£

CALIFORNIA FISH-GAME

I "CONSERVATION OF WILDLIFE THROUGH EDUCATION"

California Fish and Game is a journal devoted to the conser- vation of wildlife. If its contents are reproduced elsewhere, the authors and the California Department of Fish and Game would appreciate being acknowledged.

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Please direct correspondence, except regarding paid subscrip- tions, to:

CAROL M. FERREL, Editor California Fish and Game 987 Jedsmfth Drive Sacramento, California 95819

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VOLUME 58

OCTOBER 1972

NUMBER 4

Published Quarterly by

STATE OF CALIFORNIA

THE RESOURCES AGENCY

DEPARTMENT OF FISH AND GAME

STATE OF CALIFORNIA

RONALD REAGAN, Governor

THE RESOURCES AGENCY

NORMAN B. LIVERMORE, JR., Secretary for Resources

FISH AND GAME COMMISSION

JOSEPH RUSS III, President, Ferndale

SHERMAN CHICKERING, Vice President PETER T. FLETCHER, Member

San Francisco Rancho Santa Fe

C. RANS PEARMAN, Member TIMOTHY M. DOHENY, Member

San Gabriel Los Angeles

DEPARTMENT OF FISH AND GAME

G. RAY ARNETT, Director

1416 9th Street Sacramento 95814

CALIFORNIA FISH AND GAME Editorial Staff

CAROL M. FERREL, Editor-in-Chief - Sacramento

KENNETH A. HASHAGEN, Editor for Inland Fisheries Sacramento

MERTON N. ROSEN, Editor for Wildlife Sacramento

ROBSON COLLINS, Editor for Marine Resources Long Beach

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

HAROLD K. CHADWICK, Editor for Striped Bass, Sturgeon, and Shad ..Stockton

( 252 )

CONTENTS

Page Primary Productivity in a New and an Older California

Reservoir Lawrence L. Chamberlain 254

A Midwater Trawl for Threadfin Shad, Dorosoma pentenense

C. E. von Geldern, Jr. 268

Morphology and Variation of the Modoc Sucker, Catostonvus micro ps Rutter, with Notes on Feeding Adaptions

Michael Martin 277

Contributions to the Life History of the Piute Sculpin in Sagehen

Creek, California Albert C. Jones 285

The Effects of Diesel Fuel on a Stream Fauna_. _#. Bruce Burn 291

A Subpopulation Study of the Pacific Sardine— Kenneth F. Mais 296

Check List of Intertidal Fishes of Trinidad Bay, California, and

Adjacent Areas John B. Moving 315

Not< s

Two New Sea Urchin — Acorn Barnacle Associations

James L. Ilouk and J oh n M. Duffy 321

New Hosts and Bathymetric Range Extension for Colobomatus

embiotocae (Crustacea, Copepoda) Ernest W. Iverson 323

Southern Range Extension for the Yellowfin Goby, Acantho- gobius flavimanus (Temminck and Schlegel)

Gary E. Kukowski 326

California Condor Survey, 1971 W. Dean Carrier,

Robert D. Mallette, Sanford Wilbur, and John C. Bornemau 327

Book Reviews 329

Index to Volume 58 333

( 253 ) 2—83609

Calif. Fish ,nnl Game, 58(4) : 254-267. 1D7L'.

PRIMARY PRODUCTIVITY IN A NEW AND AN OLDER CALIFORNIA RESERVOIR1

LAWRENCE L. CHAMBERLAIN

Inland Fisheries Branch

California Department of Fish and Game

The sport fishery in new reservoirs often reaches a peak and then undergoes a marked decline a few years following impoundment. One theory attributes such declines to diminishing basic fertility. To test this theory, primary productivity in a new reservoir was measured by the C " method for 4 years. An older reservoir served as a control. High initial rates of carbon fixation in the new reservoir, attributed to flood- ing of organic material, were transitory. Subsequent patterns of primary productivity were similar in the two waters. Increases in primary pro- ductivity in both reservoirs were associated with establishment of planktivorous fish populations and demonstrated that declining primary productivity is not an inevitable result of initial reservoir aging.

INTRODUCTION

Since the 1930 's, fisheries workers have become increasingly aware that after an initial period of good fishing in new impoundments, yield to the angler and the overall production of game fish tend to decline, often dramatically. Although there is considerable variation among such waters, it appears that the long term yield of many reservoirs is half or less of that enjoyed during the first few years following im- poundment (Abell and Fisher 1953; Kimsey 1958; Jenkins 1961). Among hypotheses advanced to explain this phenomenon, major em- phasis has been accorded those which maintain either (i) that changes in fish population structures are responsible for such declines (Bennett 1947) or (ii) that these declines reflect diminishing basic fertility in the reservoir (Ellis 1937). As yet there is insufficient evidence to deter- mine whether either theory might fully explain these fishery declines, but clearly a better knowledge of those factors most important in de- termining ultimate fish yields of our freshwater reservoirs is essential to the development of sound management programs.

The completion in late 1964 of Merle Collins Keservoir in the foot- hills of the Sierra Nevada northeast of Marysville, California, provided an opportunity to study various aspects of initial aging in a reservoir. To gather information on changes in primary productivity which might influence fish yields, C14 measurements were begun in June 1964 on the partial pool forming at the reservoir and continued through De- cember 3968. In order to provide a comparative baseline to aid in the interpretation of test results from Merle Collins, limnological studies were also conducted from August 1964 through June 1967 on Folsom Lake, a large foothill reservoir formed in 1955 by the impoundment of the American Eiver near Sacramento. Concurrent studies were un-

1 Accepted for publication June 1972. This work was performed as part of Dingell- Johnson Project California F-18-R, "Experimental Reservoir Management", sup- ported by Federal Aid to Fish Restoration Funds.

( 254 )

PRIMARY PRODUCTIVITY

255

dertaken at Merle Collins to define changes in the fish populations (K. A. Hashagen, Calif. Dep. Fish and Game, MS), and to describe various aspects of the fishery (Rawstron and Hashagen 1972).

STUDY RESERVOIRS

Inherent in the experimental design of this study was the assumption that the similarity of the two reservoirs in respects other than age and size (Table 1) would allow a meaningful comparison of primary pro- ductivity in each water. However, there are also differences in the operating schedules of these two reservoirs.

TABLE 1 — Comparison of Some Characteristics of Folsom Lake and Merle Collins Reservoir, California *

Location

Surface elevation; m above m.s.l

Surface area; ha

Capacity ;m3

Maximum depth; m

Observed annual fluctuation in surface elevation; m

Maximum

Minimum

Observed surface temperature range; C

Observed pH range

Observed total alkalinity range; mg XI"1 CaC03-.

Observed range, Secchi transparency; m

Flushing ratet

Folsom	Merle Collins
lat38°42' N long 121° 9' W	lat 39° 20' N long 121° 19' W
142.0	360.6
4,633	401
1,246.3 X 106	70.3 X 106
79.3	47.2
16.2 (1964) 11.6 (1966)	14.3 (1966) 9.5 (1968)
7.8-28.3	6.1-28.6
6.2-8.1	6.2-8.5
11-32	15-52
0.27-7.92	0.19-6.25
2.59	1.18

* Data at reservoir gross stage, where applicable.

f Flushing rate = mean annual discharge -5- capacity.

Folsom Lake is a large multipurpose reservoir, operated principally for flood control but also to provide water for irrigation, domestic, mu- nicipal, industrial, and power production purposes, as well as to con- tribute to water quality control in the Sacramento-San Joaquin Delta. Releases are made through adjustable louver outlets which draw water from the upper hypolimnion or metalimnion (Rawstron 1964). Merle Collins Reservoir is a single-purpose impoundment designed to provide storage for irrigation water for the Browns Valley Irrigation District. A single-level outlet structure draws water from near the deepest part of the hypolimnion.

Both reservoirs are drawn down through the summer and minimum surface elevations occur in fall or early winter. Over half of the precipi- tation in central California occurs from December through February and produces substantially increased inflow during this period. High flows into Folsom generally persist into late spring, due to snowmelt. Relatively little snowfall occurs in the Merle Collins drainage.

256

CALIFORNIA PISH AND GAME

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PRIMARY PRODUCTIVITY

257

FIGURE 2. Typical Merle Collins Reservoir annual isotherms and Secchi depths (•) from 1966 observations.

Jan Feb ' Mar T Ap

1966

FIGURE 3. Typical Folsom Lake annual isotherms and Secchi depths (•) from 1966 observations.

Merle Collins is allowed to fill during the winter and uncontrolled surface spill generally occurs by early February and may persist through May. Depending on predicted runoff from snow accumulations at high elevations, up to 493.4 X 10° m3, or nearly half of the total

3— 83009

258 CALIFORNIA FISH AND GAME

capacity of Folsorn, may be reserved for control of flood flows through Late fall and winter. Maximum surface levels generally occur in June. Despite these differences, the total alkalinity, pH. DO, and temperature regimes of the reservoirs are quite similar (Table 1, Figures 1, 2, and 3).

METHODS AND MATERIALS

Primary productivity was measured by the C14 method of Steemann- Nielsen (1952) as modified by Goldman (1960, 1963). To allow for iso- tope discrimination, a correction factor of 6', was applied in the cal- culations.

Primary productivity was usually sampled two to five times each month while the reservoirs were thermally stratified and once monthly during the whiter. A permanent sampling station near the deepest point in each reservoir was marked by an anchored buoy from which samples were suspended. Based on Secchi transparencies observed at the start of the study, eight sampling strata were chosen at fixed depths to 15 m in Merle Collins and to 30 m in Folsom. Dark bottles were included at the upper, lower, and two intermediate depths to provide a correction for nonphotosynthetic carbon uptake. This sampling regime proved adequate to include the maximum compensation depth in each reser- voir. AYater samples were collected from the appropriate depths with a 3-liter, non-metallic Van Dorn bottle and transferred to 125-ml glass- stoppered Pyrex bottles. Each sample was injected with 0.5 ml of radio- active sodium carbonate tracer solution by means of an automatic hypo- dermic syringe, and then suspended at the depth from which that sam- ple had been drawn. Incubation was for the 4-hr period spanning local mid-day and except during actual handling for injection or filtration, samples were kept and transported in a lightproof box. fSamples were filtered in simultaneous series of four on a plexiglass multiple filtration manifold produced by Min Plastics & Supply Center, Honolulu, Ha- waii. A filtration vacuum of 10-15 inches Hg allowed an entire set of 12 samples to be filtered in approximately 15-21) min. The filtration funnels were treated with Desicote (Beckman Instruments, Inc.), which effec- tively prevented adherence of sample material to the funnel sides (C. R. Goldman, Univ. Calif., Davis, pers. comm.). Early in the study 50-ml sub-samples were filtered on 25-nmi HA Alillipore filters (porosity 0.45 ± 0.02 microns), but in later experiments the entire 125-ml sample was filtered to increase total activity of the filtered samples and reduce counting errors.

During the first year of the study a tracer solution with an absolute activity of 2.30 microcuries per milliliter (\ic/ml), obtained from Hazel- ton-Xuclear Science Corp., Palo Alto, California, in 100-ml rubber- stoppered serum bottles, was used at both reservoirs. Beginning in July 1965, a more active tracer solution was used. This solution, with an absolute activity of 3.58 ue/ml, was prepared in a single large lot by C. R. Goldman and sealed in individual, 10-ml sterile glass ampules. The higher activity decreased sample counting time; the packaging in individual sealed sterile containers insured against possible contamina- tion and loss of radioactivity.

AYater samples for both total alkalinity and primary productivity measurements were drawn from the same 3-liter sample and available

PRIMARY PRODUCTIVITY

259

carbon was determined from total alkalinity using the conversion table of Saunders, Trama, and Bachmann (1962). Ancillary limnological data, such as Secchi transparency and vertical profiles of temperature and dissolved oxygen, were collected during the primary productivity sample incubation period. Solar radiation was measured at each reser- voir on sampling days with a recording pyrheliograph (Belfort 53850), and the ratio of daily insolation to the 4-hr sample period insolation was used to expand partial photoperiod results to full-day photosyn- thesis.

Sample activity was measured by the staff of C. R. Goldman using an automatic gas-flow Geiger-Muller counter with a Micromil window. A standard sample of known activity was routinely counted with each set of experimental samples and the counting efficiency of this equipment was periodically calibrated by gas-phase assay of representative filters following a Van Slyke wet combustion of the labeled algae to C02. The resultant values were compared with gas-phase assays of National Bureau of Standards samples (Goldman 1968a).

RESULTS AND DISCUSSION

The carbon assimilation rates observed from sampling the partial pool at Merle Collins in 1964 were not only quite variable but reached levels over three and a half times the rates observed during the remainder of the study. Although experimental error may sometimes be related to the familiarity of field personnel with C14 sampling procedures (Gold- man and Carter 1965), the relatively uniform results obtained at Fol- som by the same field crews during this period indicate that experience was not an important consideration in this instance (Table 2).

TABLE 2 — Primary Productivity in the Partial Pool at Merle Collins Reservoir and in Folsom Lake During 1964

Merle Collins	Folsom
Date	mgC-m 3-day '	Date	mgC-m 3-day '
June 16	56.44 14S.71 51.30 29.34 9.67 41.57 25.97	Aug. 21	3.88
Julv 20 _ .	Sept. 8	5.14
Aug. 4	Oct. 24	2.77
Aug. 17 - - -.-	Dee. 12	5.84
Sept. l._
Oct. 14
Nov. 23

Before inundation, the basin of Merle Collins Reservoir consisted largely of grazing land interspersed with brush. Even though most of the brush was removed before basin flooding, the substantial amount of organic debris remaining undoubtedly contributed a fertilizing effect. It is also possible that, in the early stages of reservoir formation, the

260

CALIFORNIA FISH AND GAAIE

succession from lot it- to lentic plankters was reflected by these marked changes in primary productivity. During this period, the partial pool contained about 185 x 104 m3 and covered about 182 ha. Runoff from heavy precipitation in late November 1!>(>4 resulted in the rapid filling of Merle Collins Reservoir. Thereafter, such extreme and erratic fluc- tuations in primary productivity were not observed and photosynthetic rates in the two reservoirs were similar (Table 3).

TABLE 3 — Mean Annual Carbon Assimilation in mgC • m (Range of Monthly Means in Parentheses)

day ]

	} ear	Merle Collins	Folsom
I'.Mi I		53.78 (9.67-148.71)	4.28
		(2.77-5.84)
1965		10.98 (5.68-16.92)	9.44
		(3.63-16.04)
L966		13.43 (1.51-28.08)	15.32
		(4.88-23.84)
I967t		19.66 (3.56-32.29)	16.35
		(8.25-33.04)
1968...-		25.52 (6.33-38.90)

* Partial year results.

f Folsom experiments terminated in June.

Expansion of the 4-hr mid-day results by the ratio of daily insola- tion to 4-hr mid-day insolation gave an underestimate of about 8% when compared to the total observed carbon uptake during a diurnal experiment (Figure 4), which consisted of a series of 4-hr experiments

E —

0400

40	-		Carbon Assimilation
			,^--*— ^"~X
			/s	^^ ^
			//
			//	\ N
30			//	\ \ \ \ X N \ \		-
				\ \
			//	\ \
			//	\ \
			//	\ \
			//	\ \
			//	\ \
20			// / / // // // // // // //	\ \ \ \ \ \ \ \ \ \ \ \ \ \ \ \ \ \		""
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- I 2

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O8.0

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FIGURE 4. Net primary productivity and incident solar radiation during a diurnal experiment at Merle Collins Reservoir, June 21, 1967.

PRIMARY PRODUCTIVITY 261

spanning the period from dawn to dusk. The use of this expansion to obtain day-rate estimates throughout the study appeared to be a reasonable expedient since the introduced error probably remained small (Vollenweider 1965 ; Wetzel 1965) . Seasonal variation in primary productivity could not, however, be predicted from insolation, total alkalinity, or a combination of the two (Figure 1).

Although volumetric rates of carbon fixation were generally similar in Merle Collins and Folsom, except in 1964 (Table 3), the mean com- pensation depth in Folsom was 15.3 m, with a range of 8.0 to 30.0, while the mean compensation depth in Merle Collins was 6.1 m, with a range of 2.0 to 13.7, reflecting the greater transparency of Folsom Lake (Figures 2 and 3). Because of this consistently thicker euphotic zone, Folsom was actually the more productive water.

To properly account for such variations in euphotic depth, compari- sons of primary productivity are more meaningful when results are expressed in terms of unit area. For the reasons outlined below, my comparisons between primary productivity in Merle Collins Reservoir and Folsom Lake (Figure 5) represent integral photosynthesis at the sampling sites rather than mean primary productivity per unit area.

It is well established that photosynthesis may vary considerably in different parts of large lakes and that such variation can be largely influenced by the contributions of tributary inflow (Sorokin 1959; Goldman 1960; Goldman and Wetzel 1963; Goldman and Carter 1965). Rawstron (1964) found that limnological conditions in Folsom Lake were well represented by conditions measured at the same sampling location as that used in this study. Observations of transparency and of vertical profiles of temperature and dissolved oxygen at various loca- tions in Merle Collins indicated that the sampling location used there during this study was also representative of conditions in the reser- voir as a whole. However, both sampling sites are near that end of each reservoir farthest from major tributary inflow and therefore at a location least representative of overall limnological conditions during periods of high inflow. Since patterns of high inflow do not coincide exactly in the two reservoirs, results obtained at these sampling stations are not always comparably representative of average limnological conditions.

In a fluctuating reservoir, not only does the volume of the euphotic zone vary with changes in compensation depth, but also the relation- ship between euphotic zone volume and compensation depth varies with changes in surface elevation. Surface area also changes at a variable rate with respect to elevation. The method outlined by Rupp and DeRoche (1965) in describing the primary productivity of three small Maine lakes is therefore particularly appropriate for use with fluctuat- ing reservoirs. This method divides the euphotic zone into strata and the mean volumetric rate of carbon assimilation in each stratum is multiplied by the volume of that stratum. The products are summed and division by total surface area yields the desired estimate of mean productivity per unit area. In large reservoirs particularly, synoptic sampling of both off- and onshore areas should be included.

262

I M.IFOKXIA FISH AND GAME

600

'

400

1964

FOLSOM LAKE

MERLE COLLINS RESERVOIR

t

616

;

-

1965

soc

E 100

E 0

-966

: : -

1967

300

968

juTT

Sep*

Oct Nov Dec

FIGURE 5. Net primary productivity and distribution of sampling effort for all years in Merle Collins Reservoir and Folsom Lake.

PRIMARY PRODUCTIVITY 263

There are no records of elevations for Merle Collins Reservoir before mid-April 1966 ; therefore, the volumes of the sampling strata cannot be determined before that time. In August, September and October 1964, compensation depth exceeded mean depth in Folsom by as much as 5 m, or nearly 20% of the compensation depth at that time. In August and September 1966, compensation depth exceeded mean depth in Merle Collins by as much as 1.2 m, or about 9% of the compensation depth. Again, the lack of surface elevation records for Merle Collins prevents the correction suggested by Goldman (1960), of substituting mean depth for compensation depth when the average surface area productivity is limited by mean depth.

While estimates of mean areal productivity are desirable because they allow calculation of net annual production, integral photosyn- thesis is suitable for the descriptive comparisons of this study. From these comparisons it is evident that not only did the primary produc- tivity in Merle Collins Reservoir not decline, but both reservoirs ex- hibited an increase in primary productivity over the course of the investigation. The patterns of increase, however, were dissimilar.

It appears that Folsom Lake experienced a substantial increase in primary productivity each year at least through 1966, and partial year results indicated no major departure from that trend in 1967. Merle Collins Reservoir, in contrast, after demonstrating a high degree of variability during the initial stages of impoundment, exhibited the first persistent increase in net carbon fixation during the late summer and fall of 1966. Primary productivity in Merle Collins increased noticeably during 1967 relative to the two previous years and, although data are lacking for the fall of 1968, it is apparent that a dramatic increase in primary productivity occurred in that year, particularly during mid- summer. No relationships were apparent between these patterns of pri- mary productivity and any combination of elimatological and/or water withdrawal variables.

The increase in primary productivity in Merle Collins in 1966 seems to have been initiated by a single, unusual event. The structural failure of part of the dam forming Lake Mildred, a small (ca. 32 ha), pri- vately owned recreational reservoir located about 3 km upstream from Merle Collins, allowed the entire contents of the smaller impoundment to be discharged down the stream channel into Merle Collins in the late summer of 1966. A noticeable increase in turbidity, which persisted for several days in Merle Collins Reservoir, gave evidence of the con- siderable amount of bottom sediment which accompanied this discharge. Since the C14 technique is sufficiently sensitive to detect photosynthetic response to nutrient additions even below the level of ordinary chemical detectability. it is probable that the increase in primary productivity noted in 1966 was in response to nutrients contained in these sediments. The increases in Merle Collins in 1967 and 1968, and the trend of in- creasing primary productivity in Folsom, seem to be associated only with the establishment in each reservoir of planktivorous fish species.

Novotna and Korinek (1966) noted quantitative differences in the phytoplankton of two backwaters of the River Elbe. In a backwater with fHi. tlie phytoplankton was more abundant than in one without fish. More recently, Hurlbert, Zedler, and Fairbanks (1972) demon-

26 \ CALIFORNIA FISH AND GAME

strated thai phytoplankton became extremely abundant after reduction of the zooplankton by mosquitofish (Gambusia affirm). Using data on phytoplankton, zooplankton, and juvenile sockeye salmon (Oncorhyn- chus nerkd) in throe lake systems, Brocksen, Davis, and Warren (1970) developed the conceptual framework which might explain such rela- tionships.

In a given ecosystem, as the phytoplankton biomass increases from a low level, primary productivity also increases to some maximum. Fur- ther increases in biomass result in decreasing primary productivity as the effects of competition for nutrients, shading, and the accumulation of inhibitory metabolites combine to lower the growth rates of the individual components of the phytoplankton. If grazing by zooplankton in the natural environment suppresses phytoplankton abundance below that level at which maximal productivity would occur, a reduction in zooplankton should be followed by an increase in primary productivity.

The apparent year-to-year increase in primary productivity in Fol- som Lake paralleled the establishment and development of a population of kokanee, a freshwater form of the sockeye salmon. Approximately 1.000,000 kokanee swimup fry (200-240/oz) were stocked in Folsom each spring from 196-1 to 1966, and a moderately successful fishery resulted from these introductions (E. D. Beland, Calif. Dep. Fish and Game. pers. comm.). Since kokanee feed primarily on zooplankton. the increasing biomass of ihis species may have resulted in a corresponding decrease in zooplankton abundance, hence the increases in primary productivity.

Similarly, in Merle Collins Eeservoir the first successful introduction of threadfm shad (Dorosoma petenense) was made in early 1967, when approximately 15,000 juveniles and 400 sexually mature adults were stocked. By September scattered spawning activity was observed and several schools of age 0 shad were encountered during electrofishing activities in December. During 1968, shad became a significant item in the diet of several species of game fishes and gill net catches of adult shad increased dramatically (K. A. Hashagen, Calif. Dep. Fish and Game. MS). Although threadfm shad are omnivorous, they do feed heavily on zooplankton (Kimsey. Hagy, and McCammon 1957; Gerdes and McConnell 1963; Miller 1967) ."'Thus, the biological impact of threadfm shad on primary productivity in Merle Collins could have been the same as that postulated for kokanee in Folsom.

It is also possible that grazing by these planktivorous fishes hastened biological cycling of nutrient material in the reservoir. This explana- tion would not seem as appropriate for Folsom, however, since the kokanee are largely excluded from the epilimnion by high temperatures and thus the nutrients present in their metabolic by-products would not be readily available to the bulk of the phytoplankton during that part of the year when increased primary productivity was noted. Even though information concerning changes in the zooplankton in Merle Collins and Folsom is lacking, it is apparent from the known changes in primary productivity and planktivore populations that a cause-and- effect relationship, involving the zooplankters as intermediate consum- ers, did exist.

PRIMARY PRODUCTIVITY 265

The fishery in Merle Collins Reservoir developed in a manner more easily ascribable to changes in the fish population structure (Hashagen MS) than to changes in primary productivity. Initial stocking in 1964 of largemouth bass (Micropterus salmoides) as both fry and adults gave rise to an extremely strong 1964 year class which apparently inhibited reproduction of all centrarchids for several years. Over 90% of the largemouth catch each year through 1967 was from this initial year class and these fish attained a mean length of only about 10 inches by 1967. The catch rate for this species reached a peak of about 0.36 fish per angler hour in 1967 and declined to about 0.13 by 1970 (Has- hagen MS).

