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California Fish and Game is a journal devoted to the conservation of wild-
life. Its contents are not copyrighted and may be produced elsewhere provided
credit is given the authors and the California Department of Fish and Game.

Interested persons may have their names placed on the mailing list by writ-
ing to the editor. There is no charge, but subscriptions must be renewed annu-
ally by returning the postcard enclosed with each October issue. Subscribers
are asked to report changes in address without delay.

Please direct correspondence to:

AAr. Phil M. Roedel, Editor
California State Fisheries Laboratory
Terminal Island Station
San Pedro, California













Published Quarferiy by fbe







PAUL DENNY, President

LEE F. PAYNE, Commissioner HARVEY E. HASTAIN, Commissioner

Los Angeles Browley

WILLIAM J. SILVA, Commissioner CARL F. WENTE, Commissioner

Modesto Son Francisco

Director of Fish and Gome


PHIL M. ROEDEL, Editor Terminal Island

Editorial Board


FRANK KOZLIK San Francisco



Management of Chamise Brushlands for Game in the North Coast

Kegion of California H. H. Biswell, R. D. Taber,

D. AV. Hedrick, and A. M. Schultz 453

Life History of the Blue Roekfish, Sehastodes mystinus

Joseph II. Wales 485

The Tomales Bav Herrino^ Fishery W. L. Scofield 499

Life History and Productivity of a Population of Western ]\Iourn-
ing Doves in California Joiix B. Cowan 505

Tooth Development of the Nelson Bigliorn Sheep 0. V. Demixg 523

Food Habits of California Striped Bass

W. C. JoHxsox and A. J. Calhoun 531

A Sampling Program for Recovery of ]\Iarked King and Silver
Salmon Doxald H. Fry, Jr. and Eldox P. Hughes 535

The Pismo Clam in 1951 Johx E. Fitch 341

Notes on the Embryology and Behavior of the Flyingfishes {Cyp-
selurus) Off the Coast of Southern and Baja California

Daxiel J. Miller 549

Distributional Notes on Some Pacific Coast Marine Fishes

John' E. Fitch 557

The Fanfish, Pteraclis velifera, Found in California

Glenx' a. Noble and Charles 0. Blodgett 565

Variations in the Wolf Eel, Anarrliichthys oceUatiis Ayres, a Fish
Inhabiting the Eastern North Pacific Ocean

Robert II. Kanazawa 567

Food of the Pacific Sardine, Sardi72ops caeruha, from Central Baja
California and Southern California John Radovich 575

Development through the Prolarval Stage of Artificially Fertilized
Eggs of the Pacific Sardine {Sardinops caendea)

Daxiel J. ]\Iiller 587

Pheasant Cooperative Hunting Area Results, 1951

Chester M. Hart, Johx F. Davis, and Wilbur F. Meyers 597

Reviews 605

Reports 609

Index to Volume 38 611



School of Forestry, University of California, Berkeley


This stud}^ was begun in the spring of 1948 to investigate the possibility
of managing chamise brushlands for game, the primary purpose being to
determine whether game populations will build up under brushland man-
agement, and, if so, the most satisfactory way of manipulating the cover
to increase game production. The investigations have centered mainly in
Lake County, but have not been entirely limited to that area. In Califor-
nia, there are about 7,300,000 acres of chamise brushlands (Sampson,
1944). They are important in the interior Coast Range from Trinity and
Shasta Counties south to San Francisco Bay. The chief center of their
distribution, however, is in the southern Coast Range; smaller isolated
units are found in the Sierra Nevada foothills. The chamise brushlands
have been looked upon chiefly as valuable for game and watershed but
little has been done in the way of management for either purpose. The
rapid increase in population of the State, producing a corresponding in-
crease in water demands, results in a greater need for good watershed
management. Also more productive game areas are needed because there
are more hunters and a greater need for meat, and as well a greater
amount of time to be spent in recreation.

This paper should be looked upon as a progress report since the study
is still under way and the results may change with a greater accumulation
of data. It is recognized that more data are needed on nearly every phase
of the project. The investigations are being carried on cooperatively be-
tween the University of California and the California Department of
Fish and Game with funds provided by federal aid in Wildlife Restora-
tion Act, Project California 31-R. Most of the wild life studies were made
by R. D. Taber, the plant studies by the other investigators.

