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CAUFDM

FISH-GAME

"CONSERVATION OF WILDLIFE THROUGH EDUCATION"




Caiifornia Fish and Game is a journal devoted to the con-
servation of wildlife. Its contents may be reproduced elsewhere
provided credit is given the authors and the California Depart-
ment of Fish and Game.

The free mailing list is limited by budgetary considerations to
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viduals must state their affiliation and position when submitting
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scribers are osked to report changes in address without delay.

Please direct correspondence to:

JOHN E. FITCH, Editor
State Fisheries Laboratory
511 Tuna Street
Terminal Island, California

Individuals and organizations who do not qualify for the free
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individual issues for $0.75 per copy by placing their orders with
the Printing Division, Documents Section, Sacramento 14, Cali-
fornia. Money orders or checks should be made out to Printing
Division, Documents Section.



u






VOLUME 49



JANUARY 1963







V



NUMBER 1




Published Quarterly by

THE RESOURCES AGENCY OF CALIFORNIA

CALIFORNIA DEPARTMENT OF FISH AND GAME

SACRAMENTO



STATE OF CALIFORNIA

EDMUND G. BROWN, Governor

THE RESOURCES AGENCY OF CALIFORNIA

WILLIAM E. WARNE, Ac/mm/sfrafor



FISH AND GAME COMMISSION

WILLIAM P. ELSER, President, San Diego

JAMIE H. SMITH, Vice President HENRY CLINESCHMIDT, Member

Los Angeles Redding

DANTE J. NOMELLINI, Member THOMAS H. RICHARDS, JR., Member

Stockton Sacramento



DEPARTMENT OF FISH AND GAME

WALTER T. SHANNON, Director

OFFICE— FISH AND GAME COMMISSION

722 Capitol Avenue
Sacramento 14



OFFICES— DEPARTMENT OF FISH AND GAME

722 Capitol Avenue

Sacramento 14

1001 Jedsmith Drive Ferry Building 271 Tyler Street

Sacramento San Francisco Monterey

1234 East Shaw Avenue 217 West First Street 619 Second Street

Fresno Los Angeles Eureka

627 Cypress Street 511 Tuna Street Room 12, North Ramp

Redding Terminal Island Broadway Pier Building



407 West Line Street
Bishop



San Diego



CALIFORNIA FISH AND GAME
Editorial Staff

JOHN E. FITCH, Editor-in-Chief.__ Terminal Island

DAVID P, BORGESON, Editor for Inland Fisheries Sacramento

ALBERT E. NAYLOR, Editor for Game Sacramento

JOHN L. BAXTER, Editor for Marine Resources Terminal Island

DONALD H. FRY, JR., Editor for Salmon and Steelhead Sacramento



TABLE OF CONTENTS

Page
Abundance and Scarcity in California Deer

W. P. Dasmann mid E. F. Dasmann 4

Trout Survival in Taylor Creek, a Tributary
of Lake Tahoe, California Garth I. Murphy 16

Synonymy, Characters, and Variation of Gila
crassicauda, a Rare Californian IMinnow, With
an Account of Its Hybridization With
Lavmia exilicauda Robert Rush Miller 20

Mission Bay, a Review of Previous Studies

and the Status of the Sportfishery Gordon A. Chapman 30

Variant Hemoglobin and Electrophoretic Whole
Blood Studies in Two Tunas and Three
Other Fish Species Albert C. Smith 44

A New Northern Record of the

Smooth Stargazer Harry L. Fierstine and Robert G. Werner 50

Notes

The First California Record of Sierra,
Scomber omorus sierra Jordan and Starks
Bruce B. Collette, Frank H. Talbot and Richard E. Rosenblatt 53

The Schooling Behavior of Pacific Yellowfin and
Skipjack Tuna Held in a Bait Well

James Joseph and Izadore Barrett 55

Reviews 56



(3)



ABUNDANCE AND SCARCITY IN CALIFORNIA DEER'

W. p. DASMANN
U.S. Forest Service, San Francisco ,

and

R. F. DASMANN

Museum of Vertebrate Zoology
University of California, Berkeley

INTRODUCTION

Fluctuations in abundance of animals have been of interest to
wildlife managers for many decades. Success or failure of a manage-
ment program is usually judged by the number of game animals it
has produced for the hunter. The game manager's efforts often are
obscured by sudden changes in game abundance brought about by
poorly understood factors. The 10-year cycle of abundance and scarcity
in northern mammals, for instance, is yet to be comprehended or con-
trolled completely.

