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Growth and Reproduction of the Tidepool Sculpin Oligocottus Maculosus

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Growth and Reproduction of the Tidepool Sculpin Oligocottus Maculosus Japanese Journal of Ichthyology 魚 類 学 雑 誌 Vol.36, No.4 1990 36巻4号1990年 Growth and Reproduction of the Tidepool Sculpin Oligocottus maculosus Bryan E. Pierce and Keith B. Pierson (Received October 9, 1987) Abstract Growth and reproduction of tidepool sculpins (Oligocottus maculosus) from three sites in Puget Sound, Washington State, USA were characterized. One site was a sandy, gently sloping beach, the second was an exposed rock headland with minimal algal cover, and the third was an exposed rock headland with numerous tidepools containing abundant algal cover. Length- weight relationships for sculpins did not differ significantly between sexes within sites. Popula- tion age structure, growth rate and recruitment varied among the three study sites using data pooled across both sexes. Young-of-the-year sculpins comprised 91% of the population at the exposed headland site with minimal algal cover but only 6% at the headland with abundant algal cover. Selective predation by great blue herons (Ardea herodias) upon tidepool sculpins larger than 45mm total length assisted in the maintainance of these age structures. A maximum gon- adosomatic index of 28.5 was calculated; this index correlated poorly with either age, length or somatic weight. Diameters of unshed ova from preserved tidepool sculpins sampled prior to spawning fell into three classes. Comparison of the mean egg size of each mode and the number of modes present in preserved sculpins with similar data from tidepool sculpins that spawned in the laboratory suggested that spawning occurred twice yearly from at least January through August. The tidepool sculpin, Oligocottus maculosus length and weight and proposed that the fish was Girard, is a small cottid inhabiting rocky intertidal a multiple spawner. The purpose of our research areas along the northeast Pacific coast (Hart , was to examine the life history of the tidepool 1973). Although occasionally found in shallow sculpin at three diverse sites for comparison with eelgrass habitats (Read, 1968), this sculpin typ- historic data, particularly Atkinson's, and to ically dwells in tidepools where broad habitat , evaluate the environmental influence on growth temperature, salinity and food requirements rate, fecundity, population age and size structure. (Nakamura, 1976a, b; Green, 1971c) often make Knowledge of variability within these parameters it the most abundant fish species (Moring , 1976). is essential if the ecological bases for and im- Many studies on the tidepool sculpin focus plications of experimental results (such as dis- upon behaviour, for instance, Eastman's (1962) placement) are to be understood. pioneering research on homing ability following displacement. Homing occurs in 60-80% of Methods and materials 50-90mm individuals tested (Green , 1971a), depends more upon olfaction than vision (Khoo, Three field sites in Puget Sound, Washington, 1974), is most successful in age II fishes (Craik , USA, were sampled (Fig. 1). Alki Point, Seattle, 1981), and takes place relatively rapidly (generally Washington is predominantly a gently sloping within two tidal periods; Green,1971a). Green sand beach containing small shallow tidepools (1971b) showed that rhythmicity of the tidepool with little algal cover and minimal rock structure. sculpin's locomotor activity correlated with tidal Green Point, Anacortes, is an exposed rock cycles; activity decreased greatly during the low headland with a very steep slope, small tidepools, tidal phase. and minimal algal cover. Tongue Point, near Despite such advances, few data describe the Port Angeles, is an exposed headland of inter- life history and biology of the tidepool sculpin . mediate slope with many tidepools of varying Nakamura (1971) showed that tidepool sculpins depths that contain extensive algal and rock are general and opportunistic feeders while cover. Atkinson (1939) evaluated the relationship between On 5 January 1979 tidepools at Alki Point were •\4 10•\ Pierce and Pierson: Growth and Reproduction of Sculpin sampled with handnets at low tide. All visible sculpins were collected. Specimens were held briefly in aquaria, sacrificed, and frozen. Two weeks later these specimens were thawed, weighed (•}0.01g) using the light blotting technique (Parker, 1963) and their total (TL) and standard lengths (SL) measured (•}0.5mm). Neither freezing nor five day formalin preservation ap- preciably altered the mean total length, standard length or wet weight (n=48). Mean changes caused by freezing were -0.08%, -0.02% and 0.28%, respectively. Mean changes caused by formalin preservation were -0.04%, -0.05% and -0.05%, respectively. Ovaries were re- moved, weighed, and placed in individual vials containing 7% formalin. On 3 February 1979 live sculpins were again collected from Alki Point, but these sculpins were held in aquaria in 12•Ž recirculating seawater, fed grated English sole (Parophrys vetulus), and allowed to spawn. Immediately after spawning, the sculpins were sacrificed, preserved and examined as before. Tidepools at Alki Point were visited again on 14 Fig. 1. Location of the study sites. A, Alki Point, March and 2 August 1979 to search for ripe Seattle, Washington; G, Green Point, near females. Anacortes, Washington; T, Tongue Point, On 5 August 1979 one infralittoral and two near Port Angeles, Washington. midlittoral tidepools at Tongue Point were sam- pled with quinaldine after all rocks were removed. size distributions (Cassie, 1954, Harding, 1949) Tidepools at Tongue Point were again visited on rather than particular ova diameters. 