Green sunfish (Lepomis cyanellus) comprised a minor component of the fishery from 1965 through 1967 but have since been supplanted by bluegill (L. macrochirus) and redear sunfish (L. microlophus) . Angler success for these species was less than 0.01 fish/hr in 1965 but reached almost 0.33 in 1970. The increased catch rate for these sunfishes was clearly associated with the declining abundance of bass (Hashagen MS).

Although fishery data are incomplete for a like period at Folsom Lake (von Geldern 1972), there is no indication that the observed changes in primary productivity have caused changes in the fishery there.

Various studies have attempted to relate primary productivity to fish yields or standing crop (McConnell 1963 ; Rupp and DeRoche 1965 ; Nicola and Borgeson 1970) with but limited success. This is not surprising since, although the C 14 method of measuring primary pro- ductivity allows us to estimate the rate at which photosynthetic activity introduces organic energy into an aquatic ecosystem, we are not yet able to describe, in any but the most general terms, how this photo- synthate is transferred through, and contributes to, the trophic struc- ture of that system (Goldman 1968&). This study indicates that in addition to the common practice of attempting to improve the efficiency of energy transfer by introducing suitable forage and predator species into a reservoir, it may also be possible to alter the rate at which energy enters the system at the primary level by manipulating fish popula- tions. And, even though it is not possible to say how trends in primary productivity would have compared in these two reservoirs without the planktivore introductions, it is obvious that declining primary produc- tivity need not be an inevitable consequence of initial reservoir aging.

ACKNOWLEDGMENTS

Robert R. Rawstron planned and initiated this study. C. R. Goldman was retained as a technical consultant by the Department and his ad- vice, encouragement, and technical assistance were invaluable. A succes- sion of project personnel, too numerous to mention individually, ac- complished the bulk of the field work. I particularly thank Linda Fry, Thomas J. Reece, and E. Ross Thompson for their unselfish contribu- tions in data reduction and summarv. Nanci Dong; drafted the figures. Charles Goldman, Leo Shapovalov, and Charles von Geldern, Jr. pro- vided helpful criticisms of the manuscript.

4—83609

266 CALIFORNIA FISH AND GAME

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Alell, Dana L.. and Charles K. Fisher. 1953. Creel census al Millerton Lake,

California. 1945 L952. Calif. Fish Came 39(4) :463^8 1. Bennett, George W. 1017. Fish management — a substitute for natural predation.

No. Amer. Wildl. Conf., Trans. 12:276-285. Brocksen, R. W.. G. E. Davis, and C. E. Warren. 1070. Analysis of trophic

processes on the basis of density-dependent functions. Marine Food Chains, Oliver

an,! Boyd, Edinburgh, p. 108-498. Ellis. M. M. 10.".7. Some fishery problems in impounded waters. Amer. Fish. Soc,

Trans. 60 :63-75. Gerdes, J. H., and Win. ,T. McConnell. 1963. Food habits and spawning of the

threadfin shad Dorosoma petenense (Giinther) in a small, desert impoundment.

Ariz. Acad. Sci., Jour., vol. 2, p. 113-116. Goldman, C. R. 1960. Primary productivity and limiting factors in three lakes

of the Alaska Peninsula. Ecol. Mono. 30:207-230. . 1963. The measurement of primary productivity and limiting factors in

freshwater with Carbon-14. In: M. S. Doty (ed.) Proceedings of the conference

on primary productivity measurement, marine and freshwater. U.S.A.E.C. TID-

763.".. Wash., D.C. . 196Sa. The use of absolute activity for eliminating serious errors iu the

measurement of primary productivity with C u. J. du Consiel Internatl. Explor. Mer. 32(21 : 172-179.

196S6. Aquatic primary production. Am. Zool. 8 : 31-42.

Goldman, C. P.. and R. C. Carter. 1965. An investigation by rapid carbon-14 bioassay of factors affecting the cultural eutrophication of Lake Tahoe, California- Nevada. Wat. Poll. Contr. Fed., J. 37:1044-1059.

Goldman. Charles R., and Robert G. Wetzel. 1963. A study of the primary pro- ductivity of Clear Lake, Lake County, California. Ecology 44(2) :283-294.

Hurlbert, Stuart II., Joy Zedler and Deborah Fairbanks. 1972. Ecosystem altera- tion by mosquitofish ( Gambusia affinis) predation. Science 175 (4022) :639-641.

Jenkins. Robert M. 1961. Reservoir fish management — progress and challenge. Sport Fishing Institute, Washington, D. C. 22 p.

Kimsey, J. B. 1958. Fisheries problems in impounded waters of California and the lower Colorado River, Amer. Fish. Soc, Trans. S7 : 319-332.

Kimsey. J. B., R. H. Hagy, and G. W. McCammon. 1957. Progress report on the Mississippi threadfin shad, Dorosoma petenense atchajaylae [sic], in the Colorado River for 1950. Calif. Dep. Fish and Game, Inland Fish. Admin. Rep. 57-23, 48 p. (mimeo. )

McConnell, William J. 1963. Primary productivity and fish harvest in a small desert impoundment. Amer. Fish. Soc, Trans. 92(1) :1— 12.

Miller. Robert V. 1967. Food of the threadfin shad. Dorosoma petenense, in Lake Chicot, Arkansas. Amer. Fish. Soc, Trans. 96(3) :243-246.

Nicola, Stephen J., and David P. Borgeson. 1970. The limnology and produc- tivity of three California coldwater reservoirs. Calif. Fish Game 56(1) :4-20.

Novotna, Marie, and Vladimir Kofinek. 1966. Effect of the fish stock on the quantity and species composition of the plankton of two backwaters, p. 297-322. In : Jaroslav Hrbacek (ed.) Hydrobiological studies, Czech. Acad. Sci., Acidemia, Prague.

Rawstron, Robert R. 1904. Limnology of Folsom Lake, 1901-03. Calif. Dep. Fish and Game, Inland Fish. Admin. Rep. 04-13, 9 p. (mimeo.)

Rawstron, Robert R. and Kenneth A. Hashagen, Jr. 1972. Mortality and sur- vival rates of tagged largemouth bass I \Iicropterus salmoides) at Merle Collins Reservoir. Calif. Fish Game 5S(3) : 221-230.

Rupp, Robert S., and Stuart E. DeRoche. 1905. Standing crops of fishes in three small lakes compared wih C11 estimates of net primary productivity. Amer. Fish. Soc. Trans. 94(1) :9-25.

Saunders, George W., F. B. Trama, and R. W. Backman. 1962. Evaluation of a modified C14 technique for shipboard estimation of photosynthesis in large lakes. Great Lakes Research Division Publication No. 8, Univ. Michigan, Ann Arbor.

Sorokin, Y. I. 1959. Determination of the photosynthetic productivity of phy- toplankton in water using Cu. Fiziol. Rast. 6:125-1:;:;.

PRIMARY PRODUCTIVITY 267

Steemann-Nielsen, E. 1952. The use of radioactive carbon (C14) for measuring organic production in the sea. J. du Consiel Internatl. Explor. Mer. 18:117-140.

Vollenweider, R. A. 19G5. Calculation models of photosynthesis-depth curves and some implications regarding day rate estimates in primary production measure- ments, p. 425-^27. In : C. R. Goldman (ed.) Primary productivity in aquatic environments. Mem. 1st. Ital. Idrobiol., 18 Suppl., Univ. California Press, Berkeley.

von Geldern, C. E., Jr. 1972. Angling quality at Folsom Lake, California, as determined by a roving creel census. Calif. Fish Game 58(2) :75-93.

Wetzel, Robert G. 19G5. Techniques and problems of primary productivity measurements in higher aquatic plants and periphyton. p. 249-2G7. In : C. R. Goldman (ed.) Primary productivity in aquatic environments. Mem. 1st. Ital. Idrobiol., IS Suppl., Univ. California Press, Berkeley.

Calif. Fish and Game, 58(4) : 26S-27G. 1972.

A MIDWATER TRAWL FOR THREADFIN SHAD, DOROSOMA PETENENSE]

C. E. VON GELDERN, JR.

Inland Fisheries Branch

California Department of Fish and Game

A midwater trawl designed to monitor threadfin shad abundance in restricted environments was developed at Lake Nacimiento, California, in 1966 and 1967. The trawl features hydrofoils and depressors which plane at 45° angles. It dives rapidly without supplementary weights or diving doors along the bridles or towing warps.

INTRODUCTION

Midwater trawls are of comparatively recent origin and have under- gone almost continual modification and refinement since World War II. Their initial use was restricted largely to the commercial exploitation of a few species of marine fishes, most notably herrings and cods (Par- rish 1959). With the introduction of echosounding devices, the success- ful use of midwater trawls in the ocean became much more widespread (Barraclough and Johnson 1956, MeNeely 1963, Sharfe 1964, and others), and they have also been adapted for use in large inland reservoirs (Ilouser and Dunn 1967).

In 1965, a study was undertaken at Lake Nacimiento, San Luis Obispo County, California, to evaluate the effects of an experimental introduction of threadfin shad on the existing warmwater fishery. Of primary concern was the need to develop an efficient method of sampling shad populations in the pelagic areas of the lake. Lake Nacimiento has been described previously by Aron Geldern (1971), and it need be stated here only that this impoundment covers 5,300 acres and has an un- usually irregular shoreline with an abundance of long arms, sunken islands, and peninsulas which create hazards to normal midwater trawl- ing operations. This report describes the development of a midwater trawl for sampling shad in this type of environment.

PRELIMINARY INVESTIGATIONS

A 24-ft commercial type fishing vessel powered with a 185-hp gasoline engine and fitted with midwater trawing gear and an echosounder be- came available to the project in late 1965. The trawling rig was of double warp design and featured a single flat wooden cpiarter door at each of four corners of the mouth of a 10 x 10 x 50-ft trawl. Accessory weighted diving doors were added at the junctions of 100-ft bridles and towing warps at times when it was necessary to fish deep. A double drum winch powered by a 6-hp gasoline engine was used to retrieve the net.

1 Accepted for publication February 1972. This work was performed as part of Dingell-Johnson Project California F-1S-R, "Experimental Reservoir Manage- ment", supported by Federal Aid to Fish Restoration funds.

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MID WATER TRAWL 269

Preliminary nighttime sampling with this equipment was conducted to obtain information on shad abundance and distribution. The results of this initial study revealed that (i) shad were extremely abundant and (ii) the horizontal and vertical distribution of shad was distinctly nonrandom. Further efforts were then centered on finding an efficient method of sampling extremely heterogeneous populations.

Taylor (1953) demonstrated that the efficiency of sampling hetero- geneous populations is improved by reducing the size of sample units and increasing the number of samples. This finding seemed appro- priate to the situation at Lake Nacimiento and I attempted to develop a simple trawl which would dive rapidly below fish concentrations and could be immediately retrieved. The sampling program, therefore, would be one in which the sample unit consists of two diagonal hauls (one down and one up) and which would equally sample all water depths containing shad.

The trawling apparatus initially made available to the project was not designed for rapid diving. In setting out or retrieving the net, it was necessary to stop the winch at the junctions of the towing warps and bridles to add or remove diving doors. This procedure inevitably resulted in a greater share of the sample collected near the surface. In addition, the flat quarter doors created considerable drag and were expensive and difficult to duplicate.

In June 1966, I visited the South Central Reservoir Investigations of the U. S. Bureau of Sport Fisheries and "Wildlife in Fayetteville, Arkansas, and observed midwater trawling operations at Bull Shoals Reservoir by Alfred Houser and his staff. Houser was using an 8 x 8 x 45-ft trawl, of single towing warp design, equipped with hydrofoils, depressors, and aluminum otterboards (Houser and Dunn 1967). The hydrofoils and depressors opened the net vertically while the otter- boards, suspended from 30-ft pennant lines, kept the net spread in a horizontal direction. This equipment functioned quite well on 45,400- acre Bull Shoals Reservoir, but was not well suited for trawling on small waters because of the diving characteristics of the net and the presence of otterboards on pennant lines. Nonetheless, I consider Houser 's trawl as the "model" from which I was able to develop equipment better suited for trawling on small waters.

My principal need was a trawl with the following basic features : (i) it must be of double warp design; (ii) it must dive rapidly; and (iii) it must not require the addition of supplementary otterboards or diving doors at any point along the bridles or towing warps. The following sections describe a midwater trawl having these general characteristics.

DESCRIPTION OF THE NET

The body of the net is composed of four tapered sections of equal dimensions. Each section contains seven panels with graduated mesh sizes ranging from 8 inches (stretch measure) in the forward panel to 1 inch in the rear or seventh panel. Sections are joined to four rib lines of f-inch polypropylene rope. The rib lines extend from the rear of the seventh panel to about 2£ ft forward of the mouth of the net, Wire rope thimbles are spliced into the forward ends of the rib lines

270

CALIFORNIA FISH AND GAME

for attachment to hydrofoils and depressors. Top, bottom, and side lines iif polypropylene rope encompass the mouth of the net. These also extend 2\ ft forward of the webbing and are spliced to wire rope thimbles. A gang of three thimbles is therefore present at each corner of (he nel mouth. Overall net dimensions are approximately 10 x 10 x 50 ft. A 7-11 coil end of !- and J-incI) nylon mesh with a single seam for each mesh size is attached to the rear of the seventh panel (Figure 1).

SPLICED /a-INCH WIRE ROPE THIMBLES

RIB, TOP, BOTTOM, AND SIDE LINES ALL 78-INCH POLYPROPYLENE ROPE AND EXTEND 2'6" BEYOND WEBBING AT MOUTH OF NET

I \ \ 7'3" FORWARD PANEL CONSTRUCTED WITH NO.9 BONDED NYLON THREAD

ALL MESH PANELS CUT SQUARE WITH EQUAL NUMBER OF MESHES AT FRONT AND REAR OF EACH PANEL

RIB, BOTTOM, AND SIDE LINES BEHIND PANELS SHOWN AS DASHED LINES

MESHES ARE LASHED TO RIB 5'9" LINES WITH '/,-INCH NYLON THREAD

ALL REFERENCES TO MESH SIZES ARE STRETCH MEASURE

m= MESHES

PANELS 2 THOUGH 7 CONSTRUCTED WITH NO. 6 BONDED NYLON THREAD

TOP, SIDE AND BOTTOM PANELS OF EQUAL DIMENSIONS AND CONTAIN EQUAL NUMBER OF MESHES

RIB LINES END AT REAR OF 7'" PANEL

THIS SECTION CONSTRUCTED WITH BONDE.D NYLON THREAD SIZE 20/9

FIGURE 1. Sectional view of midwater trawl.

MIDWATER TRAWL

DESCRIPTIONS OF HYDROFOILS AND DEPRESSORS

271

The hydrofoils are constructed of -J-inch aluminum alloy plate. A single curved sheet of 10 x 18-inch plate representing the planing surface is welded to the top of a 16-inch curved tapered vane. The vane

<x

I

INCH NYLON ROPE

AFT I5/

FOUR 3/8-HOLES, CENTERED V.-INCH FROM SIDES

all sheetmetal 7»-inch aluminum

HEIGHT OF VANE AT 2 INCH INTERVALS FROM FORE TO AFT 6%",6 W,b",53/B",4'i/e",35/8",AND 2%"

FIGURE 2. Top, side, front, end diagonal view of left hydrofoil.

FIGURE 3. Left hydrofoil showing hookup to thimble gang and bridle. Photograph by George Bruley.

272

CALIFORNIA FISH AND GAME

is situated along the mid-axis of the planing surface at a 90° angle. Three-eighths-ineh holes are punched near the trailing edge of the vane and the trailing corners of the shearing surface for attachment to the thimble gangs. An additional hole is punched in the lower fore corner of the vane for attachment to the bridles. A split 5tj x 3^-inch urethane seine float is lashed to the top of the inner edge of the planing surface so that the hydrofoils will ride at a 45° angle (Figures 2, 3, and 4). The hydrofoils, which weigh about 5 lb. each, are shackled to the upper thimble gangs as follows: (i) the inner edge of the planing surface to the top line, (ii) the trailing edge of the vane to the rib line, and (iii) the outer edge of the planing surface to the side line (Figures 3 and 4).

FIGURE 4. View of hydrofoils from Bruley.

afterdeck of research vessel. Photograph by George

One-eighth-inch steel plate is used to construct the depressors. The planing surface consists of a single flat 16 x L5-inch sheel of plate welded at 90° angles to three 16-inch vanes tapered at each end. The middle vane is located on the mid-axis of the planing surface and the others are situated 2 inches from the inner and outer edges. Three-

MIDWATER TRAWL

273

i

V

THREE 7B-INCH HOLE S , CENT E RE D '4-INCH FROM SIDES

%-INCH HOLE, CENTERED \ INCH FROM SIDES

4-t^4t4t-f®

/.-INCH STEEL

	1	T	r
	5"	,	5	*-^ '/.
T"lr TO	I	1	/
[P"-TW0 HEX NUTS

VlNCH E rE BOLT

2 X2 X 1-INCH LEAD BLOCKS

FIGURE 5. Top, side, front, and diagonal view of left depressor.

/e INCH STEEL PLATE

FIGURE 6. Left depressor showing hookup to thimble gang and bridle. Photograph by George Bruley.

274

CALIFORNIA FISH AND GAME

eighths-inch holes are punched in the trailing corners of the planing surface and the trailing edge of the center vane for attachment to the thimble gangs. A hole is also punched in the upper leading corner of the (niter vane for attachment to the bridles. The underside of the inner edge of the planing surface is fitted with a 14-inch steel rod of J -inch diameter which is used to contain 2 s 2 x 1-inch lead blocks bored with ',-iiich diameter holes. These weights cause the depressors to plane at 45° angles (Figures 5 and 6). The depressors, which weigh about 31 lb. each, are shackled to the bottom thimble gangs as follow?^ : (i) the inner edge of the planing surface to the bottom line, (ii) the trailing edge of the center vane to the rib line, and (iii) the outer edge of the planing surface to the side line (Figure C).

The hydrofoils and depressors are attached to f-inch thimbles spliced into the ends of lUO-ft, ^-inch diameter wire rope bridles by a chain and Miller swivel assembly (Figures 3 and 6). Type 2, 3-J-inch, Miller swivels are used for the hydrofoils and depressors. The bridles and -J-inch diameter galvanized wire rope towing warps are joined with

Type

, n;

•inch Miller swivels.

OPERATION OF THE TRAWL

The trawl is normally operated with a three-man crew. One man operates the boat, a second is responsible for the operation of the winch, and the third attends the net. Before setting out the net, the depressors and hydrofoils are arranged on the afterdeck for easy access (Figure

FIGURE 7. Afterdeck of research vessel showing arrangement of trawl and accessory equip- ment prior to setting out the net. Photograph by George Bruley.

MIDWATER TRAWL 275

7). The net is then cast out between the depressors while the boat is traveling about 1 mph. When the net has cleared the afterdeck, the depressors and hydrofoils are placed in the water. The boat is then ac- celerated to 3 mph and cable is let out to a point where the hydrofoils "bite". A very brief inspection of the assembly is then made to make certain that the net is fishing properly. The desired amount of towing warp is then let out.

Trawl retrieval procedures are conducted in reverse order. The towing warps and bridles are retrieved and the hydrofoils and de- pressors taken on board. The net is then pulled in directly over the stern. A speed of 3 mph is maintained until the depressors and hydro- foils approach the stern of the boat. The vessel is then decelerated to about 1 mph and maintained at that speed until the net is retrieved.

FISHiNG CHARACTERISTICS OF THE TRAWL

This trawl solved the sampling problems encountered at Lake Naci- miento and ultimately proved useful for detecting changes in threadfm shad abundance (von Geldern 1971). The addition of floats and weights to the hydrofoils and depressors which caused them to plane at 45° jingles eliminated any need for supplementary doors to spread the net. Tangling or fouling of the gear were never serious problems. This was attributed largely to the high degree of stability provided by the thimble gangs at each corner of the net mouth (Figure 8). The trawl also dived rapidly, reaching a depth of 55 ft when towed at 3 mph with 200 ft of towing warp out.

FIGURE 8. Schematic view of mouth of midwater trawl.

In order to test the potential diving speed range of this trawl design, depressors of -|-inch aluminum alloy plate with similar dimensions to those described previously were constructed and fitted with urethane

276

CALIFORNIA FISH AXD GAME

seine floats to achieve the proper 45° planing angle. When these doors were used, the trawl dived much less rapidly, reaching a depth of only 20 ft when towed at 3 mph with 200 ft of towing warp out (Table 1). It appears, therefore, that this trawl design can be modified to operate successfully in situations where rapid diving is not required.

TABLE 1 — Depth of Trawl When Fitted With Steel and Aluminum Depressors and Fished at 3 MPH

Type of depressor	Cable out (ft)	Fishing depth (ft)
Steel . . . _. .	100 150 200	21 38
	55
Aluminum	100 150 200	8 14
	20

ACKNOWLEDGMENTS

Edward E. Miller worked closely with me through all phases of the development of this trawl and the final product is a result of our joint efforts. As previously noted. I consider the trawl used by Alfred Houser of the Bureau of Sport Fisheries and "Wildlife as the "model" from which this equipment was developed. The hydrofoils described in this report are identical to the ones used by Houser in 1966 in all respects other than size, the placement of the seine floats, and the assembly for attachment to the net. Vincent Catania (deceased) supervised the con- struction of the nets and assisted the project in various other ways.

REFERENCES

Barraclough. W. E., and TV. TV. Johnson. 1956. A new mid-water trawl for herring. Fish. Res. Bd. Canada. Bull. no. 104. 25 p.

Houser. Alfred, and James E. Dunn. 1067. Estimating the size of threadfin shad populations in Bull Shoals Reservoir from midwater trawl catches. Amer. Fish. Soc. Trans. 96(2) :176-1S4.

McNeely, R. L. 1963. Development of the John M. Cobb pelagic trawl — a prog- ress report. Comm. Fish. Rev. 25(7) :17— 26.

Parrish, B. B. 1959. Midwater trawls and their operation, p. 333-343. In: Modern fishing gear of the world, ed. by Hilmar Kristjonsson. London, Fishing Xews (Books) Ltd.

Scbarfe, J. 1964. One-boat midwater trawling from Germany, p. 221-22S. In: Modern fishing gear of the world 2. London. Fishing Xews (Books) Ltd.

Taylor, Clyde C. 1953. Nature of variability in trawl catches. U. S. Fish TVildl. Serv., Fish. Bull. 54: 145-166.

von Geldern, C. E., Jr. 1971. Abundance and distribution of fingerling large- mouth bass. Micropterus salmoides, as determined by electrofishing at Lake Xaei- miento, California. Calif. Fish Game 57(4) :228-245.

I

Calif. Fish and Game, 58(4) : 277-284. 1972.

MORPHOLOGY AND VARIATION OF THE MODOC

SUCKER, CATOSTOMUS MICROPS RUTTER, WITH

NOTES ON FEEDING ADAPTATIONS1

MICHAEL MARTIN2

Department of Biological Sciences, Sacramento State College, Sacramento, California

Occurrence of the Modoc sucker, Catostomus microps Rutter, one of the most localized species in the freshwater fish fauna of California, is reported from the type locality at Rush Creek, Modoc County. A diag- nosis of the species with data from ten recently collected topotypes and a summary of the morphometric and meristic variation of this species is reported. Catostomus microps shows distinct adaptation to swift-stream conditions in the osteoiogical features of the oromandi- bular region and in the closure of the fontanelle of the neurocranium. It differs from all other members of the Pantosteus subgenus by the absence of lateral notches in the lips and by the possession of a silvery peritoneum. Ecological information is included in the discussion of this rare species.

INTRODUCTION

The native freshwater fish fauna of California contains several spe- cies of the genus Catostomus. These belong to the subfamily Catosto- minae, tribe Catostomini; there are also two other genera of catosto- mids: Xyrauchen and Chasmistes (Bailey 1970). These three genera occupy diverse ecological habitats in western North America. This study was initiated to examine more intensively one species, Catostomus mi- crops Kutter, which is particularly adapted to a mountain-stream habitat.

The geological history of northern California has been recently re- viewed by MacDonald (1966). The northeastern corner of California, included in the physiographic provinces of the Cascade Mountains and the Modoc Plateau, is characterized by wide-spread volcanism of recent origin and fault-block mountain ranges. This extreme disruption of the Modoc Plateau and subsequent isolation has had a significant effect on the fishes of the region, principally the catostomids as well as the cyprinids and cottids (Bailey and Bond 1963).

There have been few ecological studies of the catostomid species of California, but systematic studies have been presented by Hubbs et al. (1943), Miller (1959), Weisel (1960), and Smith (1966). Koelm (1969) reported that species of the subgenus Catostomus are generally in warmer lowland habitats, but they are occasionally found at higher elevations in lakes. Smith (1966) characterized the subgenus Panto- steus as primarily associated with rapidly flowing mountain streams.

