CALIFORNIA!
FISH- GAME
t VOLUME 72
OCTOBER 1986
NUMBER 4
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California Fish and Game is a journal devoted to the conservation and
understanding of fish and wildlife. If its contents are reproduced elsewhere, the
authors and the California Department of Fish and Game would appreciate
being acknowledged.
Subscriptions may be obtained at the rate of $10 per year by placing an order
with the California Department of Fish and Game, 1416 Ninth Street, Sacra-
mento, CA 95814. Money orders and checks should be made out to California
Department of Fish and Game. Inquiries regarding paid subscriptions should be
directed to the Editor.
Complimentary subscriptions are granted on an exchange basis.
Please direct correspondence to:
Robert N. Lea, Ph.D., Editor-in-Chief
California Fish and Game
1416 Ninth Street
Sacramento, CA 95814
1
J
1
0
VOLUME 72
OCTOBER 1986
NUMBER 4
Published Quarterly by
STATE OF CALIFORNIA
THE RESOURCES AGENCY
DEPARTMENT OF FISH AND GAME
— IDA—
STATE OF CALIFORNIA
GEORGE DEUKMEJIAN, Governor
THE RESOURCES AGENCY
GORDON VAN VLECK, Secretary for Resources
FISH AND GAME COMMISSION
BRIAN J. KAHN, President
Santa Rosa
ABEL C. GALLETTI, Vice President JOHN MURDY III, Member
Los Angeles Newport Beach
ROBERT BRYANT, Member ALBERT C. TAUCHER, Member
Yuba City Long Beach
HAROLD C. CRIBBS
Executive Secretary
DEPARTMENT OF FISH AND GAME
JACK C. PARNELL, Director
1416 9th Street
Sacramento 95814
CALIFORNIA FISH AND GAME
Editorial Staff
Editorial staff for this issue consisted of the following:
Marine Resources Robert N. Lea, Ph.D. and Perry L. Herrgesell, Ph.D.
Editor-in-Chief Robert N. Lea, Ph.D.
195
CONTENTS
Page
Temporal and Spatial Patterns in Sea Otter, Enhydra lutris, Range Expan-
sion and in the Loss of Pismo Clam Fisheries 197
Frederick E. Wendell, Robert A. Hardy, Jack A. Ames,
and Richard T. Burge
Aspects of Ecology and Life History of the Woolly Sculpin, Clinocottus
analis, from Southern California 213
Alan W. Wells
Ichthyofaunal Composition and Recolonization in a Central California
Tidepool 227
Ronald H. Matson, C. Ben Crabtree, and Thomas R. Haglund
Distribution of Major Marine Macrophytes, Seasonal Estimates of Graci-
laria Standing Crop, and Spawning Activities of the Pacific Herring,
Clupea harengus pallasii, in Elkhorn Slough, California: 1979-1982 .. 232
R. E. Phillips, D. I. Gutoff, J. E. Hansen and J. E. Hardwick
Observations on the Elasmobranch Assemblage of San Francisco Bay .... 244
David A. Ebert
Book Reviews 250
Index to Volume 72 252
196
CHANGE OF EDITORSHIP
With this issue Robert N. Lea, of the Marine Resources Division, as-
sumes the duties of Editor-in-Chief of California Fish and Game. Dr. Lea's
appointment to the editorship follows the Department's policy of rotating
the editorship between staff members representing Marine Resources,
Inland Fisheries, Wildlife Management, and the Bay-Delta Project.
Dr. Lea has been with the Department for 17 years and has served as
Editor for Marine Resources the past five years. He is the co-author of
Guide to the Coastal Marine Fishes of California and has published over
thirty-five scientific papers in various journals and symposia proceedings.
Under his guidance the Journal will continue its policy of presenting the
results of scientific investigations as they relate to conservation and man-
agement programs of California's fish and wildlife resources. He will strive
to maintain the excellent reputation the Journal has achieved during its 72
year history of continuous publication.
Dr. Lea will be assisted in his duties by the following associate editors:
Jack A. Hanson — Inland Fisheries; Daniel P. Connelly, Gordon I. Gould,
and Douglas R. Updike — Wildlife Management; Peter L. Haaker, Paul N.
Reilly, and John P. Scholl — Marine Resources; Kenneth A. Hashagen, Jr.,
and Arthur C. Knutson, Jr. — Anadromous Fisheries; Donald E. Stevens —
Striped Bass, Sturgeon, and Shad; and Kim McCleneghan, Environmental
Services.
To Dr. Perry L. Herrgesell, Editor-in-Chief the past four years, we ex-
press our deep appreciation for a job well done. Jack CParnell, Director,
California Department of Fish and Game.
SEA OTTER RANGE EXPANSION AND LOSS OF PISMO CLAM FISHERIES 197
Calif. Fish and Came 72(4): 197-212 1 986
TEMPORAL AND SPATIAL PATTERNS IN SEA
OTTER, ENHYDRA LUTRIS, RANGE EXPANSION AND IN
THE LOSS OF PISMO CLAM FISHERIES1
FREDERICK E. WENDELL
ROBERT A. HARDY
California Department of Fish and Came
213 Beach Street
Morro Bay, California 93442
JACK A. AMES
California Department of Fish and Game
2201 Garden Road
Monterey, California 93940
and
RICHARD T. BURGE
Washington Department of Fisheries
600 Point Whitney Road
Brinnon, Washington 98320
Two beaches in central California that have been reoccupied by sea otters, Enhy-
dra lutris, through a southward expansion of their range once supported major
recreational fisheries for Pismo clams, Tivela stultorum. Monitoring of recreational
fishing and sea otter activity was conducted in these areas to elucidate the sea otter's
role in the loss and long term fate of these fisheries. The progressive elimination of
sections of beach from the fishery closely corresponded to the southward progres-
sion of sea otter foraging activity. Data indicate that once sea otters are established
along clam bearing beaches, any future stocks of clams will be fully utilized by sea
otters, preventing the return of a fishery.
INTRODUCTION
Sea otters, Enhydra lutris, are predators on benthic invertebrate species which
inhabit shallow coastal waters in the North Pacific Ocean. Historically sea otters
ranged as far south as Morro Hermoso on the Pacific coast of Baja California.
From that southern limit they ranged northward and westward through the
Aleutian and Commander islands to the western rim of the North Pacific Ocean
(Kenyon 1969). However, there have been major changes in sea otter abun-
dance and distribution associated with their decimation during the fur trade era,
1780 to 1900 (Kenyon 1969) and with the subsequent reoccupation of a portion
of their historic range. Considerable debate has existed concerning the effect sea
otter predation has on the structure of nearshore communities and on associated
shellfish fisheries in the reoccupied area.
Although the effect sea otter predation has on shallow sandy bottom com-
munity structure has not been studied directly, the impact on associated shellfish
fisheries has received some attention. Miller (1974) noted the loss of the recrea-
tional red crab, Cancer productus, and rock crab, C. antennarius, fishery off the
Monterey pier associated with the reoccupation of that area by sea otters. A
rapid depletion of the commercial and subsistence dungeness crab, Cancer
magister, fishery was noted in the Orca Inlet area of Prince William Sound,
1 Accepted for Publication April 1986.
198 CALIFORNIA FISH AND CAME
Alaska, as the sea otter population in that area increased (Matkin 1981 and
Kimker 1982). Wild and Ames (1974) noted that Pismo clams, Tivela stultorum,
were eaten almost exclusively when sea otters initially foraged along Pismo clam
bearing beaches in Monterey and Estero Bays, California. Although the extent
of the effect on the Pismo clam population was not determined, they indicated
it could be substantial. Stephenson (1977) subsequently indicated that sea otters
had contributed extensively to the decline of large Pismo clams within his study
area in Monterey Bay, California.
The question of the extent of the impact sea otter predation can have on a
Pismo clam fishery was first addressed by Miller, Hardwick, and Dahlstrom
(1975). Their studies indicated that sea otter foraging along Monterey Bay
beaches and at Atascadero Beach near Morro Bay precluded the recreational
Pismo clam fisheries in these areas.
Because of the possibility that over-harvest, pollution, or variable recruitment
could influence any decline in sport and commercial shellfish catches, it has
been suggested that the loss of fisheries cannot be entirely attributed to sea otter
predation (Armstrong 1979, Estes and VanBlaricom in press). Most of the pub-
lished data directed toward clarifying the sea otter's role in the loss of Pismo
clam fisheries has not included sufficient historical data to focus on pre-sea otter
trends in the fisheries.
Recent sea otter range expansion to the south has occurred along Pismo clam
bearing beaches near Pismo Beach (Figure 1). Monitoring of the intertidal
portion of that Pismo clam population and the sport harvest of clams has been
on going for many years. Data on the fishery and on sea otter distribution and
food habits along clam bearing beaches are presented to provide a historical
perspective necessary to clarify the sea otter's role in the loss of the fisheries and
the long term fate of these fisheries within the established sea otter range.
MATERIALS AND METHODS
The Pismo clam fisheries located in areas reoccupied by sea otters through
a southward expansion of their range were monitored historically by collecting
and ageing clams recovered from trench transects. The trench, 15 cm wide by
20 cm deep, was oriented perpendicular to the surf zone and extended into the
shallow (ca. 60 cm depth) subtidal during minus tides. In early years, sand was
sifted through screens; however, in most years a casting-spreading technique
was used. Probing was used to sample inundated portions of the transect. These
data were primarily used to assess incoming year-classes for management pur-
poses.
Since 1975 the fishery in the Pismo Beach area has been assessed in a direct
manner through interviewing clammers. The basic information obtained during
the interviews included the following (i) the number of active clammers in the
group, (ii) the time active, (iii) the number of legal-sized clams (114.3 mm, 4.5
in) obtained, and (iv) the number of clams within one inch of legal size which
they recalled digging and returning.
Estimates of total effort and total catch for each section of beach were gener-
ated from interview data. Analysis incorporated stratification by tide level, day
of the week, and section of the beach. Total effort for each sampling day was
estimated through an expansion, based on turn-over rates, of the number of
active clammers counted during the period from one-half hour before low tide
to low tide. The expansion used to estimate total effort was based on the
SEA OTTER RANGE EXPANSION AND LOSS OF PISMO CLAM FISHERIES
199
assumption that the interviewed clammers provided an unbiased sample of the
distribution of clamming activity about low tide. The count, therefore, was
expanded by the proportion of the interviewed clammers active during the
count period (Table 1).
Miles
50
=4
Santa Cruz
Pt. San Luis
Miles
Km
Pt. Sal
Pismo Beach
FIGURE 1. Pismo clam bearing beaches (darkened strips) within the sea otter's range. Expanded
portion of map delineates relative size and location of sampling sections along Pismo
Beach.
200 CALIFORNIA FISH AND GAME
TABLE 1. Proportion of Interviewed Clammers Actively Clamming During the Count Period
(from % hour before to low tide) for Three Tide Groups (sunrise, daylight, and
sunset) by Beach Section.
Sunrise Daylight Sunset
Beach 1 68 (n = 99) .80(n = 385) .94(n = 63)
Beach 2 63(n = 645) .85(n = 970) .81ln = 80)
Beach 3 72(n = 622) ,88(n=926) 76(n = 465)
Beach 4 65(n = 1031) .90(n = 2078) 94(n = 690)
The distribution and relative abundance of sea otters within the Pismo Beach
area was determined using both aerial and ground census techniques. Seasonal
aerial censuses, conducted since 1976, identified the location and relative abun-
dance of sea otters at the range peripheries. These censuses were supplemented
with ground counts in February 1979, south of Point San Luis, when the periph-
eral male group moved south of the point into the Shell Beach kelp beds.
Information on distribution and relative abundance of otters along Pismo clam
bearing beaches was also collected while conducting sea otter food habit obser-
vations. All ground based observations were made using a 50x-80x Questar
telescope.
RESULTS
Initial Impact — Pismo Beach
Pismo Clams
A summary of transect data was provided by Fitch (1952, 1954, and 1955)
for the years 1925 to 1954, by Baxter (1961 ) for the years 1955 to 1959, and by
Carlisle (1966, 1973) for the years 1961 to 1971. These data provide an initial
index of density, growth, and survival for clam sets by area. However, with
natural mortality and fishing pressure influencing abundance and availability, this
sampling method was inadequate to assess these parameters in older cohorts.
Trench transect data, however, have continued to provide information on the
density and survival of smaller sub-legal clams (Tables 2-5). The relative num-
ber of zero through age 2 clams (an index of recruitment), although apparently
reduced, appears to have remained within the range traditionally observed.
Interviews of clammers provide an index of legal-sized Pismo clam availability
in the intertidal and shallow subtidal portions of the beach. Over 11,800 inter-
views have been conducted on 170 interview days since November 1975. The
interviews provided catch-per-unit-effort (CPUE) data that were used to gener-
ate estimates of the total catch and effort by beach section for tides below 0.0
m ( Table 6, Figures 1 and 2 ) . Most beach sections showed considerable fluctua-
tion in the estimated total annual catch and effort as would be expected from
a heavily utilized resource based on a species with highly variable recruitment.
Clamming conditions, particularly surf height, also contribute to variation in
catch and effort. The estimated total annual catch and effort, however, never
approached zero for any beach section until 1979 or later.
Average monthly CPUE declined to near zero, beach by beach, from north
to south (Figure 2). Beach 1 dropped to near zero in May 1979, while beaches
2, 3, and 4 remained relatively high. Beach 2 followed with a drop to near zero
by May 1980. This progression continued until June 1981, when the CPUE was
near zero for all beach sections along Pismo Beach. The CPUE has remained
essentially zero through 1985.
SEA OTTER RANGE EXPANSION AND LOSS OF PISMO CLAM FISHERIES 201
TABLE 2. Number of Clams by Age Group from Trench Transects at Atascadero State Beach.
Year 0
1952 14
1953 0
1954 2
1955 3
1956 0
1957 0
1958 0
1959 0
1960 0
1961 0
1962 0
1963 0
1964 0
1965 1
1966 18
1967 6
1968
1969
1970
1971 0
1972
1973 1
1974 1
1975 0
1976 0
1977 2
1978 0
1979 0
1980 0
1981 1
AGE GROUP
/
2
3
4
5
6
7
8
Total
0
0
0
0
0
0
0
0
14
5
0
0
0
0
0
3
21
29
0
6
0
0
0
1
0
23
32
2
0
2
0
1
0
0
5
13
0
0
0
0
0
0
0
6
6
0
0
1
0
5
0
0
12
18
0
0
0
0
0
0
0
2
2
0
1
0
0
0
0
2
8
11
1
0
0
0
0
0
1
1
3
0
2
1
0
0
1
0
0
4
0
0
2
0
0
0
2
0
4
0
0
1
4
0
0
0
0
5
0
1
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
2
2
1
0
0
0
0
0
1
22
21
3
0
0
0
0
0
1
31
39
0
0
0
1
0
0
0
141
12
123
0
0
0
0
0
0
136
3
10
83
0
0
0
0
0
96
0
3
14
28
0
0
0
0
45
0
1
2
7
8
0
0
0
20
3
0
0
3
2
0
0
0
8
1
0
0
1
0
0
0
0
2
0
0
1
1
0
0
0
0
2
Sea Otters
Observations from shore indicated that the southern peripheral male group
had established a rafting site at Cayucos Point (ca. 10 km north of Atascadero
Beach) in February 1972. By January 1973, the southern male group was rafting
16 km south of Atascadero Beach at Point Buchon (Wild and Ames 1974). Sea
otters, however, were observed foraging along Pismo clam bearing portions of
Atascadero Beach in February and March 1973 (Wild and Ames 1974). Between
1973 and 1975 sea otters were frequently observed foraging along Atascadero
Beach and a small raft of four to six otters were often seen off Torro Point at
the north end of the beach (Burge 1979).
The peripheral male group moved to a rafting site south of Point San Luis at
Shell Beach in January 1979. Otters from that raft were first observed foraging
along the northernmost portion of Pismo Beach (Beach 1 ) late in January 1979.
Since that time the number of otters observed south of Point San Luis, including
areas along clam bearing beaches, has shown a typical seasonal pattern ( Figures
3 and 4).
ACE CROUP
1
7
2
3
4
5
6
7
8
Total
0
0
0
0
1
4
0
2
26
49
0
0
0
1
2
13
3
68
1
20
3
0
0
0
1
4
31
8
4
16
2
0
0
0
9
42
2
3
2
35
0
0
0
7
49
0
0
4
9
13
1
0
7
205
34
0
0
3
0
7
0
1
51
25
21
0
0
0
0
7
5
179
25
17
14
0
1
0
0
1
65
10
18
13
3
0
0
0
1
49
12
5
10
15
3
0
0
0
78
6
11
4
9
4
0
0
1
96
20
27
10
4
5
1
0
1
179
97
34
21
0
4
0
0
0
282
100
78
43
7
1
0
1
0
325
65
88
69
12
0
0
1
0
246
202 CALIFORNIA FISH AND GAME
TABLE 3. Number of Clams by Age Croup from Trench Transects at Pismo Beach (Beach
1).
Year 0
1952 19
1953 0
1954 2
1955 3
1956 0
1957 170
1958 6
1959 121
1960 7
1961 4
1962 33
1963 61
1964 111
1965 126
1966 95
1967 11
1968
1969
1970
1971 0 14 0 2 2 2 1 0 0 21
1972
1973
1974 - - ______
1975 2
1976 0
1977 12
1978 0
1979 3
1980 11
1981 23
The proportion of otters observed along sandy beaches was relatively low
during 1979, the first year the sea otter's range had extended south of Point San
Luis. During that year, much of the foraging activity occurred just north of Pismo
Beach, in the Shell Beach area. The proportion observed along sandy beaches
was noticeably higher in 1980 and 1981. It did not appear at any point that all
otters in the area occupied just areas along the clam bearing beaches (Figures
3 and 4). The May 1980 census yielded the highest count to date, 67 sea otters
along the sandy beaches and 134 total south of Point San Luis.
