Contents

Remarks

Oct. 1, 1933

Near Lancaster, Los

Angeles Co . -

Pocket mouse (Perog?ia-

thus sp.) scorpion (Scor-
pionida)

Oct. 2, 1933

Near Lancaster, Los

Angeles Co. _

Cottontail rabbit (Sylvila-

gus sp.), fragment of

snake (sp?) -

Oct. 7, 1933

Near San Bernardino,

San Bernardino Co...

2 scorpions, 1 Jerusalem

Near San Bernardino,

cricket {Stenopalmatus
sp.)

Oct. 28, 1933

San Bernardino Co —

Fragments of cricket

Nov. 4, 1933

Near San Bernardino,

San Bernardino Co...

Fragments of cricket or
grasshopper - ._

Nov. 7, 1933

Near San Bernardino,

San Bernardino Co...

Fragments of four crickets

Nov. 10, 1933

Near San Bernardino,

San Bernardino Co.-_

California quail

Boosting cover

fragment of mouse (sp?)

inadequate

Nov. 18, 1933

Near Lancaster, Los

Angeles Co ..

Jerusalem cricket, horned
lark (Otocoris alpestris)-

Dec. 6, 1933

Near San Bernardino,

San Bernardino Co...

Fragments of 2 crickets (?)

Dec. 7, 1933

Near San Bernardino,

San Bernardino Co

Fragment of cricket, rod-
ent hair (mouse?)

Dec. 7, 1933

Near San Bernardino,

San Bernardino Co

Fragments of crickets,
brush rabbit (Sylvila-

remains, many of beetles; Short-eared owl (Asio flammeus), collected
December 14, 1933, near San Bernardino, California — feathers of spar-
row (sp. ?) ; Long-eared owl (Asio ivilsonianus) , collected November
28, 1933, near San Bernardino, California — kangaroo rat (Dipodo-
mys sp.).

JAYS

Where cover is adequate jays appear to do no significant damage
to quail. Moreover, the California jay (Aphelocoma calif ornica) is of
some benefit to quail by reason of its habit of uttering a certain loud,
sharp cha! note at the approach of a Cooper or sharp-shinned hawk.
Usually, although not always, quail (and also sparrows) dive precipi-
tately into cover upon hearing this noise; however, the jays often fail
to give this note in the presence of the hawk, so that, in view of the
wariness of the quail themselves, the importance of jays to the latter in
this respect is not vital. Out of six cases in which jays were known to
have observed the approach of a Cooper hawk, the birds gave the warn-
ing note on four occasions, remaining silent two times.

CALIFORNIA FISH AND GAME

241

SNAKES

The relation of snakes to quail is discussed as completely as the
facts at hand permit in Part II. Mr. Allan McKean, formerly Research
Assistant for the Division of Fish and Game, informed the writer that
he had watched the actions of a king snake which was trying to capture
some young quail. Contrary to popular legend, the adult quail were
not at all ''hypnotized" by the snake, but ran about in front of it,
the female trying to decoy it away as she fluttered on the ground with
trailing wings.

The snake followed the adults for only a few feet and then returned
to the place where the baby quail were, but could not locate any. Pres-
ently the adults called their young over to a thicket one hundred feet
distant. For a considerable time after this, the snake remained where
it was, but just at dusk it again moved over to where the family of
quail was, as though guided by the sounds (?) made by the birds.
When dusk fell McKean collected the snake and found that its stomach
Avas empty. He believed that as a result of the activities of the reptile
some of the young had become scattered and lost, which at their age
could only prove fatal.

HOUSE CATS

Feral house cats are commonly regarded as excessively destructive
to bird life, although published records of stomach contents are rare.
The following records from the Santa Cruz mountain study area,
although admittedly meager, indicate that, like other predators, house
cats prey upon whatever is most abundant and easiest to capture — in
which category quail are not to be classed :

December 22, 1932, 2 meadow mice, 1 brush rabbit; May 16, 1933,
1 one-fourth grown brush rabbit; September 1, 1933, 1 white-footed
mouse.

BOBCATS

The effect of bobcats upon quail is discussed in Part II.
examinations bear out the conclusions reached, as follows :

Stomach

TABLE 8

STOMACH CONTENTS OF BOBCATS (LYNX RUFUS CALIFORNICUS)
TAKEN ON QUAIL REFUGES

Date

Jan. 25, 1933

Mar. 13, 1933

Feb. 8, 1933

Oct. 7, 1933

Nov. 29, 1933
Dec. 1933
Mar. 10, 1934

Mar. 12, 1934

Locality

Near San Bernardino, San Ber-
nardino Co

Near San Bernardino, San Ber-
nardino Co "-

Near San Bernardino, San Ber-
nardino Co

Near San Bernardino, San Ber-
nardino Co

Santa Cruz Mountain study area
Santa Cruz Mountain study area
Near Lancaster, Los Angeles Co..

Near Lancaster, Los Angeles Co..

Contents

Jack rabbit (Lepus californicus)

Fur of rabbit (Sylvilagus sp?)

Fur of rabbit (Sylvilagus sp.?)

Western tanager (Piranga ladoviciana)
Wood rat

Brush rabbit (Sylvilagus bachmani)
Parts of jack rabbit and cottontail rab-
bit
Fur of rabbit (Sylvilagus sp?)

212

CALIFORNIA FISH AND GAME

Mr. McKean saw a bobcat spring at a quail and miss it — an obser-
vation which has been duplicated by other field workers.

COYOTES AND OTHER PREDATORY MAMMALS

That coyotes and other predatory mammals play no significant
role in the destruction of quail is demonstrated in Part II. The results
of stomach examinations are as follows :

TABLE 9

STOMACH CONTENTS OF COYOTES (CANIS LATRANS OCHROPUS) AND
OTHER PREDATORY MAMMALS TAKEN ON QUAIL REFUGES

* —

fecal material

Date

Locality

Contents

Remarks

Feb. 1933*

Co>
Santa Cruz Mountain
study area

rotes

2 brush rabbits, 2 wood
rats, meadow mouse,
white footed mouse

Mar. 1933
Mar. 1933

Near San Bernardino,
San Bernardino Co.._

Near San Bernardino,
San Bernardino Co

Near Lancaster, Los
Angeles Co._ __

(P crom y s c u s s p. ) ,
uround squirrel.

Cottontail rabbit
Deer belly hair

Depositions of six
individuals

Carrion?

Oct. 7, 1933

1 ground squirrel (entire) ,
foot of jackrabbit.

Brush rabbit, wood rat,

Oct. 8, 1933*

Santa Cruz Mountain
study area -

Nov. 2, 19:;:;

Near Lancaster,' Los
Angeles Co.

ground squirrel, mouse
fragments (Pcromyscns
or Reithrodontomys)

Ground squirrel _ -

Depositions of three
individuals

Nov. 25, 1933

Near Lancaster, Los
Angeles Co.

Nov. 25, 1933

Near Lancaster, Los
Angeles Co -_

White footed mouse, wood
rat, ground squirrel

Fur of rabbit (sp.?)

White footed mouse,
ground squirrel frag-
ments, kangaroo rat
• fragments

Dec. 2, 1933
Feb. 7, 1934

Near Lancaster, Los
Angeles Co. __ - --

Near Lancaster, Los
Angeles Co

Near Lancaster, Los

Angeles Co..
Near Lancaster, Los

Angeles Co.

Feb. 27, 1934
Mar. 1, 1934

Half-grown brush rabbit_.
Fur of rabbit (sp.?)

Mar. 11, 1933

Near San Bernardino,
San Bernardino Co.

Badger (Taxidea taxus)

Fur of rabbit (sp.?).

CALIFORNIA FISH AND GAME

243

RODENTS

The relation of various rodents to quail nest destruction should be
more fully investigated, particularly in the case of the wood rat (Neo-
toma fuscipcs), which is exceedingly abundant in quail territory in the
Santa Cruz Mountain area.

That the meadow mouse (Microtiis calif ornicus) is not an enemy
of quail is indicated by the fact that quail eggs were left for weeks,
without ever being disturbed, near runways along which these mice
were actively foraging.

The role of the ground squirrel in the ecology of quail is discussed
in Part II.

ACCIDENTS

A surprisingly large number of deaths occur among game birds
as a result of accidents of various sorts, chief of which are collisions
with objects during flight. Errington (1933c, p. 32) believes "that
mechanical accident is a mortality cause much under-estimated by
wild life students," and that "in the aggregate perhaps [such acci-
dents] constitute a factor as lethal as any in the life equation of the
quail." The present writer is also inclined toward this opinion, to
which Table 10 lends support,

TABLE 10
ACCIDENTS TO QUAIL RECORDED DURING THE PRESENT INVESTIGATION

A Kent

Moving automobile

Fence

Tree trunk

Unknown, possibly fence
Water tank

Cactus thorn in leg

Number of
instances

small
young)

Circumstances

Birds flushed from
roadsides while
feeding

Collision while in full
flight

Collision while in full
flight

Unknown

Unknown

Prevented bird from
keeping up with
the others

Result

Three birds killed; one
uninjured?

Six birds killed: one
knocked to ground, pos-
sibly surviving ; one
(•■everely wounded but
recovered

Bird knocked to ground,

probably survived
Bird killed
Bird drowned

Removed by the observer,
Mr. Allan McKean

DISEASE

No signs of disease were noted among the quail of the Santa Cruz
Mountains, nor is it probable that an outbreak would have been over-
looked if it had occurred, especially during the year of intensive
trapping when the same birds were handled over and over again.