The generalized ecological and morphological adaptations of cato- stomids (Smith 1966) and cyprinids (Brittan 1961) apparently are

1 Accepted for publication May 1972.

2 Present address: North American School of Conservation and Ecology, 1100 Claudina

Place, Anaheim, California 92S05.

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278

CALIFORNIA I-MSII \\1» GAM]

responsible for 1 lie widespread distribution of these two groups in western North America, including California. Smith (1966) and Koehn (1969) have reported that in western montane areas members of the subgenus Caiostomus occur sympatrically with species of the subgenus Pantosti us, and there are several records of breakdown of the reproduc- tive barriers (Hubbs et al. 1943).

Catostomus microps and the Sacramento sucker, Caiostomus occi- dentalis, occur allopatrically in the upper Pit Eiver system of Modoc County, California. Kutter (1908) recognized C. microps as a small- scaled relative of the more ubiquitous largescaled C. occidentalis.

MATERIALS AND METHODS

Specimens were collected from Rush Creek, Modoc County, Califor- nia. 6 miles east of Adin on U.S. Highway 299, T. 40 N., R, 9 E. This creek is the type locality for C. microps. Five extensive collections were made on Rush Creek on 4 April 1966, 8 October 1966, 26 December 1966, 12 March 1967, and 9 April 1967. '

T 42 N

T 41 N

T 40 N

T 39 N

R 7 E R 8 E

R 9 E

R 10 E

5 mi

FIGURE 1. Map of study area, showing area of collections on Rush Creek, Modoc County. California.

MODOC SUCKER 279

Rush Creek is a mountain stream, 5 miles in length, averaging 5 to 20 ft in width and attaining a maximum depth of 6 ft. Its headwaters are located on Horsehead Mountain and Hunters Ridge, Modoc County (Figure 1). From the upper Rush Creek Campground to the upper crossing of the Adin-Canby Highway (U.S. 299), the stream is exceed- ingly swift, and has a very steep gradient. Below the upper highway crossing, the stream passes through the rather long, gently sloping Rush Creek Valley, where the greatest numbers of C. microps were captured. The flora surrounding the headwaters of Rush Creek is a yellow pine forest, which changes abruptly to a northern juniper wood- land in Rush Creek Valley. A riparian flora (Axelrod 1944) is located on the margins of the creek throughout its lower course. The entire habitat which is suitable for C. microps on Rush Creek does not exceed 3 miles. There is little or no aquatic vegetation, but some leaf litter is present during the fall and winter months. The stream bottom is rock rubble, with limited sand and gravel areas.

From this habitat, 10 C. microps were collected utilizing a 6 ft X 20 ft minnow seine in the first four collections and electrofishing ap- paratus in the final collection. All fish were preserved in the field in 10% formalin, transferred to 40% isopropyl alcohol, and were placed in the Natural History collections of Sacramento State College.

Twenty characters were utilized and tabulated for each specimen. Morphological characters and meristic data compilation follow the methodology of Smith (1966). Counts and measurements were taken from the left side of adult and juvenile specimens, utilizing dividers and ruler to the nearest 0.1 mm. On specimens less than 50 mm sl, scale counts were not analyzed due to excessive variation (after Smith 1966). Age class determinations were made by counting the number of scale annul i.

MCDOC SUCKER

• .. ' - -' .-•..-...» 1 ~J& |j, |jfn famiffltejffi

FIGURE 2. The Modo: sucker, Caiostomus microps (female; 187 mm SL).

Catostomus microps. Rutter, 1D0S : 120-121 (original description) (Rush Creek. Modoc County). Jordan. Evermann. and Clark 1930 : 106 (streams of lava beds of California). Schultz 193G : 144 (upper Sacramento River and Goose Lake drainage). Shapovalov and Dill 1'JoO : 3S6 (check-

280

CALIFORNIA FISH AND GAME

list). Eddy 1957:78 (key). Shapovalov, Dill, and Cordone 1059 (check- list). Kimsey and Fisk 1960:467 (key). Bailey 1960:17 (common name). ."Miller 1961:384 (description of habitat). Bailey 1970:24 (common name).

DIAGNOSIS

A species of Catostomus characterized by the small eyes located in the middle of the head (Rutter 1908), whence the specific name. Standard lenirtk ranging to 190 mm. Lips moderate, with two rows of papillae evi- dent on the oral surface of the upper lip, but absent from the anterior face of the upper lip ; lateral notches at the juncture of the upper and lower lips faintly evident or absent ou either side; anterior medial papillae are enlarged ; medial notch in lower lip deep, separated from the cartilaginous sheath of the lower jaw by one row of papillae on the symphysis. Fronto- parietal fontanelle reduced in young specimens and almost obsolete in speci- mens over 150 mm SL. Scales small and regular ; lateral line scales, 80 to 89, modally 81; scales above lateral line. 15 to 17. modally 16; scales below lateral' line 9 to 12. modally 10. Dorsal rays 10 or 11. Pelvic axillary proc- ess absent. Caudal peduncle depth ranges from 9.0 to 10.0% SL (Table 1).

TABLE 1 — Measurements of Catostomus microps, Collected in Rush Creek, Modoc County, California. Proportions Are Expressed as Hundredths of Standard Length. Specimen 1 = SU 9277;* 2, 3 = SSC 149-2;t 4 to 6 = SSC 162-1; 7 to 1 1 = SSC 168-3.

	Specimen number
Measurements	1	2	3	4	5	6	7	8	9	10	11
Standard length (mm)	103 23 4 10 9 16 51 15	44 -"- 6 12 10 12 56 16	'.'7 25 4 12 10 15 50 15	187 23 4 10 9 15 48 15	24 2 11 10 16 50 16	97 24 5 11 9 15 49 15	48 28 6 14 10 13 53 16	99 24 4 13 9 10 51 15	76 26 5 13 10 12 52 15	76 28 5 13 11 16 52 15	87
	27
	4
	15
	10
Caudal peduncle length	15 52
Dorsal fin base	16

* SU = Stanford University.

f SSC = Sacramento State College.

VARIATION IN C. MICROPS

The following fin ray counts, scale counts, and gill raker counts are presented for the materials included in this report, expressed as the count, followed by the number of specimens with that count. Tn the case of pectoral and pelvic fins, the two numbers represent the counts of the left and right fins, respectively. Dorsal fin rays 10 (7). 11 (3) ; anal fin rays 7 (10), typical for American Catostomus: pectoral fin rays 15-15 (3), 16-16 (5),' 17-17 (2); pelvic fin rays 9-9 (7). 10-10 (3); caudal fin rays 18 (7). 19 (3). Scales in lateral line 80 (1), 81 (3), 82 (2), 84 (1), 85 (1). 87 (1). 89 (1) ; scales above lateral line 15 (3), 16 (6), 17(1) ; scales below lateral line 9 (1). 10 (6), 11 (2), 12 (1) ; scales around caudal peduncle 20 i 1 . 22 (2), 23 (2 . 25 (3), 26 (2) ; predorsal scales 45 (1). 46 (2 . 49 1 . 50 (3), 51 (2), 53 (1). Gill rakers 18 (1). 19 (1), 22 (3), 23 (2). 24 (1).. 25 (1), 26 (1).

The life colors of C. microps have not been recorded. The back varies from greenish-brown through bluish to deep grey and olive; the sides are lighter with light yellowish below; caudal, pelvic, and pectoral fins

MODOC SUCKER 281

are light yellowish orange. There are three characteristic dark spots along the sides in the region of the lateral line. The belly region is cream-colored to white.

Breeding coloration of C. microps is similar to that of the mountain sucker, C. platyrhynchus (Smith 1966). The male (76 mm sl) pattern consists of a red lateral stripe, which intensifies in 10% formalin and fades in isopropyl alcohol. This stripe originates behind the fleshy lobe of the opercular flap and extends to the origin of the last anal ray. The fins also become brightly colored, especially the mesial and distal parts of the pectoral fins, about the bases of the pelvic fins, and in the center part of the caudal fin. Breeding tubercles on the anal fin include : four small, three medium on the first element; two medium, one large on the second element; four large on the third element; two medium, one large on the fourth element; four large on the fifth element; three medium on the sixth element; and two small on the seventh element. Small tubercles are scattered over the dorsal region of the body, about half the way down the back. Tubercles are also scattered on the caudal and pectoral fins.

Age class determination and size indicated that C. microps matures in the second year. Nuptial males as small as 75 mm sl were collected, as well as second-year mature males up to 90 mm sl. Females are gen- erally larger than males; one third-year female measured 184 mm sl.

Osteological differences are well developed above the species level in the subgenera Pantosteus and Catostomus (Smith 1966) and provide a useful taxonomic tool for differentiation of these groups. One of the most significant adaptations in the evolution of the genus has been trophically oriented and, consequently, reflected in the osteological fea- tures of the jaw bones (Smith 1966). The mandible consists of four pairs of bones. The dentary is the largest of the bones ; it has a dorsal anterior gnathic ramus which has been modified for scraping the sub- strate. The dentary is not as ventrally deflected as that of species of the subgenus Pantosteus and has become decidedly reduced in C. microps (Figure 3). The dentary of C. occidentalis is more robust than the dentary of C. microps. The dentary of G. microps shows more ridging than C. occidentalis (Figure 3D), indicating an increased musculature in the oromandibular region of C. microps. This feature seems to be correlated with adaptation of the jaws as scrapers of the substrate as Smith (1966) found in species of the subgenus Pantosteus.

The ventral part of each mandibular bone is composed of a corono- meckelian, a retroarticular, and an angular (Figure 3A) ; all except the angular are thought to be derived from Meckel's cartilage (Har- rington 1955). The configuration of the ventral mandible is similar in large and smallscale species of the Pit Eiver drainage with three ex- ceptions. First, as viewed mesially, the coronomeckelian is reduced and slightly serrate on the dorsal margin in C. occidentalis (Figure 3B), while C. microps lacks these slight serrations. Secondly, the anterior dorsal facet of the dentary is reduced in C. occidentalis, and this area is enlarged in C. microps. Thirdly, the mental foramen is pronounced in C. occidentalis and slightly reduced in C. microps.

The mandible of the Tahoe sucker, C. tahoensis, (Figure 3A) is spe- cialized toward the small stream type as shown by C. microps. The

5—83609

0S0

CALIFORNIA FISH AND GAME

rt.rononu'ckolian is slightly reduced, and the anterior dorsal facet of the dentary is enlarged as in C. microps.

10

FIGURE 3. Mesial view of the left mandibles of (A) Cafosfomus tahoensis (Lahontan Basin) from Smith, 1966; (B) Cafosfomus occidentalis (Goose Lake Basin), 84 mm SL; (C) Cafosfomus ocadenfa/i's (upper Pit River Basin), 101 mm SL; (D) Cafosfomus microps (Rush Creek), 97 mm SL. The bones of the lower jaw are the angular (AN), the coronomeckelian (CM), the dentary (D), and the retroarticular (RA).

Smith (1966) suggested that although the function of the fontanelle of the neurocranium is not known, its closure may be involved with an increase in opercular musculature and the reduction in the size of the pterotic, C. microps is definitely a small riffle type of sucker, although, heretofore, it has been considered as a member of the subgenus Catosto- mus. C. microps is separable from all Pantosteus species by the absence of lateral notches at the junction of the upper and lower lips and the lack of a black peritoneum. The lip and jaw modifications, other than the lateral notches, of C. microps show parallel adaptation with Pan- tosteus species.

FEEDING ADAPTATIONS

In addition to the extreme specialization and orientation of the oro- mandibular region, there is a behavioral modification for suctorial

MODOC SUCKER 283

feeding and other general morphological adaptations for bottom dwell- ing.

Juvenile and adult behavior was qualitatively observed: juveniles (15 to 50 mm sl) tend to remain in the shallows of large pools, free swimming above the substrate. Adult suckers remain mostly on the bottom or close to it. Adults in aquaria remained at the bottom, resting either on the pelvic fins and folded anal fin or with their ventral surface contacting the substrate. While resting on their ventral surface, the dorsal fin Avas generally elevated, with the paired fins extended to sup- port the body. Suckers do not actively forage during daylight hours unless disturbed. Feeding and foraging as well as migration usually occur nocturnally (La Rivers 1962 on Catostomus tahocnsis). The ventral surface of slow-stream inhabiting Catostomus is widened and flattened from the tip of the snout to the anal fin base. In contrast to these forms, C. microps and several members of the subgenus Pantosteus have a more rounded ventral surface, reflecting the necessity for more active swimming. The anal fin has a short base which can be folded so as not to interfere with substrate contact.

The sensory apparatus of C. microps is specialized toward the bottom feeding habit. The presence of large numbers of papillae and taste buds in Catostomus (Stewart 1926), the extremely limited eyesight, and the position of the eyes in the head are all indicators of the bottom feeding habits of this species.

MATERIAL EXAMINED

Catostomus microps. SU 9277 (1, 103) paratype ; Rush Creek, near Ash Creek. Pit River Drainage. T. 40 N., R. 9 E. ; September 1, 1898, Rutter and Chamberlain. SSC 140-2 (2, 44-97) ; Rush Creek, 6 miles E Adin, T. 40 N., R. 9 E., see. 35; October 8, 1066, M. Martin, R. B. Bury, J. Erode. SSC 162-1 (3, 80-187) ; Rush Creek, 6 miles E Adin, T. 40 N., R. 0 E., sec. 35; December 26, 1066, M. and G. A. Martin. Jr. 88C 168-3 (5, 48-00) ; Rush Creek near mouth of Johnson Creek, 7 miles E Adin. T. 40 N., R. 9 E., sec. 24; April 9, 1967, M. Martin, R. B. Bury, D. Kritsky, and R. Armstrong.

"■j >

ACKNOWLEDGMENTS

I wish to express my sincere appreciation to Martin R. Brittan for his constant encouragement, guidance, and friendship during my studies at Sacramento State College. I also sincerely appreciate the courtesy of those persons responsible for museum collections in making animals available for study : CAS, California Academy of Sciences ( W. I. Follett, L. J. Dempster, and J. Hopkirk) ; Stanford University- Division of Systematic Biology (now at CAS) (W. C. Freihofer and E. H. Neil) ; and Sacramento State College — Museum of Natural His- tory (M. R. Brittan). I thank those fellow graduate students of Sacra- mento State College, R. Armstrong, J. M. Brode, R. B. Bury, D. Krit- sky, and my brother, George A. Martin, Jr., who spent many uncomfortable days collecting. Facilities and equipment were provided by Sacramento State College. Without the constant aid and encourage- ment of my wife, Demi, this study would not have been completed.

SUMMARY

The Modoc sucker, Catostomus microps, occurs as an isolated popula- tion in Rush Creek, Modoc County, California which survives despite

284 CALIFORNIA FISH AND GAME

disruption or destruction of most of its available habitat, primarily through stream channel change. The species apparently survives in marginal undisturbed areas. The synonymy, nomenclature, and diag- nosis of the species is given. Variation in meristic and morphometric characters are given for 10 individuals captured during 1966 and 1967. The breeding coloration of C. microps is described, and observations of its feeding habits discussed.

REFERENCES

Axelrod, D. I. 1044. The Alturas flora (California). In R. W. Chaney fed.)

Pliocene floras of California and Oregon. Carnegie Inst. Washington Publ. (558). Bailey, R. M. (editor) 1960. A list of common and scientific names of fishes

from the United States and Canada. Spec. Publ. No. 2, Amer. Fish. Soc, Wash- ington, B.C. 102 p. . 1070. A list of common and scientific names of fishes from the United

States and Canada. Spec. Publ. No. 6, Amer. Fish. Soc, Washington, D.C. 150 p. Bailey, R. M. and C. E. Bond. 1903. Four new species of freshwater sculpins,

genus Cotlus, from western North America. Occ. Pap. Mus. Zool., Univ. Mich.

634 :l-27. Brittan, M. R. 1961. Adaptive radiation in Asiatic cyprinid fishes, and their

comparison with forms from other areas. Froe. 9th Pac. Sci. Congress 10 ; 18-31. Eddy, S. 1957. How to know the freshwater fishes. William C. Brown Company,

Dubuque. 253 p. Harrington, R. W. 1955. The osteocranium of the American cyprinid fish, Notro-

pis bifrenatus, with an annotated synonymy of teleost skull bones. Copeia 1955

(4) : 267-290. Hubbs, C. L., L. C. Hubbs, and R. E. Johnson. 1943. Hybridization in nature

between species of catostomid fishes. Cont. Lab. Vert. Biol., Univ. Mich. 22:1-76. Kimsey, J. B. and L. O. Fisk. 1960. Keys to the freshwater and anadromous

fishes of California. Calif. Fish Game 46 (4) :453^79. Koehn, R. K. 1969. Hemoglobins of fishes of the genus Catostomus in western

North America. Copeia 1969 (1) :21-30. Jordan, D. S., B. W. Evermann, and H. W. Clark. 1930. Checklist of the fishes

and fish-like vertebrates of North and Middle America north of the northern

boundary of Venezuela and Colombia. Rep. U.S. Comm. Fish 1928 :l-670. La Rivers, I. 1962. Fishes and fisheries of Nevada. Nevada State Fish Game

Comm. 782 p. MacDonakl. G. A. 1966. Geology of the Cascade Range and Modoc Plateau, p.

65-96. In Geology of northern California. Calif. Div. Mines Geology Bull., (190). Miller, R. R. 1959. Origin and affinities of the freshwater fish fauna of western

North America, p. 1S7-222. In Zoogeography. Amer. Assoc. Adv. Sci. Publ., (51). . 1961. Man and the changing fish fauna of the American Southwest. Pap.

Mich. Acad. Sci., Arts, and Letters 46 (1960) :365-404. Rutter, C. 1908. The fishes of the Sacramento-San Joaquin basin, with a study

of their distribution and variation. Bull. U.S. Bur. Fish. 27 (1907) :103-152. Schultz, L. P. 1936. Kevs to the fishes of Washington, Oregon, and closely

adjoining regions. Univ. Wash., Publ. Biol. 2 (4) :103-288. Shapovalov, L. and W. A. Dill. 1950. A checklist of the freshwater and anadro- mous fishes of California. Calif. Fish Game 36 (4) :382-391. Shapovalov, L., W. A. Dill, and A. J. Cordone. 1959. A revised checklist of the

freshwater and anadromous fishes of California. Calif. Fish Game 45 (3) :159-180. Smith, G. R. 1966. Distribution and evolution of the North American catostomid

fishes of the subgenus Pantosteus, genus Caiostomus. Misc. Publ. Mus. Zool.,

Univ. Mich. 129 : 1-132. Stewart, N. H. 1926. Development, growth, and food habits of the white sucker,

Catostomus commersonii Lesueur. Bull. U.S. Bur. Fish. 42:147-184. Weisel, G. R. 1960. The osteocranium of the catostomid fish, Catostomus macro-

cheilus. A study in adaptation and natural relationship. J. Morph. 106 (1) :109-

129.

Calif. Fish and Game, 58(4) : 285-290. 1972.

CONTRIBUTIONS TO THE LIFE HISTORY OF THE PIUTE SCULPIN IN SAGEHEN CREEK, CALIFORNIA1

ALBERT C. JONES

Southeast Fisheries Center

National Marine Fisheries Service

Miami, Florida

The Piute sculpin, Cottus beldingi Eigenmann and Eigenmann, is the dominant fish by number and weight in Sagehen Creek, a mountain stream on the east slope of the Sierra Nevada. Sculpins are at their greatest density in the middle part of the creek where they, along with brook trout (Salvelinus fontinalis) and rainbow trout (Sa/mo gairdneri), find good foraging for bottom dwelling aquatic insect larvae. The num- bers of sculpins are low in the precipitous headwaters of Sagehen Creek and also low in the lower reaches of the stream, which are frequented by their chief predator, the brown trout (Sa/mo trutta), and by other fishes. Sculpins in Sagehen Creek and Lake Tahoe exhibit minor differ- ences in growth and reproduction but appear to occupy a similar ecolog- ical niche in the two areas.

INTRODUCTION

The Piute sculpin lives in lakes and streams of the Lahontan and Columbia Kiver Basins of the western United States. Baker and Cordone (1969) and Ebert and Summerfelt (1969) described the biology of C. beldingi living in Lake Tahoe, an oligotrophia mountain lake. This re- port concerns the Piute sculpin in Sagehen Creek, a nearby mountain stream, and compares the biology of stream and lacustrine dwellers of this species.

Sagehen Creek is a spring-fed stream that is tributary to the Little Truckee Kiver, itself tributary to the Truckee Kiver draining Lake Tahoe. The creek rises in the Sierra Nevada at an altitude of 7,400 ft, flows through a watershed approximately 19.2 square miles in area, and after about 13 miles enters the Little Truckee River at 5,800 ft elevation. Climatic conditions are boreal. Between 1954 and 1961, mini- mum daily flows in Sagehen Creek in fall ranged from 1.0 to 2.6 cfs and momentary maximum daily flows in spring ranged from 27 to 212 cfs (Gard and Flittner MS).

ABUNDANCE

The Piute sculpin is the most common fish in Sagehen Creek and, by number and weight, is a significant part of the stream ecosystem. The population density (number of fish per acre) of sculpins in Sage- hen Creek was estimated from the number of fish collected from 10 short sections of stream (Table 1). The sections, numbered from I (up- stream) to X (downstream), totaled approximately 2,000 ft in length and were located at approximately 1-mile intervals along the course of the stream. The water flow was diverted from each section, the pools in the section drained with a pump, and the fish captured. Later the fish were returned to the stream, except for samples retained for study. Details of the collecting methods are given by Flittner (1953).

1 Accepted for publication March 1972.

(285 )

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PIUTE SCULPIN 287

Seulpins were most abundant in middle Sagehen Creek, which in- eluded sections IV- VIII, where the stream gradient was intermediate, the bottom consisted of gravel-rubble and brook trout, rainbow trout, and brown trout were the only other fish present. (A few individuals of other species were present in section VIII as noted below.) Seulpins were absent or scarce in upper Sagehen Creek (sections I-III) , which was the precipitous, boulder-strewn headwaters of the stream inhabited primarily by brook trout. Seulpins also were scarce in lower Sagehen Creek, which included sections IX and X, where the stream had a slight gradient, a bottom of gravel, mud and clay, and a fish population that included in addition to the 3 trouts, Tahoe suckers (Catostomus tahoen- sis), mountain suckers (C platyrhynchits), Lahontan redsides (Rieliardsonhis egregius), speckled dace (Rhinichthys osculus), and mountain whitefish (Prosopium ivilliamsoiii). The non-salmonid fishes in sections IX and X also occurred in reduced numbers in section VIII (Card and Flittner, MS).

The apparent success of seulpins in middle Sagehen Creek may be due to a combination of favorable conditions; gravel riffle areas which provide cover for the seulpins and a substrate for their forage (bottom dwelling aquatic insect larvae) and the general absence or scarcity of predaceous brown trout. In Lake Tahoe seulpins ranged from the lit- toral zone to 700 ft in depth, but similarly were most common in the rubble-boulder areas of intermediate depth which offered protection from lake trout (Salvelinus namaycush), the sculpin 's chief predator in the lake (Baker and Cordone 1969).

AGE COMPOSITION

Seulpins in Sagehen Creek reach 5 years of age, determined by count- ing annuli in otoliths. Calculated age distributions of the sculpin popu- lations in sections I-X are shown in Table 2. In 1953, 1-year-old fish (the 1952 year class) made up 82% of the total number of seulpins sampled, but in 1952, 1-year-old fish made up only 11% of the total number of seulpins. Seegrist and Gard (in press) reported that the severe spring floods in 1952 decimated the eggs of spring-spawning rainbow trout. Our observations indicate that the high-water conditions present in the summer of 1952 may have benefited the survival of young-of-the-year seulpins. In sections IX and X, 1-year-old seulpins were less numerous than 2-year-olds in both 1952 and 1953; this sug- gests that reproduction in lower Sagehen Creek is relatively unsuccess- ful and that the sculpin population there is maintained partly by mi- gration into the area.

REPRODUCTION

Piute seulpins spawn a small number of eggs which are relatively large in size. The fecundity of 70 fish from 65 to 86 mm tl ranged from 77 to 235 (average 132). The linear regression of fecundity on total length of fish was Y = --151.59 + 3.81X, r = 0.69, P < 0.01. Eggs taken from the ovaries of several unspawned females collected during the spawning season averaged 2.54 mm in diameter and water- hardened eggs collected from the two nests averaged 2.90 and 2.97 mm.

288 CALIFORNIA PISH AND GAME

Measurements of the eggs were made after the fish had been preserved initially in 10% formalin and then transferred to 60% ethyl alcohol.