Observations of tagged sea otters in the area documented movements
between the rafting sites in giant kelp beds, Macrocystis pyrifera, off Shell Beach
and foraging areas along the Pismo clam bearing beaches. Most sea otters, when
along the sandy beaches, foraging or resting offshore, tended to be in loose
aggregations and occupied a very limited portion of the nearshore area. Limited
radio telemetry research conducted in co-operation with the University of Min-
nesota also documented movement to and activity along those beaches during
the night.
Food habit studies were conducted along the sandy beaches to identify major
prey items. Over 500 individual food items were observed being consumed
8
20
11
10
4
0
0
0
55
0
4
2
4
0
0
0
1
11
18
2
4
4
1
0
0
0
41
5
13
1
4
0
1
0
2
26
3
4
1
0
0
0
0
0
11
2
3
2
0
0
0
0
0
18
11
4
0
1
0
0
0
0
39
SEA OTTER RANGE EXPANSION AND LOSS OF PISMO CLAM FISHERIES 203
(Table 7). Pismo clams comprised 96% of those food items. On several occa-
sions tagged sea otters were followed for extended periods during a foraging
bout. The longest continuous observation period was slightly under two hours.
During that period, the sea otter surfaced with 57 Pismo clams and six unidenti-
fied items of which 40 clams and six unidentified items were observed con-
sumed.
TABLE 4. Number of Clams by Age Group from Trench Transects at Pismo Beach (Beach
3).
Year 0
1952 8
1953 0
1954 1
1955 0
1956 0
1957 470
1958 2
1959 62
1960 10
1961 6
1962 36
1963 14
1964 200
1965 75
1966 36
1967 5
1968
1969
1970
1971 0 8 2523100 21
1972
1973
1974
1975 2
1976 0
1977 35
1978 0
1979 0
1980 5
1981 6
Area of Impact
The progression in decline in catch-per-unit-effort for the fishery at Pismo
Beach closely corresponded to the southward progression of sea otter foraging
activity.
Beach 1. The estimates of total annual catch and effort for this section of
Pismo Beach markedly decreased from 1979 to 1980. The highest monthly
count of sea otters along this section when superimposed on the mean month-
ly CPUE showed that the drop to zero in catch for the fishery closely corre-
sponded to that period early in 1979 when sea otters were actively foraging
in the area (Ffgure 5).
Age Group
/
2
3
4
5
6
7
8
Total
0
0
0
1
0
0
0
0
9
28
0
0
1
5
9
50
56
149
0
20
5
0
0
8
10
87
131
0
1
3
0
1
2
1
65
73
0
0
0
7
0
0
0
16
23
0
0
0
0
3
0
0
5
478
45
0
0
0
0
0
0
1
48
27
42
0
0
0
0
0
1
132
34
14
17
0
0
0
0
3
78
8
13
11
6
0
0
0
0
44
18
5
11
7
2
0
0
1
80
13
21
6
5
2
0
0
1
62
25
16
6
1
1
0
0
0
249
141
25
15
1
0
0
0
0
257
67
60
24
5
1
0
0
0
193
25
40
55
18
2
1
0
0
146
4
17
15
9
1
0
0
0
48
3
6
17
9
0
0
0
0
35
1
5
5
6
1
1
1 •
1
56
4
1
0
1
5
1
0
1
23
8
22
0
2
2
0
1
3
38
2
0
3
0
0
0
0
0
10
8
2
0
0
0
0
0
0
16
204
CALIFORNIA FISH AND GAME
TABLE 5. Number of Clams by Age Group from Trench Transects at Pismo Beach (Beach
4).
Year 0
1952 1
1953 0
1954 0
1955 0
1956 0
1957 13
1958
1959 23
1960 32
1961 1
1962 2
1963 4
1964 71
1965 40
1966 61
1967 43
1968
1969
1970
1971 0
1972
1973
1974
1975 32
1976 0
1977 28
1978 0
1979 0
1980 1
1981 3
Age Group
1
2
3
4
5
6
7
8
Total
0
0
0
0
0
3
1
0
5
0
0
0
0
6
4
2
2
14
0
0
0
0
0
0
0
1
1
2
0
0
0
0
0
0
6
8
0
2
0
0
0
0
0
8
10
0
0
2
1
0
0
0
7
23
3
0
0
0
0
0
2
6
34
7
3
1
0
0
0
0
2
45
7
7
3
0
0
0
0
1
19
1
6
4
4
1
0
0
0
18
6
4
5
2
0
0
0
0
21
8
2
1
7
0
1
0
0
90
41
14
0
2
1
0
0
0
98
61
50
6
0
0
0
0
1
179
26
14
13
17
3
0
0
0
116
11
28
0
0
3
1
1
0
0
0
37
14
0
8
1
0
0
0
0
23
0
2
2
1
0
0
0
0
33
14
0
2
1
1
0
0
0
18
4
0
0
0
0
0
0
2
6
0
2
5
1
0
0
0
0
9
1
0
0
0
0
0
0
0
4
TABLE 6. Estimated Annual Catch (C) of Pismo Clams and Annual Effort (E) by Beach
Section along Pismo Beach for Tides Below 0.0 m from 1975 to 1983.
Beach 1 Beach 2 Beach 3 Beach 4 Total
Year C E C E C E C E C E
1975 8,107 4,512 12,712 4,486 43,889 9,952 159,439 26,219 224,147 36,209*
1976 12,601 3,840 7,142 2,576 28,029 8,551 120,548 25,957 168,320 40,924
1977 37,606 10,157 closed 18,191 9,522 156,355 29,137 212,152 48,816
1978 26,178 5,550 109,513 16,311 36,173 9,015 171,796 24,660 343,660 55,536
1979 4,588 1,853 46,110 9,698 74,135 17,128 106,064 23,310 230,897 51,989
1980 22 478 261 1,573 9,979 6,659 137,950 31,188 148,212 39,898
1981 1 266 91 616 61 1,358 17,525 10,685 17,678 12,925
1982 0 14 0 42 3 158 19 735 22 893
1983 0 14 0 0 0 28 0 0 0 42
* An additional 13,980 (catch) and 3857 (effort) were estimated for tides between +0.5 ft and 0.0 ft during 1975.
Beach 2. The estimates of total annual catch and effort for this beach
section showed a dramatic increase in response to the management closure
of clamming from May 1976 to March 1978. The estimated total annual catch
and effort, however, were much lower from 1980 through 1983. The drop to
SEA OTTER RANGE EXPANSION AND LOSS OF PISMO CLAM FISHERIES
205
zero in mean monthly CPUE for that section of beach started late in 1979
which also corresponded to that period when sea otters had shifted their
foraging activity to Beach 2 (Figure 5).
Beach 1
Month
FIGURE 2. Mean daily catch-per-clammer of Pismo clams by month from beach sections 1 through
4, Pismo Beach, between November 1975 and January 1984.
206
CALIFORNIA FISH AND CAME
^7
Jan
77
iiiiiiiiii i| iiriiiiiiiiiMl ll|"iiiii
Jan Jan Jan Jan
78 79 80 81
Month
Jan
82
FIGURE 3. Mean number of sea otters by month south of Point San Luis showing the arrival and
seasonal fluctuation in numbers of the southern peripheral male group through July
1982.
Beach 3. Both the estimates for total annual catch and effort and mean
monthly CPUE followed the same pattern observed on Beaches 1 and 2, but
occurred slightly later (early 1980, Figure 5).
Beach 4. This beach was divided into two sub-units since it was long
enough that the influence of foraging activity on CPU E would be masked. Both
the estimate of total annual catch and effort and the means monthly CPUE
dropped for each sub-unit in 1981, when sea otters had extended their forag-
ing into those areas. There was, however, a noticable lag in the timing of the
decline for the southernmost sub-unit (Figure 5).
CPUE remained high on each beach prior to being foraged upon by sea otters,
declined to zero or near zero during concentrated foraging and remained at or
near zero after sea otters moved southward.
The Mann-Whitney non-parametric test (Sokal and Rohlf 1969) was applied
to CPUE and sea otter census data to determine whether the observed changes
associated with sea otter foraging were statistically significant. In all cases, when
otters first occupied a section of beach, the CPUE for clams remained relatively
high for a limited period of time. The data gathered during the transition from
high to low CPUE were grouped with pre-sea otter CPUE data for analysis. The
CPUE data were thereby separated into two (pre and post-otter) samples. All
beach sections had significantly lower CPUE (P >0.01 ) by the time sea otters
SEA OTTER RANGE EXPANSION AND LOSS OF PISMO CLAM FISHERIES
207
began foraging on the next beach to the south (Beach 1 n1=29, n2 = 33,
z = 6.74; Beach 2 n1=34, n2 = 34, z=4.68; Beach 3 n1=24, n2=47, z = 6.52;
Beach 4a n1=10, n2 = 65, z = 5.05).
Month
FIGURE 4. Mean number of sea otters by month along the Pismo clam bearing beaches south of
Point San Luis through July 1982.
TABLE 7. Number of Prey Retrieved and Observed Consumed by Sea Otters Foraging along
Pismo Beach from March 1979 to February 1981.
Food item Number Number
retrieved consumed
Pismo clams 559 505
Unidentified bivalves 1 1
Rock crabs 1 1
Market crabs 1 1
Unidentified crabs 1 1
Unidentified food items 15 15
Long Term Impacts — Atascadero Beach
In 1 971 , the earliest year for which interview data are available for Atascadero
Beach, 337 clammers averaged 3.3 clams/h. The southward movement of the
peripheral male group 26 km from Point Cayucos to Point Buchon (Figure 1 )
in January 1973, skipped for a short period, the Pismo clam bearing beaches
around Morro Bay, including Atascadero Beach. Sea otters, whether from the
peripheral group or from adjacent areas, were observed during that year foraging
208
CALIFORNIA FISH AND CAME
along Atascadero Beach on Pismo clams (Wild and Ames 1974; Miller, Hard-
wick, and Dahlstrom 1975; Wade 1975). Sampling of the Pismo clam fishery
indicated that the CPUE for the fishery on Atascadero Beach was zero by
December 1973. Sea otters continued foraging along Atascadero Beach after
1973.
9
7
5
3
1
9
7
5
3
1
9
7
5
3
1
9
7
5
3
1
Beach 1
Beach 2
u
Beach 3
Ms
35
h25
15
5
45
h35
25
-15
5
45
^35
C/)
<D
0)
(T>
O
O
c
3
Month
FIGURE 5. High count of sea otters by month (bar) superimposed on a measure of catch-per-unit-
effort for the Pismo clam fishery ( line ) along each beach section from November 1 975
through February 1984.
SEA OTTER RANGE EXPANSION AND LOSS OF PISMO CLAM FISHERIES
209
Transect digs in 1973 on that beach identified the 1972 year-class as the best
set of Pismo clams in almost thirty years. The 1972 year-class of Pismo clams
on that beach showed normal or better than normal survival through at least
early 1975 on all three traditional transect locations. During 1975 the observed
survival fell below the expected (Tomlinson 1968) (Figure 6). A disparity
between the expected (Herrington 1929) and observed growth of the 1972
year-class was apparent after late 1974 (Table 8). This disparity broadened after
the beach was closed to sport harvest in March 1975. The 1972 year-class clams
were expected to be 7.1 cm and 8.8 cm during the 1975 and 1976 surveys,
respectively. Instead the observed average lengths were 6.6 cm and 7.4 cm. This
disparity increased through 1978. After 1978 no 1972 year-class clams were
found.
100 n
80-
0
.o
E
3
60-
40-
20-
1973
Expected
Observed
1974
1975
1976 1977
Survey Year
1978
198
FIGURE 6.
Mean number of observed and expected Pismo clams of the 1972 year-class from al
traditionally sampled transects (three) on Atascadero Beach.
TABLE 8. Observed and Expected Mean Lengths ( ± S.D.) of 1972 Year Class Pismo Clams
from Atascadero Beach.
Observed
Year length
1972 13.9 (3.5)
1973 40.8
1974 49.8 (5.1)
1975 58.9 (6.3)
1976 65.0 (6.8)
1977 74.1
1978 80.0
Expected
n
length
n
64
15.2 (3.4)
139
433
37.0 (4.0)
109
160
55.1 (4.8)
65
154
72.8 (6.7)
41
50
89.2 (9.0)
39
18
105.8 (8.6)
32
2
113.8 (7.2)
17
210 CALIFORNIA FISH AND GAME
DISCUSSION
Human harvest of Pismo clams along Pismo Beach following the fur trade era
was sufficient to reduce high density stocks, which had grown as a result of the
removal of the otter, to a fully utilized level. A historical account of the changes
in regulations governing the Pismo clam fishery reflect this transition (Fitch
1950). Pismo clams show great temporal and spatial variability in recruitment,
yet recruitment was sufficient to maintain a sustained long-term harvest. The
harvest of legal-sized clams estimated from catch statistics appeared to be, at
least for peak use days, the same in 1 975 as it was in the late 1 940's (Fitch 1 950,
J. Fitch, Calif. Dept. Fish and Game, pers. comm.). The catch, however, may
now be divided among more diggers. The CPUE data, despite rather dramatic
fluctuations from 1975 through 1979, suggest that production continued at that
same level through 1979. Beginning in 1979, CPUE for the Pismo Beach Pismo
clam fishery dropped to zero in a stepwise fashion along the beaches progressing
from north to south.
Our observations of the decline of the Pismo Beach Pismo clam fishery, based
on otter sightings, clammer censuses, and clam transects, substantiate earlier
observations on the Morro Bay and Monterey Bay clam fisheries (Miller, Hard-
wick, and Dahlstrom 1975) and provide further evidence that sea otters are
directly responsible for the loss of these sport fisheries.
Using information available on numbers of otters south of Point San Luis
(Figure 3) and assumed consumption rates of Pismo clams (80 clams/otter/
day), it is possible to illustrate the impact sea otters had on the Pismo clam
resource in the Pismo Beach area. Even after subtracting for a percentage of
otters potentially feeding in nearby rocky habitat, the estimated number of clams
consumed still exceeded 700,000 in 1980. This number of clams is more than
double the highest yearly clammer take at Pismo Beach during the 1975 through
1979 period (Table 6). At this rate, it is easy to understand why the Pismo clam
fishery, already being fully utilized collapsed so dramatically and so completely.
What potential is there for a resurgence of a Pismo clam fishery within the
sea otter's range in the future? None of the Pismo clam fisheries which once
existed in areas now occupied by sea otters currently provide a harvestable
resource. However, several alternative views exist on the potential for a resur-
gence of a fishery in the future (VanBlaricom 1981 ).
Relatively low adult Pismo clam densities have produced successful sets in the
past and could potentially do so in an area occupied by sea otters. If sea otter
foraging pressure was low a sufficient number of clams from a major set might
survive to support a fishery. Sea otter foraging pressure does decline in an area
when the large peripheral male group moves on to new areas.
One major set of Pismo clams has occurred within the sea otters range on
Atascadero State Beach in 1972. The data collected on the growth and survival
of this year-class strongly support the contention that clam stocks will not reach
a high enough level to provide reasonable sport use. The extremely large 1972
year-class of Pismo clams on Atascadero Beach was not available as forage to
sea otters, due to the small size of the clams, when the peripheral male group
moved through that area. The survival of that year-class of clams appeared
excellent until individuals reached a size generally accepted as being available
forage for sea otters, at which point survival declined sharply. Coincidentally, a
SEA OTTER RANGE EXPANSION AND LOSS OF PISMO CLAM FISHERIES 21 1
disparity between observed and expected mean sizes became apparent, indicat-
ing that larger individuals were being selectively removed. It should be empha-
sized that Atascadero Beach was closed to sport clamming from March 1975 to
March 1978 and the clam stocks subject to predation by a low density sea otter
population. No legal sized clams have been observed in transect digs or clammer
interviews since 1973.
Data from trench transects and clammer interviews collected in Monterey Bay
through 1 986 also substantiate the contention that a sport fishery for Pismo clams
cannot coexist with sea otters. No legal-sized clams have been observed from
that area (Monterey Bay) since 1976, despite continued clam recruitment and
very low densities of sea otters (J. Hardwick, Calif. Dept. Fish and Game, pers.
comm.)
Two conclusions can be drawn from these data which have a direct bearing
on the debate concerning sea otter-shellfish fishery relationships. Firstly, sea
otter foraging, rather than human impact from legal harvest, illegal harvest,
pollution, and/or vehicle traffic on beaches, have resulted in the loss of Pismo
clam fisheries within the sea otter's range. Secondly, once sea otters are estab-
lished along clam bearing beach areas, any future harvestable stocks of clams
will be fully utilized by sea otters, preventing the return of a fishery.
ACKNOWLEDGMENTS
This study was supported in part by SECTION 6 funds pursuant to the Endan-
gered Species Act of 1973 through interagency agreement with the United States
Fish and Wildlife Service. We thank A. Baker, K. Barry, G. Coombes, T. Edel, B.
Hatfield, R. Hirano, L. Laurent, P. Lehtonen, N. Olson, S. Owen, C. Pattison, S.
Schultz, N. Siepel, L. Smith, and K. Worcester for assistance in conducting
clammer interviews. We also thank B. Hatfield for considerable assistance in
obtaining data on sea otter distribution and food habits.
LITERATURE CITED
Armstrong, ). J. 1979. The California Sea Otter: Emerging conflicts in resource management. San Diego Law Review,
16(2):249-285.