244 CALIFORNIA FISH AND GAME

On the other hand, quite the contrary condition was discovered in
the quail population of the 140-acre campus of Mills College, Oakland,
California, during the late winter months of 1932-1933. On March 20,
1933, Mrs. E. Lowell Sumner, Jr., found a sick quail standing in a
corner outside the Mills College science building. The bird was too
feeble to escape, and died twenty minutes after its discovery. It was
sent to Dr. M. Hobmaier, Pathologist of the Division of Fish and Game,
for examination, who, however, was unable to find "signs of any disease
as yet known on quail."

Subsequent to the discovery of this sick bird, the carcasses of four
others, evidently at least one month old, were found within a few hun-
dred feet of the same spot, while three additional recently-dead quail
were reported during the preceding month. This brought the total
known number of dead birds up to eight.

Four more quail were trapped from the spot where the dead birds
had been found, and were sent alive to Dr. Hobmaier, but no abnormal
condition was discovered, which indicates that the epidemic, whatever
its nature, had subsided. Although large numbers of the birds still
inhabited the region, no more dead quail were found.

PARASITES

No mortality associated with parasites was observed in the quail
of the Santa Cruz Mountains, in spite of the fact that between July
29 and October 7, 1933, at least one out of every five birds harbored
a hippoboscid fly, identified by Dr. F. C. Bishopp, United. States
Bureau of Entomology, as Lynchia hirsuta, which is known to transmit
the important blood parasite Haemoproteus lophortyx of quail
(O'Koke, 1932, p. 224). At no time other than the season mentioned
were hippoboscid flies found on quail, although the banding program,
which involved handling large numbers of birds, would have revealed
them had they been present.

POPULATION STUDIES

NUMBER OF QUAIL PER ACRE

From February 1, 1933, to February 1, 1934, as previously men-
tioned, intensive trapping operations were carried on over an area
of approximately sixty acres. The area was particularly suitable for
this work because of its natural boundaries, which reduced to an almost
negligible quantity the birds' chances of wandering into other regions
(Fig. 13).

The problem of expressing in a significant fashion the number of
quail to the acre on a given range is more difficult than might at first
appear, even wiien the number of resident birds as well as the total
land area is known. For example, in the case of the area shown in
Fig. 13, should we include in our calculations all of the heavily wooded,
deep ravines shown, which extend in certain directions for miles beyond
the limited area included by the map and which act as barriers to the
dispersal of the birds? If we do include all this wooded land, which
can never become quail environment, then our figure for number of
birds per acre must always remain almost vanishingly small, no matter

CALIFORNIA FISH AND GAME

245

bow much we may increase the number of quail locally, in the open parts
of the region, by practicing game management.

A more clearcut example would be furnished by a flock of 500 quail
which are restricted to a carefully managed 500-acre estate surrounded
by a forest comprising 50,000 acres ; should we consider only the region
occupied by the estate, together with a narrow strip of adjacent
habitable woodland and estimate the density at about one bird per acre,
which would be a high concentration, or should we include the whole
forest and consider the density as approximately one bird per 100
acres, which would be a very low concentration? In the opinion of the
writer the former figure would be more significant and therefore pre-
ferable, because the wooded portion of the area would constitute almosl
as effective a natural barrier to the birds as would a deserl or an

1 Ld-nd *^2 [_,,

Land

Fig. 13. Area comprising GO acres of territory habitable for quail where an inten-
sive banding program was carried on for more than a year. The chief head-
quarters of each covey is shown.

ocean — yet no one would include the acreage of an ocean along with
that of a game preserve which bordered upon it.

Accordingly, in the present paper, only the outer margin of forest
land, approximately 100 feet in width, has been included in the calcula-
tions. Brush land has been included to the full amount of the acreage,
in order not unduly to complicate the issue. Similarly, Nicholson
(1934, p. 145) has confined his estimates of population densities to the
areas actually inhabited, or habitable, by the animals in question.

Large open areas such as bare valley floors would also represent a
type essentially uninhabitable for quail and therefore not to be con-
sidered in bird-per-acre calculations, but in the case of this study no
such areas were present to interfere with estimates of quail per acre.

246

CALIFORNIA FISH AND GAME

In the area shown in Pig. 13, approximately 20 pairs out of a total
oL" 64 resident adults (representing a potential of 31 pairs) were suc-
cessful in raising 175 young to an age of between one and seven days;
in other words, 64 per cent of the total of 31 pairs were ultimately
successful. This figure must not be confused with the figure of 60 per
cent given on page 235, which represents the proportions of nest fail-
ures; in other words, by making a second attempt after the destruction
of the first nest, the species is able to increase its chances of success
from 40 per cent to 64 per cent.

Fig. 14 shows a considerable differential mortality between adults
and young during the months following hatching, which is in accord
with the presumption that young fall prey to enemies in much larger
numbers than adults because of the inexperience of the former.

Feb.

M&r.

Apr.

MdY

Jlme.

Ju-ly

Aug,

6ePt.

Oct.

Nov.

Dec.

Tin.

??0

rS

yon

IftO

160

V

IUO

1

\

\

170

\

\

100

^

\

Jui.\/e.rv

iles

x^_

Tote.1

"*"■**-

fto

f.0

MO

im&.tu.r

cs

20

Adults

— ••*«■»,

— -"*"

^^^

Fig. 14. Diagram showing population fluctuations during one year among quail
residing in the 60-acre territory shown in Fig. 13. Actual numbers of birds
at left.

It will be observed that, due to a combination of favorable circum-
stances, chief of which were freedom from overgrazing, an abundant
food supply, and no hunting, the quail population in the area studied
increased from 105 on February 1, 1933, to 130 on February 1, 1934,
although no predatory animal control was carried on. From Figs. 13
and 14 it can be calculated that, in accordance with the season of the
year, the number of quail varied from 1.1 to 3.9 per acre of habitable
territory.

ESTIMATES OF SH00TABLE SURPLUS

Obviously, estimates of shootable surplus in quail can only be made
upon the basis of local conditions, for the factors governing quail
abundance may vary widely within the space of a few miles. Similarly,
conditions in a given locality may differ from one year to another as a
result of weather differences or changes in human activities.

Bryant (1912, p. 138) has expressed the opinion that a certain
proportion of quail may be taken by man during the hunting season,
provided that this number does not exceed the number of birds which
would have met death during this period as a result of natural forces.

CALIFORNIA FISH AND GAME 247

It is the belief of the present writer that the number of birds harvested
may somewhat exceed the expected casualty rate for the relatively short
period of time represented by the shooting season, and that under favor-
able conditions the shootable surplus may include all birds which would
normally have died by the end of February. For example, in the case
of the quail in the area shown in Fig. 13, the number of birds dropped,
due to natural enemies and related causes, from 139 to approximately
113 between October 30 and February 28, which is a loss of 19 per
cent. In view of the fact that the quail population of this locality
showed a net increase for the year of 25 birds (Fig. 14), in spite of this
loss, there is hardly any doubt that most of the surplus of 19 per cent or
26 birds could have been taken from the area by hunters without detri-
ment, it being understood that the toll levied by natural forces would
automatically shrink below the normal figures as soon as the surplus
birds had been removed by hunters.

From Fig. 14 it can be seen that if the percentage of young birds
taken is consistently less than about 71 per cent of the total bag, it is
an indication that the breeding season was not successful, in which case
hunting should be discontinued in the region affected, in order not to
deplete the breeding stock. For example, during 1934, quail popula-
tions in drought-stricken areas such as Los Angeles and San Diego
counties, where few or no young were produced, should not be hunted
(unless supplemented by artificially liberated birds such as those raised
on game farms).

A method of distinguishing young of the year from fully adult
birds is described on page 225.

LONGEVITY

From Fig. 14, it can be calculated (after subtracting the net
increase for the year, 25 birds) that, out of 100 quail raised in a
given summer, 26.8% will be alive at the end of the first year, 7.2%
at the end of the second, 2.0% of the birds at the end of the third year,
and .5% at the end of the fourth. From this, the average life expecta-
tion is seen to be about eight months.

These longevity figures are in general agreement with supplemcn
tary survival figures obtained from 35 additional banded quail, as fol
lows: 1-llyears, 26.8% ; l|-2 years, 12.8% ; 2-2^ years, 7.35% ; 3.2 years,
1.05% ; 3.5 years and still alive, .7%.

SEX RATIO

As indicated in Table 11, there is a greater number of males than
females at all seasons, the average ratio being 53.2 to 46.8. In the
present study, the data were obtained from birds of all ages except
juveniles less than one-third grown. How far back toward hatching
time this inequality of the sex ratio extends is not known; at least it
is certain that the unequal ratio is not due to any difference between
the sexes in their readiness to enter the traps, since all the birds of each
covey were known to have been captured.

6 — 23006

248

CALIFORNIA FISH AND GAME

TABLE 11
SEX RATIO IN A POPULATION OF 280 CALIFORNIA QUAIL DURING EIGHTEEN

CONSECUTIVE MONTHS

Males

Females

Date

Number of
individuals

Percent

Number of
individuals

Perron t

Jan. 1-15

126
122

57
50

50

48

45
42

41
41

38
35

184

180

177
177

177
166

150
135

128
128

128
128

52
52

54
55

55
54

53
53

53

54

54
53

53
53

53
53

53

54

54

53

52
52

52
52

114
113

48
41

41
41

40
37

36
34

32
31

160

156

151
151

151
141

126
118

116
116

116
116

48

16-31-. .

48

Fob. 1-15 - .

46

16-28 -

45

Mar. 1-15 ._ .

45

16-31 -

46

April 1-15

47

16-30

47

May 1-15. .

47

16-31.

40

June 1-15. - - ... . .

40

16-30 .

47

July 1-15

47

16-31 - .

47

Aug. 1-15.

47

16-31.

47

Sept. 1-15. -.

47

16-30..

46

Oct. 1-15 - .

46

16-31-.

47

Nov. 1-15. .

48

16-30. -. ...

48

Dec. 1-15 ...