The spawning season of sculpins in Sagehen Creek in 1953 was short. At section VI the first spawned-out female was collected on June 2 and by June 8 all females collected in this section had spawned. Males also appeared to be in spawning condition for only a short time. Males from which milt could be extruded by applying pressure to the abdo- men were collected in section VI only from June 1 to 8. The average daily maximum and minimum water temperatures at this location were 52.2 F and 38.3 F (May 25-31) and 57.0 F and 40.1 F (June 1-7). Water temperatures are lower upstream from section VI and higher downstream (Gard and Flittner MS), so that if spawning time is dependent on water temperature, spawning probably occurs at dif- ferent times in different sections of the creek.

Sculpins in Lake Tahoe spawned primarily in May and June (Ebert and Summerfelt 1969). The report by Miller (1951) of ripe female sculpins in lake trout stomachs (which are inhabitants of the deeper, cooler water) as late as August 28 suggests that the spawning season in Lake Tahoe also varies in different parts of the lake in relation to temperature. The average fecundity of Lake Tahoe sculpins (123) (Ebert and Summerfelt 1969) was close to that of sculpins in Sagehen Creek.

The two sculpin nests observed in the spawning season of 1953 were located in riffle areas of the stream, under rocks 8 to 12 inches in diameter and in water 6 to 10 inches deep. Egg clusters were attached to the undersurface of the rocks. Each female sculpin apparently spawns only once per year and presumably deposits a single egg cluster, since the number of eggs in the two nests (122 and 160) was close to the average fecundity.

In the month previous to spawning, female sculpins contained two distinct size groups of ovarian eggs. The smaller, immature eggs were less than 0.60 mm in diameter and the mature eggs were 1.55 to 3.55 mm in diameter. ( The preserved eggs were usually misshapen ; and, as a result, their diameters had a greater than normal range.) After spawning, only immature eggs were present ; enlargement of the ova preparatory to the next season's spawning was not noticeable until October.

GROWTH

Sculpins in Sagehen Creek grew primarily from May to October. Age group 0 sculpins sampled in section VI increased from 12.0 mm in August to 25.4 mm in October. Growth slowed after October; in January the average length was only 24.5 mm. Growth from January to May is probably also slow, since age group I fish collected in January 1954 (24.5 mm) were about the same size as age group I fish collected in May 1953 (24.8 mm). The length of age group I fish increased from 24.8 mm in May to 54.4 mm in October. By January the I age group had increased to only 58.0 mm. Older sculpins also increased most in length from May to October but little from October to May.

The increased growth rate of Sagehen Creek sculpins observed in spring and summer corresponds with the higher water temperatures in

PIUTE SCULPIN 289

these months; the mean daily maximum water temperature was higher than 50 F only in the months of May through October (Needham and Jones 1959). Ebert and Summerfelt (1969) concluded that the Piute sculpin in Lake Tahoe grows primarily in the spring and early summer and found a larger volume of food in their stomachs in spring and summer compared to fall and winter. Sculpins in Lake Tahoe were generally larger at a given age than those in Sagehen Creek, especially for younger age groups. No records of water temperature were avail- able to interpret the seasonal growth patterns in Lake Tahoe as com- pared to Sagehen Creek.

The sculpins collected in August 1953 in the downstream sections of Sagehen Creek were larger than those in the upstream sections. This difference was probably because during most of the year water temper- atures are higher in the downstream portion of the stream and as a result the growth rate is more rapid. Card and Seegrist (in press) also found increased growth of brook, rainbow, and brown trout at lower elevations of Sagehen Creek.

Male sculpins grow faster than females. The difference in growth rate between the sexes was apparent in age group I individuals col- lected in August, when sexual differentiation of the gonads was first apparent from gross examination. The growth curve for males was L = 44.4 t °-5415, where L = total length (mm) and t — age (years). The growth curve for females was L = 43.4 t °-4793. Males apparently live longer than females; out of 12, 5-year-old fish collected, 11 were males.

The relationship between length (mm) and weight (g) of Sagehen Creek sculpins was W = 8.8356 X KHL3-10817.

DISCUSSION

The Piute sculpin is the most abundant fish in Sagehen Creek and is the dominant species in the gravel-rubble parts of the stream where the gradient is intermediate. Brook trout and rainbow trout coexist with sculpins throughout the stream; brown trout, which inhabit pri- marily the lower sections of the creek, are their most important preda- tor. Predation, food supply, and stream flow may be factors which limit population size. Growth and spawning time of sulpins in Sagehen Creek are related to the seasonal cycle of water temperature and are different in Sagehen Creek than in Lake Tahoe. Except for minor differences in biology, the Piute sculpin appears to occupy a similar ecological niche in the two areas.

JoJ

ACKNOWLEDGMENTS

The University of California Sagehen Creek Wildlife and Fisheries Research Station was initiated by Dr. Paul E.. Needham who directed its activities until his death in 1964. This paper is dedicated to Dr. Needham.

REFERENCES

Baker, Phillip M., and Almo J. Cordone. 1969. Distribution, size composition, and relative abundance of the Piute sculpin, G'ottus beldingii Eigenmann and Eigen- mann, in Lake Tahoe. Calif. Fish Game 55 (4) : 285-297.

290 CALIFORNIA FISH AXD GAME

Ebert. Verlyn W., and Robert C. Summerfelt. 1969. Contributions to the life history of the Piute sculpin, Coitus beldingii Eigenmann and Eigenmann, in Lake Tahoe. Calif. Fish Game 55 (2) : 100-120.

Flittner. (llenn Anion. 1953. The composition and distribution of the fish popu- lations in Sagehen Creek, Nevada-Sierra counties. MA Thesis (Zoology), Univer- sity of California, Berkeley, 150 p.

Gard. R., and D. W. Seegrist. (In press). Abundance and harvest of trout in Sagehen Creek. California. Amer. Fish. Soc, Trans.

Gard, R., and (llenn A. Flittner. (MS). A ten-year study of distribution and abundance of fishes in Sagehen Creek, California. MS, Univ. of California, School of Forestry and Conservation.

Miller, Richard Gordon. 1951. The natural history of Lake Tahoe fishes. Ph. D. Thesis. Stanford University, 100 p.

Needham, Paul R., and Albert C. Jones. 1959. Flow, temperature, solar radia- tion, and ice in relation to activities of fishes in Sagehen Creek, California. Ecology 40 (3) : 465-474.

Seegrist, 1 ». W., and R. Gard. (In press). Effects of floods on trout in Sagehen Creek, California. Amer. Fish. Soc. Trans.

Calif. Fish and Game, 58(4) : 291-205. 1972.

THE EFFECTS OF DIESEL FUEL ON A STREAM FAUNA1

R. BRUCE BURY

Museum of Vertebrate Zoology

University of California, Berkeley 94720

The spillage of approximately 2,000 gallons of diesel fuel into Hay- fork Creek, California, resulted in a large kill of invertebrates, fishes, and other life. Subsequent effects on the stream fauna are discussed. This study examines the increasing threat of pollution in remote areas due to the transportation of petrochemicals.

INTRODUCTION

"Water pollution by petroleum products is a serious environmental problem, since oily substances contain toxic components and, in general, are stable compounds that can remain in an ecosystem for a relatively long time. Oil spills have caused widespread detrimental effects in California waters and elsewhere (McCaull 1969; Mitchell, et al. 1970; Blumer, et al. 1971; and Straughan 1971).

McKee (1956) reported that petroleum products can be detrimental to aquatic organisms in the following ways: (i) free oil and emulsions may act on the epithelial surfaces of fish, thereby interfering with respiration, or may coat and destroy algae and plankton, which remove sources of food; (ii) oily substances that settle to the bottom may coat and destroy benthal organisms, and interfere with spawning areas; (iii) soluble and emulsified material may be ingested by fish and thereby taint the flavor of the flesh, or water-soluble parts may have a direct toxic action on aquatic life; (iv) organic materials may deoxygenate the water sufficiently to kill fish; and (v) heavy coatings of free oil on the surface may interfere with reaeration and photosynthesis.

Distilled petroleum substances are immediately toxic to animal life. Gutsell (1921) found gasoline had a toxic effect on rainbow trout (Salmo gairdneri) at about 100 mg/liter. McKee and Wolf (1963) reported that agitated solutions of automobile gasoline at a concentra- tion of 100 mg/liter and jet aviation fuel at 500 mg/liter is lethal to fingerling salmon (Oncorhynchus sp.). Diesel fuel is acutely toxic to rainbow trout within the range of 350 to 1,000 mg/liter (Richard Hansen, pers. comm.).

There are relatively few documented cases of oil pollution in fresh- waters. In fact, Wilbur (1969) stated that there is such limited in- formation on the effects of oily wastes in water on livestock and wildlife that any extended discussion would be futile. Swift et al. (1969) sur- veyed the literature on the biological and ecological effects following an oil spillage, noting that while some information is available on the dam- age that can occur, few quantitative and coherent data are available to assess past incidents or to predict potential effects in the future.

1 Accepted for publication March 1972.

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292 CALIFORNIA FISH AND GAME

The present study reports the detrimental effects of diesel fuel, a mod- erately toxic substance, on an unspoiled freshwater stream.

On July 28, 1970, the rear tank section fell off a truck on a sharp curve along U. S. Forest Service Road 2N01. The accident occurred about 0.5 miles upstream from the 'fish ladders,' Hayfork Creek, a tributary of the Trinity River, Trinity County, California (about 7 air miles SSE of the town of Hayfork). The 4,000-gallon tank, reported to be about half full of diesel fuel, burst when it rolled down a steep canyon. Some of the fuel evaporated or soaked into the ground, but about 2.000 Gallons entered the creek.

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MATERIALS AND METHODS

A survey of the biological effects of the diesel fuel spill in the creek was conducted from 1 to 2.5 miles downstream from the site of the accident because the pre-spill conditions in this area were well known. Field studies had been carried out along this part of the creek during the summers 1968 to 1969 and in June and July, 1970 (Bury 1972). The study area consisted of 36 pools varying in size from 5 to 50 yards long, mostly 10 to 20 yards, and 5 to 10 yards wide. The pools are 3 to 12 ft deep in the summer and are connected by long, shallow riffles 0.5 to 1 ft deep. For comparison of the effects of the diesel fuel on the stream fauna, the surveyed area was divided into 10 equal parts with each section 800 ft in length. Dead animals were counted in the study area from August 1 through 5 and then periodically until mid-Sep- tember.

EFFECTS ON THE FAUNA

The diesel fuel entered the study area about 36 hr after the accident. Initially a thin film of fuel extended entirely across the surface of the creek. The first effects were observed on the morning of July 31, 1970, when the normally clear waters turned a murky, brown color with visibility less than 1 ft and small droplets of fuel floated on the sur- face.

Most animals were adversely affected 1 to 4 days after the fuel entered the study area. Thousands of aquatic insects perished, espe- cially water boatmen (Corixidae), belostamatid water bugs, water striders (Gerridae), adult and larval diving beetles (Dytiscidae), may- fly nymphs (Ephemeroptera), and dragonfly and damselfly nymphs (Odonata). Many crayfish (Astacus sp.) were actively moving in the creek during daylight hours, a condition which had not been noticed in previous years. Ten crayfish were found dead. Hundreds of aquatic leeches (Class Hirudinae) and freshwater planarians (Planariidae) were killed.

Over 2,500 fishes were killed, including about 1,000 lamprey ammo- coetes (Entosphenus tridcntatus). 688 small-sealed suckers (Catostomus rimiculus) , 75 speckled dace (Rkinichthys osculus), and 849 rainbow trout (Table 1). Further, several fishes were seen near the surface, mouths gaping and then slowly sinking into the murky water. In one pool I observed a 7-inch trout moving slowly along the bottom upside down, and a 12-inch fish swimming on its side near the surface. Other trout in the pool remained almost motionless near the bottom, fre- quently gaping widely.

EFFECTS OF DIESEL FUEL

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Tadpoles and partly metamorphosized individuals of the foothill yel- low-legged frog (Edna boylei) were killed in large numbers. Xo adult frogs were found dead. Thirty-six western aquatic garter snakes Thamnophis couchi were killed. Several snakes were seen that ap- peared to be noticeably sluggish in their movements.

Subsequent to the initial toxic impact on the stream other effects were "1 -i rvcd. Large quantities of dead animals and algae (Spirogyra sp.. Cladophora sp.. Zygnema sp.) sank to the bottom or formed floating mats. Some of the surface masses were 2 to 4 inches thick and covered several square yards. A log across the surface of one pool caused ac- cumulation of floating material that covered an area 20 ft wide and -Id ft long. Loose aggregations of organic matter accumulated on the bottom where there was little current, and in places formed a slurry 6 to 12 inches deep. The organic matter putrefied rapidly and formed a layer of scum on the bottom of the creek. Most of the diesel fuel was flushed out of the study area 3 weeks after the spill, but some fuel remained trapped in the accumulations of dead organic matter and small slicks of fuel were observed until mid-September when observa- tions ended.

On August 8. a dead common merganser Mergus merganser) was found along the creek and its feathers were in disarray and smelled of diesel. On September 10. a pond turtle (Clemmys marmorata) measur- ing 5 inches in shell length was found dead on the bottom of a pool. Two young ones, alive, but in poor condition, were found on the shores of other pools. The eyes and necks of all these turtles were swollen. Movements of both young turtles were uncoordinated and they were unable to either swim or sink. Also. 30 pond turtles captured in early September had sloughed off pieces of epidermis on their appendages. and their necks and eyes were swollen.

DISCUSSION AND CONCLUSIONS

The large die-off of animals was a direct result of the diesel fuel pol- lution. Only rarely was a dead animal found in the creek during prior studies, and usually these were due to predation. The toxicity of the fuel killed most animals on contact and, later, caused other detrimental effects to the stream fauna due to large quantities of putrefying organic matter.

Many fishes displayed unusual behavior due to irritating or immobi- lizing effects of the diesel fuel. Adult frogs were not killed, and their survival is related to their mode of life. Frogs usually rest along banks out of water and feed principally on live insects. Hundreds of tadpoles perished since they were directly exposed to the fuel in the water and. perhaps, ingested tainted algae. Garter snakes probably died because they regularly swim in the water and prey on tadpoles and fish. Ex- posure to the fuel and ingestion of food contaminated with fuel may have killed the pond turtle and common merganser.

There was a heavy concentration and prolonged exposure to the fuel in the upstream parts of the study area, and the mortalities were greater than in sections farther downstream where the fuel was diluted. evaporated, or dispersed sufficiently To have a reduced impact on the stream fauna. There were 2,952 dead vertebrates found in sections 1

EFFECTS OF DIESEL FUEL 295

through 5, and 1,517 in the downstream sections 6 through 10 (Table 1 ) . Although no dead organisms "were found farther than 5 miles down- stream from the site of the spillage, chronic toxicity and other sub- lethal effects may have extended many miles along the creek.

Blumer et al. (1971) reported that hydrocarbons taken up into the fat and flesh of fish and shellfish are not removed by excretion or by internal metabolic processes, and that these substances remain in the animals for long periods of time, possibly for their entire lives. They state that crude oil and oil products are persistent poisons, resembling in their longevity DDT and other synthetic materials. It is expected that the diesel fuel pollution of Hayfork Creek resulted in long term effects on the stream fauna.

Caution in the transport of oily substances is obviously required to prevent accidental spills, especially in the vicinity of flowing waters because pollutants can be dispersed great distances in a relatively short time. Bonig (1965) reported that a great deal of oil pollution occurs in spite of safety measures. Persons who dispense or transport petroleum products need to be acutely aware of the great damage that these sub- stances have on fisheries and wildlife resources.

The risk of accidental spills of petroleum products is an ever present danger of pollution to aquatic ecosystems and will undoubtedly increase with rising consumption and transportation of fossil fuels. Tliis study indicates that oil pollution is a serious threat to life even in remote, unspoiled streams and rivers.

ACKNOWLEDGMEN1S

I thank Dr. Bobert C. Stebbins for his careful review and comments on the manuscript, and Mr. Biehard Hansen for providing helpful information on the pollution at Hayfork Creek.

REFERENCES

Blumer, M., H. L. Sanders, J. F. Grassle, and G. R. Hampson. 1071. A small

oil spill. Environment 13(2) :2-12. B 'nig. 1965. Danger to waters from the use and storage of oily substances in

industry and their prevention. Industrieabwasser, 1965 :51-57. Bury, R. B. 1972. Habits and home range of the Pacific pond turtle, Clemmys

marmorata, in a stream community. Ph.D. Thesis. Univ. California. Berkeley. Gutsell, J. S. 1921. Danger to fisheries from oil and tar pollution of waters. Bur.

of Fisheries. Doc. 910, Appendix to Pep., U.S. Comm. of Fisheries. 10 p. McCaull, J. 19G9. The black tide. Environment 11(9) :2-16. McKee, J. E. 1956. Report on oily substances and their effects on the beneficial

uses of water. Calif. Water Poll. Control Board, Publ. No. 16. 71 p.

— and II. W. Wolf, eds. 1963. Water quality criteria, 2nd ed. Calif. Water

Quality Board, Publ. Xo. 3-A. 548 p. Mitchell, C. T., E. K. Anderson, L. G. Jones, and W. J. North. 1970. What oil

does to ecology. J. Water Pollut. Control Fed. 42(5) :812-81S. Straughan, D. 1971. Biological and oceanographical survey of the Santa Barbara

Channel oil spill. Vol. I. Biology and bacteriology. Allen Hancock Foundation,

Univ. So. Calif. Sea Grant Publ. No. 2. 426 p. Swift, W. H., C. J. Touhill. W. L. Templeton, and D. P. Roseman. 1960. Oil

spillage prevention, control, and restoration — state of the art and research needs.

J. Wat. Pollut. Control Fed. 41(3) :392-412. Wilber, C. G. 1969. The biological aspects of water pollution. Charles C. Thomas,

Springfield, Illinois. 296 p.

Calif. Fish and Game, 5S(4) : 29G-314. 1972.

A SUBPOPULATION STUDY OF THE PACIFIC SARDINE1

KENNETH F. MAIS

Marine Resources Region California Department of Fish and Game

A subpopulation study was made of Pacific sardines inhabiting the west coast of North America and Mexico. A method of statistical treat- ment was utilized to determine the amount of overlap of single and combined meristic and morphometric characters. Results indicate the existence of three stocks centered in California, Baja California (Mexico), and the Gulf of California (Mexico). California and Baga California fish were very similar, but Gulf of California sardines appear to be a more distinctive and separate stock.

INTRODUCTION

The Pacific sardine, Sardinops caeruleus, inhabits coastal waters from British Columbia, Canada, southward to the tip of Baja Califor- nia, Mexico, including the Gulf of California. In former years, sardines were common throughout their range and were abundant in central and southern California where they supported a very large and valu- able commercial fishery. The population is at an extremely low level at present (1971). In the past 18 years, sardines have disappeared from the northern limits of the range and are generally very scarce in areas of former abundance. They are common only in the southern half of Baja California, Mexico, including the Gulf of California. The fishery was maintained at a high level in the 1930 's and early 40 's with yearly catches exceeding 400,000 tons. A drastic decline followed which even- tually resulted in a moratorium limiting the catch to 250 tons in Cali- fornia.

There have been three brief upsurges since the decline began in 1946. At least two of these, 1954-55 and 1958-59 seasons, were attributed to sardines migrating from Mexico since no large incoming year classes or adults were present previously (Ahlstrom 1959 ; Calif. Mar. Res. Comm. 1960).

These apparent migrations indicated the necessity of identifying and delineating sardine subpopulations. Does the fishery draw on one homo- geneous population or several? If more than one subpopulation exists, what proportion does each contribute to the fishery? Does a sudden upsurge in landings represent entry of a good year class or a migration of a stock from outside the normal fishery range?

Previous studies relating to sardine subpopulations have been con- ducted by a number of investigators. Clark (1947) compared vertebral counts of a large number of specimens over the species' range. Results indicated a heterogeneous group from British Columbia to Point Eu- genia central Baja California, Mexico. A second group that appeared to be separate and not mixing to any extent with the first group inhab-

1 Accepted for publication April 1972.

(29G )

SARDINE STUDY 297

ited the Gulf of California and southern Baja California, Mexico. McHugh (1950) using larval and post larval material demonstrated different rates of development and growth of various body parts by morphometrical comparisons of fish from southern California and Baja California, Mexico. Radovich and Phillips (1952) found sardines of the same year class were larger in southern California than those in central Baja California, Mexico. Felin (1951) made a similar study of fish from California and the Pacific Northwest and found different growth characteristics between the two areas.

Age and size composition of the commercial catch of central Baja California indicate sardines have a slower growth rate, a larger number of age groups, and a smaller maximum size than California fish (A\7olf and Daugherty 1964). Egg and larvae surveys by the National Marine Fisheries Service (formerly the U.S. Bureau of Commercial Fisheries) discovered a summer spawning group in central Baja California in addition to the regular spring spawners. These spawning groups may be evidence of subpopulations. A California Department of Fish and Game tagging program (Clark and Janssen 1945) indicated extensive migration between California and the Pacific Northwest, and to a lesser degree between California and central Baja California, Mexico. Blood genetic studies by Vrooman (1964) indicate three subpopulations: northern, which ranges from California to northern Baja California; southern, which ranges from Baja California to southern California ; and gulf, which is confined to the Gulf of California.

MATERIAL

Most specimens for this study were collected from 1958 to 1962 on routine fish surveys by the California Department of Fish and Game. These surveys were conducted primarily to assess sardine year class strength and covered the species' present range which is from San Francisco southward to and including the Gulf of California. Sampling was accomplished by attracting fish with a night light and capturing them with a blanket net. A few samples of adults were obtained from central and southern California commercial catches. Several samples of juveniles were obtained from southern California live bait haulers. One exotic sample originated in the Galapagos Islands off South America.

Originally only spawning adults were to be used for the study, but difficulty in capturing sufficient numbers necessitated taking all sizes and stages of sexual maturity. Although collections were made over a period of years, most samples were taken in 1958-1959. Southern Cali- fornia samples were taken over the greatest time span and during more seasons of the year. This area also was the most intensively sampled. Many Mexican samples were taken during fall and summer months. Gulf of California fish were taken on three survey cruises at various times of the year. During the later stages of collecting, several samples also were subjected to blood serology tests and classified into one of the three genetic groups reported by Vrooman (1964).

Fish sizes ranging from 110 to 209 mm sl were used for the study. The central and southern California samples contained the largest fish both in proportion and actual sizes. Central California samples con- sisted exclusively of large fish and southern California samples con-

298

CALIFORNIA FISH AND GAME

tained mostly large and medium fish. Samples from Mexican waters including the Gulf of California adequately represented small and me- dium fish but contained a low proportion of large sizes. Seventy-five samples comprised of :>.706 fish were used in this study (Figure 1). The 1956-58 year classes predominated most areas sampled.

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METHODS

Specimens were laid out straight and frozen immediately after cap- ture at sea. After thawing in the laboratory, identification tags were attached and scales taken for age determination. The fish again were laid out straight and preserved in a 10% solution of formalin for a minimum period of 3 weeks.

-Morphometric characters were selected on the basis of least likelihood • if correlation with each other. Meristie characters were limited due to the difficulty of making accurate counts on some. All morphometric measurements were made by the author and all meristie counts were double checked. The following morphometric measurements were made: standard length, head length, pectoral fin length, and postpelvic length. Standard length consisted of the distance from the tip of the snout to

SARDINE STUDY 299

the end of the hypural plates. The latter point was determined by bending the tail forward and inserting a pin where a crease appeared in the caudal peduncle. Head length was measured from the tip of the snout to posterior edge of the opercular flap. Pectoral fin length was measured from the base of the fin to the tip of the longest ray. Post- pelvic length consisted of the distance from the end of the hypural plates to the origin of the pelvic fins.

Meristic counts were made of vertebrae and gill rakers. Vertebral counts were made from X-ray photos and included the hypural. Gill raker counts were made on the lower half of the first gill arch. Age determination was made by a routine scale reading process conducted by California Cooperative Oceanic Fisheries Investigations (CalCOFI).

STATISTICAL TREATMENT

Most samples used in this study were fairly representative of areas from which they originated. The large number of samples from the major areas and the span of time over which they were collected con- tributed to their representativeness (Figure 1). One exception was the southern California area where an apparent influx of migrants from Mexico at time of sampling may have affected representativeness. The size composition of sardine samples in all areas is typical of fish taken in the past 15 years by California Department of Fish and Game sea surveys. Night light blanket net gear is equally effective in taking all sizes of fish.

Before morphometric data could be statistically treated, it was nec- essary to determine if their relation to fish size was linear. Tests were made for linearity on morphometric characters and gill rakers counts. All were linear except pectoral fin length which was only slightly curvilinear and gill raker counts which were quite curvilinear. This problem and the effects of allometric growth were minimized by strati- fying samples into different size groups. These groups designated as small, medium, and large were composed of fish 110-139, 140-169, and 170-209 mm sl respectively. Eegression of gill rakers was calculated using head length instead of standard length.

