Aspects of the Life History of Hornyhead Turbot, Pleuronichthys Verticalis, Off Southern California
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Aspects of the Life History of Hornyhead Turbot, Pleuronichthys verticalis, off Southern California he hornyhead turbot T(Pleuronichthys verticalis) is a common resident flatfish on the mainland shelf from Magdalena Bay, Baja Califor- nia, Mexico to Point Reyes, California (Miller and Lea 1972). They are randomly distributed over the bottom at a density of about one fish per 130 m2 and lie partially buried in the sediment (Luckinbill 1969). Hornyhead turbot feed primarily on sedentary, tube-dwelling polychaetes (Luckinbill 1969, Allen 1982, Cross et al. 1985). They pull the tubes from the sediment, Histological section of a fish ovary. extract the polychaete, and then eject the tube (Luckinbill 1969). Hornyhead turbot are Orange County, p,p’-DDE Despite the importance of batch spawners and may averaged 362 μg/kg wet the hornyhead turbot in local spawn year round (Goldberg weight in hornyhead turbot monitoring programs, its life 1982). Their planktonic eggs liver and 5 μg/kg dry weight in history has received little are 1.00-1.16 mm diameter the sediments (CSDOC 1992). attention. The long-term goal (Sumida et al. 1979). Their In the same year in Santa of our work is to determine larvae occur in the nearshore Monica Bay, p,p’-DDE aver- how a relatively low trophic plankton throughout the year aged 7.8 mg/kg wet weight in level fish like the hornyhead (Gruber et al. 1982, Barnett et liver and 81 μg/kg dry weight turbot accumulates tissue al. 1984, Moser et al. 1993). in the sediments (City of Los levels of chlorinated hydrocar- Several agencies in South- Angeles 1992). The highest bons several orders of magni- ern California measure the tissue levels of chlorinated tude greater than levels in bioaccumulation of trace hydrocarbons in hornyhead sediments. The objectives of metals and chlorinated hydro- turbot occur on the Palos this study were to describe carbons in muscle tissue of Verdes Shelf; concentrations the reproductive cycle and hornyhead turbot as part of decline to the north and south diet of Pleuronichthys their receiving water monitor- (Mearns et al. 1991). verticalis on the mainland ing programs. In 1991 off shelf off Orange County. 154 Fish weight-length relation- reproductive status. Male Materials and Methods ships were calculated sepa- gonads were rated based on rately for males and females the system of Grier (1981) Hornyhead turbot were from: and female gonads were rated collected off Huntington W = aLb, based on the system of Ross Beach in 30-35 m of water where: and Merriner (1983) (Table 1). approximately every six weeks W = body weight in gm and The diameters of a minimum in 1992 (Figure 1). Fish were L = standard length in cm. of 50 randomly selected collected with a 7.6 m The constants a and b were oocytes were measured for headrope otter trawl in the determined by nonlinear re- each female in a subset of morning and transported live gression technique (Marquardt females from each collection. to the laboratory. The fish 1963) in Sigmaplot™ for Only oocytes sectioned were euthanized with MS- Windows Version 1.00. through the nucleus were 222, measured to the nearest The gonadosomatic index measured. mm standard length, and (GSI) was calculated for indi- The stomach contents and weighed to the nearest 0.1g. vidual fish from: major taxa were weighed to The stomachs were removed, GSI (%) = (WG/ WB) X 100, the nearest 0.01 g. Prey placed in Davidson’s fixative where: items were identified to the for 30 days, and then trans- WG = gonad weight in gm lowest possible taxon. The ferred to 70% ethanol. The and Index of Relative Importance gonads were removed, WB = body weight in gm. (IRI) modified from Pinkas et weighed, and a 1-2 mm sec- Sections of gonads were al. (1971) was calculated tion was removed from the cut at 6 μm and stained in from : center and placed in hematoxylin and eosin. The IRI = %O (%N + %W), Davidson’s fixative for 30 histological preparations of where days. the gonads were evaluated for %O = percent occurrence, %N = percent numbers, and %W = percent weight. Figure 1. The sample size of stomachs Map of the study area off Orange County. that was sufficient to describe the diet of hornyhead turbot was determined from a plot of cumulative taxa versus num- ber of stomachs. The asymp- tote of the curve was the minimum sample size. The diel feeding pattern was determined from fish collected throughout a 24 h period in November 1992. Fish were collected by trawl- ing before and after sunrise and sunset, and around mid- day and midnight. The fish were measured, sexed, and dissected on the ship. The stomachs were placed in 155 Table 1. Reproduction Females reached sexual Rating system for female and male Plueronichthys verticalis gonads. maturity at 150-160 mm SL STAGE FEMALES MALES (Figure 3). The GSI for mature females was high (>3%) from I Immature: Ovigerous lamellae Immature: January to September (Figure composed of dense aggregations Spermatogonial of undifferentiated oogonia and proliferation. 