Baxter, J. L. 1961. Results of the 1955-1959 Pismo clam census. Calif. Fish Came, 47(2) :153-162.
Burge, R. T. 1979. The Pismo clam fishery and resource: Before and after sea otter reoccupation. Sea otter
workshop, Santa Barbara Museum of Natural History, Santa Barbara, California.
Carlisle, ). C. 1966. Results of the 1961-1965 Pismo clam census. Calif. Fish Came, 52 (3) :1 57-160.
1973. Results of the 1971 Pismo clam census. Calif. Fish Came, 59(2):138-139.
Estes, |. A. and C. R. VanBlaricom. In press. Sea otters and shell fisheries. In R. Beverton, D. Lavigne, and ).
Beddington ed., Marine Mammals and Fisheries. Allen and Unwin, London.
Fitch, J. E. 1950. The Pismo clam. Calif. Fish Came, 36(3):285-312.
1952. The Pismo clam in 1951. Calif. Fish Came, 38(4):541-547.
1954. The Pismo clam in 1952 and 1953. Calif. Fish Came, 40(2) :1 99-201.
1955. Results of the 1954 Pismo clam census. Calif. Fish Came, 41 (31:209-211.
Herrington, W. C. 1929. The Pismo clam: Further studies on its life-history and depletion. Division of Fish and Game
of California, Fish. Bull. No. 18, 1-67.
Kenyon, K. W. 1969. The Sea Otter in the Eastern Pacific Ocean. U.S. Fish. Wildl. Serv. No. AM. Fauna, No. 68,
1-352.
Kimker, A. 1982. Shellfish report to the Alaska Board of Fisheries. Dept. Fish and Game, Div. of Commercial
Fisheries. 9 p.
Matkin, C. 1981. Observations of marine mammal interactions with some Alaskan fisheries. Marine Mammal
Fishery Interaction Workshop, Vancouver, Washington. 3 p.
212 CALIFORNIA FISH AND CAME
Miller, D. ). 1974. Skindivers, abalone and sea otters. Outdoor California. Vol. 35, No. 4, 1-4.
Miller, D. ]., J. E. Hardwick and W. A. Dahlstrom. 1975. Pismo clams and sea otters. Calif. Fish and Game, Mar.
Resources Tech. Rep. 31, 49 p.
Sokal, R. R. and F. |. Rohlf. 1969. Biometry. W. H. Freeman and Co., San Francisco. 776 p.
Stephenson, M. D. 1977. Sea otter predation on Pismo clams in Monterey Bay. Calif. Fish Game, 63(21:117-120.
Tomlinson, P. K. 1968. Mortality, growth, and yield per recruit for Pismo clams. Calif. Fish Game, 54(2):1OO-107.
VanBlaricom, G. R. 1 981 . Sea otters and Pismo clams in California: some alternatives to the doomsday prediction.
Marine Mammal Conference, San Francisco.
Wade, L. S. 1975. A sea otter possible feeding on Pismo clams. ). of Mammalogy, 56:720-721.
Wild, P. W. and ). A. Ames. 1974. A report on sea otters, Enhydra lutris, in California. Calif. Dept. Fish and Game,
Mar. Res. Tech. Rept., 20:1-93.
LIFE HISTORY OF THE WOOLLY SCULPIN 213
Calif. Fish and Came 72(4): 213-226 1986
ASPECTS OF ECOLOGY AND LIFE HISTORY OF THE
WOOLLY SCULPIN, CLINOCOTTUS ANALIS, FROM
SOUTHERN CALIFORNIA 1
ALAN W. WELLS2
Department of Biology
California State University, Long Beach
Long Beach, California 90840
Approximately 3000 woolly sculpin, Clinocottus analis, were captured over a 17-
month period at Point Fermin, California, in tidepools from 1.07 m to —0.52 m
elevation from Mean Lower Low Water. An average density of 8.5 fish/m 2 and a
progressive shift of larger individuals toward lower tidal levels were noted. Spawning
was inferred to occur primarily during September through November with peak
recruitment to tidepools during November through February. Males grew faster and
to a larger size than females; von Bertalanffy growth coefficients L^, k, and t0 were
119.0, 0.71, and —0.10 for males and 96.3, 1.00, and —0.07 for females, respectively.
Maximum lifespan was 6 years for females and approximately 8 years for males. All
specimens over 60 mm tl appeared sexually mature with batch fecundity described
by the linear function F = 11.6 tl— 620.6. Overall sex ratio was not significantly
different than 1:1. Woolly sculpin preyed on copepods, isopods, gammarideans,
polychaetes, and mollusks, with the importance of each group changing markedly
with fish size. The presence of two parasites, Opecoelus adsphaericus and Ascaro-
phis sp., was noted.
INTRODUCTION
The woolly sculpin, Clinocottus analis, ranges from Cape Mendocino, Califor-
nia, to at least Punta Ascuncion, Baja California (Miller and Lea 1972). It is also
present around Coronado, Guadalupe, San Martin, Cedros and the Channel
Islands. Although the woolly sculpin is one of the most abundant intertidal fishes
throughout much of its range, only certain aspects of its life history and ecology
have been investigated. Eigenmann (1892) and Budd (1940) investigated its
larval development, while Hubbs (1966) studied fertilization, early cleavage,
and influence of temperature on hatching. Homing was studied by Williams
(1957) and Richkus (1968, 1978, 1981). Local distribution and diet were investi-
gated by Mitchell (1953), Johnston (1954), Mollick (1968, 1970) and Yo-
shiyama (1980). This study attempts to fill some of the gaps in knowledge
concerning the life history of this common intertidal fish.
MATERIALS AND METHODS
The study was conducted 1.5 km northwest of Point Fermin, Los Angeles
County, California (lat 33° 42' N, long 118° 17' W). Semi-monthly tidepool
collections were made between May 1971 and September 1972. Of the 64 total
samples, 51 tidepools were sampled once, 5 were sampled twice, and 1 was
sampled three times. Average length and width of each tidepool sampled was
recorded as well as an estimate of the degree of cover and tidal elevation. Nearly
all tidepools sampled were less than 0.5 m deep.
Collections were made using a solution of 1 0% quinaldine in ethyl alcohol and
anesthetized fish were removed either by hand or small dip net. All fish were
1 Accepted for publication February 1986.
2 Present address: Lawler, Matusky and Skelly Engineers, One Blue Hill Plaza, Pearl River, NY 10965.
214 CALIFORNIA FISH AND GAME
collected in tidepools with sparse cover; however, in extremely rocky tidepools
with numerous crevices, possibly a small proportion of the fish escaped detec-
tion. In the laboratory, fish were weighed to the nearest decigram and gonads
to the nearest milligram. Total length (tl), to the nearest millimetre, and sex
were also recorded. Unless otherwise stated, all lengths are given as tl.
Age was determined by length-frequency analysis (Petersen method) and by
counting annuli in otoliths. All otoliths were read twice and any pair of readings
not in agreement was disregarded. Von Bertalanffy growth functions were fit
using the Program BGC3 (Abramson 1971).
Approximately 20 fish per months of about equal numbers of adults and
juveniles, males and females, were chosen for stomach analysis; 248 fish
between 1 5 mm and 1 70 mm were examined. Both frequency of occurrence and
number of individual prey items were recorded. Food volumes per category and
an index of stomach fullness ( ISF ) were estimated since volumes were often too
small to be measured directly. An index of relative importance (IRI,), the
relative contribution of the , th food category to total diet, was calculated from:
N
IRIi = 1/N 2 (v„ISF,)100
i = 1
where: N — number of stomachs analyzed in the fish-size category under
consideration; Vjj = estimated volumetric proportion of total stomach content
of the | th food category in the -, th individual; and ISF -, — estimated fractional
stomach fullness of the ; th individual.
RESULTS AND DISCUSSION
Habitat and Associations
The intertidal zone at Point Fermin consists of shoreward tilted, parallel edges
of bedrock with scattered boulders. At a tide of —0.55 m elevation from Mean
Lower Low Water (MLLVV), 65 m of beach is exposed. Tides are mixed semi-
diurnal with maximum high tides about 2.13 m and extreme low tides about
-0.58 m.
Tidepool temperatures were extremely variable but generally within ± 3° C of
ambient surf temperature. Monthly mean surf temperature ranged from 12° C in
April to 20° C in August. Although pool temperatures as high as 27° C were
recorded, no woolly sculpin were taken in pools greater than 22° C. Salinity was
fairly constant at 34 %0 throughout the study. Evaporation in upper tidepools
rarely increased salinity more than a few parts per thousand. However, freshwa-
ter runoff lowered the salinity of several of them to 24 %0 on one occasion.
Predominant organisms in the study area were typical rocky shore forms.
Periwinkle, Littorina planaxis, and turban shell, Tegula funebralis, were common
in higher intertidal zones. Mussels, Mytilus californianus, and goose barnacles,
Pollicipes polymerus, were characteristic of mid-intertidal regions. Boulder sur-
faces and tidepool bottoms in lower zones were often densely covered with
coralline red algae. Several organisms, such as shore crab, Pachygrapsus cras-
sipes, and sea anemone, Anthopleura xanthogrammica, were found throughout
all zones. Fishes commonly associated with woolly sculpin in the intertidal
region included: rockpool blenny, Hypsoblennius gilberti; juvenile opaleye,
Girella nigricans; spotted kelpfish, Gibbonsia elegans; striped kelpfish, Gibbonsia
LIFE HISTORY OF THE WOOLLY SCULPIN 21 5
metzi; California clingfish, Gobiesox rhessodon; and dwarf surfperch, Micromet-
rus minimus. Species taken on rare occasions in the tidepools included: smooth-
head sculpin, Artedius lateralis; black perch, Embiotoca jacksoni; zebra perch,
Hermosilla azurea; mussel blenny, Hypsoblennius jenkinsi; rosy sculpin, Oligo-
cottus rubellio; fluffy sculpin, O. snyderi; pile perch, Rhacochilus vacca; cabe-
zon, Scorpaenichthys marmoratus; and juvenile rockfishes, Sebastes spp.
Woolly scullpin was numerically the most abundant fish in the intertidal region,
outnumbering the second most abundant species, rockpool blenny, by approxi-
mately 10 to 1.
Distribution Within Study Area
Woolly sculpin were collected from tidepools as high as 1 .07 m above MLLW,
but were occasionally observed above this level. Individuals were taken from all
tidepools sampled below 0.91 m to the lowest tidepool sampled at —0.52 m but
the region of greatest abundance was from 0.61 m to —0.30 m. Observations
made while skin diving suggest a reduced population at tidal levels below —0.61
m. This is consistent with the observation made by Hubbs (1966) at San Diego
that baited traps set below this level were seldom effective.
The average population density, excluding newly settled young-of-year, was
8.5/m2 of tidepool surface but ranged as high as 27/m2. Tidepool size, which
ranged from 0.6 m2 to 74.3 m2, was not significantly correlated with the popula-
tion density of that pool (r = —0.108, n = 62, P>0.05). However, those
tidepools with moderate to heavy cover (i.e., crevices, rocks, vegetation) ap-
peared to support higher population densities than their lightly covered counter-
parts. Although no major seasonal shift in intertidal distribution was noted,
tidepools high in the intertidal region which lacked fish during summer months
contained a few individuals during winter months.
luvenile woolly sculpin less than approximately 25 mm in length were rarely
found in pools inhabited by adults. These young fish were typically found in
small, shallow, coralline algae filled pools between 0.45 to —0.18 m. Richkus
(1968, 1981 ) also noted that the distribution of these fish differed from that of
larger individuals. He found fish less than 35 mm occurring even in small, sandy
depressions that would drain before being resubmerged.
Juvenile and adult woolly sculpin greater than 40 mm in length demonstrated
a marked tendency for larger fish to inhabit increasingly lower tidal levels ( Figure
1 ). This trend was also noted by Williams (1957) in the nearby Palos Verdes
area.
Age and Growth
Recruitment of young-of-year into the intertidal region occurred during fall
through spring as evidenced by the collection of individuals less than 20 mm in
all months from November through May, except April (Figure 2). Peak recruit-
ment, however, occurred during November through February. Based on studies
by Budd (1940) on woolly sculpin and Morris (1951) on the closely related
species Clinocottus recalvus, these 11-25 mm fish were newly settled from the
pelagic larval phase and hatched approximately 6-8 weeks prior. By April, the
1971-72 spawned fish averaged approximately 48 mm. Woolly sculpin spawned
the preceding year increased from approximately 45 mm in May to about 62 m
in September. The slightly smaller average length in May for the 1970-71 year
class as compared to April for the 1971-72 year class is likely the result of
sampling error and/or annual variation in growth rate.
216
CALIFORNIA FISH AND GAME
E
E
140-
120-
l- 100H
o
z
111
J 80H
£ J
O 60H
40-
•II-
I l I I I
.8 .6 .4 .2 0
-i r— -i
-.2 -.4 -.6
TIDAL LEVEL (m)
FIGURE 1. Relationship of woolly sculpin total length to tide level during Fall 1971. Mean, range,
standard deviation, and two standard errors are indicated.
Mean size at age for fish older than 1 year was determined from otoliths of
1 82 fish collected March through June 1 972. Maximum observed age for females
was 6+ while a minimum of seven age classes could be established for males.
Age class 1 +, 2 + , 3 + , 4 + , 5 + , and 6+ females averaged 82.5, 86.5, 93.1, 94.8,
98.3 and 100.0 mm, respectively, while males averaged 87.7, 101.3, 110.6, 112.5,
1 18.6 and 145.5 mm, respectively (Table 1 ) . The largest fish captured during the
study, a 170 mm male, appeared to be at least 8 + . The majority of specimens,
88 percent based on November through May length-frequency data, were one
year or less in age.
A growth function was calculated by combining monthly mean length for
juveniles, estimated from 0+ length-frequency modes, with the mean lengths
obtained from otolith analysis. Age was referenced from a presumed 15 Septem-
ber birthday. Since otolith samples were collected during March through June,
age was taken as the mid-point of the period, i.e., 1 May. Only otolith ages 1 +
through 5 + were included since the sample size for age 6+ and older was small.
LIFE HISTORY OF THE WOOLLY SCULPIN 217
Fitted von Bartalanffy functions (Figure 3) were:
Female
L, = 96.3(1 - e-100,,+0071)
Male
L, = 119.0(1 - e-071,,+0101)
for the general equation:
L, = U, (1 - e*(M<>»)
L, = length at time t in years, L^, = asymptotic length, k = growth constant,
and to = hypothetical age at zero length.
Average growth rate (B) at time t may be obtained from the von Bertalanffy
coefficients using Knight's equation (Ricker 1975):
B = kL00e-k,,-,°)
From this relationship the average growth rate was 6.1 mm per month during
the first month (November) after settling. By the following April this rate had
decreased to 4.3 mm per month. In subsequent months the disparity between
male and female growth rates become increasingly apparent. At age 1 year the
average growth rate for males was 3.2 mm per month while females had slowed
to an average of 2.7 mm per month.
It should be noted that these growth estimates, especially for males, likely
become increasingly biased low with increasing age. As discussed above, woolly
sculpin, with increasing size, tend to inhabit pools at increasingly lower tidal
levels. These pools are less often exposed by low tides and typically offer greater
cover. Since these fish are, therefore, more difficult to capture, a systematic bias
towards slower growing fish is probable.
Growth in weight with length for pooled sexes and juveniles was found to be:
W = 0.00001 74TL2958 (r = 0.997; n = 157; P <0.01)
where:
W = weight in grams.
For males the relationship was,
W = 0.00001 59TL2988 (r = 0.983; n = 128;/> <0.01)
while for females it was,
W = 0.0000346TL2793 (r = 0.988; n = 126; P <0.01).
Growth in length or weight with standard length (sl) may be obtained by
applying the relationship,
sl = 0.15 + 0.81TL.
Reproduction
Sexual maturity for both sexes appears to occur near the end of the first year
of life as well-developed gonads were observed in all fish over 60 mm. Mature
ovaries were found in all months; however, highest average GSI (gonad wt/
body wt X 100) values occurred in October through November (Figure 4).
Peak testicular development was observed during August through November.
Lowest average GSI values for both males and females were during February and
March. Although it appears that spawning may occur throughout much of the
year, peak reproduction, as inferred from GSI values, likely occurs during Sep-
tember through November. This is in reasonable agreement with estimates of
218
CALIFORNIA FISH AND GAME
N
87
56
227
129
20
122
129
206
359
471
203
337
MONTH
1972
120 160
TOTAL LENGTH (mm)
FIGURE 2. Monthly length-frequency distribution for woolly sculpin.
2
3
4
5
6
86.5
93.1
94.8
98.3
100.0
4.6
4.8
7.2
2.6
-
76-92
83-100
87-100
95-102
-
10
14
9
6
1
101.3
110.6
112.5
118.6
145.5
9.5
15.5
10.5
10.9
17.6
72-116
84-146
95-130
108-135
124-165
24
21
17
5
4
LIFE HISTORY OF THE WOOLLY SCULPIN 219
peak spawning deduced from the appearance of young in the tidepools when
3 to 4 weeks for hatching (Hubbs 1966) and 6 to 8 weeks for a pelagic larval
phase are allowed. Eigenmann ( 1 892 ) also indicated spawning over a prolonged
period with young occurring in tidepools 2 to 3 months later.
TABLE 1. Woolly Sculpin Total Length (mm) at Age Based on Otoliths.