The authors express gratitude to the many persons who assisted in
various wa^^s on the project. The studies were suggested by Ben Glading,
Chief of the Bureau of Game Conservation, California Department of
Fish and Game. He foresaw a need for information of this sort in develop-
ing sound game management policies for the Bureau of Game Conserva-
tion. Dan Tillotson, also of the California Department of Fish and Game,
has been in general charge of Pittman-Robertson projects and helped in
many ways. Mam^ students in the Imiversity of California helped at var-
ious times. Professor R. E. Storie of the universit}' examined the soils, and
Dr. J. Vlamis made the soil fertility tests. The food analyses of deer stom-
achs were made by Carol Ferrel and Howard Leach of the fish and game

1 Submitted for publication April, 1952. Federal Aid in Wildlife Restoration Act Project
California 31-R.

( 453 )




laboratory in Berkeley. Merton Rosen, Arthur Bischoff, John Azevedo,
and Alvin Hightower assisted in collecting the deer and making parasite
examinations. Gratitude is expressed also to ranchers in the area for their
generous cooperation, and especially to Glen Keithly for the benefit of
his keen observation and the information he gave on brushland manage-
ment. He is a pioneer in this work, having started a program of brush
control for sheep and deer years earlier. Much of the work presented here
was done on Keithly 's ranch.

Appreciation is expressed to the folloAving who critically read the
manuscript: Keith Arnold, H. F. Heady, AV. E. Howard, A. S. Leopold,
AV. M. Longhurst, R. M. Love, A. AV. Sampson, K. AY. AVagnon, University
of California ; AV. P. Dasmann, Ben Glading, Dan Tillotson, Department
of Fish and Game ; L. T. Burcham, State Division of Forestrv ; F. P.
Cronemiller, Al. AA\ Talbot, U. S. Forest Service ; R. AI. Bond, Soil Con-
servation Service; G. H. Sharrer, Bureau of Land Alanagement; E. N.
Dye, Ralph Leavers, California Farm Bureau ; X. AI. Hughes, Associated
Si)ortsmen ; Glen Keithly, rancher.


The brushlands studied are typical of western Lake County and much
of the North Coast ranges. Data on vegetation, soils, and climate are taken
from the study areas, or nearby, but are applicable to a much greater


In general the chamise brushlands in Lake County comprise two cover
types, those in which chamise (Adenostema fasciculatum) predominates
and those which contain a mixture of broad-sclerophyll shrubs and small


FIGURE 1. Close view of chamise brushland in Lake County. Chamise predominates on the south-
facing (S) exposures while a mixture of taller broad-sclerophyll shrubs and small trees ore found
on the north-facing (N) slopes. Lake County brushlands hove long been famous deer hunting
grounds. Water from springs is well distributed in the ravines.


trees. The chamise occurs mainly on sonth-facing slopes and drier sites
while the mixed chaparral is found on the more mesic north exposures
and in ravines (Figure 1). This intermixture of chamise and mixed
chaparral is especially favorable for deer. A majority of the shrubs and
small trees are good or excellent browse species. The intermixture of
browse plants in Lake County is probably as favorable for deer as most
chamise brushlands in other portions of the State. Some brushlands are
so nearly pure chamise that they furnish relatively poor browse.

On south exposures in Lake County chamise often occurs in relatively
pure stands, depending primarily upon the age of the stand and fire
history, but usuall}' it occurs with admixtures of lesser amounts of wedge-
leaf ceanothus (Ceanothus cnneatus), wavyleaf ceanothus (C. foUosus),
Stanford manzanita (Arctosfaphylos sianforcliana) , poison oak (Rhus
diversiloha) , yerba santa (Eriodicti/on calif o mi cum) , and others. The
ceanothus species above, which are nonsprouters, are in greater abund-
ance in young or recently burned chamise stands that have not been
repeatedly burned in close succession.

