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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	141
12	123	0	0	0	136
3	10	83	0	0	96
0	3	14	28	0	45
0	1	2	7	8	20
3	0	0	3	2	8
1	0	0	1	0	2
0	0	1	1	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	55
0	4	2	4	0	0	1	11
18	2	4	4	1	0	0	41
5	13	1	4	0	1	2	26
3	4	1	0	0	0	0	11
2	3	2	0	0	0	0	18
11	4	0	1	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	37
14	0	8	1	0	0	23
0	2	2	1	0	0	33
14	0	2	1	1	0	18
4	0	0	0	0	2	6
0	2	5	1	0	0	9
1	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 G = GILL NET NUMBERS INDICATE NUMBER RICHMOND OF TIMES FISHED

R5L1

R1 L2 G2 Angel Island R4

"*&r-

R8G2 L1 R3G1

Alcatraz Island

%

R10

SAN FRANCISCO

R2 L1

R2

Hunters Point R3L3 Candlestick Point 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

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

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INSTRUCTIONS TO AUTHORS

EDITORIAL POLICY

California Fish and Game is a technical, professional, and educational journal devoted to the conservation and understanding of fish and wildlife. Original manuscripts submitted for consideration should deal with the California flora and fauna or provide information of direct interest and benefit to California researchers. Authors may submit an original plus two copies, each, of manuscript, tables, and figures at any time.

MANUSCRIPTS: Authors should refer to the CBE Style Manual (Fifth Edition) and a recent issue of California Fish and Game for general guidance in preparing their manuscripts. Some major points are given below.

1. Typing — All material submitted, including headings, footnotes, and literature cited must be typewritten doublespaced, on white paper. Papers shorter than 10 typewritten pages, including tables, should follow the format for notes.

2. Citations — All citations should follow the name-and-year system. The "library style" will be followed in listing references.

3. Abstracts — Every article must be introduced by a concise abstract. Indent the abstract at each margin to identify it. Abstracts, on separate sheets of paper, should accompany "Notes".

4. Abbreviations and numerals — Use approved abbreviations as listed in the CBE Style Manual. In all other cases spell out the entire word.

TABLES: Each table should be typewritten with the heading left margin justified. Tables should be numbered consecutively beginning with "1" and placed together in the manuscript following the Literature Cited section. Do not double space tables. See a recent issue of California Fish and Game for format.

FIGURES: Consider proportions of figures in relation to the page size of California Fish and Game. The usable printed page is 1 1 7 by 1 91 mm. This must be considered in planning a full page figure, for the figure with its caption cannot exceed these limits. Photographs should be submitted on glossy paper with strong contrasts. All figures should be identified with the author's name in the upper left corner and the figure numbers in the upper right corner. Markings on figures should be made with a blue china marking pencil. Figure captions must be typed on a separate sheet headed by the title of the paper and the authors name.

PROOFS: Galley proofs will be sent to authors approximately 60 days before publication. The author has the ultimate responsibility for the content of the paper, and is expected to check the galley proof very carefully.

PAGE CHARGES AND REPRINTS: All authors will be charged $35 per page for publication and will be billed before publication of manuscripts. Reprints may be ordered through the editor at the time the proof is submitted. Authors will receive a reprint charge schedule along with the galley proof.

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39	0	0	0	1	141
12	123	0	0	0	136
3	10	83	0	0	96
0	3	14	28	0	45
0	1	2	7	8	20
3	0	0	3	2	8
1	0	0	1	0	2
0	0	1	1	0	2

8	20	11	10	4	0	0	55
0	4	2	4	0	0	1	11
18	2	4	4	1	0	0	41
5	13	1	4	0	1	2	26
3	4	1	0	0	0	0	11
2	3	2	0	0	0	0	18
11	4	0	1	0	0	0	39

0	0	3	1	1	0	37
14	0	8	1	0	0	23
0	2	2	1	0	0	33
14	0	2	1	1	0	18
4	0	0	0	0	2	6
0	2	5	1	0	0	9
1	0	0	0	0	0	4

34	103	66	156	40
31	61	69	89	23
25	51	75	68	18
53	15	47	32	8
18	27	82	33	8
44	5	56	9	2
0	2	100	2	1

39	0	0	0	1	141
12	123	0	0	0	136
3	10	83	0	0	96
0	3	14	28	0	45
0	1	2	7	8	20
3	0	0	3	2	8
1	0	0	1	0	2
0	0	1	1	0	2

8	20	11	10	4	0	0	55
0	4	2	4	0	0	1	11
18	2	4	4	1	0	0	41
5	13	1	4	0	1	2	26
3	4	1	0	0	0	0	11
2	3	2	0	0	0	0	18
11	4	0	1	0	0	0	39

0	0	3	1	1	0	37
14	0	8	1	0	0	23
0	2	2	1	0	0	33
14	0	2	1	1	0	18
4	0	0	0	0	2	6
0	2	5	1	0	0	9
1	0	0	0	0	0	4

34	103	66	156	40
31	61	69	89	23
25	51	75	68	18
53	15	47	32	8
18	27	82	33	8
44	5	56	9	2
0	2	100	2	1