Age Class
Female /
Mean 82.5
SD 4.8
Range 70-87
n 11
Male
Mean 87.7
SD 3.9
Range 80-93
n 15
Frequency analysis of ovarian egg diameter suggests the presence of at least
three modal groups of eggs. Generally, there was a single, but distinct, group of
eggs greater than 0.7 mm diameter. These eggs averaged approximately 1 .24 mm
in diameter and, in an unpreserved state, were translucent greenish-yellow to
reddish-brown in color. It appears that these are the only eggs released during
a spawn. Batch fecundity (F), the potential number of eggs produced during a
single spawn, was obtained by direct count of all eggs over 0.7 mm diameter.
The linear equation F= 11.6 tl — 620.6 (r = 0.940; n= 45; P< 0.01 ) appears to
adequately describe the relationship between the number of eggs and fish length
(tl in mm) (Figure 5). Logarithmic and semi-logarithmic transformations failed
to significantly improve the fit. The largest female examined (110 mm) con-
tained 784 eggs; the mean number was 242 eggs per average reproductive female
(74 mm). These fecundity estimates are very similar to those from Pacific Grove
specimens and substantially lower than from La Jolla specimens reported by
Hubbs (1966).
The total egg production during a single spawning season could not be deter-
mined since the number of spawnings per season is unknown. However, the
occurrence of several modal groups below 0.5 mm diameter does suggest multi-
ple spawnings. Hubbs (1966) postulated that females lay several complements
per season since over half the specimens captured and held in isolation devel-
oped ripe eggs within two weeks.
The number of males and females in the population appears to be about equal.
During the course of the study, the sex of 1776 adult sculpin was determined.
Of these, 865 (48.7%) were male and 911 (51.3%) were female. Chi-square
analysis suggests no significant departure from a 1:1 sex ratio (x2 = 1.14; df=1;
P > 0.05 ) . When analyzed by month, the percentage of females ranged from 60.5
in January to 36.7 in August (Table 2). Although sex ratios in January, May, and
June departed significantly from 1:1, no consistent or biologically meaningful
trend was discernible.
Food and Feeding
Woolly sculpin appear to feed on a wide diversity of intertidal organisms. In
the 248 stomachs examined, at least 33 prey categories could be established
220
CALIFORNIA FISH AND CAME
( Table 3 ) . Frequently encountered prey items, those with 25% or greater occur-
rence, weregammarideans (50.8%), copepods (31.5%), and the isopod Cirola-
na harfordi (26.2%). Clearly, small crustaceans formed the bulk of the diet. Less
frequently encountered items, 20-25% occurrence, were polychaetes (23.0%),
the tanaidacean Anatanais normani (22.2%), and mollusks (21.8%). Algae
(19.8%), larvae of the dipteran Paraclunio sp. (17.3%), and the polychaete
Phragmatopoma californica (15.7%) were also commonly encountered.
120
UNSEXED YEARLINGS
FEMALES
MALES
7
8
AGE (YEARS)
FIGURE 3. Size at age relationship for woolly sculpin at Point Fermin, California.
As measured by the Index of Relative Importance (IRI) diet composition
changed markedly with fish size (Figure 6). Approximately 90% of the diet of
sculpin under 20 mm was copepods. As fish increased in length (30-100 mm),
this percentage decreased and there was increased emphasis on larger items
such as amphipods, isopods, and polychaetes. Woolly sculpin over approxi-
mately 80 mm, predominantly males, placed increased reliance on relatively
LIFE HISTORY OF THE WOOLLY SCULPIN
221
large items such as mollusks, primarily chitons and Acmaea spp., and decapods,
primarily Pagurus samuelis and young Pachygrapsus crassipes. The overall pat-
tern of utilization appeared to be one of increasing prey size with increasing
body size.
MALE
1.8n
1.6
1.4
1.2<
- 1.0H
° .8-
.6"
.4<
.2-
FEMALE
10i
8
_ 6'
O 4.
J JASONDJ FMAM
MONTH
FIGURE 4. Monthly variation in gonosomatic index (CSI) for woolly sculpin. Mean, range, and
sample size are indicated.
222
CALIFORNIA FISH AND CAME
800
600«
O
LU
£ 400
00
200-
50
60
70
80
90
i
100
110
120
TOTAL LENGTH (mm)
FIGURE 5. Relationship of total length (u) to number of mature ova in woolly sculpin.
TABLE 2. Adult Woolly Sculpin Population Sex Composition by Month and Chi-Square Test
of 1:1 Sex Ratio Hypothesis
Month Total
January 258
February 385
March 105
April 103
May 141
June 336
July 182
August 30
September 12
October 93
November 52
December 79
Total 1,776
Percentage
Male
Female
Chi-square
39.5
60.5
10.9**
48.3
51.7
0.4
54.3
45.7
0.6
40.8
59.2
3.1
39.7
60.3
5.6*
57.4
42.6
7.1**
46.7
53.3
0.7
63.3
36.7
1.6
58.3
41.7
0.1
51.6
48.4
0.0
61.5
38.5
2.3
48.1
51.9
0.1
48.7
51.3
1.1
*p <
0.05
0.01
LIFE HISTORY OF THE WOOLLY SCULPIN 223
TABLE 3. Diet by Frequency Occurrence, Percent Occurrence, and Number of Organisms
of 248 Woolly Sculpin (15-170 mm tl) collected May 1971 through September
1972.
Frequency Percent Number
Protozoa
Phytomastigophora 1 0.4 1
Sipunculida 2 0.8 2
Annelida
Phragmatopoma californica 39 15.7 70
Unident. Polychaeta 23 9.3 35
Mollusca
Polyplacophora 23 9.3 35
Acaema spp 25 10.1 35
Littorina planaxis 2 0.8 2
Unident. Prosobranchia 1 0.4 1
Opisthobranchia egg mass 1 0.4 1
Arthropoda
Halacaridae 1 0.4 1
Pycnogonida 2 0.8 2
Ostracoda 15 6.0 24
Copepoda 78 31.5 3,700
Cirripedia cirri 16 6.5
Mysidacea 2 0.8 5
Anatanais normani 55 22.2 504
Cirolana harfordi 65 26.2 160
Exosphaeroma sp 1 0.4 2
Valvifera 32 12.9 135
Gammaridea 126 50.8 529
Spirontocaris sp 2 0.8 2
Bateis sp 2 0.8 2
Pagurus samuelis 9 3.6 10
Pachygrapsus crassipes 10 4.0 10
Cancer sp 1 0.4 1
Unident. Decapoda 4 1.6 4
Coleoptera 3 1.2 3
Paraclunio sp. larvae 43 17.3 115
Unident. Diptera 2 0.8
Echinodermata
Strongylocentrotus purpuratus
tube feet 20 8.1 103
Chordata
Hypsoblennius gilberti 1 0.4 1
Fish eggs 8 3.2 120
Unident. Algae 49 19.8
Empty 29 1 1.7
Although the normal mode of feeding appears to be that of engulfing whole
organisms, woolly sculpin seem to occasionally engage in a browsing-like mode
of feeding. This was inferred from the occurrence of such items as barnacle cirri,
sea urchin tube feet, and the anterior-most portion of the tube worm Phrag-
matopoma californica.
The findings of this study closely parallel those of several other workers in
demonstrating that small crustaceans play an important role in the diet of woolly
sculpin. Mitchell (1953) found that decapods, especially the shrimp Spirontoca-
ris picta, comprised the bulk of the stomach contents of 12 60-1 10 mm woolly
sculpin examined from the Palos Verdes area. Johnston (1954) found copepods
to be the most numerous and most frequently encountered food item in the
224
CALIFORNIA FISH AND CAME
LU
O
DC
o
Q.
2
hi 40«i
>
b 20
0
40
20H
0
40-i
20
0
LU
cc
LL
O
x
LU
Q
COPEPODA
I i i
POLYCHAETA
MOLLUSCA
I I I l
DECAPODA
l i l I I
100 120 140
40 60 80
TOTAL LENGTH (mm)
FIGURE 6. Index of Relative Importance (iri) of selected food items versus total length of woolly
sculpin. Points represent mean value for 20 mm size intervals.
summer diet of individuals from Monterey County while Mollick (1968, 1970)
found amphipods to be the dominant prey by mean volume and copepods to
be the most frequently encountered item during July at Bird Rock, San Diego.
Yoshiyama (1980) also found small crustaceans to be the most important cate-
gory in the diet of woolly sculpin from central California.
Yoshiyama (1980) did report, however, that in central California woolly
sculpin consume substantial quantities of algae, a resource exploited by few
other rocky intertidal fishes in California. He found algae, primarily Petalonia, in
29% of the 62 stomachs analyzed and concluded that the concomitant lack of
animal prey in some specimens indicated that algae was not taken incidentally.
He also concluded that the exploitation of this food resource served to separate
woolly sculpin ecologically from the two more carnivorous sculpin, fluffy sculpin
LIFE HISTORY OF THE WOOLLY SCULPIN 225
and smoothhead sculpin, common to the region. He was uncertain as to whether
the apparent de-emphasis of algae in the diets of southern California woolly
sculpin observed by Mitchell (1953) Johnston (1954) or Mollick (1970) result-
ed from sampling vagaries, differences in the underlying resource base, or popu-
lation differences in feeding habits.
A reduction in algae utilization by woolly sculpin in southern California rela-
tive to central California might be expected based on the occurrence of potential
competitor species. Yoshiyama (1980) found that besides woolly sculpin, the
most frequently encountered species were the smoothhead and fluffy sculpins.
These two carnivorous species were only rarely encountered during this study.
Instead, the most frequently encountered species were the rockpool blenny and
juvenile opaleye (30-100 mm TL). Both species utilize substantial amounts of
algae (Mitchell 1953, Dayneko 1975). Yoshiyama stated that opaleye were not
encountered during his study and rockpool blenny do not range north of Point
Conception (Miller and Lea 1972). Therefore, a decreased utilization of algae
by woolly sculpin in southern California would appear advantageous in lessening
interspecific competition.
It should also be noted that the occurrence of algae in the stomach does not
necessarily imply utilization. Examination of the posterior gut made during the
present study indicated that little, if any, digestion of algae had occurred. This
suggests, that while algae is ingested, it may be either coincidental with prey
capture or purposeful to obtain organisms associated with it. A concomitant lack
of animal prey may result from unsuccessful attempts at capturing prey or
differential digestive rates.
Parasites and Predators
Intestines of 21 woolly sculpin (56-116 mm), collected April 1972, were
examined for parasites. The diagenetic trematode, Opecoelus adsphaericus, was
found in 17 (81%), with an average of four trematodes per fish (range 1-15).
There is no significant correlation between fish size and number of trematodes
(r = 0.140, n = 21, P>0.05).
A nematode, Ascarophis sp., occurred in 12% of the 248 (i.e., all) stomachs
examined. It appeared most abundant during late winter and early spring. Inci-
dence was highest in woolly sculpin over 60 mm.
Although numerous avian and fish predators of woolly sculpin are suspected,
only one was confirmed; a 29 mm woolly sculpin was removed from the stom-
ach of a 140 mm spotted kelpfish. Cabezon and kelp bass, Paralabrax clathratus,
were common subtidally and were observed in the intertidal region during high
tide. Both are reported to prey on sculpins (O'Connell 1953, Smith 1970).
ACKNOWLEDGMENTS
I thank C. W. Hill for suggestions and criticisms throughout this study, J. R.
Dayneko for help with the field work, R. Appy for identification of the parasites,
E. D. Lane and J. S. Nelson for editorial assistance, and D. M. Wells for preparing
the figures. This paper was extracted from a thesis submitted to the Department
of Biology, California State University, Long Beach, in partial fulfillment of the
requirements for the degree Master of Arts.
226 CALIFORNIA FISH AND CAME
LITERATURE CITED
Abramson, N.J. 1971. Computer programs for fish stock assessment. FAO (Food Agric. Org., U.N.) Fish. Tech.
Pap. 101:1-154.
Budd, P.L 1940. Development of the eggs and larvae of six California fishes. Calif. Dept. Fish and Came, Fish Bull.
56:1-53.
Dayneko, |.R. 1 975. Life history of the rockpool blenny, Hypsoblennius gilberti ( Jordan ) , at Point Fermin, California.
Thesis, Calif. State Univ., Long Beach. 94 p
Eigenmann, C.H. 1892. The fishes of San Diego, California. U.S. Natl. Mus., Proc, 15(8971:123-178.
Hubbs, C. 1966. Fertilization, initiation of cleavage, and developmental temperature tolerance of the cottid fish
( HnocottUS analis. Copeia, 1966(11:29-42.
Johnston, R.F. 1954. The summer food of some intertidal fishes of Monterey County, California. Calif. Fish Game,
40(11:65-68.
Miller, D.J., and R.N. Lea. 1972. Guide to the coastal marine fishes of California. Calif. Dept. Fish and Game, Fish
Bull. 157:1-235.
Mitchell, D.F. 1953. An analysis of stomach contents of California tidepool fishes. Amer. Midi. Nat., 49:862-871.
Mollick, R.S. 1968. Distribution of Clinocottus analis Girard [sic] in tide pools as related to substrate preference.
Thesis, San Diego State College, Calif. 82 p.
1970. Food habits of Clinocottus analis (Girard) Calif. Fish Game, 56(2) :1 33-1 34
Morris, R W. 1951. Early development of the cottid fish Clinocottus recalvus (Greeley). Calif. Fish Game,
37(3):281-3O0.
O'Connell, C.P. 1953. The life history of the cabezon, Scorpaenichthys marmoratus (Ayres). Calif. Dept. Fish and
Game, Fish Bull. 93:1-76.
Richkus, W.A. 1968. Aspects of the ecology of the wooly sculpin (Clinocottus analis, Girard) [sic]. Thesis, Univ.
of Calif., San Diego. 75 p.
1978. A quantitative study of intertidepool movement of the wooly sculpin, Clinocottus analis. Mar. Biol ,
49:227-284.
. 1981. Laboratory studies of intraspecific behavioral interactions and factors influencing tidepool selec-
tion of the wooly sculpin, Clinocottus analis. Calif. Fish Game, 67(3):187-195.
Ricker, W.E. 1975. Computation and interpretation of biological statistics of fish populations. Fish. Res. Bd. Canada,
Bull. 191. Ottawa.
Smith, S.H. 1970. A comparative study of food and feeding in the sand bass (Paralabrax nebulifer ) and the kelp
bass (Paralabrax clathratus 1. Thesis, Calif. State Univ., Long Beach. 88 p.
Williams, G.C. 1957. Homing behavior of California rocky shore fishes. Univ. Calif. Publ. Zool., 59:249-284.
Yoshiyama, R.M. 1980. Food habits of three species of rocky intertidal sculpins (Cottidae) in central California.
Copeia, 1980(31:515-525.
ICHTHYOFAUNAL TIDEPOOL RECOLONIZATION 227
Calif. Fish and Came 72(4): 227-231 1 986
ICHTHYOFAUNAL COMPOSITION AND RECOLONIZA-
TION IN A CENTRAL CALIFORNIA TIDEPOOL n
RONALD H. MATSON, C. BEN CRABTREE AND THOMAS R. HACLUND
Department of Biology
University of California
Los Angeles, California 90024
Species composition and age class distribution data were obtained for fishes col-
lected from a tidepool at San Simeon Reef, San Luis Obispo County, California. Two
collections, taken in November and December 1983, were made by applying rote-
none to the pool. Members of the family Cottidae were the most abundant fishes
in both samples and comprised 30% of the 1004 fishes collected. Clinids were second
in abundance in the November collection whereas stichaeids were second in De-
cember. Brillouin's species diversity index (H) was calculated for the two samples.
Comparison (t-test) of diversity indices indicate significant differences between the
two samples, suggesting that the time period between collections (43d) was not
sufficiently long to allow repopulation. The age class distributions of both samples
were similar with the notable exception of Gobiesox maeandricus, for which only
age class 1 individuals were represented in the second sample.
INTRODUCTION
The high diversity of the central California rocky intertidal zone and the
accessible nature of tidepools has made tidepool fishes model organisms for the
study of a number of ecological phenonema. Each species prefers a specific
microhabitat within the intertidal zone (Yoshiyama 1980, Barton 1982) and the
actual distribution of fishes is influenced by physical factors, such as water
temperature (Thomson and Lehner 1976), as well as biotic interactions such as
competition and predation (Yoshiyama 1980, 1981; Grossman 1982). Although
there is a wide range of environmental fluctuations within the intertidal zone, the
assemblage of fishes found here tends to be both persistent and resilient. Thus,
even after a major perturbation, the community structure of the tidepool appears
to return to its original state (Grossman 1982).
Investigations of the structure of tidepool fish communities often involve
experimental defaunation (Williams 1957, Grossman 1982). It is assumed that
recolonization of a tidepool is relatively rapid after the application of an ichthyo-
cide and that the time between defaunations is sufficiently long to allow recov-
ery (Grossman 1982). While the effect of defaunation on community structure
seems to be minimal in the long term (e.g., Thomson and Lehner 1976, Gross-
man 1982), the short term effects have not been well documented. If the
recovery time is insufficient, these collections will present a biased picture of
community structure. Therefore, it is important to have an indication of the
minimum time interval needed between such collections. Herein, we describe
the ichthyofaunal composition of a central California tidepool and examine the
short-term effects of the application of an ichthyocide (rotenone) on its com-
munity structure.
METHODS
The study site is located at San Simeon Reef, roughly 8 km north of San Simeon
1 Accepted for publication March 1986.
228 CALIFORNIA FISH AND GAME
Point, San Luis Obispo County, California (approximately lat 35° 39'N, long 121°
16'W). The tidepool selected was approximately 3 X 5 m with a maximum
depth of approximately 0.5 m. It lies in the mid-intertidal approximately 40 m
offshore from the mean high tide mark. The substrate was composed primarily
of rocks interspersed with patches of sand.