48

16-31

48

WEIGHT CHANGES

Seasonal Weight Fluctuations of Adults. As shown in Fig. 15,
the adult male weighs most during the winter and early spring months,

Oct. Nov. Dec. Jan. Feb. M&r. Apr. Ma-y June. TuAy flag. Sept.

110

loo

190

ieo

HO

160
l&O
114O

Fio. 15. Weight fluctuations throughout the year shown within a population of 280

quail ; 652 weighings are represented.

CALIFORNIA FISH AND GAME

240

while the marked decrease in weight suffered during the breeding season
is almost certainly caused by the irregular feeding habits and high
state of nervous excitement resulting first from pairing activities and
later from the prolonged, arduous period of sentry duty. It may be
significant in this connection that a nonbreeding male in the large
inclosure weighed 198.2 gr. during the period when the average weighl
for breeding males was 189.4.

In the case of the female, which takes no active part in the strenu-
ous activities of courtship and is relieved of all sentry duty, no loss of
weight takes place until after the hatching of the young in July.
Indeed, between mid-April and mid-July she is actually heavier than
the male, although the reverse of this condition holds during the remain-
der of the year.

Immature birds of both sexes, since they breed at the age of one
year, show weight fluctuations similar to those of the older birds. How-
ever, since they do not reach full adult weight until the beginning of
the second year, their variations in weight follow, at a lower level,
those of the older birds.

TABLE 12

AVERAGE WEIGHTS OF VARIOUS RACES AND LOCAL POPULATIONS OF QUAIL

(Lophortyx californica)

Race

Locality

Nearest

parallel of

latitude

Average

weight

Number
weighed

L. c. californica

San Mateo Co., Calif.

'Tehama Co., Calif.

37°60'

40°

37°00'

36°60'

34°

32°60'

31°

28°-24°

189.5
180.0
100.7
185.0
159.3
151.0
157.7
150.0

052

11

L. c. californica

L. c. californica _ _

L. c. californica _ _

L. c. californica

L. c. plumbea-

2Mariposa Co., Calif

^Monterey Co., Calif. . _
Los Angeles Co.. Calif.
Lower California
2Lower California

4

7
29

8
57

L. c. achrustera . - - .

2Lower California __ _. _

32

'Grinnell, Dixon, and Linsdale, 1930, p. 208.

sWeights obtained from labels of specimens in collection of Museum of Vertebrate
Zoology.

Following the mating season, the period of molt continues the drain
upon the vitality of the birds, so that it is only after this has been com-
pleted in September that they commence to put on the weighl which
enables them successfully to endure the rigors of winter.

Daily Weight Fluctuations. Few data on daily weight fluctua-
tions were secured, due to the fact that nearly all birds trapped or
shot were taken within about two hours before roosting. Table 3 shows
the difference between late evening and early morning weights in the
case of 94 birds which were kept over night; the average loss during this
period was 5.86 per cent.

Geographic and Subspecific Weight Differences. The California
quail in its various races, and even within a given race, conforms
reasonably closely to "Bergmann's Law" (Rensch, 1929, p. 133),

250 CALIFORNIA FISH AND GAME

which states that within a given species, whose races are distinguishable
as to body size, small size is correlated with warm climate and large
size with cold climate (Table 12). Bergmann's explanation of this cor-
relation is that with increase in volume of a body, from 1 to 8, for
example, the outer surface increases only from 1 to 4; in other words,
a large animal has less surface in proportion to its size exposed to a
cold environment than does a small one, and as a result is better pro-
tected against heat loss through radiation.

In view of this circumstance, the undesirability of importing light-
weight, small-bodied races of quail from southern climates into northern
regions is apparent.

PLUMAGES

Time has not permitted the working out of molt sequences, so
that the present observations, culled from the writer's field notes, are
fragmentary.

The approximate ages at which the young undergo the transition
from natal down through the precociously developed juvenal plumage
to the first fall plumage have been given on pages 222-225. One curious
departure from the normal relation of spotted to unspotted primary
coverts (Fig. 9) was observed in a youngster which had the two distal-
most coverts mottled, instead of solid black as normally, while the two
coverts next to these were solid black instead of mottled — a condition
exactly the reverse of normal as regards these four feathers. Both
wings displayed this abnormality. A somewhat intermediate condition
was found once, in which the distalmost covert was dark, as in the
normal arrangement, but the next one to it, although also normally
dark, Avas in this case lightly mottled, like the remaining primary
coverts.

The fall molt commences about the first week of August, for both
juveniles and adults, and involves the head and neck first. From this
point it spreads posteriorly over the body, and is accompanied by the
replacement of the primary coverts. During September, molting is
at its height ; the birds present a ragged, untidy appearance, and- in
many cases the head plumes of both old and young are entirely absent.
By the middle of October the body molt is nearly complete, and the
primaries are also being replaced, while by the last week of November
only two or three pinfeathers can be found on the entire body, and the
birds are resplendent in their new plumage.

The head plumes of quail assume various positions according to
the activities, and doubtless the emotions, of the birds. This applies to
both sexes, and even to the small young, which have an abbreviated
topknot by the tenth day at least, if not from birth. When the birds
are relaxed — either wandering about aimlessly, or dozing — the plume
droops down forward, fairly overhanging the bill in the case of adult
males.

With the assumption of an alert position, however, as in the case
of a dozing bird which has been startled, the plume snaps up to a ver-
tical position. Males which are fighting depress the plume so far back-
ward that it lies almost parallel with the top of the head, as though
to avoid being injured during the scuffle. In feeding, the plume is

CALIFORNIA FISH AND GAME 251

held high- --up and someAvhat backward so that it can not touch the
ground. A similar position is also adopted when the birds are fright-
ened.

SUMMARY

FOOD HABITS AND REQUIREMENTS

The pea family provides a larger and more varied food supply
than does any other plant family (32.47 per cent of total), with bur
clover ranking as the No. 1 quail food throughout large parts of Cali-
fornia. Grasses are next in importance to the legumes as a source of
quail food (14.4 per cent of total). Damage to grain crops by quail
is relatively slight.

Other important foods are the filarees, thistles, buttercups, chick-
weeds, and docks. A great variety of food is eaten, so that very few
plants rank as major food items in the diet. A large number of impor-
tant foods now habitually depended upon by quail were not originally
native to California.

Heavy grazing results in an inferior type of quail food in addi-
tion to causing an actual food shortage.

Trees and shrubs constitute a relatively unimportant source of
food for quail, and their fruits are inferior to weed seed in nutritive
quality. Galls are eaten in considerable numbers.

Green leaf fragments comprise an extremely important food item
for quail and when obtainable render the birds independent of free
water. The seasonal fluctuation in consumption of such material runs
from 1.6 per cent of the total food for October, to 85.49 per cent in
April-May. Animal food, chiefly ants, is unimportant (.38 per cent of
total) in the diet of adult quail, although it figures more prominently
in the food of the young.

Feeding is carried on with some vigor in the early morning and
then intermittently until about 4 p.m. After this hour it increases
and reaches a climax of last-minute gorging just before the birds fly
up to roost. During the breeding season, as a result of sentry duty by
the male and incubation by the female, this schedule is largely upset.

The distance traveled by a quail covey for food each day is slight
as long as food holds out; with the decrease in available food the dis-
tance traveled per day increases. Feeding mannerisms such as scratch-
ing and jumping for food are described.

Adult quail eat about 24 grams of food per day (14.3 per cent of
their own weight), which amounts to 11310.7 cc. (.321 bushel; 10.27
qts.) per year. Considerable preference is exercised by quail as to the
food eaten. Some plant families furnishing abundant seed are avoided
entirely ; the members of some of these families are strong-scented and
perhaps distastefulf possibly others produce toxic effects.

New foods, especially seeds of large size, are usually rejected--a#
first, although they may be accepted later as a result of a more or less
protracted learning process.

In the case of seeds which are picked up singly, the upper limit
of size eaten by California quail is about £cc. ; the lower limit is about

252 CALIFORNIA FISH AND GAME

%43o cc. Certain seeds below this minimum of size are eaten in large
quantities while still inclosed in their fruit walls. Grit particles are
eaten less abundantly when hard seeds form a large percentage of the
food. Very minute grit particles are probably ingested accidentally.

WATER REQUIREMENTS

Free water is not required during the rainy season because it is
furnished by the succulent green vegetation. When the green vegeta-
tion disappears, however, water must be obtained from pools, springs,
or similar sources.

ECOLOGICAL REQUIREMENTS

Food must be present in adequate quantity within about fifty feet
of cover. Cover is of relatively little use unless well interspersed with
food, and it must include trees or dense bushes for roosting.

The California quail is limited topographically to the lower alti-
tudes, those below 4000 to 8500 feet, possibly because of its inability to
withstand the rigors of winter above these heights. Geographically
it is limited on the west by the Pacific Ocean ; on the east by the desert
regions, where the water supply is inadequate to meet its needs. To the
north it is limited in its distribution by the regions of heavy snowfall
lying north of Klamath Lake in southern Oregon. Southward it extends
to the extreme tip of the Lower California peninsula. Within its range
it is a highly successful species because of its ability to maintain itself
in regions differing widely as to food and cover types, seasonal rainfall,
and temperature range.

EFFECT OF HUMAN ACTIVITIES

Hunting is only one of several important factors in quail decrease.
It will be possible to maintain this sport on an anywhere nearly satis-
factory plane in the future only by the observance of a system of game
management.

Primitive farming methods in vogue during the early days of
settlement were somewhat favorable to quail, but the modern system
of "clean farming" has greatly reduced their numbers in certain areas.
The exclusive use by man of water sources has brought about the
virtual extinction of quail coveys in parts of the arid southwest. Irri-
gation may be locally favorable to quail, but since it is usually obtained
at the expense of surface or subsurface water supplies in other regions,
the net effect is detrimental.

