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CAUFORNIAl
FISH-GAME

"COKSKRVATION OF WILDLIFE THROUGH EDUCATION"




California Fish and Game is a journal devoted to the conservation of v/ild-
life. If its contents are reproduced elsev/here, the authors and the California
Department of Fish and Game would appreciate being acknowledged.

Subscriptions may be obtained at the rate of $5 per year by placing an
order with the California Department of Fish and Game, 1416 Ninth Street,
Sacramento, California 95814. Money orders and checks should be made out
to California Department of Fish and Game. Inquiries regarding paid sub-
scriptions should be directed to the Editor.

Complimentary subscriptions are granted, on a limited basis, to libraries,
scientific and educational institutions, conservation agencies, and on exchange.
Complimentary subscriptions must be renewed annually by returning the post-
card enclosed with each October issue.



Please direct correspondence to:

Kenneth A. Hashagen, Jr., Editor
California Fish and Game
1416 Ninth Street
Sacramento, California 95814



129



Ij













VOLUME 66



JULY 1980



NUMBER 3




Published Quarterly by

STATE OF CALIFORNIA

THE RESOURCES AGENCY

DEPARTMENT OF FISH AND GAME

—LDA—



130 CALIFORNIA FISH AND CAME

STATE OF CALIFORNIA
EDMUND G. BROWN JR., Governor



THE RESOURCES AGENCY
HUEY D. JOHNSON, Secretary for Resources



FISH AND GAME COMMISSION

SHERMAN CHICKERING, President
San Francisco

ELIZABETH L. VENRICK, Vice President ABEL GALETTI, IVIember

Cardiff Los Angeles

BERGER C. BENSON, Member RAYMOND DASMANN, Member

San Mateo Nevada City



DEPARTMENT OF FISH AND GAME
E. C. FULLERTON, Director

1416 9th Street
Sacramento 95814



CALIFORNIA FISH AND GAME
Editorial Staff

KENNETH A. HASHAGEN, JR., Editor-in-Chief Sacramento

DARLENE A. OSBORNE, Editor for Inland Fishenes Sacramento

RONALD M. JUREK, Editor for Wildlife Sacramento

J. R. RAYMOND ALLY, Editor for Manne Resources Long Beach

DAVID A. HOOPAUGH, Editor for Salmon pnd Steelhead Sacramento

DONALD E. STEVENS, Editor for Striped Bass, Sturgeon, and Shad Stockton

KIM McCLENEGHAN, Editor for Environmental Services ' Rancho Cordova



CONTENTS



131



Page



Impact of Florida Largemouth Bass, Micropterus salmoides
floridanus. Introductions at Selected Northern California
Waters, With a Discussion of the Use of Meristics for De-
tecting Introgression and for Classifying Individual Fish of
Intergraded Populations Ronald J. Pelznnan 133

Exploitation, Natural Mortality, and Survival of Smallnnouth

Bass and Largemouth Bass in Shasta Lake, California

William F. Van Woert 163

Diet and Behavioral Aspects of the Wolf-Eel, Anarrhichthys
ocellatus, on Sandy Bottom in Monterey Bay, California ..
Larry W. FHulberg and Patsy Graber 172

Decline of the Lake Greenhaven Sacramento Perch
Population C. David Vanicek 178

Notes

A Population of the Endangered Santa Cruz Long-Toed Sala-
mander, Ambystoma macrodactylum croceum, from Mon-
terey County, California Larry G. Talent and Carline L. Talent 184

Repeat Spawning of Pacific Lamprey John H. Michael, Jr. 186

A Diver-Operated Snagging Device for Capturing Lingcod,

Ophiodon elongatus James L. Houk 187

Karyotype of the Sacramento Perch, Archoplites interruptus

Craig A. Busack and Gary H. Thorgaard 189

Book Reviews 192



ERRATUM
Lesh, E. W. 1980. A head-off method of measuring chinook and coho
salmon. Calif. Fish Game, 66(1) : 59-62.
Page 60, 61. The coefficient of deterrpination should be (r^).
Page 61. The ordinate of Figure 3 should read: Fork length v^ith head off
in millimetres.