Fishes were collected during low tide on 5 November and 1 7 December 1 983
between 1500 and 1700 h. Water temperature in the tidepool on these two days
was approximately 12°C and 16°C, respectively. Derris root rotenone was em-
ployed as an ichthyocide. Following 10% formalin fixation and preservation in
45% isopropanol, specimens were identified, counted, and standard lengths
recorded to the nearest 0.5 mm. Age classes were separated through the use of
length frequency histograms (Ricker 1975). Data correlating standard length
with age were used for distinguishing age classes (Burge and Schultz 1973; Hart
1973; Fitch and Lavenberg 1975; R. D. Orton, pers. comm.). Where such data
were not available, discontinuities in the length frequency were used to assign
age class membership.
To assess the effects of defaunation on the structure of the tidepool ich-
thyofauna, a measurement of species diversity was calculated and compared
between the two samples. Brillouin's species diversity index (H) was chosen
because it ". . . is the preferred index for most problems in applied aquatic
ecology" (Stauffer, Reish, and Calhoun 1980, p. 185). This index represents the
total recoverable population of fishes within the tidepool, not just a random
sample (Thomson and Lehner 1976). Brillouin's index is:
H = 1/N In (N!/N1! N2! . . . Ns!)
where N is the total number of individuals in the sample; N1, N2 . . . Ns the
number of individuals in species 1, 2 ... s; and s the total number of species
collected (Pielou 1977). To determine if there was a statistically significant
difference between indices obtained for the two samples, a t-test was performed
( Zar 1 974, p. 1 1 5 ) . A G-test, using Williams' correction ( Sokal and Rohlf 1 981 ) ,
was utilized to determine if there were statistically significant differences in the
age class composition of certain species between the two samples. When em-
ploying the G-test on data for Artedius lateralis, age class 3 individuals were
pooled with age class 2 individuals.
RESULTS AND DISCUSSION
Species composition of the two collections is provided in Table 1. Cottids
were the most abundant fishes in both samples (28.7% and 33.1%, respective-
ly). Clinids were second in abundance in the November collection (21.8%), but
stichaeids were second in December ( 22.2% ) . The same species were obtained
in both collections with the following exceptions: Xiphister mucosus was absent
from the November sample whereas Atherniops affinis and Micrometrus aurora
were not present in the December sample. Brillouin's species diversity indicies
for the November and December collections were 2.34 and 2.04, respectively;
these value are significantly different (t = 7.85, df = 197, P < 0.05). Alterna-
tively, if the nonresident species, A. affinis and M. aurora (i.e., species not
typically found in the tidepool habitat), are omitted from the analysis, Brillouin's
species diversity index for the November sample is 2.19. However, the differ-
ence between the values for November and December remains significant (t =
3.90, df = 187, P < 0.05).
ICHTHYOFAUNAL TIDEPOOL RECOLONIZATION
229
TABLE 1. Species Composition And Relative Abundance Of Fishes Collected In The Two
Tidepool Samples. Taxonomy Follows Robins et al. (1980).
5 November 17 December
1983 1983
Taxa Number Percent Number Percent
Cobiesocidae
Cobiesox maeandricus 48 6.8 59 19.5
Atherinidae
Atberinops affinis 16 2.3
Cottidae
Artedius lateralis 79 11.2 56 18.5
Clinocottus analis 112 16.0 38 12.6
Oligocottus rimensis 1 0.1 1 0.3
Oligocottus snyderi 10 1.4 5 1.7
Kyphosidae
Cirella nigricans 36 5.1 18 6.0
Embiotocidae
Micrometrus aurora 87 12.4
Clinidae
Cibbonsia metzi 97 13.8 52 17.2
Gibbonsia montereyensis 56 8.0 1 0.3
Stichaeidae
Anoplarchus purpurescens 46 6.6 31 10.3
Cebidichthys violaceus 25 3.6 6 2.0
Xiphister atropurpureus 71 10.1 26 8.6
Xipbister mucosus 4 1.3
Pholidae
Xererpes fucorum 18 2.6 5 1.7
Totals: 702 302
Because of the small sample sizes, it was inappropriate to conduct G-tests
comparing age classes from both collections for all but three species (Table 2).
There were no statistically significant differences in the age class structure of
Artedius lateralis and Xiphister atropurpureus taken from both collections. For
Clinocottus analis there was a significant difference in the age classes represent-
ed in the two collections (G = 5.77, df = 1, P < 0.05) reflecting, in part, an
increase in the representation of age class 2 individuals from 26% in November
to 47% in December. Three age classes of G. maeandricus were found in
November, while only age class 1 individuals were present in December. A.
affinis and M. aurora collected in November were represented by only age class
1 individuals (standard lengths ranged between 28-51 mm and 48-89 mm,
respectively); these species were not obtained in December.
Our species diversity indices are within the range of those reported for other
tidepools of the eastern Pacific (Thomson and Lehner 1976). At the familial
level, composition of our tidepool is similar to that reported for other coastal
California tidepools (e.g., Yoshiyama 1981 ). However, our site, which is approx-
imately 120 km south of the San Mateo site of Yoshiyama (1981 ), has a very
different cottid and stichaeid fauna. Oligocottus snyderi, which was numerically
dominant in the tidepools sampled by Yoshiyama, is replaced by C. analis and
A. lateralis at our site. Yoshiyama (1981 ) found Anoplarchus purpurescens to
be most abundant followed by Cebidichthys violaceus, while X. atropurpureus
230 CALIFORNIA FISH AND GAME
was the most abundant stichaeid species at our site. The stichaeid composition
of our tidepool also differed from that of Barton ( 1 982 ) , whose Piedras Blancas
study site was near ours. He found that C. violaceus was most numerous among
this group of fishes. Reasons for these faunistic differences between collecting
sites are not totally clear (Yoshiyama 1981 ), although sampling bias may explain
some of them (e.g., the absence of X. mucosas in the November sample). That
fish were captured by different methods (i.e., some investigators employed
quinaldine while others used rotenone) may also account for some of the
differences between results. Furthermore, Moring (1981), Yoshiyama (1981),
and Barton (1982) have demonstrated habitat preference, in terms of vertical
distribution within the intertidal, for many of these species. Because we collected
only at one level within the intertidal zone during one season, the fact that the
species composition of our samples differ from others reported is not unexpect-
ed.
TABLE 2. Numbers Of Individuals In Each Age Class, As Determined From Length Fre-
quency Histograms, Represented In The Two Tidepool Samples.
5 November 17 December
19jB 1983
Species ' 1] 2 3 1 2 3
Gobiesox maeandricus (a)} 35 9 4 59 0 0
Artedius lateralis (a) 33 40 6 28 28 0
Clinocottus analis (a,b) 83 29 0 20 18 0
Oligocottus rimensis 10 0 10 0
Oligocottus snyderi 8 2 0 3 2 0
Girella nigricans (c) 35 1 0 18 0 0
Gibbonsia metzi (a) 91 6 0 51 1 0
Gibbonsia montereyensis (a) 47 9 0 1 0 0
Cebidicbtbys violaceus (a) 5 13 7 0 5 1
Anoplarchus purpurescens (a,d) 36 8 2 28 2 1
Xiphister atropurpureus (a,d) 36 35 0 17 9 0
Xererpes fucorum (a) 13 5 0 5 0 0
* Includes only species taken in both collections.
t Numbers refer to age classes in years: 1 (0-1 yr), 2(1-2 yr), and 3(2 yr and older).
} References for age/size class data: a) Burge and Schultz 1973, b) Fitch and Lavenberg 1975, c) R. D. Orton,
pers. comm., d) Hart 1973.
Our results, which are in general agreement with those of Grossman (1982),
indicate that representatives of all age classes are moving into the empty tidepool
habitat. Gobiesox maeandricus represents a notable exception. Three age
classes were represented in the November sample while only younger individu-
als were taken in the December sample. The absence of larger individuals in the
second sample may be an indication of the low vagility of older fishes. Other
exceptions are Xererpes fucorum, Gibbonsia montereyensis, and Girella nigri-
cans; however, due to small sample sizes, these observations may be attributable
to sampling error.
The statistically significant difference between H values suggests that a period
of 43 days is not a sufficient amount of time between artifical defaunations for
recolonization to occur such that species diversity is at pre-perturbation levels.
This is true even if the nonresident species (A. affinis and M. aurora ) are omitted
from the calculation of diversity indices (the chance capture of the two nonresi-
dent species may reflect a real biological phenomenon; not in terms of species
ICHTHYOFAUNAL TIDEPOOL RECOLONIZATION 231
composition of tidepool residents, but rather temporal changes in the use of
tidepools by these species). Williams (1957) noted that tidepools from which
he had removed fish had not fully repopulated within a period of several weeks
although Thomson and Lehner (1976) sampled tidepools at intervals as short as
three months and found no effect on the community variables which they
calculated. Thus, it would seem that a time period between 43 and 90 days is
needed to assure recovery and that assumptions of recovery within a shorter
time span may not be appropriate. To more accurately document recruitment
after defaunation, an expanded sampling strategy needs to be employed. Adja-
cent tidepools should be sampled over varying time intervals. In any case,
caution must be observed when using removal experiments in determining the
community structure of tidepools. When samples are taken within short time
periods, the total numbers may not only differ, but age class distributions may
also be affected.
ACKNOWLEDGMENTS
This study was supported in part by the Department of Biology, UCLA (Or-
ganismic Fund to CBC). Fishes were collected under a scientific collecting
permit granted to D. G. Buth by the California Department of Fish and Game
(#0562). We are grateful to R. D. Orton for providing unpublished age-size
class data for Girella nigricans. We thank D. G. Buth, T. E. Shelly, and an
anonymous reviewer for their critical evaluation of the manuscript.
LITERATURE CITED
Barton, M. C. 1982. Intertidal vertical distribution and diets of five species of central California stichaeoid fishes.
Calif. Fish Came, 68 (3): 174-182.
Burge, R. T., and S. A. Schultz. 1973. The marine environment in the vicinity of Diablo Cove with special reference
to abalones and bony fishes. Calif. Dept. of Fish and Game, Marine Resources Technical Report No. 19.
Fitch, J. E., and R. J. Lavenberg. 1975. Tidepool and Nearshore Fishes of California. Univ. of Calif. Press, Berkeley.
156 p.
Grossman, G. D. 1982. Dynamics and organization of a rocky intertidal fish assemblage: the persistence and
resilience of taxocene structure. Amer. Nat., 119 (5): 611-637.
Hart, |. L. 1973. Pacific Fishes of Canada. Fish. Res. Board Canada, Bull. 180. 740 p.
Moring, |. R. 1981. Seasonal changes in a population of the fluffy sculpin, Oligocottus snyderi, from Trinidad Bay,
California. Calif. Fish Game, 67 (4): 250-253.
Pielou, E. C. 1977. Mathematical Ecology. )ohn Wiley and Sons, N.Y. 385 p.
Ricker, W. E. 1975. Computation and interpretation of biological statistics of fish populations. Fish. Res. Board
Canada, Bull. 191. 366 p.
Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker, E. A. Lachner, R. N. Lea, and W. B. Scott. 1980. A list of
common and scientific names of fishes from the United States and Canada, 4th Ed. American Fisheries Society,
Spec. Publ. No. 12. 174 p.
Sokal, R. R., and F. ). Rohlf. 1981. Biometry, 2nd Ed. W. H. Freeman and Co., N.Y. 859 p.
Stauffer, J. R., R. L. Reish, and W. F. Calhoun. 1980. FORTRAN program for calculating Brillouin's species diversity
index. Progressive Fish Culturist, 42: 185-187.
Thomson, D. A., and C. E. Lehner. 1976. Resilience of a rocky intertidal fish community in a physically unstable
environment. |. Exp. Mar. Biol. Ecol., 22: 1-29.
Yoshiyama, R. M. 1980. Food habits of three species of rocky intertidal sculpins (Cottidae) in central California.
Copeia, 1980 (3): 515-525.
1981. Distribution and abundance patterns of rocky intertidal fishes in central California. Envir. Biol. Fish.,
6 (3/4): 315-332.
Williams, G.C. 1957. Homing behavior of California rocky shore fishes. Univ. of Calif. Publ. Zool., 59 (7): 249-284.
Zar, ). H. 1974. Biostatistical Analysis. Prentice-Hall, Inc., N.J. 620 p.
232 CALIFORNIA FISH AND CAME
Calif. Fish and Game 72(4) 232-243 1986
DISTRIBUTION OF MAJOR MARINE MACROPHYTES, SEA-
SONAL ESTIMATES OF GRACILARIA STANDING CROP,
AND SPAWNING ACTIVITIES OF THE PACIFIC HERRING,
CLUPEA HARENGUS PALLASII, IN ELKHORN SLOUGH,
CALIFORNIA; 1979-1982 *
R. E. PHILLIPS 2
Hopkins Marine Station
Pacific Grove, California 93950
D. I. CUTOFF
Marine Bioassay Laboratories
1234 Highway One
Watsonville, California 95076
|. E. HANSEN J
Hopkins Marine Station
Pacific Grove, California 93950
AND
J. E. HARDWICK
California Department of Fish and Game
2201 Garden Road
Monterey, California 93940
Marine vegetation surveys and Pacific herring spawn assessments were carried out
in Moss Landing Harbor and Elkhorn Slough, California for three consecutive herring
spawning seasons (1979-80 to 1981-82). The red alga Cracilaria sp. and the marine
vascular plant Zostera marina (eelgrass) are the major macrophyte substrata avail-
able for herring spawn deposition. Cracilaria standing crops varied considerably
among years and locations within the study area. The standing crop of Cracilaria in
Elkhorn Slough was physically reduced each winter by storm runoff and strong tidal
currents. Biomass estimates of spawning herring were calculated from each season's
egg deposits. During the 1979-80 season, an estimated 0.4 short ton of herring
spawned on Cracilaria. Biomass estimates of spawning herring for the 1980-81 and
1981-82 seasons were based on all plant substrata and were 0.1 and 0.6 short tons
respectively. The density of herring spawn deposition observed in Moss Landing
Harbor and Elkhorn Slough was consistently very light ( <0.25 egg layers).
INTRODUCTION
The Pacific herring, Clupea harengus pallasii Valenciennes, has supported a
varying portion of the California fishing industry since at least 1916. The history
of the herring fishery in California was recently summarized (Spratt 1981).
Within the past two decades commercial activities in the fishery have been
stimulated by the development of markets for herring roe in Japan.
In 1965 a fishery began in which salt-cured herring eggs-on-seaweed are
shipped to Japan where they are a delicacy. In California a comparatively small
Asian market exists and the product is served locally in restaurants. Today the
California herring eggs-on-seaweed fishery is restricted to San Francisco Bay.
From 1965 to 1977 Tomales Bay was also open to the fishery. Harvest quotas
1 Accepted for publication March 198b.
-' Current address: Monterey Bay Aquarium, 886 Cannery Row, Monterey, California 93940.
' Current address: Long Marine Laboratory, University of California, Santa Cruz, California 95064.
MARINE MACROPHYTES AND SPAWNING OF PACIFIC HERRING 233
were set at 5 short tons (wet weight of seaweed and eggs) for each bay, and
this quota is still in effect in San Francisco Bay. Harvest quotas have never been
reached in either bay. The largest recorded harvest occurred during the 1967-68
season when 6.7 short tons was harvested from both bays combined.
Much larger harvests of herring eggs-on-seaweed occur annually in Alaska
and British Columbia (Fuoco 1980, R. Hunt, pers. comm.). The most desirable
seaweed substrata from the Pacific Northwest are Macrocystis integrifolia Bory
and Laminaria saccharina (Linnaeus) Lamouroux. An even and heavy herring
spawn (approximately 80% eggs by weight) on these seaweeds may bring up
to $17.00/ lb wholesale. Other desirable seaweed substrata belonging to the
genera Laminaria, Agarum, Fucus, Ulva, and Gracilaria wholesale for $2.00 to
$8.00/lb (R. Hunt, pers. comm.). In California the major seaweeds harvested
are Laminaria sinclairii (Harvey) Farlow, Anderson & Eaton and Gracilaria sp.
(Hardwick 1973, Spratt 1981).
in 1972 the herring fishery in the Pacific Northwest was further enhanced
when Japan removed its import quota on herring. Since 1972 the herring fishery
in California has experienced a resurgence. Annual landings for the 1981-82 and
1982-83 seasons (11,615 and 10,611 short tons respectively) were more than
twice those recorded for any year prior to 1972 (Spratt 1983).
The recent enhancement of the Pacific herring fishery has prompted investiga-
tions of herring spawning activities and spawning biomass in bays and estuaries
(Hardwick 1973; Rabin and Barnhart 1977; Spratt 1976, 1981 ). In California the
present herring fishery centers around San Francisco, Tomales, and more recent-
ly, Bodega bays. Humboldt Bay and Crescent City support fisheries of 12.5 to
50.0 short tons per year. Recent surveys have estimated the spawning biomass
of herring in these areas (Rabin and Barnhart 1977, Spratt 1981).
Monterey Bay, in central California, supports a small spring and summer
herring fishery for bait and animal food from 40.0 to 340.0 short tons annually
since 1 965 ( Spratt 1 981 ) . Pacific herring have been reported to spawn in Elkhorn
Slough, Moss Landing Harbor, and the mouth of the Salinas River in Monterey
Bay (Miller and Schmidtke 1956, Nybakken et al. 1977, Spratt 1976, K. Forsyth,
pers. comm.). The extent of the herring spawn, an estimate of the spawning
biomass, or year to year variability in the Monterey Bay area are not presently
known.