Variance and Covariance

Samples were grouped by area. The areas consisted of central Cali- fornia, southern California, northern Baja California, north-central Baja California, south-central Baja California, southern Baja Califor- nia. Gulf of California, and Galapagos Islands (Figure 1). Analysis of variance and covariance tests were made of samples from the same area (within area) for each character. Significant differences at the 5% level resulted for each area and character when all samples were considered. This is quite normal when large numbers and samples are involved. Koyce (1957) concluded significant differences can always be found between very closely related stocks if large and numerous sam- ples are taken . This phenomenon has been widely experienced by tax- onomists in other investigations (Mayr, Linsley. and TJsinger, 1953). Samples stratified by year class, age, and spawning condition were sub- jected to the same tests. Similar results were obtained except for 3-year

300 CALIFORNIA FISH AND GAME

old central California samples. No significant differences were found between samples of this ape group in all characters except pectoral fin length. Age 0 fish in north-central Baja California and the Gulf of California did not differ significantly within each area in some char- acteristics. Females in spawning condition also were homogeneous in several characteristics in southern California, south-central Baja Cali- fornia, and southern Baja California. Thus it is apparent there is heterogeneity in all areas, but some strata are nearly homogeneous.

Comparisons were made between adjacent areas using all fish strati- fied by size group. Results similar to those of within areas were ob- tained. F values (F statistic used in analysis of variance) were gen- erally larger giving some indication of greater differences between areas than within areas.

Regression formulas were computed for each morphometric character in all size groups and areas. Using these formulas, mean values of each character adjusted for standard fish sizes were determined and plotted with 1 sd and 2 se's on each side of the mean (Figure 2, Appendix 1-5). Standard lengths of 126 mm, 154 mm, and 188 mm were used as standard fish sizes to represent small, medium, and large size groups. Gill rakers were adjusted to the mean head length of a standard size fish of each size category. Vertebral plots were made for only the total number of fish in each area (Figure 3). Clines are present in head length, postpelvic length, and vertebrae. Irregularities occur notably in Gulf of California large fish. Pectoral fin length and gill raker counts gave only vague indication of clines with numerous irregularities present.

The relative head length of sardines was greater in fish captured to the south with a maximum mean difference of 3 mm which occurred between medium size fish from southern California and the Galapagos Islands. The greatest difference in Pacific Coast samples was 1.94 mm between large fish from southern California and south-central Baja California (Figure 2).

Postpelvic length exhibited a well defined cline which was present in all size groups. This character decreased from north to south. A maxi- mum difference of 3.32 mm was observed between large fish from central California and north-central Baja California (Figure 2).

A definite cline was evident in vertebral counts with a decrease to the south. Although this cline was very consistent, actual differences were quite small amounting to .63 vertebra between extremes (Figure 3).

Pectoral fin lengths gave no definite indication of a cline. Tins character appears useful only in distinguishing Gulf of California fish which had longer pectoral fins in all 3 size groups. Their means differed between .68 to 1.65 mm from fish of other areas (Figure 2).

The mean number of gill rakers varied irregularly with respect to a cline. A very slight discontinuous north-south cline was discernible in the medium and large size groups which indicated a small decrease to the southward. The Gulf of California large fish mean was consider- ably less than the other areas. It differed by .91 gill rakers from southern California mean which was the next lowest (Figure 2).

SARDINE STUDY

301

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302 CALIFORNIA PISH AND GAME

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Discriminant Function and Overlap

Analysis of variance and covariance showed differences existing within and between areas. Such information is useful in preliminary analysis but does not afford a good measure of the magnitude of dif- ferences. Employing the concept of overlap described by Royce (1957), an estimate can be made of the proportion of one group having identical characteristics of another. Using Royce 's basic formula, D is the dis- tance between means in units of standard deviation:

D = - and Xi and X2 are tne means

s of the characteristic in question and s is the pooled average standard deviation. From a table of normal probability integral using the value D/2 as an entering argument, the area of half the normal curve plus the mean to the argument is obtained. This area Royce calls the relative probability. 1-P. which is the probability of correctly classifying an individual as belonging to one group or another. It varies from .5 with complete overlap to 1.0 with no overlap. The percentage of overlap is

SARDINE STUDY 303

obtained by multiplying P by 200. This value is the combined area of the overlapping tails of each distribution and measures the area or percentage of one curve which is included in the other. For example, in computing overlap of head length of large fish between southern and

central California, Xt = 15.70 + .1850X, X2 = 7.38 + .2273X (regres- sion equations from Appendix II), X = the grand means of body lengths of the two areas = 191.72. Solving the regression equations Xi = 51.17, Xo = 50.96. The pooled standard deviation from regres- sion = 1.342.

^ 51.17 - 50.96

D = 17342 = °'156

D/2 = 0.078, P = 0.469, overlaD = 93.8%

The concept of overlap thus gives an estimate of the proportion of one group having identical characteristics of another. The amount of similarity is therefore measurable and inferences concerning inter- mingling can be made. A high degree of overlap between two areas doesn't prove intermingling has occurred, but does represent a maxi- mum that could have occurred. Very low overlap values on the other hand provide good evidence of little or no intermingling.

To use this method for morphometric characters, means were com- puted from the regression equation of each group under consideration as was the grand mean of their body lengths. The pooled standard deviation was replaced by the pooled standard deviation from regres- sion.

Overlap percentages were calculated between adjacent areas for all meristic and morphometric characters using samples stratified by the size categories mentioned previously (Table 1). No single character consistently excelled in distinguishing groups. The value of each charac- ter in separating stocks varied greatly between each set of adjacent areas as well as between size categories. Overlap between the adjacent areas of north-central and south-central Baja California was large, averaging 90.17% for all characters. Comparisons of all large Cali- fornia fish with all large Mexican fish, except those from the C4ulf of California, produced overlap values of 57.1% and 67.2% in head length and postpelvic measurements. The remaining characters overlapped from 88.0% to 94.0%. These examples give some indication as to how characteristics may differ even through these differences are not large enough to support inferences.

Discriminant function and overlap of multiple characters.

This form of multivariate analysis employs the generalized distance function developed by Mahalanobis (1936) which gives a measure of distance between two groups using a combination of characters. Each character is considered only after correlation with a previous one has been excluded. The general formula is :

D2 = Wy di dj

vhich \Yij is the dj dj are differences between means. D2 is similar to D used for single

in which \Vij is the inverse of the variance-covariance matrix Wy and

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CALIFORNIA FISH AND GAME

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SARDINE STUDY 305

character overlap and can be used in the same way. At the same time D2 is computed, a linear function of the combined characters is obtained which may serve as an index for discrimination. A complete explana- tion is given by Rao (1952). This method was used by Royce (1964) in a racial study of yellowfin tuna and by Hill (1959) to distinguish races of American shad.

For this study a computer program developed at the University of California at Los Angeles was utilized to perform the computations. Output included D2, discriminant function coefficients, and discriminant function frequencies of both compared groups. A small bias in D2, due to unequal sample sizes, is removed by subtracting the value

piii + n2, nin2 '

P is the number of characters and ni and n2 are sample sizes.

Morphometric data were adjusted for use in this analysis by comput- ing a character value for each individual fish based on a standard fish size. Deviations of individuals from their computed regression means were added or subtracted from the regression mean of a standard fish size. For example, if a 175 mm sl fish had a head length of 48 mm and the computed regression mean head length for this size was 50 mm, the difference of 2 mm would be subtracted from the mean computed head length of the standard size fish. Standard fish sizes, 126 mm sl (small), 154 mm sl (medium), and 188 mm sl (large) were determined by computing the grand mean of each size category.

RESULTS, SMALL AND MEDIUM SIZES

Overlap percentages derived from D2 values were calculated (Table 2). Small and medium size fish from southern California were compared with each area to the south (Figure 4). Small fish differed relatively little from southern California to southern Baja California with over- lap ranging from 56.6% to 64.3%. This overlap range also was found between samples from the same school group so it cannot be considered low enough to infer separate stocks. Small fish from the Gulf of Cali- fornia and Galapagos Islands overlapped the southern California group by 25.0% and 24.6% respectively This low degree of overlap is a clear indication of different stocks.

Medium size fish were compared in the same manner (Figure 4). A great similarity was apparent between fish from southern California southward to and including south-central Baja California. A great degree of multiple character overlap ranging from 70.4% to 79.3%, was found between these areas. This may have been due to the migra- tion of medium size fish into California during the period of data collection. From 1957 to 1960 there was an appearance of large quanti- ties of medium size fish in areas where none had occurred for many years.

Fish from southern Baja California, the Gulf of California, and the Galapagos Islands overlapped southern California fish by 46.0%, 34.7%, and 10.8% respectively. These differences are large enough to suspect separate stocks in these areas.

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SARDINE STUDY

307

tOO 90

80

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SIZE I (SMALL)

PERCENT OVERLAP 60 50

SIZE II (MEDIUM)

SIZE III (LARGE)

SOUTHERN CALIFORNIA

NORTH CENTRAL BAJA CAL. SOUTH CENTRAL - SOUTHERN BAJA CAL. GULF OF CAL. GALAPAGOS ISLANDS

SOUTHERN CALIFORNIA

NORTHERN BAJA CAL. NORTH CENTRAL BAJA CAL. SOUTH CENTRAL GULF OF CALIFORNIA GALAPAGOS ISLANDS

CENTRAL CALIFORNIA SOUTHERN CALIFORNIA NORTHERN BAJA CAL. NORTH CENTRAL BAJA CAL. SOUTH • » "

GULF OF CALIFORNIA

FIGURE 4. Overlap percentages derived from multivariate analysis of Pacific sardine morpho- metric and meristic characters. Small, medium, and large size fish from central and southern California compared with successively southward sampling areas.

RESULTS, LARGE FISH

Central California large fish, which analysis of covariance indicated was the least heterogeneous group, were compared with large fish from each of the remaining areas for which samples were collected (Figure 4). Fish from central California, southern California, and northern Baja California differed only moderately. Fish from the latter two areas overlapped those from the first by 63.6% and 61.07c- North- central and south-central Baja California fish were very similar to each other and substantially different from the areas to the north. Stock differences are indicated by the relatively low overlap values (43.5% and 41.8%) of these fish with those from central California. Gulf of California fish differed greatly from all other areas and most certainly comprise a separate stock. Overlap between fish from this area and those from central California was 29.4%.

Von Bertalanffy growth parameters, L infinity and K, were added to the morphometric and meristic characters for each fish 3 or more years old. Very little was gained by using these extra characters as D2 values increased only slightly even when large growth rate differences were apparent. This was due to the extremely large variance in the growth parameter values.

Overlap comparisons were made using five samples of large fish upon which blood serology tests were performed by the National Marine Fisheries Service, La Jolla. These samples were classified into the north-

308 CALIFORNIA FISH AND GAME

era or southern groups as postulated by Vrooman (1964). All but one were classified northern. The southern sample originated in San Pedro Bay and the northern ones at Santa Catalina Island, Santa Cruz Island, and Todos Santos Bay. These locations are in southern California except the last -which is in northern Baja California, Mexico.

Two samples were taken from the same school group at Santa Cata- lina Island. One Santa Catalina Island northern sample was compared with each of the other samples. Overlap with the three other northern samples was 65.4%, 58.9%, and 36.4%. The highest was between sam- ples from the same school group and the lowest with the Todos Santos Bay sample. The southern sample overlapped 48.2%. The overlap of the same school samples indicates how variable very closely related stocks can be and affords some idea of the amount of overlap necessary to differentiate stocks.

Differentiation using the overlap method agreed favorably with the blood serology method except the Todos Santos Bay northern sample differed more from other northern samples than did the southern sample. A possible explanation is a sampling error could have occurred either in the blood serology test or the morphometric-meristic multi- variate analysis due to a small sample size of 72 fish.

DISCUSSION

Results of this study indicate three stocks of sardines occur in the eastern temperate Pacific. These stocks are located in California, central Baja California (Mexico), and the Gulf of California (Mexico). Cali- fornia and central Baja California stocks are quite closely related and undoubtedly intermingle to a considerable degree. Small and medium size fish from these areas overlapped so greatly that no inference of separate stocks can be made. This large overlap (56.6% to 79.3%) was probably due to a heavy influence of migrants from Mexico and a paucity of native born fish in the California samples. Large fish from these regions overlapped considerably less (41.8% to 43.5%), but would have probably differed even more if the California samples had not been influenced by migrants from Mexico.

At first glance the overlap between large fish from California and Mexico indicated a considerable difference, but when fish of the same school and blood group overlapped by 65.4%, an overlap of 41.8% to 43.5%.. between these areas indicates a relatively small difference. The actual physical differences between stocks are extremely small thereby precluding any differentiation of individual fish or assessing the degree of intermingling.

Gulf of California fish appear to be a more distinct stock. Analysis of all size groups in this area indicated the least similarity to fish of other California areas with overlap values ranging from 25.0% to 34.7%. Small and medium sized southern Baja California fish were quite similar to those of central Baja California. No large fish from southern Baja California were available but they probably resemble central Baja California fish.

Galapagos Island fish differed greatly from those of all other areas as indicated by the low overlap of 10.8% to 24.6%.

SARDINE STUDY 309

These fish are separated from the others by extensive geographic and oeeanographic barriers which exclude intermingling.

SUMMARY

A subpopulation study was made of the Pacific sardine population inhabiting the eastern Pacific Ocean. Seventy-three samples comprised of 3,706 fish were collected from California, Baja California, Gulf of California, and the Galapagos Islands off South America. Morpho- metry measurements and meristic counts consisting of : standard length, head length, pectoral fin length, postpelvic length, gill rakers, and vertebrae were made. Samples were stratified by fish size and area of catch.

Analysis of variance and covariance indicated a high degree of heter- ogenity of fish from both within and between geographic areas. A method of statistical treatment which computes the proportion of one group having identical characteristics of another (overlap) was used for single characters to compare samples from adjacent areas. A more sophisticated method employing the same concept was used with all characters combined in aggregate. Single character comparisons gave rather large overlap values with over three-fourths of all comparisons exceeding 70%. Head length was the most useful distinguishing char- acter and number of vertebrae the least.

Overlap comparisons using all characters in aggregate were made between California samples and those from each successive area to the south. This analysis produced much lower overlap values ranging from 10.8% to 79.3%. Small and medium size fish from California to south- ern Baja California, Mexico, overlapped so greatly that no inference of separate stocks can be made. Fish of these size groups from the Gulf of California and the Galapagos Islands overlapped California fish 10.8% to 34.7% and must certainly be separate stocks, botli from each other and all other areas.

Similar comparisons of large fish produced overlap values ranging from 29.4% to 63.6%. California samples overlapped each other and northern Baja California 63.6% and 61.0% respectively. They over- lapped Baja California samples from 41.8% to 43.5%, and those from the Gulf of California 29.4%. These results indicate California and northern Baja California sardines are probably a separate stock but are vary similar to a second stock off southern and central Baja Cali- fornia. Gulf of California fish are a third more distinctive stock.

REFERENCES

Ahlstrom, E. A. 1959. Distribution and abundance of eggs of the Pacific sardine,

1952-1956. U.S. Dep. of Interior, Fish Wildl. Serv., Fish. Bull. 60(165) :

185-213. California Marine Research Committee. 1960. Review of the partial resurgence of

the sardine fishery during 1958-59. Calif. Coop. Oceanic. Fish. Invest. Reports,

7 :S-10. Clark, Frances N. 1947. Analysis of populations of the Pacific sardine on the

basis of vertebral counts. Calif. Dep. Fish and Game, Fish Bull. (65) :l-26. Clark, Frances N., and John F. Janssen. 1945. Movements and abundance of the

sardine as measured by tag returns. Calif. Dep. Fish and Game, Fish. Bull. (61) :

7-42. Felin, Frances E. 1954. Population heterogeneity in the Pacific pilchard. U.S.

Dep. of Interior, Fish Wildl. Serv., Fish Bull. 54(86) : 201-225.

310

CALIFORNIA PISH AND GAME

Hill, Donald R. 1959. Some uses of statistical analysis in classifying races of

American shad (Alosa sapidisima) . U.S. Dep. of Interior, Fish Wildl. Serv.,

Fish. Bull. 59(147) :269-284. Mahalanobis, P. C. 1930. On the generalized distance in statistics. Proceedings of

the National Institute of Sciences (India) 2(1) :49-55. Mayr. Ernst, E. G. Lindsley, and It. L. I'singer. 1953. Methods and principles of

systematic zoology. McGraw-Hill, New York :l-328. McHugh, J. L. 1950. Variations and population in the clupeoid fishes of the

North Pacific. Doctoral thesis, University of California, Los Angeles : 1-116. Radovich, John R., and J. B. Phillips. 1952. Distribution and abundance of

young sardines. Calif. Dep. Fish and Game, Fish. Bull. (87) :l-63. Rao, C. R. 1952. Advanced statistical methods in biometric research. Wiley, New- York : 1-390. Royce, William F. 1957. Contributions to the study of subpopulations of fishes.

U.S. Dep. of Interior, Fish Wildl. Serv., Spec. Sci. Rep.-Fish. (208) :7-28. — . 1904. A morphometric study of yellowfin tuna, Thunnus albacdres (BOn-

naterre). U.S. Dep. of Interior, Fish Wildl. Serv., Bur. of Com. Fish., Fish. Bull.

63(2) :395-443. Vrooman, A. M. 1904. Seriologically differentiated subpopulations of the Pacific

sardine, Sardinops caerulea. Res. Bd. of Can., J. Fish. 21(4) :691-701. Wolf, Robert, and Anita E. Daugherty. 1904. Age and length composition of the

sardine catch off the Pacific Coast of the United States and Mexico in 1901 and

1902. Calif. Fish Game 50(4) :241-252.

APPENDIX I— Vertebrae AH Fish

Area	X	X	SD	SE
	258 898 285 1,007 483 240 492 43	51.85 51.51 51.36 51.33 51.36 51.17 51.17 51.05	0.60 0.61 0.64 0.58 0.59 0.54 0.56 0.49	0.04
	0.02
	0.04
North-Central Baja California- South-Central Baja California	0.02 0.03 0.04
	0.03
	0.07

N = Sample Size

X = Mean

sd = Standard Deviation

se = Standard Error

SARDINE STUDY

311

APPENDIX II— Head Length Small (126 mm SL)

Area	N	Regression Equation	Y	Sy	SE
Central California- -	0 87 0 325 202 112 154 28	3.68 + .2533 X 4.55 + .2416 X 4.60 + .2412 X 2.11 + .2661 X 5.63 + .2466 X 3.94 + .2654 X	35.60 34.99 34 . 85 35 . 64 36.70 37.38	1.01 0.91 0.77 1.43 0.88 0.62
Southern California Northern Baja California - North-Central Baja California South-Central Baja California Southern Baja California Gulf of California	0.11 II ().-, 0.06 0.14 0.07
Galapagos Islands	0.01

Medium (154 mm SL)

Central California

Southern California

Northern Baja California

North-Central Baja California South-Central Baja California.

Southern Baja California

Gulf of California

Galapagos Islands

0 258 211

i;;.s 104 128 228 15

-1

4

11

4

8.59 + 0.41 + 32 + 96 + 67 + 24 +

10.01 +

.2190 X .2741 X . 2877 X .2427 X .2089 X .2569 X .2292 X

42.32 42.62 42.98 42.34 43.84 43.80 45.31

1.10 0.97 1.37 1.20 0.95 1.02 0.77

0.07 0.07 0.07 0.12 0.09 0.07 0.20

Large (188 mm SL)

Central California

Southern California

Northern Baja California

North-Central Baja California South-Central Baja California.

Southern Baja California

Gulf of California

Galapagos Islands

258 553 74 244 177

0 110

0

15.70 + 7.38 + 9.51 +

13.52 + 2.70 +

.1850 X .2273 X .2216 X . 2045 X .2625 X

0.75 + .2708 X

50.48 50.11 51.17 51.97 52.05

51.66

01

46

I .06

64 39

1.11

0.06 0.06 0.12 0.10 0.10

0.11

N = Sample Size

X = Standard Length

Y = Mean Character Length

Sy = Standard Deviation from Regression

se = Standard Error

312

CALIFORNIA FISH AND GAME

APPENDIX III— Post Pelvic Length Small (126 mm SL)

Area	N	Regression Equation	Y	By	SE
Central California	0 87 0 325 202 112 154 28	-0.24 4- .5011 X -8.38 + .5667 X -0.06 + .4998 X 0.18 + .4928 X -4.47 + .5265 X -7.01 4- .5430 X	62.89 63.02 62.91 62.27 61.87 61.41	1.87 1.50 1.17 1.37 1.59 1.23
	0.20
Northern Baja California North-Central Baja California South-Central Baja California Southern Baja California.. Gulf of California -	0.09 0.08 0.13 0.13
Galapagos Islands	0.23

Medium (154 mm SL)

Central California

Southern California

Northern Baja California

North-Central Baja California South-Central Baja Californa_

Southern Baja California

Gulf of California

Galapagos Islands

0 258 211 438 104 128 228 15

-2.37 + 4.52 + 4.05 + 4.18 +

-8.41 +

5.31 +

-14.36 +

.5167 X .4698 X .4713 X .4752 X . 5489 X .4593 X . 5834 X

77.20 76.86 76.63 77.36 76.12 76.04 75.48

1.84 2.46 1.94 2.10 1.67 1.85 1.58

0.11 0.17 0.09 0.21 0.15 0.12 0.41

Large (188 mm SL)

Central California

Southern California

Northern Baja California

North-Central Baja California South-Central Baja California.

Southern Baja California

Gulf of California

Galapagos Islands

258 553 74 244 177

0 110

0

- 1 1 . 85 + -4.69 +

-12.23 +

-3.29 +

5.22 +

.5727 X .5289 X .5654 X .5095 X . 4686 X

-4.29 + .5200 X

95.82 94.74 94.07 92.50 93.32

93.47

2.32 2 .22 2.56 2.39 2.13

1.91

0.14 0.10 0.30 0.15 0.16

0.18

SARDINE STUDY

313

APPENDIX IV— Pectoral Fin Length

Small (126 mm SL)

Area

Central California

Southern California

Northern Baja California

North-Central Baja California

South-Central Baja California

Southern Baja California

Gulf of California

Galapagos Islands

N

0 87

0 325 202 112 154

Regression Equation

-1.31 + .1809 X

15 + 99 + 24 + 32 +

93 +

1466 X 1554 X 1615 X 1347 X 1902 X

21.48

21.62 21.57 21.59 23.29 22.03

Sy

1.07

0.75 0.71 0.93 0.75 0.80

0.11

0.04 0.05 0.08 0.06 0.15

Medium (154 mm SL)

Central California

Southern California

Northern Baja California

North-Central Baja California South-Central Baja California.

Southern Baja California

Gulf of California

Galapagos Islands

0 258 211 438 104 128 228 15

4.54 + 3.98 4- 2.33 + 2.43 + 9.34 3.50 0.79

+ +

. 1386 X .1469 X .1543 X .1522 X .1112 X . 1540 X . 1669 X

25 . 88 26.60 26.09 25.87 26.46 27 .22 26.49

0.99 0.95 1.09 0.89 0.93 0.96 0.79

0.06 0.07 0.05 0.09 0.08 0.07 0.20

Large H88 mm SL)

Central California

Southern California

Northern Baja California

North-Central Baja California South-Central Baja California.

Southern Baja California

Gulf of California

Galapagos Islands

258 553 74 244 177

0 110

0

6.65 + 3.45 + 9.79 + 10.23 + 7.88 +

.1281 X .1451 X .1161 X .1110 X .1211 X

-3.46 + .1902 X

30.73 30.73

31

31

62 10

30 . 65

32.30

.39 .28

.11 .22 .27

1.08

0.09 0.06 0.13 0.08 0.10

0.10

::i I

CALIFORNIA FISH AND GAME

APPENDIX V— Gill Rakers

Small (36 mm HL)

	X	Regression equation	Y	Sy	SE
i 'entraj ('alifornia __	0 S7 0 325 202 112 15 1 28	15.92 + .6157 X 15.15 + .6132 X 19.67 + .4884 \ 8.06 4- .7608 X 20.19 + .4838 X 19.16 + .4634 X	37.38 36 . 52 36.69 35.45 37.05 35.31	1.73 1 . 52 1.39 2.00 1.51 1.44
Southern California . Northern Baja California North-Central Baja California ^-Central Baja California Southern Baja California. Gulf of California	0.19 0.09 0.10 0.19 0.12
	0.27

Medium (43 mm HL;

Central California

crn California

Northern Baja California

North-Central Baja California South-Central Baja California.