The north-facing slopes are vegetated chiefly by interior live oak
{Quercus ivislizenii), scrub oak (Q. dumosa), Eastwood manzanita
{Arctostaphylos glandidosa), California laurel {VmhelUdaria cali-
fornica), toyon {Photinia arhuti folia), birchleaf mahogany (Cerco-
carpus hettdoides) , deerbrush {Ceanothus iniegerrimus) , chamise, and
others, roughly in that order of abundance. Interior live oak frequently
makes up half of the cover.

Other species may predominate or become more important in special
sites. In wetter situations, madrone {Arl^uius menziesii) and canyon live
oak {Quercus chrysolepis) are common. Knob-cone pine {Pinus tuhercu-
lata) is frequent on certain soil types. On ridgetops chaparral pea
' {Pickeringia montana) is more abundant than on either north- or south-
facing slopes.

The brush canopy on all sites is usually so complete as to preclude
much herbaceous vegetation as understory.


The soils on south exposures where chamise occurs in the study areas
are derived from consolidated sedimentarv rocks. The shallower and least


productive ones have developed on parent materials which have under-
gone the least metamorphosis. The Maymen series is typical of this group
and occurs most extensively. These soils are less than 12 inches deep and
are light brown in color, moderately acid, and vary from clay to clay
loam in texture. Thev are relativelv low in fertilitv. Soils of the Dorado

ft- «, «.

series, which usually occupy the lower edge of the chamise slopes, are
formed on sediments having undergone an intermediate amount of
metamorphosis. They are deeper than the Maymen, 12 to 18 inches, less
acid, and light reddish-brown in color.

On north exposures the soils are generally deeper and darker in color
than those on south exposures. Some are a deeper phase of Ma^^men,
ranging in depth from 12 to 24 inches. The Los Gatos series, inter-
mingled with the Maymen and usually supporting a mixed stand of
brush with occasional knob-cone pine trees, are soils developed on the
more highly metamorphosed sediments and are general underlain by more



massive rocks. The}^ are 18 to 24 inches in depth, moderately acid on the
surface, and are reddish-brown in coh)r. Laughlin soils may occur where
chamise contacts or invades into Avoodland-grass. This series is charac-
terized by depths of 24 to 30 inches, medium to fine texture, and slight
acidity. The soils are light brown to light reddish-brown in color and are
formed on residual parent material. They are relatively fertile soils and
support fairly heavy stands of herbaceous vegetation where the brush
is removed or thinned out.

Soils from the study areas were tested for fertility by using pot tests
conducted in the greenhouse by the Soils Division of the University of

FIGURE 2. Chamise brushlands in Lake County. Some are not excessively steep while others are

rugged and somewhat inaccessible.


California at Berkeley. Results of these tests indicate that chamise-
covered soils are generally low in nitrogen even when the woody vegeta-
tion has not been disturbed over a long period of time.

It is generally recognized that a majority of chamise brushland soils
are low in fertility. On the study areas the Maymen soil was the most
extensive and least productive ; perhaps it forms a larger area than any
other chamise brushland soil in California. Probably the second most
extensive chamise brushland soil in California is Los Gatos. This is con-
sidered more productive than Maymen. Another frequently occurring
chamise soil, perhaps the third in rank, has not yet been named but con-
tains a large amount of granitic material. The fertility of the latter soil
is considered about the same as that of Maymen. Serpentine soils are
about the poorest of all. Chamise also occurs on several other soils, a
majority of which are more fertile than those named above. Some are
relatively deep and have been cleared for agricultural crops.

Slopes where chamise occurs are generally rather steep (Figure 2). On
the study areas in Lake County they averaged about 20 to 25 degrees. In
some chamise brushlands the average slope is greater,


The weather station nearest to the study areas is at Lakeport, on the
west shore of Clear Lake, about three to eight miles away. The mean
annual temperature recorded there is 57 degrees F., the extremes varying
generally from about 20 degrees to 110 degrees.

Mean annual precipitation, based on a 30-year average, is 28 inches.
Practically all of this comes as rain between late September and April,
inclusive; consequently, the summer months are extremely dry. Since
the vegetation becomes very dry during summer, the possibility of large
acreages of brush being destroyed by fire is high. Water becomes less
abundant along drainage ways as the dry season advances but many of
the springs maintain their flow throughout the summer.