This study was part of a collaborative effort between the California Depart-
ment of Fish and Game and a California Sea Grant project (R/F-58 'Multiple
Species Utilization of the Herring Eggs-On-Seaweed Fishery', Abbott and Hans-
en 1 981 ) to evaluate the potential for integration of two marine fisheries: ( i ) the
existing winter herring eggs-on-seaweed fishery, and (ii) Gracilaria mariculture
for an agarweed resource. Reported here are distributions of the major marine
macrophytes, seasonal estimates of Garcilaria standing crop, and spawning ac-
tivities of the Pacific herring in Moss Landing Harbor and Elkhorn Slough for
three consecutive spawning seasons (winters 1979-80 to 1981-82).
MATERIALS AND METHODS
Vegetation Surveys and Mapping
At the onset of the 1979-80 herring spawning season, intertidal and subtidal
vegetation surveys were conducted throughout Moss Landing Harbor (Lat
36°48.T N, Long 121°47.9' W) and Elkhorn Slough (Figure 1 ). The distribution
234
CALIFORNIA FISH AND CAME
of each major vegetation type was determined by observations during low tides
for intertidal populations (The term 'population' is used here to describe macro-
phyte beds which are spatially distinct; it does not imply any genetic distinc-
tions.) and by boat using a Miller Aquatic Vegetation Sampler or SCUBA for
subtidal populations. A Miller Vegetation Sampler consists of two heavy duty
garden rakes (3 tines/10 cm) welded back-to-back and fastened to a line (Miller
and Schmidtke 1956, Hardwick 1973).
1 Harbor
2A Yacht Club Channel
2B Yacht Club
3 Zostera Bed
4 Experimental Population
5 Dairy
6 Mussel Bed
7 Duck Club
FIGURE 1. Location of sampling stations in Moss Landing Harbor and Elkhorn Slough.
Seasonal changes in standing crop within one representative intertidal Craci-
laria population (approximately 325 m2) were measured at Station 4 (Figure 1 ).
The population was mapped and divided into a 0.5 m2 grid pattern using lines
MARINE MACROPHYTES AND SPAWNING OF PACIFIC HERRING 235
and permanent anchors. Fifteen new quadrats were chosen randomly once
every three months and algal biomass was harvested using sheepshears. Samples
were washed in fresh water, cleaned of debris, drained, weighed fresh, and then
dried to constant weight at 60°C
Herring Spawn Assessments and Estimates of Spawning Biomass
Herring spawn assessments were carried out at eight major populations of
intertidal and subtidal vegetation (Figure 1 ). Vegetation was sampled from these
stations every 3-7 days during the 1979-80 and 1980-81 spawning seasons (18
Dec. 1979 - 28 Feb. 1980, 15 Dec. 1980-5 March 1981 ) and every other day
during the 1981-82 season (15 Dec. 1981-29 Feb. 1982). Observations of the
mud bottom and other substrata (pilings, shell beds, etc.) indicated that herring
in the study area spawn primarily on vegetation, rarely on other substrata.
During the 1979-80 and 1980-81 spawning seasons, intertidal vegetation was
assessed for the presence of herring spawn at all stations during low tide periods.
When herring spawn was found, samples of the vegetation and spawn were
collected and returned to the laboratory for processing. Intertidal spawns were
sampled by harvesting the vegetation within several 0.5 m2 quadrats. The total
area of an intertidal spawn was determined from distance measurements around
the perimeter of the spawning location. Subtidal spawns were sampled using the
Miller Vegetation Sampler. The vegetation sampler was towed along the bottom
behind a boat for a distance of approximately 20 m and then retrieved with
adherent vegetation. Alternatively, the sampler was tossed from the boat, al-
lowed to sink to the bottom, and then retrieved. The distribution of subtidal
spawns was determined by successive bottom tows around the perimeter of the
plant population, and the total area of spawn was calculated from distance
measurements between sighted landmarks and/or visual estimates of distance.
During the 1981-82 spawning season, herring spawn found on intertidal vege-
tation was assessed by subsampling the entire station area at low tide. Twenty
or more 0.25 m2 quadrats were chosen randomly and all vegetation within the
quadrats was harvested. Subtidal spawns were assessed by randomly sampling
a similar number of 0.25 m2 quadrats using SCUBA. Distance measurements
were taken to determine the total area of all spawns.
Each eggs-on-seaweed sample was sorted into major vegetation type and wet
weights were determined. Representative subsamples were taken from each
vegetation type, wet weighed, and the herring eggs adhering to the subsample
were counted. The number of herring eggs/g wet wt of sample and the number
of eggs/m2 of vegetated area were calculated.
Estimates of spawning herring biomass were made based on the number of
eggs spawned. The total number of eggs spawned for each season was convert-
ed to short tons of herring by multiplying by 0.966 X 10" 8. This factor is derived
from studies on Pacific herring fecundity. The calculation assumes a 1 :1 sex ratio
in the spawning herring population (Hardwick 1973; Rabin and Barnhart 1976;
Spratt 1976, 1981).
Data collected during the 1979-80 season did not include quantitative esti-
mates of Cracilaria or Zostera marina standing crops at all spawning locations.
The standing crop of Cracilaria at each spawning location was estimated by
visual inspection and comparison to the known standing crop at Station 4 (see
Vegetation Surveys). The estimates for Gracilaria standing crops and the number
of eggs/g wet wt of seaweed were used to calculate the number of eggs spawned
236 CALIFORNIA FISH AND GAME
on Cracilaria. No estimates were made of Zostera marina standing crops and,
therefore, the total number of eggs spawned on Zostera was not calculated.
During the 1980-81 and 1981-82 seasons, plant standing crops were quantita-
tively determined at each station when a spawn was found. The number of
eggs/g wet wt of vegetation was counted for each spawn. Direct calculations
of number of eggs/m2 could then be made.
RESULTS
Floristic Studies
In Moss Landing Harbor and Elkhorn Slough the major low intertidal and
subtidal flora is composed of the red alga Gracilaria sp. and the marine vascular
plant Zostera marina (eelgrass). The forms of Gracilaria that occur in Moss
Landing Harbor and Elkhorn Slough are combined in this study. The taxonomy
of local Gracilaria species is presently under study (I. A. Abbott and M. Hoyle,
in prep.). Other algal genera that may contribute ephemeral or minor substrata
for spawning herring are Enteromorpha and Polysiphonia. Enteromorpha spp.
are especially prevalent on the mud flats of Elkhorn Slough during spring and
summer. Enteromorpha was sparse during the 1979-80 and 1981-82 spawning
seasons but was fairly common during the 1980-81 season. Relatively small
populations of the filamentous red alga Polysiphonia sp. (predominately P.
mollis Hooker & Harvey) were also observed among populations of Gracilaria
sp. and Zostera marina.
Populations of Z marina were found in low intertidal and shallow subtidal
areas of Moss Landing Harbor and the mouth of Elkhorn Slough (Figure 2). On
an area basis, Gracilaria was the dominant macrophyte found in the study area.
Substantial populations occurred in the intertidal and shallow subtidal areas of
the Harbor (Station 1 ) and the Yacht Club (Stations 2A & 2B). Gracilaria was
also intermixed with the Zostera population at the mouth of Elkhorn Slough
(Station 3; Figures 1 and 3). Discrete populations occurred in the upper reaches
of Elkhorn Slough along the southern bank ( Stations 4,5,6, and 7 ) . These popula-
tions were associated with areas that: (i) were semi-protected from strong tidal
currents (Station 7), or (ii) consisted of mollusc shell deposits and/or mussel
beds (Mytilus edulis Linnaeus) (Stations 4,5, and 6; Figures 1 and 3).
The standing crop of Gracilaria at most stations varied substantially over the
study period. Quarterly biomass estimates for the experimental population at
Station 4 (Figure 1 ) give an indication of the three year seasonal variability for
most Elkhorn Slough Gracilaria populations. The results (Figure 4) show a dra-
matic decrease in the standing crop of Gracilaria during the 1979-80 herring
spawning season. Storm runoff, high tides, and rapid currents (23-25 Dec. 1979
and 15-17 Feb. 1980) were probably responsible for the decline in standing
crop. This population had recovered to about 10% of the 1979 standing crop
by the beginning of the 1980-81 herring spawning season (Dec. 1980), but again
showed a sharp decline by March 1981 (Figure 4). The seasonal trend of
regrowth followed by physical removal is also apparent in the 1981 data; howev-
er the decline occurred prior to our December 1981 sample. By the beginning
of the 1981-82 spawning season, Gracilaria biomass was approximately 4% of
the standing crop measured in December 1979. The other Gracilaria populations
in Elkhorn Slough (Stations 5 and 6) displayed similar fluctuations in standing
crop, except for Station 7 which is semi-protected from strong tidal currents.
Populations in Moss Landing Harbor (Stations 1,2A,2B and 3) were less severely
affected by winter storm runoff and tidal currents.
MARINE MACROPHYTES AND SPAWNING OF PACIFIC HERRING
237
Zostera morinq
fc!
1
FIGURE 2. Distribution of Zostera marina in Moss Landing Harbor and Elkhorn Slough.
Gracilaria collected from Elkhorn Slough was predominately non-reproduc-
tive; rarely were tetrasporangial plants found. However, cystocarpic plants were
common in the adjacent Gracilaria populations of Moss Landing Harbor.
Herring Spawn Assessments and Estimates of Spawning Biomass
The areas where herring spawns were found during the three-year study
period are illustrated in Figure 5. Three discrete herring spawning runs were
observed during the 1979-80 season (Table 1 ). On the basis of vegetated area,
the spawns were primarily on Gracilaria. Herring spawns were found only in the
Harbor, the Yacht Club, and the mouth of Elkhorn Slough (Stations 1,2A,2B and
3; Figure 1). Egg deposits were very light and the total spawning area was
estimated at 3,730 m2. Based on our estimates of Gracilaria standing crops,
approximately 41 million eggs were spawned on Gracilaria by 0.4 short ton of
herring.
We were informed that a fourth herring spawn occurred during the 1979-80
season at the head of Elkhorn Slough near Hudson's Landing. We attempted to
document this report and found no evidence of herring spawning activity.
However, herring eggs hatch in 6-11 days (Hardwick 1973), and we may have
been too late.
238
CALIFORNIA FISH AND CAME
FIGURE 3. Distribution of Gracilaria in Moss Landing Harbor and Elkhorn Slough. The maximum
area occupied by Gracilaria is illustrated.
During the period from 15 December 1980 to 5 March 1981, three herring
spawns were recorded in Elkhorn Slough (Table 2). On 15 Dec. 1980 a small
herring spawn was found on Enteromorpha at Kirby Park (Station 8). All other
herring spawns occurred on the Gracilaria population at Station 7 ( Figure 1 ) . The
deposition of herring eggs was generally very light and the total spawning area
was estimated to be only 480 m2. Estimates from standing crop and egg count
data for the 1980-81 season include 6 million eggs spawned on Enteromorpha
and 4 million eggs spawned on Gracilaria (Table 2). The meager total for the
1980-81 season was 10 million eggs spawned by 0.1 short ton of herring.
MARINE MACROPHYTES AND SPAWNING OF PACIFIC HERRING
239
Seven herring spawns were recorded in Moss Landing Harbor and Elkhorn
Slough during the 1981-82 spawning season (Table 3). Herring spawned on an
estimated 80,1 18 m2 of vegetated area, including nearly all of the Cracilaria and
Zostera populations. The deposition of herring eggs was very light for all spawns.
Based on standing crop and egg count data, total estimates for the 1981-82
season include 58 million eggs spawned on vegetation by 0.6 short ton of herring.
600
500-
E
m
400
o
^
300
•
*
<0
E
o
200
I00-
A 3,168
±354
X±S.E.
r^H
DEC. MARCH JUNE SEPT. DEC. MARCH JUNE SEPT.
1979 1980 1981
DEC
FIGURE 4. Seasonal variation in the standing crop of Cracilaria at Station 4.
DISCUSSION
We consider our estimates of spawning herring biomass to be conservative.
Predation on herring eggs by birds and fishes can be quite extensive within the
first several days after a spawn (Hardwick 1973; Spratt 1976, 1981 ). Bird preda-
tion (predominantly gulls) was noted on several intertidal spawns in Elkhorn
Slough. We have not included egg loss via predation in our calculations of total
eggs spawned per season. It is also possible that our estimates of spawning
herring biomass are low due to: (i) herring spawns which occurred before or
after our survey periods, and (ii) small spawns which were not detected.
The results of our surveys show that a very small biomass of herring ( < 0.6
short ton) was responsible for the egg deposits observed in Moss Landing
Harbor and Elkhorn Slough during the 1979-80, 1980-81, and 1981-82 spawning
seasons. Compared to the results of other spawning surveys performed in Cali-
fornia (Spratt 1981 ) and British Columbia (Humphreys and Hourston 1978), the
egg densities found in our study area were consistently very light ( < 0.25 egg
layers). While the desirable herring eggs-on-seaweed species Cracilaria pro-
vided abundant substratum for herring spawn deposition, the very light egg
densities were below commercial standards.
240
CALIFORNIA FISH AND GAME
FIGURE 5. Location of herring spawn deposition in Moss Landing Harbor and Elkhorn Slough
during the 1979-80, 1980-81, and 1981-82 spawning seasons.
Based on our estimates of spawning herring biomass it appears that a large
portion of the herring caught in Monterey Bay during spring and summer must
come from spawning grounds other than Moss Landing Harbor and Elkhorn
Slough. However, the consistent year-to-year presence of spawning and larval
herring populations in Moss Landing Harbor and Elkhorn Slough is indicative of
the historical importance of this area as a herring nursery grounds.
MARINE MACROPHYTES AND SPAWNING OF PACIFIC HERRING
241
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MARINE MACROPHYTES AND SPAWNING OF PACIFIC HERRING 243
ACKNOWLEDGMENTS
This study was supported by the California State Department of Fish and
Game in cooperation with the California Sea Grant Program (R/F-58; I. A.
Abbott, Hopkins Marine Station, Stanford University). Special thanks are ex-
tended to B. Dyer, J. Slumpberger, F. Henry, and S. Danick for their extensive
support both in the field and laboratory. We are indebted to C. Vierra, L.
Calcagno, and L. Simpson for providing access to study sites and boat mooring
facilities. We thank J. Spratt for advice regarding the herring assessment studies.
LITERATURE CITED
Abbott, I. A., and |. E. Hansen. 1981. Multiple species utilization of the herring eggs-on-seaweed fishery. Calif. Sea
Grant College Program (R/F-58). 1980-81 project summary.
Fuoco, S. W. 1980. Herring roe's future may lie in kelp-filled Canadian ponds. National Fisherman, April 30:
160-162.
Hardwick, ). E. 1973. Biomass estimates of spawning herring, Clupea harengus pallasi, herring eggs, and associated
vegetation in Tomales Bay. Calif. Fish Game, 59(1): 36-61.
Humphreys, R. D., and A. S. Houston. 1978. British Columbia Herring Spawn Deposition Survey Manual. Fish, and
Mar. Serv. Spec. Publ., 38: 1-40.
Miller, D. J., and |. Schmidtke. 1956. Report on the distribution and abundance of Pacific Herring {Clupea
pallasi) along the coast of Central and Southern California. Calif. Fish Game, 42(3): 163-187.
Nybakken, )., G. Cailliet, and W. Broenkow. 1977. Ecological and Hydrographic studies of Elkhorn Slough, Moss
Landing Harbor and nearshore coastal waters )uly 1974 to June 1976. Moss Landing Marine Laboratories, Moss
Landing, Calif. 465 p.
Rabin, D. )., and R. A. Barnhart. 1977. Fecundity of Pacific Herring, Clupea harengus pallasi, in Humboldt Bay.
Calif. Fish Game, 63(3): 193-196.
Spratt, j. D. 1976. The Pacific Herring resource of Tomales and San Francisco Bays: its size and structure. Calif.
Dept. Fish and Game. Mar. Res. Tech. Rep., 33: 1-44.
1981. Status of the Pacific Herring, Clupea harengus pallasii. Resource in California 1972 to 1980. Calif.
Dept. Fish and Game, Fish Bull. 171: 1-107.
1983. Biomass estimate of Pacific Herring, Clupea harengus pallasi, in California from the 1982-83
spawning ground surveys. Calif. Dept. Fish and Game, Mar. Res. Admin. Rep. 83-3: 1-23.
244 CALIFORNIA FISH AND CAME
Calif. Fish and Game 72(4): 244 249 1986
OBSERVATIONS ON THE ELASMOBRANCH ASSEMBLAGE
OF SAN FRANCISCO BAY 1
DAVID A. EBERT
Moss Landing Marine Laboratories
P.O. Box 450
Moss Landing, California 95039
Elasmobranchs were collected over a 19 month period using several types of
fishing gear. The information gathered suggests a possible shift in the elasmobranch
species composition in San Francisco Bay when compared with previous data. A
difference in the sex ratios of some species was also noted. The population of at least
one species appears to have declined.
INTRODUCTION
An increasing interest in elasmobranch fishes as a food source has raised
concern about the stability of this fishery, primarily because most elasmobranchs
are slow growing and have a low reproductive rate. A problem with managing
this growing fishery is the lack of information concerning their population struc-
ture. The shark and ray population of San Francisco Bay is one of the few
elasmobranch fisheries where historical data, albeit limited, is available concern-
ing species abundance.
Herald and Ripley (1951 ) reported on the abundance of sharks and rays in
San Francisco Bay based on studies by the California Department of Fish and
Came and catch records from annual shark derbies. They presented their data
as a preliminary working basis for future population studies in the bay. Later, de
Wit (1975) reported possible changes in the shark species composition of south
San Francisco Bay. With an increasing interest in elasmobranchs as a food source
more current information is required to better assess their populational status.
The objective of this research was to investigate the elasmobranch assemblage
in San Francisco Bay and to compare these data with historical catch records.