Southern Baja California

Gulf of California

Galapagos Islands

0 258 211 438 104 128 228 15

28.43 + .2984 X 22 . 37 + . 4332 X

19.78 + 20.07 + 26.96 + 23.60 + 31.12 +

4970 X 4718 X 3294 X 3942 X

181'. i X

41.26 40.99

41.14 40.35 41.12 10.55 38.94

1.74 1.59 1.60 1.73 1.69 1.96 1.50

0.11 0.11 0.08 0.17 0.15 0.08 0.39

Large (51 mm HL)

( !entral California

Southern California

Northern Baja California

North-Central Baja California South-Central Baja California.

Southern Baja California

Gulf of California

Galapagos Islands

553

74 I'll 177

0 110

0

19.34

.4949 X

23.14 4- .4049 X 12.76 + .6222 X .417C X

.4259 X

23 . 22.29 -

_':;. II -

.3822 X

44.90 44.06 44.90 44.96 44.01

43.15

1.58 1.79 1.56 1.87

1.73

1 .66

0.10 0.08 0.18 0.11 0.13

0.16

HL = head length.

Calif. Fish and Game, 58(4) : 315-320. 1972.

CHECK LIST OF INTERTIDAL FISHES OF TRINIDAD BAY, CALIFORNIA, AND ADJACENT AREAS1

JOHN R. MORING2

Department of Fisheries, Humboldt State College,

Areata, California 95521

Intertidal fishes of Trinidad Bay, California, were sampled from May 1965 to May 1970. The 1,517 fishes collected represented 20 species. Three additional species were collected intertidaily near Point St. George, Del Norte County. As no check lists of tidepool fishes of Hum- boldt and Del Norte counties are currently available, a check list is provided for Trinidad Bay species, with supplemental notes for adjacent regions.

INTRODUCTION

There have been few reviews and descriptions of intertidal fishes of the northern California coast, Most descriptions have been included in regional monographs and studies. Jordan and Evermann (1896, 1898) and other workers included intertidal fishes in early monographs. But- ter (1899) studied intertidal fishes of Kodiak Island, Alaska, Hubbs (1926) reviewed the intertidal cottid genera of the Pacific coast, and later discussed several intertidal blennioid fishes (Hubbs, 1927). Schultz (1936) included many intertidal fishes in his key to fishes of Washing- ton, Oregon, and adjacent regions. Bolin (1944, 1947) discussed inter- tidal cottids, and Clemens and Wilby (1961) reviewed intertidal fishes of northern California in discussing fishes of the Pacific coast of Can- ada.

Several workers included brief descriptions of dominant northern California intertidal fishes in their reviews of fishes of other regions. Starks and Morris (1907) and Barnhart (1936) included intertidal fishes in their descriptions of fishes of southern California, and Ever- mann and Goldsborough (1907) included intertidal fishes in their re- view of fishes of Alaska. Intertidal residency of juvenile northern Cali- fornia fishes has been noted for Scorpaenichthys marmoratus (O'Con- nell, 1953) and Sebastes mystinus (Wales, 1952).

Gersbacher and Denison (1930), Williams (1957), and Green (1971) studied fish movement in the intertidal zone. Morris (1960) and Graham \ 1970) analyzed temperature sensitivity of several species, and Mitchell (1953), Johnston (1954), and Nakamura (1971) reviewed food habits of many dominant northern California tidepool fishes. Other studies, including those by Hubbs and Barnhart (1944), Schultz (1944), and Briggs (1955), are helpful in considering distributional patterns.

Two workers have concentrated solely upon enumerating and describ- ing California tidepool fishes (Greeley, 1899; Bolin, 1964). However, both based their discussions upon central California intertidal fishes. There have been no reviews of tidepool fishes of Trinidad Bay, Cali- fornia, or adjacent regions along the coasts of Humboldt and Del Norte

1 Taken in part from a Master of Science thesis submitted to the faculty, Humboldt

State College. Accepted for publication January 1972.

2 Present address : Fisheries Research Institute, University of "Washington, Seattle,

Washington.

( 315 )

316

CALIFORNIA FISH AND GAME

counties. It is the purpose of this paper to provide a check list of the intertidal fishes of Trinidad Bay, and furnish notes on other intertidal species from adjacent regions.

MEMORIAL LIGHTHOUSE TIDEPOOLS

BARE ROCK TIDEPOOLS

124° 9' 0"

DOUBLE ROCK TIDEPOOLS

8

BAKER TIDEPOOLS

NORTH

SOTSIN PT

LUFFENHOLTZ TIDEPOOLS

TEPONA TIDEPOOLS

0

SCALE (km)

FIGURE 1. Trinidad Bay, California: site of intertidal fish sampling, May 1965 to May 1970.

FISHES OF TRINIDAD BAY 317

COLLECTION

Trinidad Bay, California, is located approximately 14 miles north of Humboldt Bay, at lat 41° 31' N; long 124° 8' W (Figure 1). It is semi-protected, and characterized by rocky shores with scattered tide- pools. Tidepools were sampled at several areas within the bay for inter- tidal fishes : Tepona, Luffenholtz, Baker, Double Hock Tidepools, Bare Rock Tidepools, Memorial Lighthouse and Little Head. Tidepool areas, in most cases, have been identified herein by their proximity to certain geographical landmarks in Trinidad Bay.

Between May 1965 and May 1970, 1,517 intertidal fishes were col- lected and measured during 53 collecting trips. Twenty species were identified in Trinidad Bay. Fishes ranged from 10 to 180 mm tl, aver- aging 49.4 mm. Of the 20 species collected, six occurred intertidally only as juveniles (Citharichthys stigmaeus, Hemilepidotus spinosus, Hcxagrammos decagrammus, Scorpaenichtliys marmoratus, Sebastes melanops, and 8. mystinus), and these were generally seasonal in ap- pearance.

Specimens were collected with a variety of hand nets and small seines. Fishes were anesthetized with quinaldine for ease in handling during measurement (Moring, 1970).

Intertidal fishes were also collected and measured from April 1967 to March 1970 from tidepools near Cape Mendocino, Pewetole Island (north of Trinidad State Beach), Patricks Point State Park, and Point St. George. Such sampling provided opportunities for examining fishes from varying intertidal environments.

SPECIES COLLECTED

The twenty intertidal species noted for Trinidad Bay, and the addi- tional species collected in Humboldt and Del Norte counties, by no means complete the check list for intertidal species in northern Cali- fornia. TJndescribed fish species may exhibit restrictions of habitat, low density in tidepools, or seasonal availability. The check list included in Table I attempts to provide a basis for further collection and enumera- tion of Humboldt and Del Norte county species. Reference to keys and descriptions by Schultz (1936), Bolin (1944, 1964), and Clemens and Wilby (1961) will supplement the check list.

Other Species

Additional intertidal species were collected from tidepools along the coasts of Humboldt and Del Norte counties. Some or all of these species may occur in Trinidad Bay, but are either uncommon, restricted in habitat, or seasonal in appearance.

Embiotocidae : Three juvenile Embiotoca lateralis (77, 78, and 79 mm tl) were collected in July 1969 in tidepools near Point St. George, Del Norte County.

Scorpaenidae : A single Sebastes paucispinis (88 mm tl) was col- lected in July 1969 near Point St. George. It was schooling in a deep tidepool with juvenile S. melanops and S. mystinus.

Cottidae : Several additional species of cottids may be found in Trinidad Bay. Bolin (1944) reported Ascelichthys rhodorus and Oligo- cottus rimensis from Crescent City. He noted the latter species was

318

CALIFORNIA FISH AND GAME

TABLE 1 — Cheek List of Intertidal Fish Species Collected, and Their Size Ranges in Trinidad Bay, California, During May 1965 to May 1970.

Species

Gobiesocidae: Gobiesox maeandricus.

Stichaeidae:

Anoplarchus purpurescens.

Cebidichthys violaceus

Xiphister atropurpureus

Pholidae:

Apodichthys flavidus.

Pholis ornata

Xcrerpes fucorum

Scorpaenidae:

f Sebastes mclanops.

f .Sebastes myslinus.

Hexagrammidae :

f Hexagrammos decagram)

Cottidae:

Artedius fenestralis

Artedius lateralis

Clinocottus acuticeps

Clinocottus globiceps

t Hemilepidotus spinosus

Oligocottus maculosus

Oligocottus snyderi

t Scorpaenichthys m i

Cyclopteridae:

Liparis florae

Bothidae:

: arichthys stigm

X	X
X
X	X
X	X
	X
X	X
X	X
	X

Tidepool areas

	o
	X
	X
•— --. >>	r*
-	_
m	P

-

— «

•r c

"3

21

*

38

11

1

15

74

4

to

c

25-91

35-110 29-180

IS 17(1

61 46-155

42 65

05-75

1	47
2	119-120
5	22-41
5	42-130
2	34-30
1,031	10-95
191	13-101
24	33-134

_v, lis

47 62

* One specimen, approximately 1 m tl, not measured, t Juveniles.

"uncommon. Leptocottus armatus, a common subtidal species in Trinidad Bay, has been reported, intertidally in central California (Bolin. 1964). Tomales Bay (Jones, 1962), and British Columbia (Clemens and Wilby, 1961)/

Pleuronectidae : A juvenile Parophrys vetulus (26 mm tl) was col- lected in May 1969 from a sand-mud bottomed tidepool near Point St. George. A larger juvenile (43 mm tl) was collected in the same area in July 1969. The only apparent previous literature record of young P. vetulus found in tidepools was by Villadolid (1927) concerning

fly collections by Hubbs. Hubbs (pers. comm.) collected one juvenile /'. vetulus (20 mm sl) in June 1923 from tidepools near Point St. George. Hubbs reported other intertidal collecting of P. vetulus juve- niles along the San Luis Obispo County coast.

FISHES OF TRINIDAD BAY 319

ACKNOWLEDGMENTS

Special thanks are due David Misitano and my wife, Kathleen, for assisting me on many collecting trips over the past years.

Carl L. Hubbs of Scripps Institution of Oceanography kindly sup- plied field notes and other information concerning his intertidal collections. George H. Allen, of Humboldt State College reviewed the manuscript, and offered helpful suggestions throughout the course of the study.

REFERENCES

Barnhart, P. S. 1936. Marine fishes of southern California. Univ. California Press,

Berkeley. 209p. Bolin. R. L. 1944. A review of the marine cotticl fishes of California. Stanford

Ichthyol. Bull. 3(1) : 1-135. . 1947. The evolution of the marine Cottidae of California with a discussion

of the genus as a systemic category. Stanford Ichthyol. Bull. 3(3) : 153-168.

1964. Key to intertidal fishes. In S. F. Light, R. I. Smith, F. A. Pitelka.

D. P. Abbott, and F. M. Weesner, Intertidal invertebrates of the central California coast. Univ. California Press, Berkeley : 313-322.

Briggs, J. C. 1955. A monograph of the clingfishes (Order Xenopterygii ) . Stan- ford Ichthyol. Bull. 6. 224p.

Clemens, W. A., and G. V. Wilby. 1961. Fishes of the Pacific coast of Canada. Fish. Res. Bd. Canada Bull. (68) 2nd ed. : 1-443.

Evermann, B. W., and E. L. Goldsborough. 1907. The fishes of Alaska. U.S. Bur. Fish. Bull. 26 (1906) : 219-360.

Gersbacher, W. M. and M. Denison. 1930. Experiments with animals in tide- pools. Publ. Puget Sound Mar. Biol. Sta. 7:209-215.

Graham, J. B. 1970. Temperature sensitivity of two species of intertidal fishes. Copeia 1970(1) : 49-56.

Greeley, A. W. 1S99. Notes on the tide-pool fishes of California, with a description of four new species. Bull. U.S. Fish. Comm. 19 : 7-20.

Green, J. M. 1971. High tide movements and homing behaviour of the tidepool sculpin Oligocottus maculosus. J. Fish. Res. Bd. Canada 2S(3) : 3S3-3s:t.

Hubbs, C. L. 1926. A revision of the fishes of the Subfamily Oligocottinae. Occ. Papers Mus. Zool., Univ. Michigan 7(171). 18p.

. 1927. Notes on the blennioid fishes of western North America. Papers

Michigan Acad. Sci., Arts, and Letters 7 : 351-394.

and P. S. Barnhart. 1944. Extensions of range for blennioid fishes in

southern California. Calif. Fish and Game 30(1) : 49-51. Johnston, R. F. 1954. The summer food of some intertidal fishes of Monterey

County, California. Calif. Fish and Game 40(1) :05-68. Jones, A. C. 1962. The biology of the euryhaline fish Leptocottiif; armatus >n mains

Girard (Cottidae). Univ. California Publ. Zool. 67(4) : 321-368. Jordan, D. S. and B. W. Evermann. 1*96. A checklist of the fishes and fish-like

vertebrates of North and Middle America. Rep. U.S. Comm. Fish. (1895), Appen- dix 5 : 207-5S4.

— . 1S98. The fishes of North and Middle America. Bull. U.S. Nat. Mus 47

(Part II) :1241-21S3. Mitchell, D. F. 1953. An analysis of stomach contents of California tide poo]

fishes. Amer. Midi. Nat. 49: 802-871. Moring, J. R. 1970. Use of the anesthetic quinaldine for handling Pacific coast

intertidal fishes. Trans. Amer. Fish. Soc. 99(4) : 802-805. Morris, R. W. 1960. Temperature, salinity, and southern limits of three spec

of Pacific cottid fishes. Limnol. and Oceanogr. 5(1) : 175— 179. Nakamura, R. 1971. Food of two cohabiting tide-pool Cottidae. J. Fish. Res. Bd

Canada IS (6) : 928-932. O'Connell, C. P. 1953. The life history of the cabezon, Scorpacnichthys marmora-

tus (Ayres). Calif. Dep. Fish and Game, Fish Bull. (93) : 1-76. Rutter, C. 1899. Notes on a collection of tide-pool fishes from Kadiak (sic) Island

in Alaska. Bull. U.S. Fish. Comm. 18(1898) :189-192. Schultz, L. P. 1936. Keys to the fishes of Washington, Oregon, and closely ad- joining regions. Univ. Washington Publ. Biol. 2 ( 4 ) : 103-22S. ■ . 1944. A revision of the American clingfishes, Family Gobiesocidae. with

descriptions of new genera and forms. Proc. U.S. Nat. Mus. 96(3187) : 47-77.

320 CALIFORNIA FISTT AXD GAME

Starks, E. C. and E. L. Morris. 1907. The marine fishes of southern California.

Univ. California Publ. Zool. 3(1) : 159-251. Villadolid, D. V. 1927. The flatfishes (Heterosoruata) of the Pacific coast of the

United States. Ph.D. dissertation, Stanford University, Stanford. 332p. Wales. J. H. 1952. Life history of the blue rockfish, Scbaslodes mystinus. Calif.

Fish and Game 38(4) : 4S5-19S. Williams, G. C. 1957. Homing behavior of California rocky shore fishes. Univ.

California Publ. Zool. 59 ( 7 ) : 249-2S4.

NOTES

TWO NEW SEA URCHIN-ACORN BARNACLE ASSOCIATIONS

On August 25, 1970, John Duffy. Bob Hardy, and Jack Ames col- lected a purple sea urchin, Strongyloccntrohis purpuratus (Stimpson), off Mussel Shoal, Ventura County, California. This 66 mm urchin, (Figure 1) living on a rocky substrate at a depth of 15 ft, had an acorn barnacle, Balanus concavus pacificus Pilsbry, attached to the surface of its test.

On October 12, 1970, Reinholt Banek, Fish and Wildlife seasonal aid, collected a red sea urchin, Strongylocentrotus franciscanus (Agassiz), 1 mile south of Davenport Landing, Santa Cruz County,

FIGURE 1. The acorn barnacle Balanus concavus pacificus attached to the purple sea urchin, Stronglyocentrotus purpuratus. Photograph by John Duffy and Jack Ames.

(321)

322

CALIFORNIA FISH AXD GAME

California. The urchin lest measured 324 mm (204 mm including spines), and was collected in a rocky, sandy area at a depth of 40 ft. All ached to the urchin was a large barnacle Balanus nubilis Darwin measuring 41 mm basal width (Figure 2).

FiGURE 2. The large barnacle, Balanus nubilis attached to the red sea urchin, Strongylo- cenfrotus franciscanus. Photograph by John Geibel.

To our knowledge only two reports of growths on urchins have been published. Strachan (1969) reported Balanus tintinnabulum cali- fornicus Pilsbry living on Lytechinus an am < sit* II. L. Clark. Boolootian (1958) reported the same species of barnacle living on the red sea urchin. St ran g ylocentrotus franciscanus (Agassiz).

The attachment of B. cmu-am* pacificus to S. purpuratus represents Hie first report of any barnacle attaching to 8. purpuratus. It also represents the first report of B. concavus pacificus attaching to any urchin. Strongyloci ntrotus franciscanus has been reported in the litera- ture as harboring the barnacle B. tintinnabulum califomicus. However, the attachment of B. nubilis to S. franciscanus represents the first re- port of B. nubilis attaching to any urchin.

NOTES 323

Since this note was submitted, two Department of Fish and Game biologists have called other urchin-barnacle associations to our atten- tion. K. A. Hardy noted three, S. purpuratus encrusted with unidenti- fied barnacles at Point Fermin San Pedro, and M. W. Odemar reported a large number of both S. purpuratus and 8. franciscanus harboring barnacles at Point Dume, Los Angeles County.

REFERENCES

P>oolootian, R. A. 1958. Notes on an unexpected association between a common

barnacle and eehinoid. So. Calif. Acad. Sci.. Bull. 57(2) :91-92. Light. S. F., li. I. Smith, F. A. Pitelka, D. P. Abbott and F. M. Weesner. 1964.

Intertidal invertebrates of the central California coast. U. C. Press. Berkeley.

446 p. Strachan, A. R. 1970. A white sea urchin-acorn barnacle enigma. Calif. Fish

and Game 56(2) :134-135.

James L. Houk and John M. Duffy. Marine Resources Region, Cali- fornia Department of Fish and Game. Accepted for publication February 1972.

NEW HOSTS AND BATHYMETRIC RANGE EXTENSION

FOR COLOBOMATUS EMBIOTOCAE

(CRUSTACEA, COPEPODA)

In July and August, 1971, the R/V Searcher, while otter trawling in Monterey and Bodega bays, for the California Academy of Sciences recovered 131 pink seaperch, Zalembius rosace us, and five spotfin surf- perch, Hyperprosopon anale. Eleven, or 13.4%, of the 82 adult pink seaperch were parasitized by the philichthyid copepod, Colobomatus cmbiotocae Nobel, Collard and Wilkes, 1969. Two of the five spotfin surfperch were parasitized by the copepod. The occurrence of C. em- biotocae on the pink seaperch and spotfin surfperch represents two new host records for this parasite, which was previously known from nine other species of embioticids (Nobel et al. 1969).

C. embiotocae is found under the thick cartilaginous skin covering the bony ridges of the head, and in the cephalic sensory canal system (Nobel et al. 1969). In this study, female copepods were recovered only from the preopercular section of the preopercular-mandibular canal (terminology follows Freihofer, pers. comm.) ; see Figure 1.

The pink seaperch occupies a habitat distinct from the characteristic intertidal and shallow subtidal inshore habitat of other embioticids. It occurs commonly between 15 to 50 or more fathoms, and rarely enters shallow water (Roedel, 1953). De Martini (1969) reported that the pink seaperch is a benthonic feeder, with the principal foods being gastropods and gammarid amphipods; pelecypods and small fish make up a minor portion of the diet. Examination of the stomach contents from six of the pink seaperch revealed a similar dependence on the benthos. Identi- fiable molluscan fragments included in the gastropods Nassarius meu- dicus, Olivcila sp. and Turbonilla sp., the scaphopods Cadulus fusi- formis, and Dentalium pretiosum, and the pelecypods Nucidana taphria

324

CALIFORNIA FISH AND GAME

and Lasaca cistula. Based on the feeding of the pink seaperch on ben- thonic molluscs, it seems reasonable to conclude that these fish were trawled on or near the bottom, thus extending the bathymetric range of C. cmbiotocac from near-shore shallow water (Nobel et al. 1969) to a depth of 40 fathoms.

FIGURE 1. Colobomatus embioiocae in the preopercular section of the preopercular-mandib- ular canal. Copepod is a little longer than normal.

Young of the year pink seaperch, perhaps 2 or 3 months old, were found at every station producing pink seaperch, except Station 3 (Table 1). At Station 2 (20 fathoms), young of the year pink seaperch composed over two-thirds of the pink seaperch taken at that station. Pink seaperch in this age group were not parasitized, and therefore have not been included in the calculation of the rate of infection. All of the copepods recovered were ovigerous females, except one from the pink seaperch and two from the spotfin surfperch. Male copepods were not observed. Equal infection rates were found at Monterey Bay (Sta- tion 1) and a month later at Bodega Bay (Station 3 and 4).

Spotfin surfperch are found typically along sandy beaches of the outer coast. Isaacson and Pool (1965) reported spotfin surfperch be-

NOTES

325

tween 25 and 37 fathoms in the Bodega Bay region. The R/V Searcher trawled five specimens in one tow off the mouth of Tomales Bay in 10 fathoms of water. Pink seaperch were not taken at this station.

Twelve copepods have been deposited in the Department of Inverte- brate Zoology, California Academy of Sciences, Golden Gate Park, San Francisco, California and three specimens in the Smithsonian Insti- tution.

TABLE 1 — Collection Data and Infection Rates of the Parasitic Copepod Co/obomafus emb/ofocae on the Pink Seaperch.

Station number	Number of pink seaperch trawled	Number of young of the year	Number of adults	Number of adults parasi- tized	Rate of infection %
Station 1 Monterey Bay, 4-6 miles WNW of Elkhorn Slough, 30-40 fathoms .	35 54 17 25	5 39 0 5	30 15 17 20	4* 2 2 3*	13.3
Station 2 Bodega Bay, 2.5 miles W of Dillon Beach 20 fathoms ..	13.3
Station 3 Off Bodega Bay, 3-4 miles W of Tomales Bay, 30 fathoms ._ . .	11.8
Station 4 Off Bodega Bay, 3 miles SW of Bodega Head, 40 fathoms ..	15.0
Total	131	49	82	11	13.4

* A bilateral infection was found on one fish from these stations.

ACKNOWLEDGMENTS

I am indebted to the Janss Foundation, Thousand Oaks, California, and to Dr. Earl Herald, Steinhart Aquarium, California Academy of Sciences, for the ship time aboard the R/V Searcher. Mr. Dustin Chi- vers and Mr. James Carlton offered many helpful suggestions through- out the preparation of this paper, and Mr. Allyn G. Smith kindly identified the molluscan material. Dr. "Warren C. Freihofer provided the name of the cephalic sensory canal.

REFERENCES

De Martini, Edward E. 1969. A correlative study of the ecology and comparative

feeding mechanism morphology of the Embiotocidae (surf -fishes) as evidence of

the family's adaptive radiation into available ecological niches. Wasman J. Biol.

27(2) : 177-247. Isaacson, Peter A. and Richard L. Pool. 1965. New northern records for the spotfin

surfperch. Hyperprosopon anale. California Fish Game 51(1) : 47. Nobel, Elmer R., Sneed B. Collard, and Stanley N. Wilkes. 1969. A new phili-

chthyid copepod parasitic in the mucous canals of surfperches (Embiotocidae).

J. Parasitol. 55(2) : 435-442. Roedel, Phil M. 1953. Common ocean fishes of the California coast. Calif. Dep.

of Fish and Game, Fish Bull. (91) : 1-184.

— Ernest W. Iverson, Skaggs Island, Sonoma, California 95476. Ac- cepted for publication April 1972.

326 CALIFORNIA FISH AND GAME

SOUTHERN RANGE EXTENSION FOR THE YELLOWFIN

GOBY, ACANTHOGOB1US FLAVIMANUS

(TEMMINCK AND SCHLEGEL)

Four specimens of the yellowfin goby, Acanthogo'bius flavimanus have been collected from Elkhorn Slough, Monterey County, California. Brittan, et al. (1963) first reported its occurrence in California waters from the San Joaquin River. Since this report, the yellowfin goby has spread throughout the San Francisco Bay-Delta area and has recently been reported from Bolinas Lagoon. (Brittan, et al. 1070).

The first specimen from Elkhorn Slough was collected by L. J. Hen- dricks of San Jose State College, 17 July 1970. It was collected with a beach seine in the northern arm of Elkhorn Slough near the Elkhorn Yacht Club. It has a standard length of 155 mm (196 mm tl) and is deposited in the fishes collection at San Jose State College (SJSC no. ES-39).