There is evidence that California deer populations have been under-
going marked fluctuations during the past several decades. These
changes in abundance bear no apparent relationship to hunting pres-
sure, management effort, or any other single factor that could be de-
fined. A consideration of these changes can be of value toward under-
standing the behavior of animal populations in general, and can be of
immediate significance in evaluating deer management programs.

Where hunting is regular and sustained, the annual kill should reflect
game population abundance. In California, deer hunters have been
required to report the deer they kill, by returning deer tags, since 1927.
While the deer tag return may not represent total kill, there is no
reason to believe that the percentage of people failing to return tags
has fluctuated greatly over the years. Hunting, since 1927, has largely
been confined to bucks with forked antlers. The buck kill will therefore
be considered here as an index to deer abundance.

Many factors other than game abundance may affect the total kill.
Most obvious is the number of hunters. This has increased markedly
since 1927, and along with it has come an upward trend in bucks killed.
This does not necessarily reflect an increase in deer numbers, but is at
least partially the result of the buildup in hunting pressure, coupled
with easier access to hunting territory. Other factors which may affect
deer kill are length and time of hunting season, weather during the
season, and weather cycles as they modify water and forage distribu-
tion. However, none of these factors explain the major fluctuations in
the California buck kill. Changes in deer abundance appear to be the
principal cause of the kill fluctuations.

1 Submitted for publication September 1962.



(4)



DEER ABUNDANCE AND SCARCITY



THE STATEWIDE PICTURE

Based on hunting information, major peaks and troughs in the buck
kill occur approximately in 10-year intervals (Figure 1).

Although deer tag sales showed a fourfold increase during the 1927-
1960 period, there were sales declines during the middepression (1932-
33), during World War II (1942 in particular), and in 1957-1958.
These hunting-effort decreases could explain the low deer kill, were
it not for the lack of correspondence in the hunting-success ratio. Thus,
the 1940 peak in deer kill was followed by an increased number of
hunters in 1941. In 1941, however, the hunting-success ratio and deer
kill fell off, suggesting that deer had either decreased or were harder to
kill. During 1942 and 1943, the much-reduced number of hunters did
not find deer easier to kill. Instead, the hunting-success ratio continued
to fall to a low in 1943, paralleling the low in deer kill. Reduced deer
abundance is again suggested. Following the 1954 peak in deer kill
and hunting success, a similar pattern is shown. The numbers of
hunters increased for two more years, while the deer kill and the suc-
cess ratio declined.



NUMBER OF DEER TAGS SOLO




FIGURE 1. Relation of statewide buck kill with deer tag sales and hunting success ratios.
Solid vertical lines show years of high kill; broken vertical lines indicate years of low kill.

However, if the statewide buck kill is examined critically, county
by county, it does not show any marked synchrony throughout the
various regions of California. Deer do not increase and decrease in all
areas simultaneously. In the years of major statewide peaks, 1931, 1940,
1954, a high percentage of the counties did produce peak kills and
most of the other counties peaked either one year before or one year
after. There is thus an indication that throughout the State, condi-
tions have favored high deer kills in certain series of years, and low
deer kills in others, but local or regional exceptions occurred.



CALIFORNIA FISH AND GAME



Vegetation, climate and conditions for deer vary too greatly through-
out California to permit more detailed analysis of the statewide kill.
In order to carry this consideration further, a group of counties was
selected in the north coastal region of the State within which climate,
vegetation and hunting conditions are generally similar.

DESCRIPTION OF AREA AND DEER

The area selected embraces Marin, Napa, Sonoma, Lake and Mendo-
cino Counties lying just north of San Francisco Bay. Two other coun-
ties, Humboldt and Del Norte, lie between Mendocino and the Oregon
border. All of the counties lie within the coastal mountain ranges and
are characterized by fairly large valleys (mostly devoted to agricul-
ture), rolling foothills, and rather steep mountains rising to elevations
up to 7,500 feet. Vegetation ranges through grassland, savannah and
oak woodland in the valleys and low foothills. Chaparral brush species
grow on higher foothills and a conifer cover is found in the moun-
tains. Redwood (Sequoia sempervirens) stands occur on north slopes
and in some of the more humid bottoms.

The deer in north coastal California belong to the Columbian black-
tailed race of mule deer (Odocoileus heniio7ius columhianus) . Those
found in Marin, Napa, Sonoma, southern Lake and western Mendocino
Counties tend to be nonmigratory, although some may move up to a
mile or so between summer and winter ranges. In northern Lake and
eastern Mendocino Counties, where more mountainous terrain and
higher elevations occur, definite migratory patterns exist. Here, popula-
tion ceilings for the separate herds may be set by their more limited
seasonal ranges.