METHODS AND MATERIALS
During the course of a study on the biology of the sevengill shark, Notoryn-
chus maculatus, ( Ebert in press) I gathered data on the elasmobranch fishes that
were caught in San Francisco Bay. Sampling was conducted between the Rich-
mond-San Rafael Bridge and the San Mateo Bridge (Figure 1).
Shark samples were collected by several methods, including gill net (3.1 m
x 275 m x 20 cm), long-line (6 to 335, 12/0 hooks per set), and rod-and-reel
(one 12/0 hook-per-pole). Shark fishing was conducted year-round using long-
line and rod-and-reel. Gill nets were used only during the winter months.
Data were recorded monthly for all species and used to provide an estimate
of the species composition. Catch-per-unit-effort (CPUE), using catch per hook-
hour, was calculated for long-line and rod-and-reel. Gill net CPUE was calculat-
ed by catch-per-hour.
Accepted for publication May 1986
SAN FRANCISCO BAY ELASMOBRANCHS
245
i i i i
i
5 mi
3 10 km
Rich
Sa
"**«**,
9e
R = ROD and REEL
L ■ LONG-LINE
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NUMBERS INDICATE NUMBER
RICHMOND OF TIMES FISHED
R5L1
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R4
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SAN FRANCISCO
R2 L1
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R3L3
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R14L1
Oyster Point
R1G1 L1
South San Francisco Bay
Coyote Point
^P^
\e°
&
\o9e
FIGURE 1. Map of San Francisco Bay showing gear type and areas fished.
RESULTS
Using all gear combined, a total of 441 sharks and rays, representing seven
species, was caught in San Francisco Bay from November 1981 through May
1983. These species included: the brown smoothhound shark, Mustelus henlei;
leopard shark, Triakis semifasciata; soupfin shark, Galeorhinus zyopterus; spiny
246
CALIFORNIA FISH AND CAME
dogfish, Squalus acanthias; sevengill shark, Notorynchus maculatus; bat ray,
Myliobatis californica; and big skate, Raja binoculata.
The type of fishing gear employed was dependent on commercial fishing
practices; and rod-and-reel was the most prevalent and effective method (Table
1 ). Long-line gear was too time consuming and expensive (bait) for the yield.
One fishermen who long-lined exclusively was financially forced to suspend
operations. Winter storms limited my use of gill netting to only six sets between
December 1982 and March 1983.
The most productive fishing months for all elasmobranch species using rod-
and-reel in San Francisco Bay were May through October 1982 and May 1983
(Table 2). There was no fishing effort reported for February through April 1982,
and February and April 1983 (Table 2). Fishing during these months was pre-
cluded by inclement weather.
TABLE 1. Elasmobranch Catch-and-Effort Data, According to Gear Type, From San Fran-
cisco Bay.
No. No. Hours
Gear type hooks sets fished Catch
Set line
(range 6-335 hooks per set) 1736 10 63 44
Gill net 6 32 8
Rod-and-reel
(one 12/0 hook per pole) 158 57 125 389
Leopard shark were the most abundant elasmobranch caught in San Francisco
Bay and represented 40% of the rod-and-reel catch (Table 3). Brown smooth-
hound and spiny dogfish also ranked high representing 23% and 22% of the
elasmobranch catch, respectively. The soupfin shark was the least abundant
species caught, representing 1%.
The rod-and-reel catch categorized according to sex revealed that the seven-
gill shark was the only elasmobranch caught in which males outnumbered
females. The sevengill shark was caught at a 1.1:1 (male:female) sex ratio.
Females dominated in the remaining elasmobranch catch, with ratios ranging
from 1:2 for leopard shark to 1:4.5 for bat ray.
DISCUSSION
Results of this study indicate that the dominance of leopard shark in San
Francisco Bay suggests that a shift may have occurred in the elasmobranch
assemblage, at least between two major species. Herald (1953) and de Wit
(1975) both reported the brown smoothhound as being the most abundant
elasmobranch in the bay, comprising over 41% of the recorded catch. I found
the brown smoothhound to represent only 23% of the elasmobranchs caught,
while the leopard shark accounted for 40%. De Wit (1975) reported the brown
smoothhound as being most numerous in the catch from June through Septem-
ber 1972, while I found them to be dominant in the catch only during August
and October 1982 (Table 2). My data represents only a rough estimate of the
elasmobranch assemblage in San Francisco Bay and any observed differences
may reflect a bias in the various sampling methods employed. However, the
common denominator in Herald's (1953) and de Wit's (1975), and my own
sampling methods was that the fishing gear (gill nets, long-line, and rod-and-
reel) employed was specifically targeted toward elasmobranchs. Furthermore,
although de Wit (1975) conducted his study south of the San Mateo Bridge, his
SAN FRANCISCO BAY ELASMOBRANCHS
247
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248 CALIFORNIA FISH AND GAME
TABLE 3. Number, Percent, and Sex for Seven Elasmobranch Fishes Caught in San Francisco
Bay on Rod-and-Reel, November 1981 through May 1983.
Male Female Total
Species No. % No. % No. %
leopard shark 53
brown smoothhound 28
spiny dogfish 17
sevengill shark 17
bat ray 6
big skate 4
SOupfin shark 0
results compared favorably to those of Herald (1953) whose sample area was
similar to mine (Figure 1 ).
The sex ratios I found for some species were different from that reported by
Russo and Herald (1968), who based their estimate on a shark mortality that
occurred along the east bay. I found the male-female sex ratio for brown
smoothhound to be 1:2.2. Russo and Herald (1968) reported that the brown
smoothhound had a sex ratio of 1:4.5 (male : female). Susan Smith (National
Marine Fishery Service, Tiburon, Calif., pers. comm.) found a 4:1 male-female
ratio for brown smoothhound from a long-line study conducted near Hunter's
Point in September 1979. Herald and Ripley (1951 ) noted a similar reversal in
the sex ratio of sevengill sharks between 1943 and 1950, but could offer no
explanation. I also found a difference in the sex ratio of leopard shark (1:2,
male:female) and bat rays (1:4.5) compared to what Russo and Herald (1968)
reported which was 1:1 for both species. I recorded a 1 :1 .6 male-female sex ratio
for leopard shark at a San Francisco Bay shark derby held in September 1983.
Therefore, these differences between Russo and Herald (1968), Susan Smith
(pers. comm.), and my own data may not be significant due to differences in
sampling methods.
There may be a general population decline of several elasmobranch species
occurring in San Francisco Bay. Particularly evident was the low abundance of
soupfin shark when compared to that reported by Herald (1953), Russo and
Herald (1968), and Smith and Kato (1979). The reasons for this apparent
change are unclear, but may be related to: (i) a combination of lower salinities
in San Francisco Bay from large amounts of rainfall ( Reilly and Moore 1 982 ) and
unusually warmer open coastal waters (Smith 1983) that caused sharks to seek
a cooler, more suitable habitat, or (ii) over-exploitation. Increased fishing pres-
sure was evident in San Francisco Bay during the mid- to late 1970's (L.J.V.
Compagno, Tiburon Center for Environmental Studies, pers. comm.) and al-
though the California Department of Fish and Game obtains commercial shark
catch data, the principal take of elasmobranchs is by recreational fishermen.
Catch data are not available for this type of fishing. Due to time constraints I was
unable to measure size frequency, but it appeared that most sharks caught were
at the minimum size for maturity. Further evidence for this came from the 1983
shark derby where I recorded total length (tl) measurements for leopard shark.
I found the mean tl for male (98.9 cm tl) and female (106.9 cm tl) leopard
shark to be under the approximate size of maturity (males — 107 cm tl and
females — 118 cm tl). This derby had a minimum check-in size of 70 cm tl. The
soupfin shark fishery in California collapsed from overfishing (Ripley 1946).
SAN FRANCISCO BAY ELASMOBRANCHS 249
Holden (1974) cites other instances where over-exploited elasmobranch fisher-
ies have collapsed. If the reduction in shark numbers in San Francisco Bay is due
to reduced salinity or warm water then the decline may be temporary. However,
if the decline is due to over-exploitation this may have long term effects and a
management program should be implemented to sustain this fishery.
ACKNOWLEDGMENTS
I would like to thank the following persons for their time and consideration
throughout this research. D. Kittredge and B. Van Gorp for their fishing efforts.
G. Cailliet, E. Ebert, R. Lea, M. Foster, J. McCosker, and S. Smith for their many
helpful suggestions and comments in reviewing this manuscript. L. Compagno
provided many helpful ideas and suggestions. M. Kittridge for illustrating the
figure. General assistance in various portions of this study was given generously
by T. Ebert, K. Hauge, and S. Willis.
LITERATURE CITED
de Wit, L A. 1975. Change in the species composition of sharks in south San Francisco Bay. Calif. Fish Came,
61(21:106-111.
Ebert, D. A. In press. Life history aspects of the sevengill shark, Notorynchus maculatus Ayres 1855, in two northern
California bays. Calif. Fish Came.
Herald, E. S., and W. E. Ripley. 1951. The relative abundance of sharks and bat stingrays in San Francisco Bay.
Calif. Fish Came, 37(3): 315-329.
Herald, E. S. 1953. The 1952 shark derbies at Elkhorn Slough, Monterey Bay, and at Coyote Point, San Francisco
Bay, Calif. Fish Game, 39(2): 237-243.
Holden, M. ). 1974. Problems in the rational exploitation of elasmobranch populations and some suggested
solutions. Pages 117-137 in: F. R. Harden-Jones eds., Sea fisheries research, John Wiley and Sons, NY.
Reilly, P. N„ and T. O. Moore. 1982. Pacific herring, Clupea harengus pallasii studies in San Francisco Bay,
December 1981 to March 1982. Marine Resources Administrative Report no. 82-8: 43.
Ripley, W. E. 1946. The soupfin shark and the fishery. Calif. Div. Fish and Came, Fish Bull. 64: 7-37.
Russo, R. A., and E. S. Herald. 1968. The 1967 shark kill in San Francisco Bay. Calif. Fish Game, 54(3): 215-216.
Smith, R. L. 1983. Peru coastal currents during El Nino: 1976 and 1982. Science, 221 (4618): 1397-1398.
Smith, S. E., and S. Kato. 1979. The fisheries of San Francisco Bay: past, present, and future. California Academy
of Sciences, Pacific Division, AAAS. 445^168.
250 CALIFORNIA FISH AND CAME
BOOK REVIEWS
ARIZONA WETLANDS AND WATERFOWL
By David E. Brown. Illustrations by Bonnie Swarbrick Morehouse. University of Arizona Press,
1985. 169 p. $24.95 cloth.
It is difficult to characterize this book. Too general to be considered a reference
work, it has too few color illustrations (only seven species covered) to make
a good "coffee table" book. It is directed at the general public, amateur natural-
ists and waterfowl hunters who are most interested in waterfowl and wetland
matters as they pertain to Arizona conditions.
The book is divided into five chapters with an additional section of reference
materials (appendices and a bibliography). The introduction includes material
on early history, uses and users and the need for wetland and waterfowl conser-
vation. A chapter entitled "Environmental Setting" covers in detail the wetlands
and waterfowl resources of the main physiographic regions or Arizona, both
historically and as they exist today. The chapter on Waterfowl Biology discusses
taxonomy, anatomy (including molts and plumages), life history, migration,
foods, sexual dimorphism and sex ratios, and diseases (including lead poison-
ing). Much data on Arizona situations are presented in this chapter.
Chapter four focuses on waterfowl management, beginning with an historic
overview of hunting and early conservation laws. This is followed by discussions
of the refuge system (both state and federal), banding and the flyway concept,
surveys and bag checks, regulating the harvest, species identification by the
hunter and habitat enhancement. The remainder of the book almost a third of
it, is devoted to species accounts, which are thorough and informative. They
cover distribution and status in Arizona, descriptions, habitat preferences and
miscellaneous information "of interest to the waterfowler and habitat manager".
The author's style is relaxed and easy to read, there is an absolute minimum
of typographical and technical error and a large variety of subject matter cov-
ered, though not in great depth. The book will probably prove attractive and
useful tc Arizona residents and those non-residents with a special interest in
Arizona wetlands and waterfowl management. Others may well wish to send
the $25 on one of the recent works by Paul Johnsgard or Frank Bel I rose.— Bruce
E. Deuel
HUMMINGBIRDS: THEIR LIFE AND BEHAVIOR
By Esther Q. Tyrrell, with photographs by Robert A. Tyrrell; Crown Publishers, Inc., New York
City; 1985; 224 p; $35.00.
"Nature has favored the Americas with a sparkling bird whose jewellike colors
and fascinating aerial acrobatics make it unique." So begins the first chapter, "An
Introduction to Hummingbirds", in this oversized book whose subtitle is "A
Photographic Study of the North American Species." At first glance one might
believe that this is another coffee table book. However, a closer look finds much
scientific detail to accompany the 235 full-color photographs. The photographs
themselves may be worth a good deal of the price of the book. Many stop-action
pictures are found throughout the third through ninth chapters, which cover the
topics of anatomy, feathers, flight, courtship and nesting, food and metabolism,
behavior, and wildflower pollination.
In the second chapter, "A Portfolio of North American Hummingbirds", the
16 hummingbirds which breed in the United States are each described on a
two-page spread with two color photographs and a discussion of field marks,
BOOK REVIEWS 251
range, nesting period, migration times, and habitat. This book definitely is orient-
ed toward U.S. breeding species. The authors do list the additional seven hum-
mers which have been recorded in the U.S. north of Mexico, and provide a list
of the 338 species (in 116 genera) of hummingbirds in the world.
The most fascinating part of the book to this reviewer is the chapter on
wildflower pollination. Illustrated with many photographs of hummingbirds at
flowers and showing pollen on the bills, crowns, and chins of the birds, this
chapter discusses that group of plants known as hummingbird flowers, since they
are pollinated only by these birds. At least 129 of these species of plants are
known from the American southwest, and 20 others are in the northeast. There
is even a discussion of the hummingbird flower mites, which live, mate, and
reproduce in the flowers but are transported between flowers in the nostrils of
hummingbirds. A list of hummingbird-pollinated wildflowers by family is includ-
ed.
The authors, a husband and wife team, claim to have traveled over 30,000
miles in several years to find hummingbirds to photograph. They depended on
a number of birders to guide them to appropriate sites. The text of the chapters
was submitted to knowledgeable ornithologists for verification.
Photographer Robert Tyrrell states that "The largest problem I encountered
while attempting to photograph hummingbirds was 'stopping' their wings." The
photos in the book attest to his success in overcoming the problem. Humming-
birds are the jewels in the avian crown, and this book illustrates that superbly.