The second specimen was collected by Larry Wade of Moss Landing Marine Laboratories on 13 July 1971, by hand from Bennett Slough (a northern extension of the northern arm of Elkhorn Slough) near the locality of capture of the first specimen. The standard length is 177 mm (231 mm tl). It has been deposited in the fishes collection at Moss Landing Marine Laboratories (MLML no. ES-27).

«&>

Slpppp : ; mm

m^Jf !2! r3* '4' lV !6! '?' '8' 9! W hf V '13s W V:

I ' : :

FIGURE 1. Yellowfin goby, Acanthogobius flavimanus, 186 mm SL, collected from Elkhorn Slough near Kirby Park, Monterey County, by G. Victor Morejohn. Photograph by the author, March 1972.

The other two specimens were collected by G. Victor Morejohn of Moss Landing Marine Laboratories on 8 October 1971, by seine in the upper reaches of Elkhorn Slough near Kirby Park. Their standard lengths are 186 mm and 132 mm (235 mm and 162 mm tl) and also have been deposited at Moss Landing Marine Laboratories (MLML nos. ES-29 and T-75). Figure 1 is a photograph of the larger specimen (MLML no. ES-29).

NOTES 327

Brittan, et al. (1970) discussed three possible methods of introduc- tion of the yellowfm goby into Bolinas Lagoon from San Francisco Bay: migration, discard of baithsh and transport in a ship's sea water system. One or a combination of these methods is probably responsible for the introduction of the yellowfin goby into Elkhorn Slough. Since no vessels visiting the Orient anchor near the mouth of Elkhorn Slough, the source of the introduction may be a natural dispersion of tbe species population of San Francisco Bay.

ACKNOWLEDGMENT

Support for this note was provided by NOAA Office of Sea Grant, Department of Commerce at the Moss Landing Marine Laboratories of the California State Universities.

REFERENCES

Brittan, Martin R., Arnold B. Albrecht, and John B. Hopkirk. 19G3. An oriental goby collected in the San Joaquin River Delta near Stockton, California. Calif. Fish Game 49(4) : 302-304.

Brittan, Martin R., John D. Hopkirk, Jerrold D. Conners, and Michael Martin. 1970. Explosive spread of the oriental goby Acanthogobius flavimanus in the San Francisco Bay-Delta region of California. Proc. Calif. Acad. Sci., ser. 4, 38(11) : 207-214.

— Gary E. Kuhowski, Moss Landing Marine Laboratories, Moss Land- ing, California 95039. Accepted for publication April 1972.

CALIFORNIA CONDOR SURVEY, 1971

The seventh annual California condor (Gymnogyps calif ornianus) survey was conducted on October 13 and 14, 1971. Survey methods and evaluation procedures were essentially the same as has been re- ported in past surveys (Mallette, et al., 1966, 1967, 1970, 1972, Sibley, et al., 1968, 1969). Eighteen observation stations were manned during the survey, an increase of two over 1970 ; however, the number of observers remained the same at 45. Observation stations were manned from noon until condor activity ceased, normally around 5 :00 pm. No baiting was attempted. Weather on October 13, was low overcast and fog in the Tehachapi Mountain area and clear skies in the Sespe Condor Sanctuary and vicinity. Winds were calm. Temperatures ranged from 68 F on Frazier Mountain to 98 F in the Sespe Condor Sanctuary. Weather on October 14, was clear with gusty winds to 20 mph in the Tehachapi Mountain area with visibility of zero in the Sespe Condor Sanctuary due to fog. Temperatures ranged from 60 F on Frazier Mountain to 77 F on Grapevine Peak.

An analysis of the sightings was made to eliminate duplication and indicated a minimum of 29 individual condors were seen on October 13, and 34 on October 14. The age structure for October 14, was 28 adults, 4 immatures and 2 unclassified. Only the Tehachapi portion of the population was counted as no sightings were reported from the Sespe Condor Sanctuary on October 14. It is believed no exchange of

::in

CALIFORNIA FISH AND OA.MK

birds occurred between these two areas on this day. Other raptors observed were :

Numbers

Species

Turkey vulture (Cathartcs aura)

Golden eagle (Aguila chrysaetos)

Sharp-shinned hawk (Ascipiter striatus)

Cooper's hawk (A. cooperi)

Red-tailed hawk {Buteo jamaicensis)

Swainson's hawk (B. swainsont)

Ferruginous hawk (B. regalis)

Sparrow hawk (Falco spanerius)

Prairie falcon (F. mexicanus)

Peregrine falcon (F. peregrinus)

Marsh hawk (Circus cyaneus)

A comparison of the data collected over the past seven surveys indi- cates that the condor population has remained fairly constant during this time. Fluctuations of the total number seen during the 1065 through 1971 surveys is more an indication of weather conditions during the survey days than it is of any major changes in the actual population structure.

REFERENCES

Mallette, Robert D. and John C. Borneman, 1966. First cooperative study of the

California condor. California Fish Game 52(3) : 1S5-203. Mallette, Robert IX, John C. Borneman, Fred Sibley and Raymon S. Dalen. 1967.

Second cooperative survey of the California candor. California Fish Game 53(3) :

132-145. Mallette, Robert D., Fred C. Sibley, W. Dean Carrier and John C. Borneman, 1970.

California condor surveys, 1969. California Fish and Game 56(3) : 199-202. Mallette, Robert D., Sanford Wilbur, W. Dean Carrier and John C. Borneman, 1972.

California condor survey, 1970. California Fish Game 58(1) : 67-68. Sibley, Fred C, Robert D. Mallette, John C. Borneman and Raymond S. Dalen,

1968. Third cooperative survey of the California condor. California Fish Game 54(4) : 297-303.

Sibley, Fred C, Robert D. Mallette, John C. Borneman, and Raymond S. Dalen,

1969. California condor surveys, 1968. California Fish Game 55(4) : 298-306.

-W. Dean Carrier, U.S. Forest Service; Robert D. Mallette, Califor- nia Department of Fish and Game; Sanford Wilbur, Bureau of Sport Fisheries and Wildlife; John C. Borneman, National Audubon Society. Accepted for Publication July 1972. Supported by Federal Aid to Wildlife Restoration Project W-54-R "Special Wildlife In- vestigations." Prepared for and with approval of the Condor Tech- nical Committee.

BOOK REVIEWS

Mountain Sheep: A Study in Behavior and Evolution.

by Valerius Geist; University of Chicago Press, 1971; 383 p. $14.50

Valerius Geist used the behavior of mountain sheep as a tool to study the animal; thus this in not just another case of using the animal to study behavior. It is a book that deals with the evolutionary forces that have shaped the sheep's behavior pat- terns and developed the animal to what it is today.

There is something about mountain sheep that is extremely fascinating and chal- lenging to study. Once exposed to this animal in the wild you are "hooked," something akin to gold fever. Whether you are a "hooked" student, photographer, or hunter of mountain sheep or not. I recommend this book. You will find his approach unique. He has deliberately slighted some ecological factors and management or conservation considerations, instead concentrating on developing a comprehensive theory of mountain sheep evolution. Yet the information developed will be very useful in giving administrators the understanding of the animal necessary in devel- oping and implementing necessary management programs.

If you are not a professional wildlifer and you think the graphs, charts, and formula are beyond your comprehension, do as the author suggests : first read Chapter 1 and the conclusions in Chapter 12, then Chapters 5 and 11, and then the introductions to the remaining chapters.

I waited anxiously the availability of this book for I had the good fortune of discussing sheep with Dr. Geist in Bishop, California, wliere he was the invited speaker of the Desert Bighorn Council in April 1970. I also got to view some of the 16 mm film taken during his studies. He mentions these films in the preface where information is given on how to purchase or borrow them. I believe these films should be viewed to augment the knowledge you can gain from the book. The 89 black and white plates in the book give you an indication of the quality of the pictures taken during his 3^ years in the field with the Stone, the Dall, and Rocky Mountain bighorn. The pictures, all taken at close range, will also let you appreciate the incredible feat of conducting a study of wild free-ranging mountain sheep at all seasons of the year.

This book will be the sheep biologists' bible for many years to come. His methods for recording quantitative behavioral data with a minimum of subjective evaluation will be copied by many. For those of you who will buy the book and use it as reference there is a very good index.

Dr. Geist has answered for me in Chapter 5, Tradition and Evolution of Social System, a paradox that I have been at a loss to explain. Wild sheep as a group have been very successful, spreading during the Pleistocene through most of the mountain ranges of the northern hemisphere. However, today sheep maintain their distribution as a living tradition and rarely will they fill empty suitable habitat. Today sheep in the western United States survive in a situation far different from that of the ideal bighorn habitat of posl glacial period during which they evolved and developed behavior patterns. Sheep habitat is not as continuous today as it once was. Young sheep follow adults and adopt their habitat. Natural selection would be against dispersing animals wandering into unsuitable habitat.

— R. A. Weaver

Fishes of Montana

By C. D. J. Brown; Big Sky Books, Montana State University, Bozeman, Montana; 1972. 207 p. Illustrated. $4.50 paper.

There is a wealth of information in this book, of use to the professional fisheries worker as well as the interested angler or student. The information is logically presented. Short introductory sections describe the history of fish collections in Montana and the rivers, lakes and reservoirs and present a map of Montana show- ing major waters, and notes on the preservation of fish and the use of keys. There is a brief but complete glossary which identifies the few scientific terms used in language easily understood by the layman. Diagrams of a cutthroat trout and a largemouth bass are used to identify the major characters used in the keys.

t :!2!t I

330 CALIFORNIA FISH AND GAME

There are keys to both family and genus, including all known Montana fishes. The keys arc easy to use. with sketches showing outstanding features.

Eighteen families and 50 genera are included in the book. Information for each species includes: ii) both common and scientific names, (ii) a drawing or black- and-white photograph, i iii > a map of Montana showing collection locations, (iv) a brief description, (v) native range and. if not native to Montana, details of intro- ductions, i vi I life history information (age and growth, spawning information, and food habits) and, (vii) a brief paragraph on the species abundance and importance for sport or forage. The book concludes with a reference section, an index of com- mon and scientific names, a map of the major drainages of Montana, and a listing of all species discussed.— IT. A. Hashagen, Jr.

Remembrances of Rivers Past

By Ernest Schweibert; The McMillan Company, N.Y., 1972. 287 p., illustrated. $6.95.

This is the book for those long winter evenings next to the fire, when trout season is several months away. "Remembrances of Rivers Past" is a collection of 25 short stories about fly fishing for trout and salmon on rivers and streams throughout the world. The stories are Schweibert' s reminiscences of past fishing experiences, some from his childhood, some quite recent. The waters he has fished are among the most famous waters of the world: the Little Manistee. Schoharie. Beaverkill, Neversink, Esopus, the Firehole, and the Letort. Salmon were taken in Norway. Labrador, and Iceland and huge trout in Tierra del Fuego and Argentina. The stories are well written, describing not only the fish caught or lost, but also his companions, native customs, and the history of the area. The reader can feel he is right with Schweibert. and considering some of the fish taken, certainly must wish he was. Many of the stories have been published previously in sporting magazines under different titles. In almost every story the author expresses concern for the environment, citing examples of prime waters destroyed by pollution, dams, or poor management. His obvious knowledge of trout, stream entomology, and tackle add to the authenticity of his recollections. — K. A. Hashagen, Jr.

A Trout and Salmon Fisherman for Seventy-Five Years

By Edward R. Hewitt; Van Cortlandt Press, Croton-On-Hudson, N.Y., 1972; XXIV + 338 p. illustrated. $8.50.

Seventy-live years of a life devoted to fishing and. as quickly becomes apparent, little time spent on matters not related to fish or fishing. Obviously wealthy and opinionated. Mr. Hewitt records his observations on fish and fishing. He conducted detailed experiments on fish vision, color perception, and physiology. Tackle and techniques were developed, tested and modified. Anecdotes from his fishing diary point out the success of his methods and. usually, the deficiencies of other methods.

A Trout and Salmon Fisherman for Seventy-Five Years was originally published as two volumes. Tilling on the Trout and Secrets of the Salmon, but they were com- bined in lit is and revised and updated with knowledge and anecdotes from the 20 years between publication dates. It was reprinted in 1966 and now again in 1972. Much of the information, particularly on tackle, is now outdated but still interest- ing from a historical standpoint. Some of Mr. Hewitt's observation on behavior and life history have been modified by the research of professional biologists: how- ever, there is much information in this hook, much to think about, many new tech- niques to try next time the fish aren't rising. Most of the book pertains to fly fishing and much to dry fly fishing for Atlantic salmon. I think that any fisherman reader will find the book informative and interesting. — K. A. Hashagen, Jr.

The Dry Fly and Fast Water and The Sclmon and the Dry Fly

By George M. L. LaBranche; Van Cortland Press, Croton-On-Hudson, N.Y., 1972. 252 p. $6.95.

Tfn Dry Fly and Fust Water was first published in 1014; The Salmon and the Dry Fly in 1924. In 1951 they were combined into one volume and reprinted. Now. over 20 years later, another unrevised edition has been printed.

Tt is necessary to know the past history of these two short books in order to appreciate them. In The Dry Fly and Fast Water, LaBranche tells of his experiences

REVIEWS 331

in learning to fish the dry fly. He fished with large rods, silk lines, and gut leaders and began using the dry fly when it was considered a novelty in the U.S. His ob- servations on fish behavior, stream conditions, and fly presentation are valid today — and are often touted as "new" information by more recent authors.

The Salmon and the Dry Fly is less than a hundred pages long and tells how he and his associates pioneered techniques and tackle to successfully take Atlantic salmon on dry flies. He relates pertinent experiences to illustrate his points and concludes with a comprehensive chapter on "Casting the Curve", which includes de- tailed instructions for this rather difficult maneuver.

Both books are written in a rather stilted, wordy fashion but the information is there. Both are considered classics in the history of fly fishing, and I feel they are a welcome addition to any fisherman's library.- — K. A. H ash a gen, Jr.

Hardy's Book of Fishing

By Patrick Annesley (Editor); E. P. Dutton and Co., Inc., N.Y., 1972; 304 p., illustrated. $16.50.

As any serious fisherman knows. Hardy Bros, of England is one of the most famous tackle manufacturers in the world and has been for over 100 years. During this period they have issued catalogs describing their fine rods, reels, and other fishing tackle. In addition to the product information, there have been helpful articles for the angler and testimonials from satisfied customers.

Hardy's Book of Fishing is the best of the catalogs. Loosely arranged in three sections — "The Equipment", "Fish and Fishing", and "The Angler at Large" — with each section further arranged chronologically, the book provides a history of English angling and the development of fishing tackle and delightful fishing stories and advice. The book is attractively illustrated with reproductions of illustrations of fish and tackle and "how to" diagrams. Much of the advice given is still valid today.

The language for the most part, is delightfully wordy. The readers of early Hardy Bros, catalog were told "How to Fish" and "How to Tell a Salmon Pool". A detailed article describes "Spinning and Prawning for Salmon". In 1SSS there was the question "Wet Fly or Dry?" A question which is still debated today. The rods ranged from "light" trout rods of 9 ft to double-handled, 20-ft salmon rods. Gillies "grassed" fish and the catch was recorded in hundreds of pounds.

The "Angler at Large" section describes fishing in the late 1800's and early 1900's in Norway, Finland, Tasmania, Kashmir, New Zealand, North America and other countries.

I found the book very entertaining and spent many enjoyable evenings reading Hardy's Book of Fishing. — K. A. Hashagen, Jr.

The Art and Science of Fly Fishing (revised edition)

By Lenox H. Dick; Winchester Press, New York; 1972. 169 p., illustrated. $6.95.

Both novice and advanced fly fishermen will benefit from reading "The Art and Science of Fly Fishing". As in the first edition, the book begins with a section on "Fundamentals", with chapters on basic tackle, casting, fly presentation, reading water, entomology, and flies. The second part has four comprehensive chapters on "Stream Tactics", well illustrated with figures and black and white photos. In the second part the author utilizes the information presented in the first chapters to take the reader on four fishing trips where all basic stream situations, water condi- tions, and casting techniques are encountered. This method is effective and for the most part the information is clearly and interestingly presented. The final section, new in this edition, consists of eight chapters on "Salmon, Steelhead, and Others".

Most of the information is accurate; however, experienced anglers will take excep- tion to some of the author's opinions and statements. Occasionally, I felt basic terms, such as "drag", were not explained sufficiently for the novice. The author also describes 4X tippet as "quite fine", which doesn't help the beginning fly fisher- man. Chapters on "Cutthroat Trout", "Silver Salmon", and "Jacks or Orilse Fish- ing" are so brief they make the reader wonder why they were included. In addition to these negative comments, I must mention the numerous typographical errors, deletions, incorrect references to figures, plates and page numbers, and occasional misspellings. They definitely detract from what otherwise is an interesting and informative book. — K. A. Hashagen, Jr.

332 CALIFORNIA PISH AND GAME

World Dynamics

By Jay W. Forrester; Wright-Allen Press, Inc., 238 Main Street, Cambridge, Massachusetts 02142, 1971; 142 p.

This book is likely the most advanced treatment of the Malthusian argument. Professor Forrester, unlike Malthus, has cybernetics, computer technology and a greater arraj of information with which to predict the future. This hook's greatest value will be in stimulating further development of predictive models of the world social system. Such models will hopefully assist the world in making the transition from growth and "progress" to economic and social stability.

Forrester and his colle-i-ues at MIT constructed a world model of ."» system level variables which are Population. Pollution. Capital Investment. Agriculture Capital Investmenl Fraction, and Natural Resources. Each system level is controlled by other system levels and assumed rate functions which have negative or positive [backs. Secular trends under various assumptions and inputs are lucidly presented by graphs and the accompanying text.

The results are generally depressing. Current trends of population growth will mosl likely only be altered by a dramatic rise in the death rate as the limits of the earth are exceeded in terms of carrying capacity. A pollution crisis, dwindling natural resources, crowding or food shortages may bring the population explosion to a catastrophic halt. Birth control programs are not likely to have enough everage to effectively forestall disaster only delay it.

This book should be mandatory reading for those concerned about the future and man's fate on this planet. This should include all resource biologists and espe- cially the resource administrators who set policies, priorities and budgets. The mate- rial is intellectually stimulating and helps to quantify what many of us observe to be happening to the planet. Although Forrester's model is simplistic it will un- doubtedly serve as a framework for constructing more sophisticated models as we increase our knowledge of social systems. — Lev W. Miller.

The New York Aquarium Book of the Water Worid

By William Bridges; American Heritage Publ. Co., N.Y., 1970; 287 p., illustrated. S6.95.

Bridges gives a brief look at some of the numerous animals that live in or are closely associated with the aquatic environment. The book includes extensvie. pro- fessional photographs with non-technical descriptions of one-celled animals, amphib- ians, reptiles, birds, mammals and invertebrates (excluding insects).

The main theme concerns animals (mainly fishi that can be kept in aquari. Starting with the Chinese, credited as the first to keep fish for observation, a condensed history of fish keeping is covered. During the Sung dynasty. 960— 127S, fancy goldfish and carp were held in porcelain vessels. The ancient Romans confined marine fish in ponds connected to the ocean.

Among the oddities covered i> the walking catfish, C. hairachus and its establish- ment info Florida's waters. Bridges presents this as a rather casual observation without pointing out the real dangers involved to native species with the introduc- tion of exol ics.

In a book of this nature it was disappointing not to find real concern with the continued premature extinction of many species. 1'nless the problem is recognized and meaningful accomplishments are made, there won't be many species left to view, even in public aquari. — James A. si. Amant.

INDEX TO VOLUME 58

AUTHOR

Alton. Miles S. and Christine J. Black- bum : Diel changes in the vertical distribution of the euphausiids, Thy- sanoessa spinifcra Holmes and Eu- phausia pacifica Hansen, in coastal waters of Washington, 179-190

Azevedo, John A., Jr., Eldredge G. Hunt and Leon A. "Woods, Jr. : Melanistic mutant in ringneck phea- sants, 175-178

Barlow. George W.. and Victor L. De Vlaming: Ovarian cycling in long- jaw gobies, Gillichthys mirabilis, from the Salton Sea. 50-57

Barnhart, Roger A.: see Kesner and Barnhart. 204-220

Benville, Pete E.. Jr.: see Earnest and Benville, 127-132

Blackburn, Christine J. : see Alton and Blackburn, 179-190

Borneman. John C. : see Carrier. Mal- lette, Wilbur and Borneman. 327- 328; see Mallette, Wilbur. Carrier and Borneman, 67-68

Briggs, Kenneth T.. and C. William I >avis : A study of predation by sea lions on salmon in Monterey Bay, 37^3

Bury, C. Bruce : The effects of diesel fuel on a stream fauna. 291—295

Cappueei. D. T., Jr. and W. M. Long- hurst : Rabies in deer, 111-144

Carrier, W. Dean : see Mallette, Wil- bur, Carrier and Borneman, 67-68

Carrier. W. Dean. Robert D. Mallette. Sanford Wilbur and John C. Borne- man : California condor survey, 327- 328

Castle, William T. and Leon A. Woods, Jr. : DDT residues in white croakers, 198-203

Chamberlain. Lawrence L.: Primary productivity in a new and older Cali- fornia reservoir, 254-267 ; see Lasker, Tenaza and Chamberlain, 58-66

Chen, Lo-Chai: see Rosenblatt and Chen, 32-:!(i

Clark, William E.: see Espinosa and Clark, 149-1 .".2

Davis. C. William: see Briggs and Davis, 37-43

Day. John S. : see St. Amant and Day, L54 -155

Demory. Robert L. : Tailless Dover sole from off the Oregon coast. 147- 148

De Vlaming, Victor L. : see Barlow and De Vlaming, 50-57

Dexter, Deborah M. : Molting and growth in laboratory reared phyllo- si unes of the California spiny lobster, Panulirus interruptus, 107-115

Duffy, John M. : see Honk and Duffy,

::2i-323

Earnest, Russell D. and Pete E. Ben- ville. Jr. : Acute toxicity of four or- ganochlorine insecticides to two species of surf perch, 127-132

Elliott, George V. and T. M. Jenkins, Jr.: Winter food of trout in three high elevation Sierra Nevada lakes, 231-237

Espinosa. Lawrence R. and William E. Clark : A polypropylene light trap for aquatic invertebrates. 149-152

Farley. David G. : A range extension for the logperch, 248

Fitch. John E. : A case for striped mul- let. Mugil cephalus, spawning at sea, 246-21S; The cottonmouth jack, Uraspis secunda, added to the marine fauna of California. 245-24(!

Franklin, George W. : see Schneegas and Franklin. 133-140

Grover, Charles A.: Population differ- ences in the swell shark Cephalos- cyllium ventriosum, 191-197

Haaker, Peter L. : First record of a reversed butter sole. Tsopsetta isol-e- pis. 244-245

Hanson, Jack A.: Tolerance of high salinity by the pileworin. Neanthes succinea, from the Salton Sea, Cali- fornia. 152- lot

Hashagen, Kenneth A., Jr. : see Raw- stron and Hashagen. 221-230

Hawthorne. Vernon M. : Coyote food habits in Sagehen Creek basin, north- eastern California. 5-12

Hobson, Edmund S. : The survival of Guadalupe cardinalfish Apogon gua- dalupensis at San Clemente Island. 68-(i!»