The abscissa of Figure 3 should read: Fork length with head on in mil-
limetres.



132 CALIFORNIA FISH AND GAME



IN MEMORIAM

J. BRUCE KIMSEY

J. Bruce Kimsey was born in Portland, Oregon on 18 July 1921 and died on 24
January 1980 at the Kaiser Hospital in Sacramento after a lengthy illness. Bruce
leaves a wife and two grown sons and a legacy of devotion to his family and his
career. He had a long and productive career as a professional biologist in fisheries
research and management that spanned about 32 years.

Bruce received a B.A. degree from Chico State University in 1948 and an M.A.
degree from the University of California at Berkeley in 1951. He served with the
Armed Forces in the South Pacific during World War II. His first permanent
position in fisheries was with the California Department of Fish and Game as a
Junior Aquatic Biologist in 1948.

Bruce had wide ranging responsibilities with the Department on matters con-
cerning inland fisheries. Probably the most challenging position during the 13
years he worked for the Department was as a leader of statewide warmwater
fisheries coordination and research. It was during this period that most of his
publications appeared.

His publications numbered about 45. Most appeared in either California Fish
and Game or the Inland Fisheries Administrative Report series. Bruce's interest
and enthusiasm for all aspects of natural history were reflected in his publications,
some which concerned birds and appeared in the Condor.

Bruce's expertise in fisheries matters led to a long involvement and much
overseas travel as a consultant for a number of foreign aid organizations. Bruce
and his family spent an entire year at Lakes George and Edward in Uganda on
an assessment of the fish stocks plus the training of African fisheries workers. This
was just the begining and throughout the remainder of his career, Bruce took part
in numerous short-term overseas assignments. Besides Uganda, Bruce traveled to
Kenya, Tanzania, Rhodesia, Cameroon, Brazil, Colombia, El Salvador, Nicaragua,
Philippines, and Indonesia.

After leaving the Department in 1961, Bruce went to work for the old U. S.
Bureau of Commercial Fisheries as leader of a shrimp research project with
headquarters at Galveston, Texas. He later moved to the Bureau's main office in
Washington, D.C. where he assessed fisheries developments in various countries
around the world. He transferred in 1 961 to the old U. S. Bureau of Sport Fisheries
and Wildlife where he became Chief of the Branch of Ecosystem Research. In this
capacity he supervised six laboratories engaged in reservoir and marine sportfish
research. Bruce returned to Sacramento in 1971 where he assumed the position
of Regional Environmental Quality Officer for the Mid-Pacific Region of the Water
and Power Resources Service. He remained at this post until he died.

Bruce was a fellow of the American Institute of Fishery Research Biologists and
a member of various honorary and professional societies. He was President of the
California-Nevada Chapter of the American Fisheries Society in 1976 and served
as chairman or member of numerous national and regional committees of this
organization.

Bruce will be sorely missed by his many friends and co-workers from around
the world. — Almo J. Cordone



FLORIDA LARGEMOUTH BASS ELECTROPHORETIC STUDIES 133

Calif. Fish and Game 66 ( 3 ) : 1 33-1 62

IMPACT OF FLORIDA LARGEMOUTH BASS, MICROP-

TERUS SALMOIDES FLORIDANUS, INTRODUCTIONS AT

SELECTED NORTHERN CALIFORNIA WATERS WITH A

DISCUSSION OF THE USE OF MERISTICS FOR DETECTING

INTROGRESSION AND FOR CLASSIFYING INDIVIDUAL

FISH OF INTERGRADED POPULATIONS ^

RONALD J. PELZMAN

California Department of Fish and Game

Inland Fisheries Branch

1701 Nimbus Road

Rancho Cordova, CA 95670

Florida largemouth bass, Micropterus salmoides floridanus, had a notable genetic
impact following their introduction into five northern California waters containing
northern largemouth bass, M. s. salmoides, populations: Folsom Lake, New Hogan
Reservoir, Lake Amador, Lake Isabella, and Clear Lake. Analysis of malate dehy-
drogenase isozyme patterns of fish systematically collected in years subsequent to
the introduction indicated that intergraded populations developed at each of the
waters. Incidence of the Florida allele at the study waters, based on malate dehy-
drogenase analyses, eventually ranged from 0.35 at Lake Amador to 0.52 at both New
Hogan Reservoir and Clear Lake.