Overgrazing is one of the most important causes of quail decrease
on many range lands because of the resulting destruction of available
food, and possibly because of the depletion of the mineral supply.

Predator control as practiced by private individuals has done
more harm than good to quail ; as practiced by government agencies it
has been largely neutral in results as far as quail are concerned.
Rodent control, when involving the use of thallium, has been destructive
to this species,

CALIFORNIA PISH AND GAME 253

Lumbering- has tended to increase the numbers of quail by creating
more "edge" environment. Fire control has probably had an unfavor-
able influence on an average by promoting the growth of impenetrable
stands of tall brush, with the. consequent choking out of food plants.

Roadways have in some cases furnished food and shelter not other-
wise available, but their net effect has been distinctly harmful because
they have enabled hunters to reach the most remote strongholds of
the birds.

"Protection" has consisted of (1) laws, which have reached
their highest development short of a continuous closed season; (2)
predator control, largely futile as so far practiced; (3) the establish-
ment of refuges, which, although representing a step in the right
direction, have realized only a fraction of their potentialities because
of the lack of a system of game management.

CALLS

The calls of quail are varied but the significance of a given call
is rather constant; ten distinct types are described.

SOCIAL ORGANIZATION

The California quail is strongly gregarious; many of the reactions
of the birds are cooperative, although instinctive. The instinct to
imitate other quail is present from the first, and aids the inexperienced
young in acquiring the wariness of the adults ; it continues to serve
as the basis for mass action throughout life.

Leadership within the flock is determined by chance. The larger
the flock, the greater the number of potential leaders, which tends to
prevent the permanent association of large, aggregations of quail.

Sentry duty is performed by the adult male, and by him alone,
from the time incubation commences until several weeks after the
young have attained nearly full size and the family group has merged
with other groups to form a covey in late autumn.

COVEY FORMATION AND RANGE

Coveys which have been reduced by enemies to less than ten birds
usually unite with other coveys, so that in a given locality covey size
is maintained, even though the number of coveys dwindles. Splitting,
recombining, and interchanging of individuals readily occur among
coveys whose ranges intersect or adjoin, rendering the chances of
inbreeding remote.

The distance traveled daily by a covey varies from about 100 feet
to about a quarter of a mile, depending upon the availability of food.
No definite route is taken, the flock moving by impulse. During the
reproductive season the range of the individual pairs is even more
circumscribed.

Although the individual members of a covey may scatter widely
during the breeding season, often to join some other covey in the fall,
the covey as such usually remains permanent since its size, and its
location is governed by the available food and shelter rather than by
the fortuitous presence of an}- given individual.

254 CALIFORNIA FISH AND GAME

COURTSHIP BEHAVIOR

The pugnacious stage of the males occurs in early spring before
the covey has split up. Successful males attach themselves to their
prospective mates and keep all other males at a distance.

Following pairing an impulse to wander considerable distances in
search of a nest site results in the gradual dissolution of the covey.
Male birds which have been unsuccessful in securing mates set up
territory and give the cow! note, which means announcement that they
are ready to pair. Successful males are silent and lose most of their
pugnacity.

NESTING HABITS

No male quail were observed to share in the function of incuba-
tion. The female is guarded by the male during her recess periods
away from the nest. No indication was observed of a pair raising two
broods in a season.

CARE OF YOUNG

The small young, which are highly susceptible to chilling, are
brooded on the ground by the female from about 4.30 p.m. until about
9.00 of the following morning, or even much later if the day is cold or
damp. The solicitude displayed by the parents for their young,
although well developed, is instinctive and unreasoned ; it is expressed
largely by vocal warning or encouragement, and by active defense
against various enemies, but does not extend to physical aid of the
young if they are for any reason unable to keep up with the family
group. Greater solicitude is shown by the male than by the female.

GROWTH AND BEHAVIOR OF YOUNG

Behavior at time of hatching, initial fear and feeding reactions,
play activities, voice development, and weight increase are described.
Wariness is learned from the parents by imitation of their behavior ;
however, the process is only gradual, so that many young perish as a
result of inexperience. Birds of the year may be distinguished until
their second fall molt from adults by the presence of mottled primary
coverts.

GENERAL BEHAVIOR AND MISCELLANEOUS HABITS OF QUAIL

Roosting behavior is described. Trees or tall bushes with dense
foliage are preferred as roosting sites, but in certain semidesert regions
where such vegetation is largely absent, the birds are forced to roost
in low, open brush, which renders them more than usually vulnerable
to owls. Instances of ground roosting are rare. Power of vision at
night is poor. The hour of roosting and of awakening is determined
within remarkably close limits by light intensity rather than by an
absolute time unit; accordingly, the duration of roosting varies from
8 hours 18 minutes in midsummer to 13 hours 33 minutes in midwinter.

The warm hours of the forenoon and early afternoon are usually
spent in resting or dust bathing.

CALIFORNIA FISH AND GAME 255

The wariness of quail results from an innate timidity and nervous-
ness, together with a retentive memory for fear experiences, rather
than from any unusual mental development.

Hearing is relatively unimportant as a means of detecting danger.

MORTALITY

Nest failures in the region studied amounted to about 60 per cent
of all nests, irrespective of whether they represented first, second, or
third attempts. Mortality of young during the first week of life
appeared to be approximately 21.6 per cent; for the first month prob-
ably about 39.6 per cent.

Stormy weather is usually an unimportant cause of mortality, but
extreme drought, by curtailing or entirely preventing breeding, is
occasionally serious in its results.

Natural enemies constitute a slow drain on quail, but mortality
from this cause is not serious so long as the food supply lasts. Cooper
and sharp-shinned hawks are the most important enemies of quail ;
horned owls prey on quail to a significant extent only in localities
where roosting cover is inadequate ; jays do no significant damage
where cover is adequate ; the precise status of snakes as enemies of quail
is unknown, but certain species may possibly be important locally; the
status of various other predators, none of which is important as an
enemy of quail, is discussed.

Accidents, chiefly collisions with objects while in flight, result in
a considerable number of deaths to quail.

Disease was not observed in the Santa Cruz Mountain area, but an
outbreak the cause of which could not be ascertained was detected in
Oakland, California.

POPULATION STUDIES

In a restricted area representing nearly ideal environmental con-
ditions the number of quail varied from 1.1 to 3.9 per acre, according
to the season of the year. On 60 acres, 64 per cent of a total of 31
potential pairs were ultimately successful in raising broods.

A shootable surplus of approximately 19 per cent apparently
existed in the population studied.

On the area covered by this study, which was not shot over,
26.8 per cent of the birds were alive at the end of the first year, 7.2
per cent at the end of the second, 2.0 per cent at the end of the third,
.5 per cent at the end of the fourth.

Males outnumbered females in the ratio of 53.2 to 46.8.

WEIGHT CHANGES

Maximum weight in the male occurs during the winter and early
spring months, while marked decrease takes place during the breeding
season as a result of nervous excitement and the irregular feeding
habits imposed by sentry duty. The female is relieved of strenuous
courtship activity and sentry duty, and is at this period heavier than
the male, showing no loss of weight until after the young have hatched.

256 CALIFORNIA FISH AND GAME

Neither sex regains lost weight until after the completion of the fall
molt. Immature birds show similar weight fluctuations, but do not
attain full adult weight until the beginning of the second year.

An average loss in weight of 5.86 per cent takes place between
the time when quail go to roost with full crops and the time of
awakening with empty crops in the morning.

California quail decrease in size from the northern part of their
range southward to the southern extremity of Lower California, in
accordance with "Bergmami's Law."

PLUMAGES

Plumage changes, including those of the young, are described;
birds of the year can be distinguished from full adults until the second
summer, by the mottling of the first six (proximal) upper primary
coverts which are vestiges of the juvenal plumage.

CALIFORNIA FISH AND GAME 257

CALIFORNIA FISH AND GAME

A publication devoted to the conservation of wild life and published quarterly by
the California Division of Fish and Game.

The articles published in California Pish and Game are not copyrighted and
may be reproduced in other periodicals, provided due credit is given the California
Division of Pish and Game. Editors of newspapers and periodicals are invited to
make use of pertinent material.

All material for publication should be sent to J. O. Snyder, Division of Fish
and Game, 450 McAllister Street, San Francisco, California.

Vol. 21 JULY, 1935 No. 3

EDITORIAL

This and the following number of California Fish and Game
will be mostly devoted to two parts of a paper by E. Lowell Sumner,
Jr., entitled "A Life History Study of the California Quail, with
Recommendations for its Conservation and Management." The paper
is a final report of an investigation carried on by Mr. Sumner, under
the general supervision of Dr. Joseph Grinnell of the Museum of
Vertebrate Zoology, Berkelev, who has also edited the manuscript. —
J. 0. S.

REORGANIZATION OF THE DIVISION OF FISH AND GAME

The Fish and Game Commission, at its meeting of July 6, 1935,
approved the following reorganization plan of the Division of Fish and
Game.

The reorganization was done to create a smoother working program
and this was brought about by a combination of the functions which
referred to fish, and a separate unit was made of those functions which
referred to game.

The Bureau of Fish Culture name was changed to that of the
Bureau of Fish Conservation, and the scope of its work will cover
fresh and salt water fishes. The reasons for this change in name are :
First, to have the name of the bureau more clearly describe its activities,
and secondly, it is expected that the scope of this bureau's work will
be extended to cover all fish on the sporting list, both fresh water and
salt water. Dr. J. 0. Snyder will remain in charge of this bureau.