FIGURE 2. Comparison of deer kills in six north coastal counties. Solid vertical lines show

years of general high kill; broken ve'-ticai lines indicate years of general low kill. There was

no open season in Lake, Marin, Mendocino, Mapa or Sonoma Counties during 1943.



DEER ABUNDANCE AND SCARCITY 7

TABLE 1

Reported Buck Kill *

Six Counties in Northwestern California

Year Marin Sonoma Lake Mendocino Napa Total Humloldt Status

1927 367 751 901 1,475 442 3,936 821

1928 444 753 1,038 1,468 569 4,272 777

1929 394 732 841 1,355 523 3,845 689

1930 403 865 885 1,483 536 4,172 917

1931 449 903 726 1,706 488 4,272 1,069 High

1932 376 709 524 1,273 304 3,186 807

1933 301 748 481 1,234 285 3,049 842

1934 341 704 419 1,185 288 2,937 877 Low

1935 328 554 570 1,207 278 2,937 921

1936 411 536 757 1,372 415 3,491 796

1937 482 744 1,418 2,072 544 5,260 940

1938 534 948 1,803 2,700 795 3,780 1,284

1939 649 1,094 2,103 2,967 927 7,740 932

1940 581 1,242 2,328 3,517 907 8,575 1,791 High

1941 615 1,005 1,805 3,460 962 7,847 2,200

1942 403 489 1,042 1,652 557 4,143 1,101 Low

1943 — No open season 1,068

1944 579 685 1,787 2,297 932 6,280 945

1945 438 787 1,674 2,365 809 6,073 1,133

1946 ^_ 520 1,102 1,768 2,980 887 7,257 1,459

1947 488 1,151 1,835 3,067 899 7,440 1,335

1948 634 1,505 2,120 3,627 1,027 8,913 2,083 High

1949 597 1,269 2,182 3,354 1,020 8,422 1,962

1950 544 1,138 1,942 2,927 952 7,513 1,770 Low

1951 767 1,447 2,155 3,665 983 9,017 2,313

1952 832 1,553 2,056 4,252 1,220 9,913 1,792

1953 885 1,679 2,146 4,394 1,161 10,265 2,323

1954 1,048 1,979 2,549 5,232 1,386 12,194 3,055 High

1955 901 1,586 2,479 4,587 1,329 10,882 3,408

1956 707 1,303 2,030 4,051 1,144 9,235 3,393

1957 548 1,267 1,585 3,847 1,045 8,292 3,631

1958 613 1,172 1,568 3,754 994 8,101 3,047

1959 407 1,016 1,684 3,655 997 7,759 3,649 Low

1960 659 1,483 1,706 4,426 1,306 9,580 4,214

1961 749 1,883 1,763 4,585 1,356 10,336 4,486

* From Game Management Handbook, California Department of Fish and Game.

The deer kill in five of the California north coastal counties (Table 1)
shows a marked upward trend related to the increase in hunters, and
also a marked periodicity. The high and low kills occur simultaneously-
over a broad area, although sometimes parts of the area will reach a
high or low one year before or after the balance. A review of the data
reveals that low kills have occurred every eight or nine years and high
kills at six- to nine-year intervals (Figure 2),

HUNTING PRESSURE

We have no way of knowing how many people hunted annually in
the five counties during the 35-year period, 1927-1961, under consid-
eration. We can only assume that the number of hunters seeking deer
in this part of California yearly was proportional to the total people
purchasing deer tags in the State. Accepting the assumption that state
deer tag saJes are indicative of annual hunting pressure in the five

2—74951



8



CALIFORNIA FISH AND GAME



counties, these figures are compared with the total five-county deer kill



in Figure 3.



480



420



360 -



Q

< 300
CO

o

X



240



<



<



cc

LiJ



180



120



60



NUMBER OF DEER TAGS SOLD




J — I I I I i—i I I I I I I I I I I I I I I I I I I I

1930 1940 1950

YEARS



J-J I I I I I I



14,400



12,600



10,800



9000



7200



5400



UJ

I-
z

o
o

UJ

>



2
Q






UJ

o



UJ

m



3600



1800



i960



FIGURE 3. The relation of buck kill to hunting pressure in the five counties under study.

The immediate reaction is that a close correlation between the ups
and downs of kill and of hunting pressure has existed. However, a
closer inspection will show, with the exception of 1931, the decline in
hunting pressure has followed the reduction in deer kill by one or more
years. It is suggested that the news deer are in a decline results in
fewer people purchasing tags. It would also appear that preseason
information spurs tag sales once the deer population shows signs of
recovery.