I recommend the book also for its scientific fact. The price is not prohibitive in
today's book market. — John R. Gustafson
252 CALIFORNIA FISH AND CAME
INDEX TO VOLUME 72
AUTHORS
Ames, Jack A.: see Wendell, Hardy, Ames, and Burge, 197-212
Asson-Batres, Mary Ann: The Feeding Behavior of the luvenile Dungeness Crab, Cancer magister Dana, on the
Bivalve, Transennella tantilla (Could), and a Determination of Its Daily Consumption Rate, 144-152
Barnhart, Roger A.: see Rabin and Barnhart, 4-16
Barrett, Reginald H.: see Cogan, Thompson, Pierce, and Barrett, 47-61
Bartonek, James C: see Gilmer, Hicks, Bartonek, and McCollum, 132-143
Becker, Dennis: see Botti, Warenycia, and Becker, 62-63
Blahm, Theodore H.: see Emmett, Miller, and Blahm, 38-46
Botti, Fred, Dee Warenycia, and Dennis Becker: Utilization by Salt Marsh Harvest Mice, Reithrodontomys raviven-
tris halicoetes, of a Non-pickleweed Marsh, 62-63
Bowyer, R. Terry: Habitat Selection by Southern Mule Deer, 153-169
Burge, Richard T.: see Wendell, Hardy, Ames, and Burge, 197-212
Compagno, Leonard J.V.: see Seigel and Compagno, 172-176
Crabtree, C. Ben: see Matson, Crabtree, and Haglund, 227-231
Cross, Jeffrey N.: Epidermal Tumors in Microstomas pacificus (Pleuronectidae) Collected Near a Municipal
Wastewater Outfall in the Coastal Waters off Los Angeles (1971-1983), 68-77
Dinnel, Paul A., and Christopher W. Rogers: Northern Range Extension for California Tonguefish, Symphurus
atricauda, to Washington State, 119-121
Ebert, David A.: Observations on the Elasmobranch Assemblage of San Francisco Bay, 244-249
Emmett, Robert L., David R. Miller, and Theodore H. Blahm: Food of Juvenile Chinook, Oncorhynchus tshawyt-
scha, and Coho, O. kisutch, Salmon off the Northern Oregon and Southern Washington Coasts, May-
September 1980, 38-46
Gilmer, David S., Jane M. Hicks, James C. Bartonek, and Earl H. McCollum: Waterfowl Harvest at Tule Lake
National Wildlife Refuge, 1936-^11, 132-143
Gogan, Peter J. P., Steven C. Thompson, William Pierce, and Reginald H. Barrett: Line-Transect Censuses of Fallow
and Black-tailed Deer on the Point Reyes Peninsula, 47-61
Graham, T.W.: see Lavoipierre, Graham, Walters, and Howarth, 78-82
Gutoff, D.I.: see Phillips, Gutoff, Hansen, and Hardwick, 232-243
Haglund, Thomas R.: see Matson, Crabtree, and Haglund, 227-231
Hansen, J.E.: see Phillips, Gutoff, Hansen, and Hardwick, 232-243
Hardwick, J.E.: see Phillips, Gutoff, Hansen, and Hardwick, 232-243
Hardy, Robert A.: see Wendell, Hardy, Ames, and Burges, 197-212
Herder, Michael: see Stein, Herder, and Miller, 179-181
Hicks, Jane M.: see Gilmer, Hicks, Bartonek, and McCollum, 132-143
Howarth, J.A.: see Lavoipierre, Graham, Walters, and Howarth, 78-82
Juarez, Stephen M.: see O'Farrell, Juarez, and Uptain, 187-189
Langenwalter, Paul E. II: Indigenous Muskrats, Ondatra zibethicus, in Coastal Southern California, 121-122
Lavoipierre, M.M.J., T.W. Graham, L.L. Walters, and J. A. Howarth: The Occurence of Two Nematodes, Spirocerca
lupi and Dirofilaria immitis, in Wild Canids of the Lake Berryessa Area, Northern California, 78-82
Lea, Robert N., and Lawrence F. Quirollo: First Record of Hemitripterus bolini, the Bigmouth Sculpin, from
Californian Waters, 117-119
Littrell, E.E.: Mortality of American Wigeon on a Golf Course Treated with the Organophosphate, Diazinon,
122-124
Littrell, E.E.: Shell Thickness and Organochlorine Pesticides in Osprey Eggs from Eagle Lake, California, 182-185
Matson, Ronald H., C. Ben Crabtree, and Thomas R. Haglund: Icthyofaunal Composition and Recolonization in
a Central California Tidepool, 227-231
Matthews, Kathleen R.: Movement of Two Nearshore, Territorial Rockfishes Previously Reported as Non-Movers
and Implications to Management, 103-109
McCollum, Earl H.: see Gilmer, Hicks, Bartonek, and McCollum, 132-143
Miller, David R.: see Emmett, Miller, and Blahm, 38—46
Miller, Kathy: see Stein, Herder, and Miller, 179-181
INDEX TO VOLUME 72 253
O'Farrell, Michael J., Stephen M. Juarez, and Curt E. Uptain: An Addition to the Known Range of Stephens'
Kangaroo Rat, Dipodomys stephensi, in San Diego County, California, 187-189
Phillips, RE., D.I. Gutoff, ).E. Hansen, and I.E. Hardwick: Distribution of Major Marine Macrophytes, Seasonal
Estimates of Cracilaria Standing Crop, and Spawning Activities of the Pacific Herring, Clupea harengus pallasii,
in Elkhorn Slough, California; 1979-1982, 232-243
Pierce, William: see Gogan, Thompson, Pierce, and Barrett, 47-61
Rabin, Douglas )., and Roger A. Barnhart: Population Characteristics of Pacific Herring, Clupea harengus pallasi,
in Humboldt Bay, California, 4-16
Raquel, Paul F.: Juvenile Blue Catfish in the Sacramento-San Joaquin Delta of California, 186-187
Rogers, Christopher W.: see Dinnel and Rogers, 119-121
Roletto, Jan, and Robert Van Syoc: The Occurrence of Lepas anatifera on Zalophus californianus and Mirounga
angustirostris, 1 24-1 26
Seigel, Jeffrey A. and Leonard J.V. Compagno: New Records of the Ragged-tooth Shark, Odontaspis (erox, from
California Waters, 172-176
Sikkel, Paul C: Intraspecific Cleaning by Juvenile Salema Xenestius californiensis (Pisces: Haemulidae), 170-172
Stein, Janet L., Michael Herder, and Kathy Miller: Birth of a Northern Fur Seal on the Mainland California Coast,
179-181
Sunada, John S.: Growth and Reproduction of Spot Prawns in the Santa Barbara Channel, 83-93
Swift, Camm C: First Record of the Spotted Scorpionfish, Scorpaena plumieri, from California: The Curtain Falls
on "A Comedy of Errors.", 176-178
Thompson, Steven C: see Gogan, Thompson, Pierce, and Barrett, 47-61
Uptain, Curt E.: see O'Farrell, Juarez, and Uptain, 187-189
Walters, L.L.: see Lavoipierre, Graham, Walters, and Howarth, 78-82
Warenycia, Dee: see Botti, Warenycia, and Becker, 62-63
Welker, Heather J.: Fawn Mortality in the Lake Hollow Deer Herd, Tehama County, California, 99-102
Welker, Heather J.: Fawn Rearing Habitat of the Lake Hollow Deer Herd, Tehama County, California, 94-98
Wells, Alan W.: Aspects of Ecology and Life History of the Woolly Sculpin, Clinocottus analis, from Southern
California, 213-226
Wendell, Frederick E., Robert A. Hardy, Jack A. Ames, and Richard T. Burge: Temporal and Spatial Patterns in Sea
Otter, Enhydra lutris, Range Expansion and in the Loss of Pismo Clam Fisheries, 197-212
White, James R.: The Striped Bass Sport Fishery in the Sacramento-San Joaquin Estuary, 1969-1979, 17-37
Winter, Brian D.: A Method for the Efficient Removal of Juvenile Salmonid Otoliths, 63-64
Zielinski, William J.: Relating Marten Scat Contents to Prey Consumed, 110-116
SUBJECT
Bass, striped: Sport fishery in the Sacramento-San Joaquin estuary, 1969-1979, 17-37
Behavior, feeding: Of the juvenile Dungeness crab, Cancer magister Dana, on the bivalve Transennella tantilla
(Gould), and a determination of its daily consumption rate, 144-152
Catfish, blue: Juvenile, in the Sacramento-San Joaquin Delta of California, 186-187
Clam, Pismo: Temporal and spatial patterns in sea otter range expansion and in the loss of, fisheries, 197-212
Cleaning: Intraspecific, by juvenile salema, 170-172
Censuses, line-transect: Of fallow and black-tailed deer on the Point Reyes Peninsula, 47-61
Crab, Dungeness: The feeding behavior of the juvenile, on the bivalve, Transennella tantilla (Gould), and a
determination of its daily consumption rate, 144-152
Deer, black-tailed: Line-transect censuses of fallow and, on the Point Reyes Peninsula, 47-61
Deer, fallow: Line-transect censuses of, and black-tailed deer on the Point Reyes Peninsula, 47-61
Deer: Fawn mortality in the Lake Hollow, herd, Tehama County, California, 99-102
Deer: Fawn rearing habitat of the Lake Hollow, herd, Tehama County, California, 94-98
Deer, southern mule: Habitat selection by, 153-169
Elasmobranch: Observations on the, assemblage of San Francisco Bay, 244-249
Growth: And reproduction of spot prawns in the Santa Barbara Channel, 83-93
Habitat: Selection by southern mule deer, 153-169
Habitat: Fawn rearing, of the lake Hollow deer herd, Tehama County, California, 94-98
Herring, pacific: Distribution of major marine macrophytes, seasonal estimates of Cracilaria standing crop, and
spawning activities of the, in Elkhorn Slough, California, 1979-1982, 232-243
254 CALIFORNIA FISH AND CAME
Herring, Pacific: Population characteristics of, in Humboldt Bay, California, 4-16
Lepas anatifera: The occurrence of, on Zalophus califomianus and Mirounga angustirostris, 124-126
Life history: Aspects of ecology and, of the woolly sculpin, Clinocottus analis, from southern California, 213-226
Macrophytes, marine: Distribution of major, seasonal estimates of Cracilaria standing crop, and spawning activities
of the Pacific herring, in Elkhorn Slough, California, 1979-1982, 232-243
Marten: Relating, scat contents to prey consumed, 110-116
Mice, salt marsh harvest: Utilization by, of a non-pickleweed marsh, 62-63
Muskrats: Indigenous in coastal southern California, 121-122
Nematodes: The occurrence of two, in wild canids of the Lake Berryessa area, northern California, 78-82
Organophosphate, Diazinon: Mortality of American wigeon on a golf course treated with, 122-124
Osprey, eggs: Shell thickness and organochlorine pesticides in, from Eagle Lake, California, 182-185
Otoliths, salmonid: A method for the efficient removal of juvenile, 63-64
Otter, sea. Temporal and spatial patterns in, range expansion and in the loss of Pismo clam fisheries, 197-212
Pesticides, organochlorine: Shell thickness and, in osprey eggs from Eagle Lake, California, 182-185
Population characteristics: Of Pacific herring, in Humboldt Bay, California, 4-16
Prawns, spot: Growth and reproduction of, in the Santa Barbara Channel, 83-93
Rat, Stephens' kangaroo: An addition to the known range of, in San Diego County, California, 187-189
Recolonization: Ichthyofaunal composition and, in a central California tidepool, 227-231
Reproduction: Growth and, of spot prawns in the Santa Barbara Channel, 83-93
Rockfishes: Movement of two nearshore, territorial, previously reported as non-movers and implications to
management, 103-109
Salema: Intraspecific cleaning by juvenile, 170-172
Salmon, chinook: Food of juvenile, off the northern Oregon and southern Washington coasts, May-September
1980, 38-46
Salmon, coho: Food of juvenile, off the northern Oregon and southern Washington coasts, May-September 1980,
38-46
Scat: Relating marten, contents to prey consumed, 110-116
Scorpionfish, spotted: First record of the, from California: The curtain falls on "A Comedy of Errors.", 176-178
Sculpin, bigmouth: First record of, Hemitripterus bolini, from Californian waters, 117-119
Sculpin, woolly: Aspects of ecology and life history of the, from southern California, 213-226
Seal, northern fur: Birth of, on the mainland California coast, 179-181
Shark, ragged-tooth: New records of the, from California waters, 172-176
Shell thickness: And organochlorine pesticides in osprey eggs from Eagle Lake, California, 182-185
Tonguefish, California: Northern range extension for, to Washington state, 119-121
Tumors, epidermal: In Microstomas pacificus (Pleuronectidae) collected near a municipal wastewater outfall in
the coastal waters off Los Angeles (1971-1983), 68-77
Waterfowl: Harvest at the Tule Lake National Wildlife Refuge, 1936-41, 132-143
Wigeon, American: Mortality of, on a golf course treated with the organophospate, Diazinon, 122-124
INDEX TO VOLUME 72
255
SCIENTIFIC NAMES
Acanthogobius flavimanus: 17
Acmaea spp.: 221
Agarum: 233
Aix sponsa: 133
Ammodytes hexapterus: 41, 42, 43
Anas acuta: 134
Anas americana: 122-124
Anas clypeata: 134
Anas platyrhynchos: 123, 134
Anatanais normani: 220
Anoplarchus purpurescens: 229
Anser albifrons: 134
Anthopleura xanthogrammica: 214
Aquila chrysaetos: 47
Artedius lateralis: 215, 228, 229
Ascarophis sp.: 225
Atherinops affinis: 171, 228, 229
Atylus tridens: 41, 42, 43
Aves: 111, 113, 114
Axis axis: 48
Aythya americana: 133
Aythya collaris: 140
Aythya spp.: 134
Aythya valisineria: 133
Bos taurus: 47
Branta canadensis: 123
Branta canadensis hutchinsii: 134
Branta canadensis minima: 134
Branta canadensis spp.: 134
Bucephala albeola: 133
Bucephala spp.: 134
Callorhinus ursinus: 179
Cancer antennarius: 197
Cancer magister: 41, 42, 43, 119, 144-152, 197
Cancer oregonensis: 41, 42, 43
Cancer productus: 197
Cam's latrans: 47, 78, 100, 155
Canis lupus: 110
Capra spp.: 47
Carcinus maenas: 150
Cebidichthys violaceus: 229, 230
Cerastoderma edule: 150
Cervus elaphus: 47
Chen rossii: 133
Cirolana harfordi: 220
Cirripedia: 42, 43
Clinocardium nuttali: 145, 146, 148, 149, 150
Clinocottus analis: 213-226, 229
Clinocottus recalvus: 215
Clupea harengus pallasii: 4-16, 41, 42, 43, 232-243
Crangon sp.: 17
Cryptomya sp.: 149
Dama dama: 48
Dipodomys stephensi: 187-189
Dirofilaria immitis: 78-82
Dorosoma petenense: 17
Embiotoca jacksoni: 215
Engraulis mordax: 17, 41, 42, 43
Enhydra lutris: 197-212
Enteromorpha: 236, 238
Equus caballus: 47
Eutamias spp.: 111, 113, 114, 115
Felis concolor: 47, 100
Fucus: 233
Fulica americana: 123
Galeorhinus zyopterus: 245
Cibbonsia elegans: 214
Cibbonsia metzi: 214, 229
Cibbonsia montereyensis: 229, 230
Girella nigricans: 171, 214, 229, 230
Claucomys sabrinus: 111, 112
Cobiesox maeandricus: 229, 230
Gobiesox rhessodon: 215
Gracilaria: 232-243
Haliaeetus leucocephalus: 47
Hemitripterus bolini: 117-119
Hemitripterus marmoratus: 117
Hemitripterus marmoratus: 117
Hermosilla azurea: 171, 215
Heterostichus rostratus: 170
Hippoglossoides elassodon: 74
Hyperoche medusarum: 42, 43
Hypsoblennius gilberti: 214
Hypsoblennius jenkinsi: 215
Ictalurus furcatus: 186
Laminaria: 233
Laminaria saccarina: 233
Laminaria sinclairii: 233
Lepas anatifera: 124-126
Leptocottus armatus: 17
Lepus califomicus: 114
Limacina sp.: 41, 42, 43
Littorina planaxis: 214
Lynx rufus: 155
Macoma sp.: 149
Macrocystis integrifolia: 233
Macrocystis pyrifera: 202
Martes americana: 110-116
Martes pennant i: 116
Micrometrus aurora: 228, 229
Micrometrus minimus: 215
Microstomus pacificus: 68-77, 1 17
Microtus spp.: 111, 113, 114, 115
Mirounga augustirostris: 124-126
Morone saxatilis: 17-37
Mustela nivalis: 115
Mustela putorius furo: 1 10
Mustelus henlei: 245
Myliobatis califomica: 246
Mytilus californianus: 214
Mylilus edulis: 236
Nautilus pompilius: 124
Neomysis kadiakensis: 42, 43
Neptunea pribiloffensis: 1 18
Notorynchus maculatus: 244, 246
Odocoileus hemionus: 47-61, 94, 99
Odocoileus hemionus califomicus: 167
Odocoileus hemionus columbianus: 94-98, 99-102,
167
Odocoileus hemionus fuliginatus: 153
Odocoileus hemionus hemionus: 167
Odontaspis ferox: 172-176
Odontaspis herbsti: 173
Oligocottus rimensis: 229
Oligocottus rubellio: 215
Oligocottus snyderi: 215, 229
Oncorhynchus kisutch: 38—46
Oncorhynchus spp.: 38
Oncorhynchus tshawytscha: 38—46
Ondatra zibethicus: 121-122
256
CALIFORNIA FISH AND GAME
Ondatra zibethicus bemardi: 121
Ondatra zibethicus mergens: 121
Opecoelus adsphaericus: 225
Osmeridae: 41, 42, 43
Ovis aries: 47
Oxyjulus californica: 170, 171
Oxyura jamaicencis: 133
Pachygrapsus crassipes: 214, 221
Pagurus samuelis: 221
Pagurus spp.: 41, 42, 43
Pandalus borealis: 89
Pandalus jordani: 42, 89
Pandalus platyceros: 83-93
Pandion haliaetus: 182
Paraclunio sp.: 220
Paralabrax clathratus: 171, 225
Paralichthys californicus: 34
Parathemisto pacifica: 42, 43
Parophrys vetulus: 74
Peromyscus: 1 14
Peromyscus maniculatus: 111, 112, 113
Petalonia: 224
Phragmatopoma californica: 220, 223
Pinnixa spp.: 43
Platichthys stellatus: 74
Pleuronectidae: 42
Pollicipes polymerus: 214
Polysiphonia: 236
Porcellanidae: 41, 42, 43
Raja binoculata: 246
Reithrodontomys raviventris halicoetes: 62-63
Reithrodontomys raviventris raviventris: 62
Rhacochilus vacca: 215
Sagitta sp.: 42
Salicornia pacifica: 62
Salmo gairdneri: 63
Sardinops sp.: 17
Scapanus: 114
Scapanus latimanus: 115
Scorpaena: 176
Scorpaena guttata: 176, 177
Scorpaena histrio: 176
Scorpaena mystes: 176
Scorpaena plumieri: 176-178
Scorpaenichthys marmoratus: 215
Sebastes: 103, 118
Sebastes carnatus: 103-109
Sebastes chrysomelas: 103-109
Sebastes sp.: 34
Sebastes spp.: 41, 42, 43, 173, 215
Seriola lalandi: 171
Sorex spp.: 113, 114
Spermophilus lateralis: 111, 113, 1 14
Spirinchus thaleischthys: 42
Spirocerca lupi: 78-82
Spirontocaris picta: 223
Squalus acanthias: 246
Sus scrofa: 47
Symphurus atricauda: 119-121
Tamiasciurus: 114, 115
Tamiasciurus douglasii: 111, 113
Tegula funebralis: 214
Thomomys: 1 14
Thomomys monticola: 113, 115
Thyssanoessa spinifera: 41, 42, 43
Tivela stultorum: 198
Transennella tantilla: 144-152
Triakis semifasciata: 245
Ulca bolini: 117
Ulca marmorata: 117
Ulva: 233
Urocyon cinereoargenteus: 78
Ursus americana: 100
Ursus arctos: 47
Vibilia cultripes: 42
Vulpes vulpes: 47, 110
Xenistius californiensis: 170-172
Xererpes focurum: 229, 230
Xiphister atropurpureus: 229
Xiphister mucosus: 228, 229, 230
Zalophus californianus: 124-126
Zoster a marina: 5, 235, 236, 237
I'hotoclcctronic composition by
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