The Bureau of Game Conservation was formed to include the
bureau formerly known as Game Refuges and that formerly known as
Game Propagation. It will be composed of two units — one dealing with
birds and one dealing with mammals. It was thought advisable to have
the Bureau of Game Propagation represented in the main office of the
division where inquiries might be answered without having to refer
them to the Game Farm at Yountville, as was formerly the case.
J. S. Hunter will be in charge of the new bureau, and August Bade
will serve under the title of Superintendent of State Game Farms.

The name of the Bureau of Education and Research has been
changed to the Bureau of Research and Engineering. All research
work of the division will be handled bv this bureau and other bureau

258 CALIFORNIA FISH AND GAME

heads will act as an advisory council. This bureau is to be headed by
Herbert C. Davis.

The positions in the Bureau of Patrol have been reorganized as
follows: There are to be six civil service ratings — Chief, Inspector,
Captain, Sergeant, Warden and Deputy Warden. The examination for
Deputy Warden is to be open to the public and will serve as an
apprentice grade. The Deputy Warden will serve an apprenticeship
in the various bureaus, thoroughly learning the scope of work in the
division and to what phase of the work he is best suited. After a
fixed period of years' service as Deputy Warden, promotion can be
obtained by taking and passing the examination for Warden, which
examination, and subsequent promotional examinations will be open
only after the successive probation periods have been successfully
passed.

It is sincerely hoped that under this reorganization, new interest
and confidence will be instilled in the personnel of the division and
that the work will be efficiently handled. — Herbert C. Davis.

FORECAST FOR THE 1935-1936 SARDINE SEASON

This report comprises the third prediction of the abundance and
sizes of sardines to be expected in the immediately succeeding sardine
season. The results of the first forecast, which were very satis-
factory, were discussed when the second prediction was published
in 1934.* The success of the second prediction is indicated at the
close of this article.

PREDICTIONS FOR THE 1935-1936 SARDINE SEASON

The outlook for the coming sardine season is not promising.
The fishery should draw chiefly from three super-abundant year
classes, which entered the fishery in 1929, 1930, and 1933, respectively.
The groups which entered in 1929 and 1930 should furnish the main
support for the winter fishery and the one which entered in 1933 should
supply the major portion of the fall fishery, but due to the intensive
fishing from 1933 to 1935, especially in the fall months, the super-
abundance of these three groups has been wiped out. The groups
entering in 1929 and 1930 are now below normal in abundance and the
1933 group is reduced to normal or perhaps below. We have no
indications that a new superabundant group will enter the fishery in
the fall of 1935.

All of the year classes from which the 1935-1936 fishery will draw
are, therefore, no more abundant than average and many of them
are below average. This means that the entire sardine population is
seriously reduced in numbers. An absolute measure of the extent of
this reduction is not available and we can not state definitely how
scarce sardines will be in the coming season. Schools may be only
moderately difficult to locate and they may be extremely so. This
will hold for both the fall and winter fisheries.

♦The 1934-1935 sardine season. Calif. Fish and Game, vol. 20, pp. 298-300,
1934.

CALIFORNIA FISH AND GAME 259

For each locality the predict ion is as follows:

FALL FISHERY

San Francisco and Monterey (August-November).

Sardines will be less abundant than in 1934-1935, and the sizes
will be mixed. San Francisco fish will be somewhat larger than those
taken at Monterey. The sizes at Monterey will vary from 8h to 11
inches, total length, and approximately one-half will be from 9h to 10.1
inches.

San Pedro (November-December).

In this region also, sardines will be less abundant than in the fall
of 1934—1935, and the sizes will be mixed. The range will be from
8^ to 11 inches, total length, and about half will be between 9 and 10
inches.

WINTER FISHERY

All Localities.

During the winter months sardines will be less abundant than
in former winters and a mixture of sizes will be found in the fishery.
Fish as small as 8^ and as large as 13 inches will be taken, but probably
the majority will be between 10 and 1H inches.

2G0

CALIFORNIA FISH AND GAME

SUCCESS OF THE 1934-1935 PREDICTIONS

To facilitate comparisons, the predictions which were made for
1934—1935 are listed on the left and the actual occurrences on the right
of the following tabulation :

Prediction

Actual Occurrence

SIZES OF SARDINES

Fall Fishery
Average would be between 9i and 10 Monterey fish averaged 10.0 inches,

inches, total length. San Pedro fish averaged 9:9 inches.

Winter Fishery

About 25 per cent would approximate
10 inches, total length, and the majority
would be 11 inches or greater.

For the combined Monterey and San
Pedro fisheries, 20.3 per cent were be-
tween 9^ and 10J inches, and 59.6 per
cent were 11 inches or greater.

ABUNDANCE OF SARDINES

Fall Fishery

Quite abundant, and boats would not Sardines were satisfactorily abundant at

have difficulty in making catches. all ports, but perhaps less so at San

Pedro than at Monterey.

Winter Fishery

Fish would be no more, and perhaps
less, abunant than in the winter of
1933-1934.

The winter fish appeared at Monterey
in December and at San Pedro in Feb-
ruary. During each of these months at
each port the winter sardines were
numerous and the fishermen made their
catches with ease. At Monterey in Jan-
uary and February and at San Pedro in
March, winter fish were scarce and on
many nights much of the fleet failed
to make any catch or brought in only a
small tonnage.

The predictions for the sizes of sardines and the abundance of fish
in the 1934-1935 season proved to be accurate. For the coming season,
1935-1936, all available information points to no increase in abundance
and to the probability of an increasing and possible decided scarcity.
Our predictions refer to abundance or scarcity of fish in the ocean
but not to variations in the amount of total catch. — Contribution No.
149 from the California State Fisheries Laboratory, May, 1935.

CALIFORNIA FISH AND GAME

261

WHALE CAUGHT IN CARQUINEZ STRAITS

A whale of a species not previously known to fishermen in the
Carquinez Straits was caught in a net set for salmon off Dillon's Point,
near Benicia, California, on May 26, 1935.

Humpback whale caught in the Carquinez Straits.

May 27, 1935.

Photograph by E. A. Stoner,

The presence of a mammal of this kind in these waters, especially
at a season when the water is practically fresh, is considered rather
surprising. Assumed evidence of the freshness of the water is the
fact that the California-Hawaii Sugar Refinery was still taking on
water in their water barge near this point.

The whale was identified by the writer as the Pacific Humpback
(Megaptera versabilis) using Paul Bonnot's "The Whales of
California," published in the July, 1929, California Fish and' Game
as a guide. It was thirteen feet in length and conservatively esti-
mated as weighing fifteen hundred pounds.

Lawrence Gando and George Marinos, who brought the specimen
to the shore at Capello's Beach, Benicia, reported a four-hour fight
with the animal before finally subduing it with the aid of an axe
when it had finally become worn out in the struggle. The net was
badly torn, and the boat many times threatened during the battle to
get it to shore. Mr. Joseph Capello estimated that five hundred people
came to his wharf to view the stranger.

The only other whale reported as having been seen in these
straits was a killer whale brought in about five years ago. — Emerson A.
Stoner, Benicia, California.

262 CALIFORNIA FISH AND GAME

A LARGE NORTHERN HALIBUT IN MONTEREY

On the morning of May 3, 1935, a northern halibut (Hippoglossus
hippoglossus) , weighing 89 pounds and measuring 5 feet 3 inches
from tip of snout to tip of tail, was caught in 30 fathoms of water,
two miles north of the town of Monterey. The specimen was caught
on a roekfish long line by commercial fishermen, M. Silveria and B.
Silveria, fishing from a skiff. From available records, this is the
largest specimen of northern halibut taken so far south.

The northern halibut (right halibut or genuine halibut as it is
sometimes called) is a north Pacific form, ranging as far north as
Bering Sea. Only occasional specimens are taken as far south as
Monterey Bay. To the northward, some specimens attain a weight
of 500 pounds.

The California halibut (Paralichthys calif ornicus), sometimes
called chicken halibut or southern halibut, ranges from San Francisco
southward to the Gulf of California, and rarely reaches a weight of
60 pounds.

In California, the largest commercial landings of northern halibut
are at the port of Eureka, and the largest landings of California
halibut are at the ports of Los Angeles, Santa Barbara and San Luis
Obispo. The annual poundage of California halibut landed is usually
two to three times that of the northern halibut. — J. B. Phillips, Cali-
fornia State Fisheries Laboratory, May 14, 1935.

NEW MACKEREL CANNERIES

In view of the rapidly expanding market for canned mackerel,
it is not surprising that new canneries are being built in southern
California.

With the 1934 pack of 1,230,000 standard cases completely sold
out, practically every fish cannery from Monterey to San Diego is at
present packing mackerel. Six plants at Monterey, seven at San Pedro
and four at San Diego are canning all the mackerel they can obtain.
Two additional San Pedro canneries are preparing to recommence
activities after five years' idleness as far as mackerel are concerned.

In addition to the activity of the established plants, there are
three new plants either ready or nearly ready to start mackerel
packing. All three are located at Newport. Two of these are com-
pletely rebuilt fruit and vegetable canneries. Both of them have
already packed on an experimental scale and are ready to pack in
earnest as soon as mackerel appear in abundance. The third cannery
is new in every respect. At the present time the building itself is
not quite completed, so production can not commence before late
summer.- — Richard S. Croker, California State Fisheries Laboratory,
May 17, 1935.

A NEW NET DEVELOPMENT

During the past winter (1934—1935) a somewhat new type of net,
or rather a new adaptation of an old type, has been developed at San
Pedro. So far, but one boat has used this net.

CALIFORNIA FISH AND GAME 263

The net is a miniature purse seine made of tanned webbing. It
is 100 fathoms long, 10 fathoms deep, of 3-inch mesh (stretched meas-
ure). It can be handled and hauled by four or five men. This rapid-
action net is used in shallow water in the small open spaces between
kelp beds. The fish most often caught is the half moon (Medial una
calif orniensis) which is in fairly good demand at the San Pedro mar-
kets. The halfmoon can be observed in small schools swimming leis-
urely near the surface, so the small'net can be laid around them without
much trouble. The largest single haul so far made was somewhat
over one ton. — Richard S. Croker, California State Fisheries Labora-
tory, May 17, 1935.