DEER ABUNDANCE AND SCARCITY 9

Legal hunting in the five connties has been confined to bucks, forked-
horn or better, throughout the 35-year period, with two exceptions.
In 1955, 452 antlerless deer were taken in Napa County during a spe-
cial hunt (Ferrel, 1956) and in 1956 when that county was open for
three days to either-sex deer hunting, 461 antlerless deer were removed
(Dasraann et al., 1958). Removing these antlerless deer appears to
have been insufficient to affect the kill cycle in Napa County. This con-
tinued to behave in close correlation with other counties where no
antlerless deer removals were made.

Since deer are polygamous, removing a portion of the males should
not affect annual production. At least half of the buck deer taken from
coastal ranges in California are more than two years old and many
are not taken until four or five years old (California Department of
Fish and Game, 1955). When such a high percentage of males escapes
the gun each year, hunting removals cannot prevent a deer population
from increasing. There are more than enough left for breeding. We
can safely assume heavy buck removal in peak years has not been the
cause of subsequent prolonged declines in the take.

HUNTING SEASONS

All five counties have enjoyed the mixed blessing of one of the
earliest deer seasons in the United States. Deer are hunted in mid-
summer, when temperatures may reach 100° F. or more, and success-
ful nimrods perspire in efforts to prevent meat spoilage. During the
time hunting seasons were established by the State Legislature (1927-
1945), the season for Marin, Napa and Sonoma Counties invariably
was set for August 1 to September 14 or 15. In Lake and Mendocino
Counties, however, the season was periodically reshuffled. Occasionally,
these two counties were open to hunting at the same time as the others,
but often, northern Lake County and eastern Mendocino were opened
two weeks later (August 16) yet were closed on September 15. For a
four-year period (1935-38), deer were hunted in Lake County a full
six weeks later than the other four counties. During this period, it
was included with the largest part of California, and with Humboldt
and Del Norte Counties to the north, in a September 16 to October 15
hunting season.

After the authority to establish hunting seasons was granted to the
Fish and Game Commission, the five counties, along with a consider-
able block to the south and east, were opened and closed at the same
time. The area still has an early season, but the opening dates have
ranged from July 25 to August 11, and the closing dates from Septem-
ber 12 to 25.

The tradition of the midsummer hunt has developed from the be-
lief that deer meat will be tainted and bucks will become too vulnerable,
if the hunting season extends into the period of rut. It is true that
the breeding season is quite early in parts of some counties imme-
diately adjacent to the seacoast and most notably in Marin (Cook
et al., 1949). Here, breeding may start in September or early October,
and neck swelling in bucks occurs a week or two before that. But
on most of the blacktailed deer range, the breeding periods, according
to Bischoff (1957) occur from late October to early January. The
shifting of the hunting seasons in the five counties concerned, and



10 CALIFORNIA FISH AND GAME

particularly the late season in Lake County during the 1939-1942
period, does not appear to have affected the periodicity of high and
low kills. In fact, the Lake County kill reached a high simultaneously
with other counties in 1940, and a low in 1942 apparently without
respect to the placement of the hunting season.

EFFECT OF WEATHER

August and the first half of September are normally hot and dry
in the five counties. A check was made to determine if the occasional
rain in August might show a correlation with high and low deer kills.
Some rain occurred in one or more of the counties in 18 out of the
35 years. Usually only a trace occurred, but in eight years measurable
precipitation fell. When these years of rainfall were compared with
years of high and low kill, the following lack of relationships was
revealed :

1929 — Kill lower than that of the year preceding or following.

19.35 — Kill at a low part of the cycle.

1941 — Kill relatively high, but lower than the peak.

1951 — Kill just starting up from a low to a peak which occurred three years

later.
1953 — Kill at a high level, but lower than peak.
1954 — Kill at a record peak.
. 1959 — Kill at a low.
1961— Kill at a high level.

No general rains occurred during the other years of peak kill. One
general rain occurred in 1935, an extended low, and one in 1959, but
none in the other years of low kill. To have the most effect on deer
take, rain must occur just prior to and during the opening days of
the season when most hunters are afield and a large part of the total
kill is made. Because of the lack of correlation shown above, and
because a similar comparison of July rainfalls was equally nonproduc-
tive, we decided not to pursue the matter further. The occurrence of
rain during the opening month of the hunting season apparently has
no significant effect on either the peaks or the troughs of kill fluc-
tuations.