Discriminant function analysis of meristic data for fish of known electrophoretic
phenotype showed that meristic values were not reliable for classifying individual
fish from mixed populations as to Florida, northern, or hybrid bass categories. This
was supported by meristic data for known F-, hybrids. Hybridization could not
necessarily be detected by an increase in mean meristic value or by unimodality of
a frequency distribution of meristic values. The mode value of lateral line scale
counts appeared to be the best meristic indicator of hybridization.

Information from this study and from a similar study at southern California waters
indicates that introductions of Florida bass into northern bass populations have
generally been beneficial through reducing high exploitation rates, increasing the
mean size of bass in the catch, and providing exceptional fishing for trophy-sized
bass at some waters.

Results of this study indicate that current largemouth bass populations at the study
waters possess a wider spectrum of performance capabilities through the inclusion
of desirable traits attributed to Florida bass. This is particularly advantageous in the
reservoir setting where heavy angling pressure, water level manipulation, competi-
tion of prey species with small bass, and other factors work against the maintenance
of a bass population.

TABLE OF CONTENTS

Page

ABSTRACT 133

INTRODUCTION 134

METHODS AND MATERIALS 138

RESULTS AND DISCUSSION 141

Electrophoretic Analysis — Malate Dehydrogenase 141

Frequency of the Florida Allele Based on Malate Dehydrogenase

' This work was performed as part of Dingell-Johnson Project F-18-R, "Coldwater Reservoir and Special Experi-
mental Reservoir Program," supported by Federal Aid to Fish Restoration funds. Accepted for publication.



134 CALIFORNIA FISH AND GAME

Analyses 144

Electrophoretic Analysis — Tetrazolium Oxidase 147

Meristic Analysis 148

Use of Meristic Data in Evaluating Study Populations 151

Population Sampling 154

Rancho Seco Reservoir 155

Maladaptive Genes 156

Performance Capabilities of Fish of Intergraded Populations 157

Management Implications 158

ACKNOWLEDGMENTS 160

REFERENCES 160

INTRODUCTION

Northern largemouth bass were widely distributed to California's low- and
mid-elevation waters in the years following their introduction from Quincy,
Illinois^ in 1891 (Shebley 1917). Florida largemouth bass were not introduced
until 1959 when about 20,400 fingeriings from Holt State Fish FHatchery, Pen-
sacola, Florida were liberated at Upper Otay Reservoir, San Diego County,
which had been chemically treated to eradicate all fish and closed to public
access (Sasaki 1961). These fish and their progeny were stocked at many
southern California waters under the concept that Florida bass superiority in
growth rate and longevity and possible lower vulnerability to angling would
provide bass stocks with more large fish than northern bass were providing.
While Florida bass were stocked into northern bass populations at most of these
waters, Lake Hodges, San Diego County, which served as a source for some
Florida bass plants in northern California, was dewatered and chemically treated
prior to receiving Upper Otay fish in 1969 (L. Bottroff, Fishery Biologist, Calif.
Dept. Fish and Game, pers. commun.).