4 September 1979 to search for ripe females; no Age estimates were obtained by both length other sculpins were kept. and weight frequency analysis and verified by On 10 August 1979 tidepools were sampled at reading growth rings on otoliths (n=29; Jearld, Green Point employing the same procedures used 1983; Bagenal and Tesch, 1978) and vertebral at Tongue Point. centra (n=11; Menon, 1950). The total length Paired ovaries from 104 individual sculpins to weight relationship was calculated after loge (52 from Alki Point, 48 from Tongue Point, and transformation of both variables. The ELEFAN1 4 from Green Point) were broken up and ova computer program (Pauly and David, 1980) was examined in water under a dissecting microscope used to calculate length-frequency-based param- equipped with a calibrated ocular micrometer. eters (L•‡=theoretical maximum length, K=growth All oocytes (except those less than 0.10mm coefficient, D=correction for mean environmental which could not be consistently removed from temperature, ESP/ASP ratio=data variance the stroma without damage) were counted and ratio, winter point=time of coldest part of the diameters measured (•}0.01). Similar examina- winter on an annual scale from zero to one) for tions were conducted using eggs and ovaries from the generalized von Bertalanify growth formula sculpins that had spawned in the laboratory (n= (Pauly 1981): 2). Total annual fecundity of each female was Lt=L•‡[1-e-KD(t-t0)]1/D. defined as the number of albuminous ova present prior to spawning. Ova diameters from individ- On 4-7 and 28 August 1979, predation by the ual fish were highly variable depending on the great blue heron (Ardea herodias) on tidepool state of maturity of the fish; hence, spawning sculpins was observed from cover at Tongue modes were characterised as statistically discrete Point during low tide. Binoculars allowed iden- •\ 411•\ 魚類学雑誌 Japan. J. Ichthyol. 36 (4), 1990 Fig. 2. Weight frequency distribution of tidepool sculpins, Oligocottus maculosus, from Tongue Point (5 August 1979), Green Point (10 August 1979) and Alki Point (5 January 1979). tification of prey, timing of foraging and estima- Fig. 3. Length frequency distribution of tidepool tion of the length of captured sculpins by com- sculpins from Tongue Point (5 August 1979), parison with heron bill length. Green Point (10 August 1979) and Alki Point (5 January 1979). Results weight regressions for the male and female sculpins The relationship of standard length to total collected after the peak spawning period at Green length was: SL=0.813 (TLmm). This relation- Point (Ts=0.036, Ti=0.033) or Tongue Point ship did not vary with sex, size (36.5-72mm TL) (Zs=0.20, Zi=-0.37). After pooling across or sampling locality. sexes, significant differences were not found be- The length-weight regression for female Alki tween sites (using the corrected Alki Point female Point tidepool sculpins in ripe spawning condition data; Alki X Green Point, Zs=-0.69, Zi=0.77; was significantly different from that of males (s= Alki X Tongue Point, Zs=-0.58, Zi=0.49; Green slope, i=intercept, Zs=2.33, P<0.05, Zi=-2.08, X Tongue Point, Zs=0.15, Zi=-0.39). There- P<0.05, df=52). However, when the weight of fore, the pooled length-weight regression for all the unspawned ovaries was subtracted from the 256 Puget Sound tidepool sculpins sampled was: total weight of each female to simulate non- Ln(W)= -12.1+3.23 (Ln(TL)) where W=body breeding condition, no significant difference weight in g. existed between the slope and intercept of the The ages of mature sculpins (ages I, II and III) regressions for each sex (Zs=1.28, Zi=-1.27). were most easily estimated by weight-frequency Similarly, no significant differences existed analysis (Fig. 2), but the ages of young-of-year between the slope and intercept of the length- (YOTY) sculpins were more easily determined •\ 412•\ Pierce and Pierson: Growth and Reproduction of Sculpin Fig. 4. The age-weight relationship for tidepool sculpins from three sites in Puget Sound. Data for each location were pooled across sexes. The crosses indicate Alki Point speci- mens, the solid circles Tongue Point fish and the open circles Green Point sculpins. using length-frequency analysis (Fig. 3). Length- frequency and weight-frequency age estimates agreed in 96% of cases. All weight-frequency- derived age estimates agreed with age estimates derived from otolith and vertebral growth data; however, in one case (5%) length-frequency analysis disagreed with both otolith and vertebral age estimates.
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