WEATHER CYCLES

"Weather cycles may influence deer abundance and kill through their
effect on forage production and water distribution. Because the data
for Mendocino County are incomplete, the averages in Figure 4 are
for the other four counties. Precipitation trends in Mendocino County,
as indicated by the partial record, were in conformity with the four-
county trends.

One would expect a comparison of precipitation with deer kill from
a stable population to show higher kills during drought periods when
deer tend to concentrate around permanent water than during wet
periods when they are scattered. But the opposite generally has oc-
curred in the five counties.

The coincidence of peaks and troughs of precipitation with high
and low buck kills was fairly close from 1927 to 1949. Major and minor
peaks for both kill and precipitation occurred in 1931, 1937, 1940 and
1948, and lows for both elements occurred simultaneously in 1929,
1934 and 1943. However, the relationship loosened after 1949. The



DEER ABUNDANCE AND SCARCITY



11



66



60



54



48



42



5: 36

o

LlI

tr

Q.



u.
o



30



24



18



12 -





FOUR-COUNTY AVERAGE




^35 YEAR
AVERAGE



I I I I



_L_I I I I L



_L_I L



I I I I I I I ' I



1930



1940



1950



I960



YEARS
FIGURE 4. Annual precipitation in the five-county area.



record kill of 1954 was coincident with a minor precipitation peak
and the low kill of 1959 occurred simultaneously with a moderate pre-
cipitation low. Otherwise, a close relationship is not evident after 1949.
No explanation for the change from a close to loose relationship is
offered; however, the earlier period of close relationship was one of



12 CALIFORNIA FISH AND GAME

rather violent fluctuation in precipitation. Both the wettest and the
driest years fell within this period. In contrast, the period since 1949
has been one of relatively moderate fluctuation and characterized by
a higher average annual rainfall (36 inches). If the deer kill is a reflec-
tion of population abundance, one could expect fairly close relationship
of kill to precipitation if deer are crowding the upper limits of range
carrying capacity, particularly during a period of extremes. Such a
relationship appears to have occurred within the five counties.

DEER POPULATION FLUCTUATIONS

Because the periodicity of deer-kill highs and lows in the five counties
appear unrelated to the various factors we have considered, except
precipitation, it is believed they reflect animal abundance. Possibly the
population cycles reflected in these kills are a product of habitat supply
and deer demand. Starting in a period of high habitat supply and low
animal demand, with no shortages to affect health, thrift or reproduc-
tion, the deer multiply. After a period of years, the animals more fully
occupy the range and reach a population peak imposed by the habitat
ceiling. Now there is a high animal demand for a short habitat supply.
Depressant factors are brought into play. Only the strongest are able
to fill their needs, and one animal must die to make room for another.
The deer are living along the margin of disaster. A drop in food pro-
duction, appearance of disease, spread of parasites among unthrifty
animals, accumulating stress of competition for proper food, water,
cover and living space — any or all of these factors may be the cause of
snowballing mortality, resulting in a sharp population decline. With
deer numbers once again at a low, the cycle repeats itself.

That some such mechanism may be fundamental to population fluctu-
ations in the five counties is indicated by the record of what has hap-
pened in the sixth county, i.e. Humboldt. The deer in Humboldt County
showed close relationship with the other five counties from 1927 to 1950
(Figure 2). After 1950, this kill showed general correspondence with
that of Lake County until 1954, when it broke away from the general
pattern and started a sustained climb. In an effort to discover a
possible cause of this phenomenon, lumber production records of the
counties were examined (Figure 5).

Humboldt County supports largely a conifer cover. Areas having
unbroken mature stands of conifers usually make poor deer habitat,
because of the low volume and poor quality of herbaceous and shrubby
cover growing in the shade beneath them (Einarsen, 1946). But when
the forests are opened by logging, a profusion of palatable undergrowth
will often appear on the site and good-quality deer habitat is created.

Dasmann and Hines (1959) found that virgin redwood and mixed
coniferous forests supported few deer. During the first five years after
logging, these animals increased to a level 7 times greater, and in the
next five years to a peak population over 20 times greater than existed
in the virgin forests. Deer population densities of 11 to 66 animals
per square mile were found in one cutover area 10 to 13 years after
logging. About 10 years after logging, deer numbers began to decline
and after 20 years the population was not significantly higher than
before logging.



DEER ABUNDANCE AND SCARCITY



13



1600






il (


1400


-


.


A


1200






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1000
800


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HUMBOLDT CO.-

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1940



1950



I960



YEARS


1 3 4 5 6 7

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