The same general pattern of temperatures and rainfall characterizes
all chamise brushlands in California. The brushlands usually occur where
the precipitation is between 14 and 40 inches. Where rainfall is 14 inches
and below the chamise becomes open and desert-like in appearance and
above 40 inches it generally gives way to forest growth. In areas receiving
between 14 and 40 inches of rainfall the soil is apparentl}^ more impor-
tant than climate in delimiting chamise brushlands.


Chamise brushlands have been generally looked upon as valuable for
game and as watersheds. Some are used in sheep production. They are
more suitable for sheep than cattle because of steepness of slopes and
the predominance of browse. Poisonous plants occur on many chamise
brushlands. Little effort has been made to manage these lands.

Usually the intention of public agencies has been to protect chamise
brushlands from fire, but, in spite of this, wildfires occur frequently
(Figure 3) . Some of the wildfires have been large and destructive (Figure
4). Some people wonder if large fires are not to be expected where total
protection is attempted. As a result of protection the brush is permitted





FIGURE 3. The shaded portions Indicate areas burned by wildfires. Much of the unshaded por-
tion is agricultural land. It appears that a majority of the brushlands in the southern half of Lake
County burn at least one time in each 20 years. Fire records were furnished by the State Division
of Forestry. The map does not show all of the fires but attempts to show all the area burned at

least once by wildfires in each of the 10-year periods.

to grow lip uniformly and within about 25 years much of it becomes dead
fuel; this material greatly increases the fire hazard. AVlien fires are
started in this kind of cover they are difficult to control and are likely
to be large and destructive. Limited "spot burning" in the spring
months has been suggested as a means of reducing the hazard of such fires.
In fact, burning of this nature was initiated in Lake County in 1950-
1951 by personnel of the county board of supervisors, State Division of
Forestry, county farm advisor. Fish and Game, and private landowners.
Plans call for limited burning in the spring to reduce fire hazard around
cultural developments and also to decrease the possibility that large
wildfires will occur during summer. Another purpose of spot burning is
to improve conditions for game.

Areas burned by wildfires are usually not reseeded and the soil is
nearly bare the following winter. Tn the second Avinter the soil is fairly
Avell covered with sprouts and brush seedlings. Then within 8 to 12
years the brush stand is well developed and becomes nearly impenetrable.
During these years little dead wood or litter is produced and fire presents
no particular problem (Buck, 1951). If it were not for this period of



FIGURE 4. Wildfire burned area of 6,000 acres in Lake County. Wildfires are frequent and often
large and destructive. In this case the fire burned so intensely that insufficient unburned brush

remained as cover for game.

reduced fuel following wildfire burns in dense brush, fires would probably
occur more frequently than at present and would also be larger and more
uniform in intensity.

Game populations in or around wildfire burns usually build up while
the brush is young and palatable but then decrease or move out when the
brush grows up. Of recent date controlled burning and other manipula-
tion of brush to improve conditions for game has gained popularity. Some
areas are reseeded to herbaceous plants and managed following brush
removal ; some are not.

There has been some question as to watershed conditions under chamise
brush and the effects of fire on erosion. Investigations by Yeihmeyer
(1951) have shown that small plots burned annually convert largely to
grass and in this condition there is less erosion than from undisturbed
areas of chamise. Studies by Colman (1951) and others have shown that
erosion accelerates following occasional brush fires in chamise and may
continue for at least eight years. The latter studies were in areas not
reseeded to grasses, where little herbaceous vegetation occurred. Both
sets of studies would indicate that chamise is a relatively poor watershed
cover, especially since such brushlfinds burn frequently. More studies are
needed on the feasibility of converting chamise brushlands to a cover
better adapted to prevent erosion. Both grass and trees offer considerable
promise for this purpose but studies on the possibilities of manipulating
vegetation to improve watershed conditions have scarcely begun in

It has been observed many times that the surface soil is loose following
fires in dense brush and is easily moved downhill by deer feeding over
slopes. On steep slopes considerable soil may gradually be moved to the
bottoms of drainagewavs. This is washed awav during winter storms. On
the other hand, the soil is usually firm in areas of burned grass and little
movement takes place. Again, the observations indicate that grass shows


promise of lessening soil erosion in the presence of fires. Differences be-
tween grass and brush covers may be summarized as follows: Fires in
grass are less intense than those in brush ; the grass seeds and residue are
not all destroyed ; the grass begins growth following fall rains and soon
covers the soil ; the soil is firm following fire. In dense brush, on the other
hand, the fires are intense ; the soil is loose and bare the following winter ;
the shrubby vegetation recovers slowly.