Florida bass were first stocked in northern California (that portion of California
north of the Tehachapi Mountains) in April 1969 at Clear Lake and Hidden
Valley Reservoir, both in Lake County, from Upper Otay Reservoir (R. Wood,
Fishery Biologist, Calif. Dept. Fish and Game, pers. commun.). Northern bass
were present at Clear Lake, while newly impounded Hidden Valley Reservoir
was devoid of bass at the time the introductions were made. From 1970 through
1973, Florida bass were stocked at a limited number of northern California
waters containing northern bass of Illinois origin, including the five waters exam-
ined by this study ( Figure 1 ) . During that period, Florida bass were also stocked
at a small number of waters that were devoid of bass including a farm pond in
the San Joaquin Valley and Rancho Seco Reservoir, Sacramento County. These
two waters served as sources for plants made later at the study waters (Figure
1).

The largemouth bass present at the study waters were considered descendants
of northern bass brought to California from the northern part of their range in
the United States. Based on a review of fish stocking records and communica-
tions with knowledgeable hatchery personnel and fishery biologists, northern

' The earliest documentation of largemouth bass shipments into California gives 1891 as the year of introduction
(Shebley 1917). According to Shebley the United States Commission brought largemouth bass and warmouth
"bass" here for stocking at Lake Cuyamaca and the Feather River. While the source for largemouth bass is
not given it was very likely Quincy, Illinois since this is given as the warmouth "bass" source.



FLORIDA LARCEMOUTH BASS ELECTROPHORETIC STUDIES



135



HOLT HATCHERY
PENSACOLA, FL.



1959




■^969-



UPPER OTAY
RESERVOIR



1972
1973



RANCHOSECG
RESERVOIR




REGION 4^
FARM POND




HIDDEN

VALLEY

RESERVOIR



1970
1971



FOLSOM
LAKE



LAKE
AMADOR



HOGAN
RESERVOIR



LAKE
ISABELLA



1969

u



CLEAR
LAKE



FIGURE 1. History of Florida largemouth bass stocking as related to the study waters (1-5).^
Region 4 is one of six geographical areas of California designated by the Department
for administrative and mangement purposes. It includes largely the San Joaquin Valley
and adjacent foothills.



bass from other portions of their range were not stocked at the five study waters
prior to the period of this study.

Apparent intolerance of Florida bass to 4°C in Missouri (Johnson 1975) and
concern that maladaptive genes possibly related to this intolerance would be
transmitted to northern bass populations (Childers 1975), prompted the Depart-
ment to establish a moratorium in May 1974 on further stockings of the subspe-
cies in northern California. Consequently, this study was initiated in July 1975 to
evaluate the survival and genetic impact of Florida bass at northern California
waters. Largemouth bass populations at Folsom Lake, Sacrdmento County; New
Hogan Reservoir, Calaveras County; Lake Amador, Amador County; Clear Lake,
Lake County; and Lake Isabella, Kern County were selected for evaluation. The
largemouth bass population at Rancho Seco Reservoir was also analyzed since
it served as a source for Florida bass plants at four of the study waters.

Identification of Florida and northern largemouth bass and their hybrids was
critical to evaluating mixed populations at the study waters. Bailey and Hubbs
( 1 949 ) first used scale counts for separation. Workers since have counted scales,
pyloric caeca, and vertebrae, and made body measurements (Bottroff 1967;
Buchanan 1968; Bryan 1969; Addison and Spencer 1971; Buchanan 1973; Chew
1975; Johnson 1975; and Bottroff and Lembeck 1978).



136 CALIFORNIA FISH AND CAME

Physiological differences between Florida and northern largennouth bass were
reported by Hart (1952). Bryan (1964) noted differences between serum elec-
tropherograms of the two subspecies. Differences in the electrophoretic mobility
of isozymes (different molecular forms of enzymes) from tissues of largemouth
and smallmouth bass and their hybrids were described by Whitt, Childers, and
Wheat (1971). Chew (1975) reported that Dr. William Childers of the Illinois
Natural History Survey, Urbana, Illinois, was able to separate Florida bass,
northern bass, and fish thought to be their hybrids by isozyme analysis. Childers
( pers. commun. ) utilized starch gel electrophoresis and separated the fish on the
basis of their different isozyme patterns of the enzyme malate dehydrogenase
(MDH).