INCREASED UTILIZATION OF FISH LIVERS

The value of various fish liver oils as sources of vitamins A and D
has recently been realized, following the research work of several com-
mercial, university and government laboratories.

For years, cod liver oil has been recognized as beneficial in the
treatment of various deficiency diseases. Several years ago the liver
oil of the halibut was found to be much more potent than the cod
liver oil. More recently it has been found that the livers of many
other species of fish produced oils relatively high in vitamin content.

At the present time several of the large medicinal firms have
buyers in California, purchasing fish livers from fish markets and
canneries. They have found that the livers from some species are
worthless, yielding very little oil and being poor in vitamin content.
Other species yield very good quantities, relatively high in either
vitamin A or D content. The white sea-bass, barracuda, swordfish,
totuava, mackerel, bonito, tuna, and several other species have livers
worth saving. The marketmen have found that cleaning costs can be
defrayed by saving and selling the livers of the fish they handle.
Tuna canners realize a considerable saving in the cost of producing
canned tuna by selling the livers. The utilization of mackerel livers
has resulted in an increase in employment in the canneries, and at the
same time the canners have found that the cost of producing a case of
mackerel has been reduced 5 to 10 cents since the practice started. As
production costs in general have risen notably the last few months, the
sale of tuna and mackerel livers often marks the difference between
selling canned fish at a profit and selling at cost. The fish cleaners
separate the livers from the offal and place them in five-gallon cans
which have a capacity of 40 pounds. The cans are packed in ice and
are picked up daily by the buyers. During the past two years the
liver cans have become a familiar sight in practically every market
and cannery in California. — Richard S. Croker, California State
Fisheries Laboratory, May 18, 1935.

-23006

L'GI CALIFORNIA FISH AND GAME

NOTES FROM THE STATE FISHERIES LABORATORY

The tagging of tuna was begun on an experimental scale in May,

1934, and during the summer something over 100 fish were marked
on the preopercle with a metal strap tag. The tag carries a brightly
colored celluloid disc, so as to be readily recognized by fishermen.
The first tag to be returned was by a San Diego fisherman on February
1, 1935.

Much may be learned about the behavior of sardines and mackerel
by tagging, but with these two fishes it is probable that an internal
tag carried in the body cavity would be more successful, especially so
in the case of sardines where large numbers are dumped into reduction
plants without being handled as individual fish. If internal tags are
used the tags will have to be recovered in the reduction plants either
by some mechanical device or by the use of magnets which will pick
the metal tag from the fish meal. Initial experiments in the use of
such tags are being conducted at the laboratory.

H. C. Godsil and Richard S. Croker made a trip on the patrol
boat Bluefin to Turtle Bay, Mexico, on January 29 — February 13,

1935. for the purpose of tagging tuna and securing samples of the
bait sizes of sardines from that area.

On April 21, 1935, II. C. Godsil sailed on the fishing boat Reliance
for Mexican waters to carry on tuna studies, including the tagging of
yellowfin tuna.

During April, 1935, D. H. Fry, Jr., made a trip on the Bluefin to
Magdalena Bay, Mexico, to scout for adult mackerel during the time
they are so scarce in the waters of southern California. Adults were
not found in any quantity in that region, and the eggs and larval
stages of the mackerel were almost absent from Mexican waters at that
time of year.

During the spring of 1935, several trips by different members of
the staff were made to San Diego in order to determine the relative
abundance of the sardine quarter-oil sizes that may be expected to
enter the San Pedro commercial catch in the fall of 1935. This next
entering age-class of sardines appears to be below normal in abundance,
so we can not expect that our present sardine supply will be heavily
recruited in the fall.

During April, L. A. Walford, a former member of our staff,
returned on the yacht Haida from a two months' cruise in Mexican
waters where he collected specimens and notes for issuing a book on
some of the sport fishes of that area. An artist accompanying the
expedition prepared color paintings of the more important sport
fishes. We look forward with great interest to the appearance of this
volume.

Through the summer of 1935, the statistical clerks under the direc-
tion of Geraldine Conner were unusually busy with special reports in
addition to the routine work of tabulating current fish catch data.
Many of these special reports were for use at Sacramento during the
session of the State Legislature and others were for use of the laboratory
research staff.— W. L. 8., May 16, 1935.

CALIFORNIA FISH AND GAME 265

SIZES OF SARDINES TAKEN BY GEAR AND AREA

The California State Fisheries Laboratory recently issued Fish
Bulletin 43, "The sizes of California sardines eaughl by the differenl
fishing gear and in the different localities of the Monterey and San
Pedro regions," comprising two reports by J. B. Phillips and (I. H.
Clark. The authors present the results of an investigation made to
determine whether or not the system of sardine sampling as carried
on in the past is still adequate since changes have occurred in the
fishery. The results are of importance to the industry because they
show that lampara, purse seine and ring nets all take the same sizes of
fish, that these sizes comprise the sardines available to the fishermen,
and that no type of gear exhibits size selection differing from any
other type. Of greater importance to the industry is the evidence
indicating that large fish first appear each winter to the north of
Monterey and gradually become disseminated southward throughout
the entire Monterey region, whereas in the San Pedro region no con-
sistent distribution of differential sizes within the region is evident.
The results are of value to the sardine investigation because of the
demonstration of the continual reliability of our sardine sampling-
system and the furnishing of additional evidence of a southward
movement of sardines along the California coast during the wilder
months.— G. II. C.

SEIZURES OF FISH AND GAME

January, February, March, 1935

Game: ,,

l),.rr _ _ 11)

D eer mea t , pounds 2,809

Deer hides - ^

Ducks, geese, mudhens - 4'5-'

Doves J

Non-game birds i

Pheasants..'

Pigeons \

Quail ?6

Rabbits "

Tree squirrels ■-

Bird nets A

Guns 10

Fish: .

Abalones, pounds 07

Bass, striped -- l'

Crappie, sunfish

Catfish, pounds --- „ J^r.

Clams ----- 2'^

Halibut, pounds --

Lobsters :,

Trout .'-'-

Trout, steelhead, pounds - '■' "'

Traps, fish

266

CALIFORNIA KISII AND GAME

GAME CASES
January, February, March, 1935

Offense

Deer; closed season; killing of does

Ducks; closed season; selling of

Doves ; closed season

Hunting, no license.

Firearms in refuge

Nongame birds; killing of

Pheasants; closed season

Pigeons; closed season

Quail ; closed season

Rabbits; closed season

Shorebirds; killing of

Spotlight hunting

Trespassing on posted grounds

Tree squirrels; killing of

Shoot ing from highway

Bird nets; illegal

Miscellaneous game cases

Totals

Number
arrests

91

59
2

41
3

16
6
2
5
1
4
3
2
1
3
1
4

244

Fines
imposed

$1,902 00

2,860 00

25 00

400 00

50 00

500 00

275 00

25 00

225 00

25 00

100 00

225 00

2 00

50 00

10 00

15 00

15 00

$6,764 00

Jail

sentences

(days)

1,441
31

217^

1.6S9K

FISH CASES
January, February, March. 1935

Offense

Angling; no license

Abalones; small; drying of

Bass, striped; overlimit

Commercial fishing; no license

Clams; over limit; small

Catfish; selling small

Cockles; over limit

Crappie; closed season

Lobs t ers ; small

Night fishing

Pollution of streams

Spears; illegal use of

Salmon; spearing of; closed season.
Trout; closed season; spearing of _ .

Nets and set lines;illegal

M iscellaneous fish cases

Totals.

Number
arrests

19

28

1

41

79

1

4

5

3

2

9

7

2

14
4
7

226

Fines
imposed

$295 00

397 50

100 00

45 00

1,275 00

25 00

75 00

60 00

50 00
150 00
160 00

50 00
155 00
150 00

50 00

$3,037 50

Jail

sentences
(days)

47H
162H

54
134^

12H

55
120
12H

598^

CALIFORNIA FISH AND GAME

267

STATEMENT OF EXPENDITURES
For the Period July 1, 1934, to March 31, 1935, of the Eighty-sixth Fiscal Year

Function

Salaries

and

wages

Materials

and
supplies

Service

and
expense

Property

and
equipment

Total

Administration:

Executive . .„ _ ..

$5,504 77
4,140 00

$5,504 77

Clerical and office . __

$693 77
779 48

2,416 83
353 40

$158 19

$49 02

5,040 98
779 4S

Printing, Fish and Game Magazine

Printing, general ..

2,416 83
731 75

378 35
1,872 99
3,059 26
2,760 06

550 32
9,824 18
5,261 62

Traveling . ...

1,872 99
3,059 20
2 760 06

Postage .. ... __ _ _.

Telephone and telegraph

Freight, cartage and express

550 32

Rent

9,824 18

Accident and death claims, .. _

5,261 62

Accounting pro rata __

3,991 13

3,991 13

1,958 75

50 00

Legal . .

1,942 75

50 00

954 37

16 00

Premiums on bonds ... .

Publicity _ .

954 37

Total Administration

$13,635 90
$1,440 00

$4,243 48

$9 90
422 32

$26,812 09

S2.S 62
101 47
1,431 38
52 39
52 07
38 90
61 38

!*65 i)2

$44,756 49
$1,478 52

Bureau Education and Research:
Clerical and office . ___ __

Automobiles,. .. .. . .. _ _ _

523 79

Traveling _ . _

1,431 38

Photography ___ .

11 84
14 94
87 42

64 23

Library..

810 00
2,400 00
1,381 84

$109 33
12 30

986 34

Research .

2,538 62
1,443 22

Publicity ...