4). More females with stage primary oocytes. four and five oocytes were collected between January II Resting: Primarily early and Early recrudescence: previtellogenic oocytes with <1% Spermatogonia and and March (Figure 5) and the early vitellogenic oocytes. spermocytes present. largest oocytes occurred in March. Ripe and running III Developing: Previtellogenic Mid-recrudenscence: All oocytes numberically dominant with stages of sperm females (stage 6) were not some early and active vitellogenic development present. collected during the study. oocytes. Males reached sexual matu- rity at 90-100 mm SL (Figure IV Well developed: Vitellogenic Late recrudescence: oocytes predominant and evenly Tubules filled with 6). The GSI for mature males distributed over a large range sperm and number of was highest between July and developing cysts are September (Figure 7). Males declining. with sperm (stage 5) were V Ripe: Broad distribution of Functional maturity: collected between October vitellogenic oocyte, with a mode of Tubules filled with and January (Figure 8). very large vitellogenic oocytes with sperm, very little, if some atresia. any spermatogenisis is occurring. Food Habits The plots of cumulative VI Spent: No evidence of Post spawn taxa became asymptotic at vitellogenisis, stage IV and V oocytes are atretic, a few stage III about 40 stomachs in the oocytes occur. annual study and in the 24 h study (Figure 9). Hornyhead turbot fed on bivalves and Davidson’s fixative and polychaetes almost exclu- treated as before. Individual sively (Table 2). Bivalves fish weight was estimated Results composed 44% of the total from a weight-length regres- IRI in the annual study and sion for males and females Size 57% in the 24 h study. Poly- previously collected. The The relationship between chaetes composed 56% of stomach contents weight was length and weight was differ- the total IRI in the annual normalized to fish weight ent for females and males study and 40% in the 24 h from: (Figure 2). Females weighed study. Turbots ate only the more than males at the same WNS = (WS / WB) X 100, tip of the bivalve siphon and where: length and also grew to a not the whole bivalve. Solen larger size than males. The WNS = normalized stomach roseaceous was the most contents weight, weight-length relationships common bivalve in the diet. W = stomach contents were: Polychaetes were generally S 2.145 weight, and females W = 0.3027 L consumed whole. Diopatra W = fish body weight and ornata, Pista alata, and B 1.767 predicted from regression. males W = 0.8915 L . Paraprionospio pinnata were 156 the most frequently occurring polychaetes. Echinoderms, Figure 2. Weight-length regression for female and male Pleuronichthys nemerteans, nematodes, and verticalis off Orange County. cnidarians were also con- sumed, but made up less than 3% of the total IRI. Hornyhead turbots had diel 500 and annual feeding patterns. In the 24 h study, fish col- FEMALES 400 MALES lected around sunrise had the highest stomach contents weight, which declined to a 300 low in the evening (Figure 10). In the annual study, the 200 median stomach contents WEIGHT (G) weight of fish collected in the 100 summer (2.64 g) was more than double the weight of fish 0 collected in the winter (0.98 50 100 150 200 250 300 g). STANDARD LENGTH (MM) Discussion Males and females grew at Figure 3. the same rate up to about Stage of development of female Pleuronichthys verticalis by 150 mm, about the size of size class. sexual maturity. After 150 mm, females grew proportion- STAGE 1 STAGE 2 STAGE 3 ately more than males. STAGE 4 STAGE 5 STAGE 6 Hornyhead turbot appeared to spawn mainly from late winter 100 to spring, but Goldberg (1982) suggested that they spawn 80 through the year. Their larvae occur in the plankton all year, 60 but abundance is highest between June and August 40 with a smaller peak between PERCENT February and April (Moser et 20 al. 1993). The larvae occur predominately inshore (Gruber et al. 1982, Moser et al. 0 1993) in depths between 5- 70 m (Barnett et al. 1984). 80-89 90-99 We found the largest concen- 100-109 110-119 120-129 130-139 140-149 150-159 160-169 170-179 180-189 190-199 200-209 210-219 220-229 230-239 240-249 250-259 260-269 270-279 280-289 trations of hornyhead adults STANDARD LENGTH (MM) 157 and juveniles off Huntington Figure 4. Beach in 30-35 m of water. Stage of development of male Pleuronichthys verticalis size class. Spent males were collected in January, but spent females were not encountered during STAGE 1 STAGE 2 STAGE 3 the study.