Game populations were studied in Lake County under three different
conditions of chamise brushland : (1) HeaYj brush cover protected from
fire ; (2) Wildfire burn, in which there were a few unburned islands the
first winter, followed by the presence of brush sprouts, seedlings, and a
small quantity of herbaceous cover the second year; (3) Opened brush,
consisting of small burned patches within dense brush seeded to suitable
herbaceous plants (Figure 5). Area of each condition was about 1,000

Detailed investigations were made on the Columbian blacktailed deer
{Odocoileus hemioniis colum'hianus) . During the course of this work ob-
servations were made of the California jackrabbit {Lepus calif ornicus),
brush rabbit (Sylvilagus hachmani) , mountain quail {Oreortryx picta),
valley quail {Lophortyx calif ornica) , and mourning dove {Zenaidura

Deer Populations

The deer of this region are resident rather than migratory, but they
do make short seasonal movements which depend on weather conditions
and food supplies. Population densities were determined by pellet
censuses checked by sight records. The number of deer on the study areas
in heavy brush ranged from averages of 10 to 30 to the square mile, in
the wildfire burns from 5 to 160, and in the opened brush from 40 to
110. AVhere surrounding food conditions are poor, a wildfire burn of
newly sprouting brush will attract large numbers of deer. However, all
of these deer may move to better cover in bad weather ; also, the burned
area loses its attractiveness very rapidly as the sprouts grow up and
become less palatable. Over a period of years extensive wildfire burns
support lower average deer populations than opened brush because there
may not be enough deer to keep the sprout growth in a palatable stage.

Search for basic factors governing deer population densities in the
different brush covers included the investigation of ia^vn production, the
weights or relative condition of individual deer on the three study areas,
and food habits and comparative nutrition.

Fawn Production

Studies on collected does indicate that fawn production is governed
largely by ovulation rate. 0\Tilation rates in adult does (18 months and
older) were approximately as follows: In hea^y brush, 84 percent; on
wildfire burn. 116 percent; in opened brush, 147 percent (Taber, 1953).
Although the doe collection consisted of only 42 individuals, reliance on
these figures is strengthened by the fact that they correspond closely to



FIGURE 5. Areas of opened brush produced by spot burning. The interspersion of grass and

brush forms a favorable habitat for deer. During late winter and early spring the deer graze

heavily on the grasses. The spots of dense brush form necessary cover.

the ratio of fawns to 100 adult does following the riit. This ratio varies
somewhat from year to year, but the following values seem to be repre-
sentative for the areas under studv : in heavv brush, about 60 to 85 fawns
per 100 adult does; on the wildfire burn about 100 to 110 fawns per 100
adult does ; in opened brush, about 115 to 140 fawns per 100 adult does.
A change in the f aAvn : doe ratio between the fawn drop and the rut obvi-
ously may be brought about by either fa^^1l or doe mortality. Therefore
the ratios do not necessarily represent an extremely high fawn survival.
Actually there is evidence of some mortality among both does and fawns
during the summer.



Deer Weights

Skeletal size differences between deer of the same age for the different
brush cover conditions seem to be either small or absent. Therefore, com-
paring weights of deer of the same age gives an indication of relative
condition rather than comparative size.

The weights of 23 bucks taken from the heavy brush and the opened

brush during the hunting season are compared in Table 1. The bucks are

grouped by antler beam diameter, the first class (15-19 mm.) containing

mostly young bucks and the second class (19.1-23 mm.) containing those

of medium age. AVeights were obtained with the paunch out but all other

organs present.


Field-dressed Weights of 23 Bucks From Two Conditions of Brusli Cover, 1949

(.2 Level of Significance)

Heavy brush

Opened brush

Beam diameter

Weight, pounds

of deer

Weight, pounds

of deer

15-19 mm. _ - _



7.5.. 5 ±4. 99


19. 1-23 mm.


Bucks from opened brush tend to be heavier than those from heavy

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