Total Bureau Education and Research

Bureau Patrol and Law Enforcement:
Chiefand assistants

$6,031 84

$8,325 00
2,385 00

$546 42

$1,766 21

$121 63

$8,466 10
$8,325 00

Clerical and office

$28 20

19,792 17

$62 75

9,745 03

31,514 47

457 11

1,286 55

6 45

547 71

891 43

763 67

1,759 61

232 50

$30 00
5,583 65

2,505 95

Automobiles ...

35,120 85

Traveling _ ... . ... .. __

31,514 47

Postage .

457 11

Telephone and telegraph

1,286 55

Freight, cartage and express

6 45

Rent .. ._

547 71

Captains and wardens,.

138,502 42

716 99

1,328 17

492 82

26 51

310 27

12 19

140,197 35

Launches

2,402 11

Fish planting

3,465 00

5,729 62

Premiums on bonds _

232 50

Cooks.. .. _

1,016 12

2,070 00

10,117 10

8,756 63

8,041 32

1,016 12

Commercial fisheries patrol:
Chief

12 52
323 16

2,070 00

Captains and wardens .. __

6 60
3,162 69

8 00
2,180 28

10,144 22

Launches ._

Fish cannery inspectors, seasonal _ .

14,422 76
8,041 32

Traveling ..

3,937 83
658 00
410 42

3,937 83

Rent... . _

658 00

Automobiles..

538 57

948 99

Temporary help. .

237 92

237 92

Total Bureau Patrol and Law Enforcement .

Bureau Commercial Fisheries:
Chief and assistants.. ..

$182,976 51

$10,215 00
7,200 00

$26,006 21

$54,461 81

$6,298 30
$97 83

$269,802 83
$10,215 00

Clerical and office .

$58 91
393 10

$74 45
229 88
4,098 21
474 83
76 40
106 48
273 23

7,431 19

Automobiles.

622 98

Traveling

4,098 21

Telephone and telegraph . .

474 83

Freight, cartage and express. .

76 40

Rent. .

106 48

Heat, light, water and power

273 23

Research

1,710 00
17,280 00

12 72
490 36

105 93

376 32

1,828 65

Laboratory. _ _

514 74
450 00

18,661 42

Hydro-Biological Survey —
Monterey Bay

450 00

Temporary help

75 87

75 87

Statistics. ....

487 66

1,552 50

29 42

2,069 58

Total Bureau Commercial Fisheries... . .

Less Indirect Abatements:
Deer tag regulation

$36,4S0 87

$1,442 75

$7,850 72
$328 60

$609 50

$46,383 84
$328 60

Total Bureau Commercial Fisheries

Adjusted

$36,480 87

$1,442 75

$7,522 12

$609 50 '

$46,055 24

268

( AI.IFOKXIA I-'ISH AND CAME

STATEMENT OF EXPENDITURES Continued
For the Period July 1, 1934, to March 31 , 1935, of the Eighty-sixth Fiscal Year

Function

Bureau Fish Culture:

Chief and issistants

Clerical and office

Automobiles

Traveling

I'os tage

Telephone and telegraph

Freight, cartage and express

Rent

Heat , light and power

Hatcheries

Fish cars

Blueprinting

Cooperative research

Temporary help

Fish hatchery assistant, seasonal.

Hydraulic engineering

Special field

Fish rescue:

Miscellaneous supplies

Heavy truck mileage

Travel .

Rent and water service

Motorized eiuipmen t

Chief and iisistants

Total Bireau Fish Culture-

Bureau Game Propagation:

Chief and is^iitants

Clerical and office

Automobiles

Traveling

Postage

Telephone and telegraph

Freight, cartage and express

Heat.li'ht, water and power _

Maintenance

Temporary help

Refuge maintenance

Quail trapping and expansion of quail program
Purchase of quail and pheasants

Total Bureau Game Propagation-

Bureau Game Refuges:

Chief and assistants

Clerical and office

Automobiles

Traveling

Freight, cartage and express

Blue -minting

Research

Lion hunters

Refuge posting

Predatory animal control

Refuge maintenance

Predatory animal hunters and trappers,

seasonal

Temporary help, seasonal

Total Bureau Game Refuges-

Bureau Licenses:

Clerical and office

Printing, licenses and applications.

Traveling

Postage

Freight, cartage and express

Premiums on bonds

Identification license buttons

Total Bureau Licenses .

Total 86th fiscal year expense paid from
support appropriations

Prior yearexpensef or support, 85th fiscal year. . .

Total 85th and 86th fiscal year expense paid
from support appropriations

Salaries

and

wages

55,445 00
3,968 39

81,231 17
2,655 00

1,558 39

809 00

14,619 49

1,800 00

5,859 05

1,800 00

$119,745 49

$4,095 00
630 00

8,703 00

33 55

1,269 00

3,465 00

$18,195 55

$6,374 97
1,440 00

4,077 00
675 00

6,115 32

4,500 00
2,820 50

$26,002 79
$9,855 00

$9,855 00

$412,923 95

Materials

and
supplies

$2 89

17 72

5,125 88

38,957 28
231 81

545 52

17 76
52 46

47 IS

124 82

$45,123 32

$S00 9S

6,533 48

788 78
4,134 60

$12,257 S4

$11 83
392 87

1,482 16

$1,886 86

$83 29
1,360 28

5,191 00

$6,634 57

$98,201 45

Service

and
expense

$5 00

12 25

2,499 30

5,978 56

160 01

777 43

548 59

2,564 00

1,204 91

660 25

2,081 89

51 40

697 56

653 76
9 40

336 5 1

1,211 Ss

59 00

26 10

235 53

$19,773 33

$260 50

1,413 95

It 04

141 55

5 36

1,226 52

181 70

1,9S1 3S

$5 222 00

$11 50

229 67

3,133 79

4 88

4 18

328 60

4.8S0 00
1,016 93

$9,619 55

$14 45

413 62

893 58

188 58

1,063 25

$2,573 48

$127,750 59

Property

and
equipment

$73 82
6,346 48

1,009 34

224 II

4 46
13 72

5 98

551 25

■-._'_'> I.',

$963 66

4 55

10 55

$978 78

$115 51

$115 51

$93 79

163 79

$16,512 47

Total

$5,452 89

4,072 LS

13,971 66

5,978 56

160 01

777 43

548 59

2,564 00

1,204 91

121,858 04

4,968 70

51 40

3,026 38

809 00

14,619 49

2,475 98

5,934 63

53 16

336 51

1,211 SS

59 00

702 17

2,035 53

$192,872 10

$4,095 00

630 00

2,025 14

1,413 95

U 04

141 55

5 36

1,226 52

15,422 73

33 55

1,269 00

6,245 71

4,134 60

$36,654 15

$6,374 97

1,463 33

622 54

3,133 79

4 88

4 18

328 60

4,077 00

675 00

4,890 00

8,729 92

4,500 00

2,820 50

$37,624 71

$10,046 53

1,360 28

413 62

893 58

188 58

1,063 25

5,191 00

$19,156 84

$655,388 46
17,423 20

:- -. ^ii 66

CALIFORNIA FISH AND GAME

269

STATEMENT OF EXPENDITURES- Continued
For the Period July 1, 1934. to March 31, 1935, of the Eighty-sixth Fiscal Year

Function

Special items:
86th fiscal year-
License commissions

Construction Russian River Jetties,

Chap. 989-33.

California Code Commission for expenses,

Chap. 645-33

Electro Metal Co.'s claim, Chap. 599-31 - - -

Total 86th fiscal year

85th fiscal year:

License commissions —

Construction Russian River Jetties,

Chap. 989-33

Total 85th fiscal year

Total special items _

Permanent improvements:
Construction, improvements and equipment:

86th fiscal year

85th fiscal year

Total permanent improvements.

Bureau of Commercial Fisheries — Chap. 825-
33 — Fresh fish marketing:
86th fiscal year:

Chief and assistants

Clerical and office

Printing—

Automobiles

Traveling

Postage -

Telephone and telegraph

Freight, cartage and express

Rent

Exhibits

Temporary help

Pro rata fish and game administration

Salaries

and

wages

$5,172 50

Materials

and
supplies

$4,718 96

Service

and
expense

$33,312 54

419 46

3,198 21
18,750 00

$55,680 21

$7,704 28

Property
and

equipment

$315 65
30 00

$4,52(1 50
990 00

Total 86th fiscal year.
85 th fiscal year

Total Bureau of Commercial Fisheries —
Chap. 825-33— Fresh fish marketing,
85th and 86th fiscal years

Contributions to retirement system

Total from current biennium

212 37

$5,728 87

$192 00

3,244 00

95 51

295 29
13 01

$3,839 81

$93 72

1,753 60

484 50

2 95

433 32

70 70

2,224 60

Prior biennium appropriations:
85th fiscal year —
Permanent improvements:
Construction, improvements and equip-
ment

84th fiscal year:

Support _--- --

Total from prior biennium appropriations

Grand total proprietary group.