As related to this study, gel electrophoresis is a method for observing genetic
variation of mixed populations by examining variant proteins (isozymes) manu-
factured by different individuals. A tissue sample from each individual to be
studied is homogenized to release its cell contents, including isozymes. These
are introduced into a gel made of starch and subjected to an electric current for
a few hours. Each isozyme in the sample migrates through the gel in a direction
and a rate that depends primarily on its net electric charge and, to some extent,
on its molecular size and conformation. The gel is then treated with a solution
containing a specific substrate, which is cleaved by the enzyme to be observed,
and a salt, which couples with the cleavage products. This process yields a
colored band at the zone to which the enzyme has migrated.

Because isozymes that are specified by different alleles may have different
molecular structures and charges (and hence different mobilities in an electric
field), the genetic makeup at the gene locus coding for a given enzyme can be
established for each individual from the number and position of the electropho-
retic bands (Ayala 1978). The advantage of data obtained through gel electro-
phoresis is that genetic interpretations can be made directly. Most variant alleles
show codominant expression. This permits designation of the genotypes of
individual samples based on staining patterns (Utter, Hodgins, and Allendorf
1 974) . When animals are crossed that are homozygous for different codominant
alleles at the same locus, their offspring are heterozygous, receiving one allele
from each parent. Because each allele codes for a slightly different protein,
heterozygosity can be inferred from the presence of two variants of a given
protein in a single individual (Ayala 1 978) . The simplest form of diploid variation
is when two codominant alleles are present in a population, one specifying a
fast-moving band and the other a slow-moving band. An individual homozygous
for either allele will show a single band, whereas the heterozygote will have both
bands (Gottlieb 1971).

Childers (pers. commun.) found that the MDH isozyme patterns (pheno-
types) of Florida largemouth bass and northern largemouth bass (from the
northern part of their geographic range) differed (Figure 2). Florida bass and
northern bass were both homozygous for supernatant MDH-A and MDH-B.
However, because of mutational differences in the makeup of the B gene locus,
the most anodal band (B„ B„,; m == fast) of the Florida bass showed greater
mobility than the most anodal band (B, B^; s = slow) of northern bass (Figure
2). The A gene locus and B gene locus of the northern bass each code for the
production of a different subunit (A and B^, respectively). These translated
subunits randomly combine to form the various dimers which will migrate



FLORIDA LARCEMOUTH BASS ELECTROPHORETIC STUDIES



137



through starch gel differing distances in response to an electric current to pro-
duce a three-banded pattern. The Florida bass pattern is similarly produced by
an A gene locus and a B gene locus, each coding for the production of a different
subunit (A and B„, respectively). When northern and Florida bass are crossed
the genotype AA B„B. translates subunits that randomly combine to form six
dimers (AA, AB„ B,B., AB„, B„B„ and B„B„). This appears as a five-banded
pattern, hov^ever, since two dimers (B,B, and ABn,) migrate the same distance
in starch gel. The F, hybrid shows all bands found in Florida and northern bass
plus an additional band ( B„BJ . For discussion of MDH phenotypes, MM is used
for Florida bass, SS for northern bass, and MS for the hybrid.



[ NORTHERN
I UARGEMOUTH
BASS



FLORIDA LARGEMOUTH BASS



F, HYBRID




FIGURE 2. Malate dehydrogenase and tetrazolium oxidase phenotypes of northern and Florida
largemouth bass and their F, hybrid. ' Results from study waters showed all MS fish
by MDH pattern to show only MM or MS by TO.

The frequency of alleles in a given population can be measured by direct
counts from the electrophoretic expression of a representative sample (Utter
and Allendorf 1977). In this study, the number of M alleles (coding for the B„B„
band) and S alleles (coding for the 83, band) in a population was determined
(see Figure 3 for example). The number of MDhI genes in a particular collection
is twice the number of individuals sampled because a complementary pair of
MDH genes are coded in each individual. A homozygous individual codes in
duplicate for a given allelic form and the heterozygous individual codes for two
different allelic forms.