Honk. James L. and John M. Duffy : Two new sea urchin — acorn barnacle associations, 321-323

Hunt. Eldredge G. : see Azevedo. Hunt and Woods, 175-1 7,s

( 333 )

334

CALIFOKM \ PISB AM) GAME

[verson, Ernesl AN'.: New hosts and bathymetric range extension for Colo- bomatus > mbiott ,,■„,■ (Crustacea, Co- pepoda I . 323 325

Jenkins, 'I'. M.. Jr.: see Elliott and Jenkins, 233 2::7

Jones, All.. 11 C. : Contributions to the life history of the Piute sculpin in Sagehen Creek, California, 285 290

Kesner, William I ». and Roger A. Barn- hart: Characteristics of the fall-run '<! troul i Salmi, gairdneri gairdneri) of the Klamath River sys- tem with emphasis on the half- pounder, 204-220

Knaggs, Eric II.: xre Parrish and Knaggs, L3 2] ; Southern California Pacific mackerel fishery and age com- position of commercial landings dur- ing the 190K-09 and 1069-70 seasons, 116 L20

Kukowski, Gary E. : Southern range ex- tension for the yellowfin gohy, Acan- tkogobius fiavimanus I Temminck and Schlegel), 326-327

Lasker, Reuben, Richard H. Tenaza, and Lawrence L. Chamberlain : The response of Salton Sea fish eggs and larvae to salinity stress, 58-66

Lea, Robert X.: Southern geographical records for four surfperches, family Embiotocidae, with notes on a popu- lation resurgence of the sharpnose seaperch, 27-.'!1

Longhurst. W. M. : see Cappucci and Longhurst. 141-144

Mais. Kenneth F. : A subpopulation study of the Pacific sardine. 296-314

Mallette. Robert D. : see Carrier, Mal- let te, Wilbur and Borneman, 327- 328

Mallette, Robert D.. Sanford Wilbur, W. Dean Carrier and John C. Borne- man : California condor survey, 1970 67 68

Martin. Michael: Morphology and variation of the Modoc sucker, Catos- tomus microps Rutter. with notes on feeding adaptations, 277-284

Miller, Lee W. : Migrations of sturgeon la — ed in the Sacramento-San Joa- quin estuary. 102-100; White stur- geon population characteristics in the Sacramento-San Joaquin estuary as measured by tagging, 94-101

Moring, John R. : Check list of inter- tidal fishes of Trinidad Bay, Cali- fornia, and adjacent areas. 315-320

Morrell, Stephen: Life history of the San Joaquin kit fox. 162-174

Odenweller, Dan Bowman: A new range record for the umbrella crab, Crypto- lithodes sitchensis Brandt. 240-243

Parrish, Richard II.: Symbiosis in the blacktail snailfish, Careproctus me- I" minis, and (he box crab. Lopho- lithodes foraminatus, 239-240

Parrish, Richard II.. and Eric II. Knaggs: The southern California Pacific mackerel fishery and age com- position of the catch for the 1964- 65 through 1967-68 seasons, 13-21

Penhale, Leonard B. : Reproductive failure of pelagic cormorant, San Luis Obispo County, California, 1970, 238

Rawstron, Robert R. : Harvest, sur- vival, and cost of two domestic strains of tagged rainbow trout stocked in Lake Berryessa, Califor- nia, 44—49; Nonreporting of tagged largemouth bass, 1966-1969, 145-147.

Rawstron, Robert R. and Kenneth A. Hashagen, Jr. : Mortality and sur- vival rates of tagged largemouth bass ( Hicropterus salmoides) at Merle Collins Reservoir, 221-230

Rosenblatt, Richard IL. and Lo-Chai Chen : The identity of Sebastes bab- cocki and Sebastes rubrivinctus, 3°- 36

Schneegas, Edward R. and George W. Franklin : The Mineral King deer herd, 133-140

Shaw, Stanton B. : DDT residues in eight California marine fishes, 22-26

Spratt, Jerome D. : A.ge and length composition of northern anchovies, ]'n ani ulis mordax, in the California anchovy reduction fishery for the 1969-70 season, 121-126

St. Amant, James A. and John S. Day : Range extension of Palaemonetes paludosus (Gibbes) in California, 154-155

Tenaza, Richard H. : see Lasker, Tenaza and Chamberlain, 58-66

Varoujean, Daniel H. : The reoccur, rence of the California scorpionfish, Scorpaena guttata Girard, in Mon- terey Bay, 238-239

von Geldern, C. E., Jr.: A midwater trawl for threadfin shad, Dorosoma petenense, 26S-276 ; Angling quality at Folsom Lake, California, as deter- mined by a roving creel census, 75-93

Wilbur. Sanford: see Carrier, Mallette, Wilbur and Borneman. 327-328; see Mallette, Wilbur, Carrier and Borneman, 67-68

Woods, Leon A., Jr. : see Azevedo, Hunt and Woods, 175-178; see Castle and Woods, 198-203

INDEX

335

SCIENTIFIC NAMES

Acanthogobius flavimanus: 326-327 Accipiter cooper ii: 07

striatus: G7 Acipenser medirostris: 102-106

transmontanus: 94—101, 102-106 Alces alces: 141 Amphistichus argenteus: 27 Anisotremus davidsoni: 58-66 Anoplopoma fimbria: 22—26 Apogon guadalupensis: 6S-69

retrosella: 68 Aquila chrysaetos: 67 Arctostaphylos sp.: 137 Artemia sp.: 107-115 Astacus sp.: 202 Bairdiella chrysura: 65

icistia: 58-66 Balanus concavus pacificus: 321-322

nubilis: 321-322 Buieo jamaicensis: 67 Cadulus fusiformis: 323 ('a a is luf runs: 5-12 Capreolus capreolus: 111 Careproctus melanurus: 239-240 Cathartes aura: 67 Catostomus microps: 277—284 platyrhynchus: 287 rimiculus: 292 tahoensis: 287 Cephaloscyllium ventriosum: 191-107 Cervus canadensis: 141

elaphus: 141 Chaenogobius urotaenia: 54 Chara, sp.: 154

Chasmichthys dolichognathus: 54 Chromis punctipinnis: 30 Circus cyaneus: 67 Citellus beecheyi: 5-12 beldingi: 5—12 lateralis: 5-12 Citharichthys sordidtis: 22-26 Cladophora sp.: 294 Clemmys marmorata: 294 Clostridium botulinum: 149 Colobomatus embiotocae: 323-325 Coryphaeiioides acrolepis: 22-26 Ooi!fMS beldingi: 285-290 Crago sp.: 94

Cryptolithodes sitchensis: 240-243 Cymatogaster aggregata: 2S. 127-132 Cynoscion xanthulus: 5S-66, 152 Dama dama: 141 Dentalium pretiosum: 323 Dorosoma petenense: 44. 92, 268-276 Echidnophaga gallinacea: 170 Engraulis mordax: 121-126 Entosphenus tridentatus: 292 Eopsetta jordani: 22-26 Erethizon dorsatum: 5-12 Euarctos americanus: 137 Eumetopias jubata: 38

Eupliausia pacifica: 179-100 I', ii !a niias sp.: 5-12 Falco mexicanus: 07

sparverius: 07 Fasciola hepatica: 142 Fundulus confluentus: 55 Genyonemus lineatus: 22-26, 19S-203 GUlichthys mirabilis: 50-57 Glaucomys sabrinus: 5-12 Gobius paganellus: 51 Gymnogyps calif ornianus: 67-68, 327-

328 ' Hippcampus zosterac: 55 Hyperprosopon anale: :V23

ellipticum: 27 Ictalurus catus: 78-93 melas: 7N-93 iiihiilosus: 7s '.)."> Isopsetta isolepis: 244-245 Lasaea cistula: 324 Lepomis cyanellus: 78-93 macrochirus: 75-93 microlophus: 7*-93 Lepus americanus: 5-12

sp.: 5-12 Lipoptena depressa: 142 Lopholithodes foraminatus : 239-240 Lytechinus sp.: 109 Macrocystis sp.: 6S Marmota flaviventris: 5-12, 136 Medialuva californiensis: 30 l/< phitis mephitis: 143 Mergus merganser: 294 Micrometrus minimus: 127-132 Micropterus dolomieu: 75-93

salmoides: 75-03. 145-147. 221-230 Microstomas pacificus: 147-14S Microtus sp.: 5-12 J///-//'/ cephalus: 246-248 Mi/til us sp.: 107-115 Nassarius mendicus: 323 Neanthes succinea: 152-154 Neolipoptena ferrisi: 142 Nuculana taphria: 323 Odocoihus In in ion us: 141 californicus: 133-140 ruin mhiunus: 141—143 virginianus: 141 Olivella sp.: 323 Oncorhynchus nerka: S7

sp.: 37—13 Ophiodon elongatus: 22-26 Palaemonetes paludosus: 154-155 Panulirus intcrruptus: 107-115 Parophrys vetulus: 22-26 Percina caprodes: 248 I'crom i/si-us sp.: 5 12 Phalacrocorax pelagicus: 23S Phanerodon atripes: 2S-31

furcatus: 30 Phasianus colchicus torquatus: 175-178

336

CALIFOKMV FISH AND GAME

Phoradendron villosum: 136 /'"( cilia latipinna: 15 I

I 'am n.ris mi a a la ris: S6

n igromaculatus: 86 Prosopium williamsoni: 287 /'/( rogobius < lapoid< s: 5 I

I'll It. r si HI II III us: 1 12

/.'(/// 1/ boylei: 29 I

I'n ntii it r tarandus: 1 1 1

Rhacochilus toa otes: 28

rtit-fti: .",11

It'll in ifhtln/s osculus: 2S7, 202 Richardsonius egregius: 2V7 Sti!i,i„ clarki henshawi: 44

gairdneri: 44 49, 7^ 93, 231-237, 2S5, 292

p. gairdneri: 204-220

/,■»//</: 2S5 Salvelinus fontinalis: 231-237, 285 Sardinops caeruleus: 29G-314 Scomber japonicus: 13-21, 116-120 Scorpaena guttata: 238-239

Sri/lhi rus americanus: 107 Sebastes babcocki: ::2-36

rubririnrl ns: .'!"_' .",0 Spirogyra sp.: 294

Strongylocentrotus franciscan us: 321- 322

p ti rp ii nil us: 321-322 Sylvilagus sp.: 5—12 Tamiasciurus douglasi: 5—12 Thamnophis couchi: 294 1 In hr.iti calif or niensis: 142 Thysanoessa spiiiifcru: 179 T.in Tilapia mossambica: 154 Trachurus symmetricus: 1'\. 22-26. 116 Tubifex sp.: 109 Turbonilla sp.: 323 ? raspis secunda: 245—246 L'rocyon rinrrroiirticntiiis: 143 \ iil/its macrotis mutica: 162-174 Zalembius rosaceus: 323 Zalophus California n us: 37—43 Zygnema sp.: 2! 1 1

SUBJECT

Auf: of anchovies taken the 1969 70 season, 121—126; of Klamath River fall-run steelhead, 204-220; of Paci- fic mackerel during 1964-6o through 1967-68 seasons, 13-21; of Pacific mackerel in southern California fishery. 1968 69, 1969-70, 116-120

Aldrin: toxicity to surfperch, 127- 132

Anchovy, northern: fishery during 1969-70 season, 121-126

Angling: qualitj al Folsom Lake, 75—93

Bairdiella: effect of salinit,\ mi • u_- and larvae. 58-66

Barnacle, acorn: association with sea urchins, 321-323

Bass: fishery at Folsom Lake. S3

Bass, largemouth: mortality and sur- vival at Merle Collins Reservoir, 221- 230; nonreporting of tagged fish, 1 15 1 17

Birds: as fund for coyotes, 9

Cardinalfish, Guadalupe: survival at San Clemente Island. 6S-69

Catfish: fisherj al Folsom Lake, S4— 85

< !attle: as feud for eoj otes, 7

Census, roving creel: of fishery at Folsom Lake 75- 93

Check list: intertidal fishes of Trinidad Bay and vicinity. 315-320

Chlorinated hydrocarbons: residues in California marine fishes, 22 26

Coleman Kamloops trout strain: per- formance at Lake Berryessa, 11 19

Condor, California: 1970 population . 67 68 ; population survey of 1971, 327-328

Copepod, parasitic: on embiotocid fish, 323 325

Cormorant, pelagic: reproductive failure in San Luis Obispo County coastal area. 23S

Costs: of planting two domestic trout strains at Lake Berryessa, 44—49

Coyote: food habits in Sagehen Creek basin, 4—12

Coyote, juvenile: food habits at Sage- hen Creek basin, 9-11

Crab, box: case of symbiosis with blacktail snailfish, 239-240

Crab, umbrella: range extension, 240- 243

Croaker, white: DDT residues. 22-26. 198-203

Cycling, ovarian: of gobies at the Salton Sea, 50-57

DDT: in the diet of pheasants produc- ing melanistic mutants, 17-1-17S; residues in California marine fishes. 22-26 : residues in white croakers, 198 203; toxicity to surfperch. 127- 132

Deer: as food for coyotes, 0—7; occur- rence of rabies, 141-144; study of .Mineral King herd, 133-140

Dieldrin: toxicity to surfperch, 127-132

Disease: occurrence of rabies in deer, 141-144

Distribution, geographic: appearance of California scorpionfish in Monterey Bay. 23s 239 ; extension of range of yellowfin goby, 320-327; of flag and redbanded rockfishes, 32-36; of the cottonmouth jack. 245-246; of the logperch, 248 : of the umbrella crab, 240-243 ; of white and green stur- geons, 1(12 100; range extension for four surfperches, 27-31; range ex-

INDEX

337

tension of Palaemonetes paludosus, 154-155

Distribution, vertical: of euphausiids off the coast of Washington. 179-190

Eggs, fish: effect of salinity on Salton Sea fish, 5S-G6

Elkhorn Slough: yellowfin goby occur- rence, 320-327

Embiotocidae: geographical records for four species, 27-31

Endrin: toxicity to surfperch, 127-132

Estuary, Sacramento-San Joaquin: white sturgeon population study, 94- 101 ; migration study of white and green sturgeons, 102-100

Euphausiids: diel changes in the ver- tical distribution, 179-190

Fauna, stream: effects of diesel fuel, 291-295

Feeding: of spiny lobster phyllosomes in the laboratory, 107-115

Fishery: characteristics at Folsom Lake, 75-93; for northern anchovy during 1909-70 season. 121-126; for trout at Lake Berryessa, 44-49; for white sturgeon, 99-190 ; of Pacific mackerel in southern California 1964—65 through 1967-68 seasons, 13-21 ; of Pacific mackerel 1968-69 and 1969 1970 seasons. 116-120

Fishes, intertidal: of Trinidad Lay and vicinity. 315-320

Folsom Lake: angling quality study. 75-93; site of primary productivity studies, 254-267

Food habits: of coyotes in Sagehen Creek basin. 4-12; of Klamath River fall-run steelhead, 217-218; of San Joaquin kit fox, 167-169; of trout in high elevation Sierra lakes in winter, 231-237

Fox. San Joaquin kit: life history, 162- 174

Fuel, diesel: effects of spilling into stream. 291-2115

Goby, long.jaw: ovarian cycling at the Salton Sea, 50-57

Goby, yellowfin: southern range exten- sion. 326-327

Growth: characteristics of Klamath River steelhead, 207-211 ; of spiny lobster phyllosomes in laboratory, 107-115; of Piute sculpin. 2*7

Half-pounder: steelhead of the Klamath River. 2(14 220

Harvest: of two domestic trout strains at Lake Berryessa, 44—19

Insecticides, oiganochlorine: toxicity to surfperch, 127-132

Insects: as food for coyotes, 9

Invertebrates, aquatic: capture by use of polypropylene light trap, 149-152

Jack-, cottonmouth: added to marine fauna of California, 245-240

Klamath River : characteristics of fall- run steelhead, 204-220

Lake Berryessa: study of planting two domestic strains of rainbow trout, 44— 49

Larvae, fish: effect of salinity on Salton Sea fish, 58-66

Length: composition of anchovies taken 1909-70 season, 121-126

Life history: of Piute sculpin. 285-290; of San Joaquin kit fox, 102-174

Lingcod: DDT residues, 22-26

Lions, sea: predation on salmon, 37-43

Pollster. California spiny: molting and growth of phyllosomes in laboratory, 107-115

Logperch: range extension, 248

Mackerel, jack: DDT residues, 22-20

Mackerel, Pacific: age composition of the catch 1964-65 through 1967-68 seasons, 13-21; southern California fishery during 1968-69 and 1969-70 seasons, 116-120

Merle Collins Reservoir: bass tagging study. 145-147 ; site of largemouth bass mortality and survival study. 221-230; site of primary productivity studies. 254-207

Methods: aging Pacific mackerel 1964- 65 through 1967-68 seasons. 13-21 ; constructing polypropylene light trap for aquatic invertebrates, 149-152; coyote food habits study, 4-12; deter- mining DDT residues in marine fishes.' 22-20; determining DDT residues in white croakers, 100; de- termining effects of salinity on eggs and larvae of Salton Sea fish. 58-66; determining organochlorine toxicity to surfperch. 127-1M2; ovarian cycling of longjaw goby, 50-57; rearing spiny lobster phyllosomes in labora- tory. 107-115; separation of Sebastes babcocki and 8'. nihririiicfiis. .">2 .3d; studying life history of the kit fox, 162-174; studying mortality and sur- vival of bass at Merle Collins Reser- voir. 221-230 studying vertical dis- tribution of euphausiids, 170-100; study of Mineral King deer herd, 133-140; study of fall-run steelhead of the Klamath River, 204-220; study of life history of Piute sculpin, 285—290; study of Modoc sucker, 277- 284 ; study of predation by sea lions on salmon, 37-43; study of subpopula- tions of Pacific sardines, 296-314; study of reservoir primary productiv- ity, 254-207: study of trout fishery at Lake Berryessa. 44-49; study of variation in the swell shark, 191- 107 ; tagging sturgeon to provide migration data, 102-100; tagging to determine white sturgeon population,

3

c M.ii ORN] \ PISH AND QA Ml.

:'i KM : tesl ing tolerance of pile w orm to salinitj . L52 L5 I : use of roving creel census at Folsom I - .- 1 K * • . 75 93

Migration: of Klamath River steelhead, •_M l 21 I : of Mineral King deer, 133 1 in : of v. hite and green si urgeons, L02 L06

Mineral King: deer study, 133 1 10

Molting: of spiny lobster phyllosomes in laboratory, LOT L15

Monterej Bay: studj of predation by lions mi salmon, '■>' 13

Morphology: of the Modoc sucker, 277— I'M

Mortalitj rates: of largemouth bass at Merle Collins Reservoir, 221 230

Mt. Whitnej troul strain: performance at Lake Berrj essa, 1 1 19

Mullet, striped: apparent spawning al sea, 246 248

M itant: melanism in ringneck pheas- ants, IT-" 17s

Net: midwater trawl for taking thread- fin shad, 268 276

Perch, dwarf: pesticide toxicity studies, 127 132

Perch, shiner: pesticide toxicity studies. 127-132; southern range record, 2S

Pesticides: chlorinated hydrocarbon resi- dues in California marine fishes. 22 I'll; toxicity of organochlorines to surfperch, L27 L32

Pheasant, ringneck: appearance of melanistic mutant, 17o-178

Phyllosomes: growth of spiny lobsters under laboratory conditions, "MH-115

Pileworm: tolerance of high salinity, 152-154

Pollution: effects of diesel fuel on a stream fauna. 291-295

Population: of Mineral King- deer herd. 133 L40; resurgence in sharpnose sea- perch, 28 31; study of variation within subpopulations of Pacific sar- dines. 296 314; survej of condors, 1971, 327 328; white sturgeon in Sacramento-San Joaquin estuary, 94- 101

Predation: bj sea lions on salmon. .".7 13

Productivity, primary: in a new and old California reservoir, 254—267

Rabbits: as food for coyotes, S

Rabies: occurrence in deer, 141-144

Raptors: numbers seen during 1970 condor survey, <17 -6S : seen during 1971 condor survey. 327 328

Rattail, roughscale: DDT residues, 22 2G

failure of pelagic cormor ed, 238 : of Iongjaw gobj i Sea, 50 57; of Piuti in, J^-7

Reviews: A trout and salmon fisherman for seventy-five years, 330; Biology

and water pollution control. 249; British Columbia game fish. 71 : Come wade the river, 71; Fishes of Mon- tana. :;l!'.i 330; Fishless days, angling nights, 250; Hardy's book of fishing, 331; Hikers and hack-packers hand- hook. 70: If deer are to survive. 157; Kamloops, 71 : Life and death in a coral sea. 250; Mountain sheep: a study in behavior and evolution, 329; Remembrances of rivers past, 330; Round river. 250; Sea shells of tropical west America; marine mol- luscs from Baja California to Peru. 249; Systematics, variation, distribu- tion, and biology of rock fishes of the subgenus Sebastomus CPisces, Scor- paenidae, Sebastes), l"><i l."»7; The art and science of fly fishing, 321 ; The dry fly and fast water and the salmon and the dry fly, 330-331; The ecology of running waters, l.~>6; The New York aquarium book of the water world, 332; The vanishing jungle — the story of the "World Wild- life Fund expeditions to Pakistan, 70-71 ; "Wildlife of Mexico: the game birds and mammals, 250 ; World dy- namics, 332

Rockfish, flag: identity and range, 32-36

Rockfish, redbanded: identity and range, 32-36

Rodents: as food for coyotes, 8

Rusk Creek: site of study of Modoc suckers, 277-284

Sablefish: DDT residues, 22-26

Sagehen Creek: site of coyote food habits study, 4-12 ; study of life his- tory of Piute sculpin, 2S5-290

Salinity: tolerance by pileworm, 152- 154

Salmon: study of predation by sea lions. 37 13

S.ilton Sea: ovarian cycling of Iongjaw goby, "><i 57 : study of salinity stress on fish eggs and larvae, os <»6: tests on pileworm tolerance of salinity. 1 52 -154

San Cleniente Island: survival of Guadalupde cardinalfish, 68-69

Sanddab, Pacific: DDT residues, 22-26

Sardine. Pacific: subpopulation study. 296- ::i I

S.-iruo: effect of salinity on eggs and larvae, 58 66

Scorpionfish, California: reoccurrence in Monterej Bay, 238-239

Seal]. in. Piute: life history. 285-290

Seaperch, pink: host for parasitic copepod, 323-325

Seaperch. rubberlip: southern range record, 28

INDEX

339

Sea perch, sharpnose: southern range

records and notes on population

resurgence, 2S-31 Sea urchin: in association with acorn

barnacles, 321-323 Shad, threadfin : trawl designed for

catching. 268-276 Shark, swell: population differences off

California and Baja California, 191-

197 Sheep: as food for coyotes, 7 Snailfish blacktail: case of symbiosis

with box crab. 239-24(1 Sole, butter: case of reversal. 244. 245 Sole, Dover: tailless specimens taken off

Oregon, 147-148 Sole. English: DDT residues. 22-26 Sole, petrale: DDT residues, 22-26 Spawning: of striped mullet at sea,

246-248 Stress, salinity: on Salton Sea fish eggs

and larvae, 58-66 Sunfish: fishery at Folsom Lake, 84 Sturgeon, green: migration, 102-106 Sturgeon, white: migration, 102-106;

population in Sacramento-San

Joaquin estuary, 94-101 Sucker, Modoc: morphology, variation,

feeding adaptations, 277-284 Surfperch: pesticide toxicity studies,

127-132 Surfperch. silver: southern range

records, 27 Surfperch, spotfin: host for parasitic

copepod, 323-325

Survey: 1970 California condor popula- tion, 67-68

Survival: of Guadalupe cardinalfish at San Clemente Island. 68-69; of two domestic trout strains at Lake Ber- ry essa, 44-49

Survival rates: of largeniouth bass at Merle Collins Reservoir, 221-230

Symbiosis: between blacktail snailfish and box crab, 239-240

Tagging: of largemoufh bass at Merle Collins Reservoir. 145-147; to deter- mine sturgeon migration. 102-106; to determine white sturgeon populations, 94-101

Telemetry: use in studying kit foxes. 165; use in studying Mineral King deer, 138-140

Toxicity: of organochlorines to surf- perch, 127-132

Trap: for aquatic invertebrates, 149- 152

Trawl, midwater: for threadfin shad, design and operation. 268 276

Trinidad Bay: intertidal fishes, 315-320

Trout, rainbow: fishery at Folsom Lake, 85 ; performance of two domestic strains at Lake Berryessa, 44—49

Trout, steelhead: characteristics of fall- run in Klamath River, 204-220

Tii i ut: winter food habits in high eleva- tion Sierra lakes. 231-237

Vegetable matter: in coyote food habits, 9, 11

83609 — 800 7-72

printed in California office of state fkintinc 5,300

Notice is hereby given that the Fish and Game Commission shall meet on October 6, 1972, at 9:00 a.m., in the Auditorium of the Resources Building, 1416 Ninth Street, Sacramento, California, to receive recommendations from its own officers and employees, from the Department and other public agen- cies, from organizations of private citizens, and from any interested groups as to what, if any, regulations should be made relating to fish, amphibia, and reptiles, or any species or subspecies thereof.

Notice is hereby given that the Fish and Game Commission shall meet at 9:00 a.m., on November 3, 1972, in the Board of Supervisors' Chambers, County Courthouse, Redding, California, for public discussion of and presentation of objections to the proposals presented to the Commission on October 6, 1972, and after considering such discussion and objections, the Commission, at this meeting, shall announce the regulations which it proposes to make relating to fish, amphibia and reptiles.

Notice is hereby given that the Fish and Game Commission shall meet on December 8, 1972, at 9:00 a.m. in Room 1138 of the New State Building, 107

5. Broadway, Los Angeles, California, to hear and consider any objections to its determinations or proposed orders in relation to fish, amphibia and reptiles or any species or subspecies thereof for the 1973 sport fishing season, such determinations and orders resulting from the hearings held on October

6, 1972 and November 3, 1972.

FISH AND GAME COMMISSION

Leslie F. Edgerton Executive Secretary

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