$5,063 39

$1,822 39

{6 35

Total

$33,312 54

411! Hi

3,198 21
18,750 00

$55,680 21

345 65

$56,025 86

$19,418 13
46 32

$6 35

$19,464 45

$4,526 50

1,188 35

3,244 00

189 23

1,753 60

484 50

2 95

433 32

70 70

2,519 89

212 37

13 01

$14,638 42
11 55

$14,626 37

11,451 01

774,379 85

583 19

853 65
270 46

$774,650 31

270 CALIFORNIA FISH AND GAME

STATEMENT OF REVENUE

For the Period July 1, 1934, to March 31, 1935, of the Eighty-sixth Fiscal Year

Revenue for Fish and Game Preservation Fund, Current Year-
License sales*

Angling licenses, 1934... $306,498 50

Anglinglicenses, 1935 8,545 50

Commercial hunting clublicenses, 1934-35 1,225 00

Commercial hunting club operators' licenses, 1934-35 265 00

Deertags, 1934 10S.909 00

Fish breeders' licenses, 1934 25 00

Fish breeders' licenses, 1935 300 00

Fish importers' licenses, 1934 35 00

Fishimporters'licenses, 1935 75 00

Fish packers' and wholesale shellfish dealers' licenses, 1934-35 900 00

Game breeders' licenses, 1934 115 00

Game breeders' licenses, 1935 737 50

Hunting licenses, 1933-34 3,931 50

Hunting licenses, 1934-35 205,255 34

Kelplicenses, 1934 20 00

Kelplicenses, 1935 20 00

Market fishermen's licenses, 1934-35 28,790 00

Market fishermen's licenses, 1935-36 520 00

Trapping licenses, 1934-35 . 1,470 00

Total license sales . $667,637 34

Other income:

Court fines S29.9S3 51

Dividends California National Bank 7,418 03

Dividends Trinity County Bank at Weaverville. 71 00

Fish packers' tax 268,548 57

Fish tag sales 1 1,986 55

Game tag sales 127 95

Importers' contributions 230 00

Interest on bank balances 5,009 96

Kelp tax 67 01

Lease of kelp beds 1,309 20

Miscellaneous sales 3,068 11

Publication sales 446 20

Total other income $318,266 09

Total Fish and Game Preservation Fund S9S5,903 43

Revenue for the General Fund (part of):
Unclaimed checks and deposits 168 15

Grand total all funds $986,071 58

CALIFORNIA FISH AND GAME

271

FRESH FISH IMPORTATIONS BY POINT OF ORIGIN, JANUARY, FEBRUARY AND MARCH,
Compiled by the Division of Fish and Game, Bureau of Commercial Fisheries

1935

Species

Gulf of
California

West Coast

Lower
California

International

waters south

U. S. boundary

(definite

origin

unknown)

Mexican

Mainland

and Central

and South

America

Japan

Total

pom

39,312
3,481

39,312

9,603

59,694

394

1,290

204,327

1,429

1,862

70,564

1,832

405

Cabrilla

6,103
59,544

19
150

Corbina, Mexican...

Cultus, Pacific

394

500

183,073

1,429

1,862

70,564

1,832

405

100,435

Grouper

790
21,254

Halibut, California..

Mackerel, Pacific

Rock Bass

Rockfish...

Sablefish

Sea-bass, Black

729
908,808

35,159

136,323

908,808
691

Sea-bass, Totuava. .

Sea-bass, White

649
733
577
325
2,011
139
511

42

Shark

733

Sheepshead

577

Skate

325

Smelt

2,011
139

Sole

Swordfish, Broadbill

511

Tuna, Albacore

581,269

581,269

38,707

1,232,690

11,217,772

Tuna, Bonito .

36,939

142,208

1,169,237

7,118

659,637

441,776

1,768

219,939

7,422,505

Tuna, Skipjack

30,925
2,626,030

839,618

Tuna, Yellowfin

Whitefish

7,118

Yellowtail - ..

46,063

705,700

Crustacean:

Lobster, Spiny _

441,776

Total pounds...

975,184

2,865,147

7,747,689

2,656,955

1,420,887

15,665,862

FRESH FISH IMPORTATIONS* FROM FOREIGN COUNTRIES, FOR JANUARY, FEBRUARY AND MARCH, 1935
Compiled by the Division of Fish and Game, Bureau of Commercial Fisheries

Species

Landed in
Region 70,
Los Angeles

Landed in
Region 80,
San Diego

Total pounds

8,153

8,748

58,849

31,159

855

845

394

70

148,000

1,429

891

70,554

1,832

405

40,473

24,596

559

602

223

325

1,811

64

511

39,312

9,603

59,694

Cultus, Pacific

394

1,220
56,327

1,290

Halibut, California - - - - -

204,327

Mackerel, Pacific

1,429

971
10

- 1,862

Rockfish -. .

70,564

1,832

405

95,850

884,212

132

131

354

136,323

908,808

691

Shark - -

733

577

325

Smelt --

200
75

2,011

Sole --

139
511

581,269
9,992

876,616

3,557,350

1,903

324,582

51,789

581,269

28,715

356,074

7,660,422

5,215

381,118

389,987

38,707

1,232,690

11,217,772

Whitefish

7,118

705,700

Crustacean:

441,770

6,518,733

9,147,129

15,665,862

* These importations are included in tables of landings.
8- 23006

272

CALIFORNIA FISH AND GAME

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BUREAU OF PATROL

E. L. MACAULAY, Chief of Patrol San Francisco

CENTRAL DISTRICT (Headquarters, Sacramento)

S. H. Lyons, Inspector in Charge Sacramento

Jos. H. Sanders, Captain Northern Division Sacramento

S. R. Gilloon, Captain Southern Division Fresno

John O'Connell, Captain Modesto

O. P. Brownlow, Captain Bakersfteld

A. H. Willard, Captain Nevada City

, Sergeant Redding

J. E. Newsome, Captain, Newman 1

S. J. Carpenter, Captain, Maxwell j- Special Duty

E. W. Smalley, Captain, Hanf ord J

WARDENS (Northern Division)

Roy W. Anderson Red Bluff

W. J. Black Vallejo

L. W. Dinsdale Yuba City

C. O. Fisher Susanville

C. L. Gourley Westwood

Alvin Granstrom Loyalton

Brice Hammack Yreka

Earl Hiscox Grass Valley

Wm. Hoppe Lodi

A. A. Jordan Alturas

Paul Kehrer Fall River Mills

Taylor London Orovllle

L. W. Longeway Sonora

Charles Love Redding

Leslie Mercer Quincy

Nelson Poole Sacramento

Albert Sears Placerville

Chas. Sibeck Sacramento

R. L. Sinkey Woodland

Fred R. Starr Macdoel

Vernon Sutton Weaverville

R. A. Tinnin Camptonville

E. C. Vail Willows

H. S. Vary Walnut Grove

Edwin O. Wraith Chico

(Southern Division)

Lester Arnold Bakersfield

H. E. Black Madera

Clarence Brown Mariposa

F. A. Bullard Reedley

Ray J. Bullard Porterville

M. S. Clark Merced

Cliff Donham Coalinga

Wm. Hoppe Lodi

F. F. Johnston Auberry

R. J. Little Jackson

Walter I. Long Woodlake

Geo. Magladry Modesto

C. S. Donham Angels Camp

J. W. Thornburg Markleeville

Roswell Welch Kernville

COAST DISTRICT (Headquarters, San Francisco)

K. P. Allred, Inspector in Charge San Francisco

Wm. J. Harp, Captain Northern Division Ukiah

Wm. Lippincott, Captain Southern Division San Francisco

J. D. Dondero, Captain Eureka

Henry Lencioni, Captain Santa Rosa

Ralph Classic, Captain Monterey

WARDENS (Northern Division)

W. C. Blewett Crescent City

Earl Caldwell Eureka

Ray Diamond Willits

Scott Feland Fortuna

E. R. Greenleaf Monterey

J. H. Groves Cloverdale

J. W. Harbuck Napa

Ovid Holmes Fort Bragg

E. J. Johnson Garberville

Wm. F. Kaliher Fortuna

Bert Laws San Anselmo

Earl Macklin Ukiah

Leo Mitchell Point Arena

K. J. Ransdell Nice

Victor Von Arx Santa Rosa

L. J. Weseth Monterey

R. J. Yates San Rafael

(Southern Division)

F. J. McDermott Santa Cruz

Tate Miller Moss Landing

C. R. Peek San Mateo

Orben Philbrick Pacific Grove

Fred H. Post Salinas

Lee C. Shea San Francisco

Geo. Smalley Richmond

Paul L. Turner Paso Robles

J. p. Vissiere Watsonville

C. M. Bouton San Francisco

C. L. Bundock Oakland

Ed Clements San Rafael

T. K. Duncan Concord

Chas. England San Rafael

Fred W. Hecker San Luis Obispo

C. E. Holladay San Jose

John Hurley Pismo Beach

Mansfield Joy San Francisco

McPherson Lough Palo Alto

SOUTHERN DISTRICT (Headquarters, Los Angeles)

C. S. Bauder, Inspector in Charge Los Angeles

LaRue Chappell, Captain Western Division Los Angeles

E. H. Ober, Captain Eastern Division San Bernardino

L. T. Ward, Captain San Diego

C. H. Groat, Captain Terminal Island

Sergeant Santa Barbara

Sergeant Indio

Sergeant Bishop

SOUTHERN DISTRICT (Western Division)

A. R. Ainsworth Santa Maria

R. E. Bedwell Ventura

E. A. Chan Terminal Island

Ray Ellis Newport Beach

Walter Engelke Terminal Island

E. H. Glidden San Diego

W. L. Hare Santa Ana

H. C. Jackson Santa Barbara

Carmi Savage Los Angeles

C. L. Towers Los Angeles

(Eastern Division)

A. F. Crocker Bridgeport

J. H. Gyger Perris

Geo. Johnson El Centro

Theo Jolley Idyllwild

W. C. Malone San Bernardino

R. C. O'Connor — Bishop

W. S. Talbott Forest Home

C. J. Walters Independence

E. L. Walker Independence

LAUNCH PATROL

Motor Vessel "Bluefin," Terminal Island

Launch "Albacore," Monterey

Launch "Shrapnel," Sacramento

Launch "Quinnat," San Rafael

Launch "Hunter," Vallejo

Launch "Rainbow," Walnut Grove

Launch "Silverside," Eureka

CALIFORNIA STATE PRINTING OFFICE

GEORGE H. MOORE, STATE PRINTER

SACRAMENTO, 1935