Childers (pers. commun.) used an additional enzyme system, tetrazolium
oxidase (TO), which permitted partial separation of Florida and northern bass
(Figure 2). He found that northern bass were homozygous, showing a single



138



CALIFORNIA FISH AND CAME



fast-moving band (MM), whereas Florida bass showed either the fast-moving
band (MM), or a single slow-moving band (SS), or a heterozygous three band-
ed pattern (MM, MS, and SS). Fish thought to be first filial generation hybrids
between Florida and northern bass showed the MM or MS pattern. Tetrazolium
oxidase was used in this study to provide supportive data.



PHENOTYPE


SS


MS


MM


NO. OF FISH

SHOWING

PHENOTYPE


25


50


25


NO. OF FLORIDA
BASS ALLELES
IN SAMPLE




2X25^=^50






Wms


100


NO. OF NORTHERN
BASS ALLELES
IN SAMPLE


2X253=50

"■ 50n,s
100






FREQUENCY OF

FLORIDA
ALLELE (M)


^00=0.50
200


FREQUENCY OF

NORTHERN
ALLELE (S)


100 -0.50
200



FIGURE 3. Calculation of gene frequency for a collection of 100 fish composed of 25 fish showing
the SS phenotype, 50 showing the MS phenotype, and 25 showing the MM pheno-
type.

METHODS AND MATERIALS

Fish from purportedly pure California sources of Florida bass ( Upper Otay and
Hidden Valley reservoirs) and northern bass (Central Valleys Warmwater
Hatchery, Sacramento County; Shasta Lake, Shasta County; and Merle Collins
Reservoir, Yuba County) were examined meristically and electrophoretically.
Results for fish from Central Valleys Warmwater Hatchery were of particular
interest. This hatchery had been involved for many years with stocking northern



FLORIDA LARGEMOUTH BASS ELECTROPHORETIC STUDIES 139

largemouth bass at northern California waters, including Folsom Lake, New
Hogan Reservoir, and Lake Amador ( M. Cochran, Fish Hatchery Manager, Calif.
Dept. Fish and Game, pers. commun.). Also, hatchery brood fish had been
obtained from several northern California sources.

Fish from mixed populations were analyzed using meristic and electrophoretic
information for pure populations as baseline data. The study plan was to assess
the degree of hybridization of Florida and northern bass by analyzing 100
young-of-the-year bass from each of the study waters annually. Electrophoresis
was the primary method used; however, since no largemouth bass study involv-
ing both meristic and electrophoretic evaluations of the same fish was found in
the literature, meristic data were collected through 1977 for fish analyzed elec-
trophoretically. This provided a means to assess the value of meristic data for
classifying individual fish from mixed populations and to determine if meristics
could be used by fishery managers in their assessment of the degree of hybridiza-
tion of mixed populations.

Fish were collected by electroshocking each fall from 1975 through 1978. So
that samples would be representative of populations being evaluated, collections
were made using the following guidelines:

(1 ) Sample at each of the four major compass directions.

(2) Sample representative cover types.

(3) Collect young-of-the-year of all sizes.

(4) Collect no more than 10 fish when encountering heavy concentrations
of fish such as at the apex of coves, in brushy areas, etc.

Fish were sacrificed, individually enclosed in plastic bags, and transported in
crushed ice to prevent breakdown of enzyme systems. At the laboratory they
were retained in ice until each received an identifying tag and tissue was
removed for analysis. When time did not permit this, fish were frozen and later
thawed a few at a time for processing. Skeletal muscle tissue, used for MDFH,
was removed from an area just below the dorsal fin on the right side of the fish
to facilitate scale counts. Liver tissue was taken for TO analysis. All utensils and
the worker's hands were thoroughly cleaned after each fish was processed. Each
fish and all samples removed from it received an identifying code which includ-
ed information as to the year and water of collection. For example, F-lll-1 was


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