Quantitative studies on gonad cycle,fecundity, reproductive output and recruitment in a baltic Mytilus edulis population

Since Mytilus edulis L. is a biomass dominant in the Baltic much interest is focused on the ecology of the species. In this paper an attempt is made to quantitatively cover the reproductive cycle of a Baltic M. edulis population in order to provide data for energy flow models and to discuss aspects of recruitment in this species. Histological preparations of gonads showed that gametogenesis started with declining temperatures in autumn and proceeded very slowly through winter. At the beginning of March when food was supplied during the spring phytoplankton bloom, rapid maturation took place. This was also revealed by an increase in meat weight of the mussels. Only one spawning period was recorded, from the middle of May until the beginning of June, due to food being strongly limited to the population during the rest of the year. The length of the larval period was estimated as being 5 to 6 wk and settlement was registered from the end of June through July. In general the large annual variations found in the Baltic with regard to temperature and food abundance give rise to a more marked annual pattern in the reproductive cycle than is encountered in other seas. Fecundity was assessed for two populations from 4-and 15-m depths from studies covering two annual cycles of the changes in the relation of shell length — meat weight. The size-related fecundity was found to be equal in both populations and related to food abundance and not to growth or age. Fecundity, expressed as weight loss at spawning, ranged from 0% in 2-mm mussels and increased from 38 to 52% in 10-to 30-mm mussels. The fecundity as percentage of biomass in full-grown Baltic M. edulis is of similar magnitude as in full-grown mussels from other areas despite the smaller size of Baltic mussels. The reproductive output for the total 160 km² research area was calculated as being 1 200 tons dry weight or 80% of the standing stock, which, due to the particular features of the Baltic M. edulis population probably represents the larger part of the total mussel production. This reproductive output, calculated as 8·10⁷ eggs·m^-2 and corresponding to 50% of the total annual zooplankton production, may thus consitute an important food source for herring larvae and carnivorous zooplankton. Recruitment was divided into two phases: (1) Recruitment of juveniles (=settlement of larvae), and (2) recruitment to breeding stock. Monitoring studies of settlement on ropes and the year round presence of high abundances of mussels <2 mm indicate that settlement is in excess of the demands for maintaining population size and that most settled mussels form a pool of competitively suppressed non-growing individuals. Not until death of an already established mussel will these become recruited to the breeding population. Thus recruitment is possible throughout the year which stabilizes the population and maintains it near the carrying capacity of the area with regard to food and space availability.     

Seasonal changes in pyloric caecum and gonad indices during the annual reproductive cycle in the seastar Asterias forbesi

Seasonal changes in pyloric caecum and gonad indices were studied in a population of the seastar Asterias forbesi (Desor) from East Rockaway Inlet (Long Island, New York) during two annual cycles (1980/81, 1982/83). Pyloric caeca indices increased during fall, reached a maximum about April, and declined sharply to a minimum in mid-summer. Gonad indices increased during fall and winter and reached a maximum about May. Judging from gonad size analysis, spawning occurred in late June to early July, at bottom temperatures of 16° to 18°C. There was no long-term inverse relationship between pyloric caecum and gonad indices. Since nutrients and energy ingested during the important fall feeding period are utilized simultaneously for body growth and gonad development, it is unnecessary for the pyloric caeca to store nutrients over long periods as occurs in A. rubens and many other species. The specific caeca-gonad relationship is probably an adaptation to the extreme seasonal thermal fluctuations of the NW Atlantic, which constrain the feeding activity of A. forbesi and limit the possibility of long-term storage. In this population, most individuals grow rapidly in their first year (including first two summers) and spawn in their third summer. Few seastars survive to spawn again in their fourth summer. However, gametogenesis may take place in some individuals in their second summer. Year-to-year variability in mean size at spawning may reflect temporal variation in environmental conditions (weather, food availability) at this site.     

Reproductive biology of <Emphasis Type="Italic">Cosmasterias lurida</Emphasis> (Echinodermata: Asteroidea) an anthropogenically influenced substratum from Golfo Nuevo,Northern Patagonia (Argentina)

The starfish Cosmasterias lurida was sampled from the pier piles of Almirante Storni in the Golfo Nuevo, northern Patagonia, from November 2001 to November 2003. In this anthropogenically modified site there is an unusually abundant food supply of mussels and large aggregations of starfishes. The reproductive cycle was studied by organ indices and histological examination of gonads. The highest gonad index (GI) values were found in the summer (February–March), followed by a drop in GI values, indicating spawning. Histological analyses indicated that gametogenesis takes place from June to January, sexual maturity occurred during the summer and spawning occurred in April. The GI and pyloric caeca index (PCI) tended to have an inverse relationship, but this was only statistically significant in males. However, biochemical analyses of the starfish indicated no transfer of nutrients between pyloric caeca and gonads in either sex. In fact, simultaneous accumulation of carbohydrates was found in the pyloric caeca and gonads of both sexes.     

Seasonal variations in condition,reproductive activity and biochemical composition of the flat oyster,Ostrea edulis,from San Cibran (Galicia,Spain)

The influence of environmental parameters (temperature, salinity and available food) on the condition, reproductive activity and biochemical composition of a native population of Ostrea edulis L. in San Cibran (Galicia, Spain) was studied between September 1988 and December 1989. Histological preparations of gonads showed that gametogenesis started when temperature was at its lowest in winter. The water temperature in San Cibran never fell low enough to interrupt it. Gametogenesis proceeded slowly and spawning took place in May–June, although the predicted time of ripening was early March. Salinity in San Cibran was relatively stable throughout the year; it did not seem to have any influence on gametogenesis. Available food appeared to be a very important factor in controlling gonad growth, once gametogenesis was initiated. The major concentration of suspended organic particulate matter was present in the spring at the time of rapid gonadal maturation. Only one spawning period was observed. Larvae were released when the food in the water was high. Seasonal changes in the main biochemical components of this oyster were determined for a standard individual. Lipids and carbohydrates presented a similar time-course whereas proteins were constant. When food was abundant, energy reserves were built up. Spawning produced a decrease in biochemical constituent levels, and recovery coincided with the phytoplankton bloom. The stored reserves, mainly lipids, were used to overcome a state of energy imbalance in late autumn associated with low food availability. Results show this oyster to be an opportunist organism which concentrates its reproductive effort during a short period of favourable conditions and which is directly dependent on nutritive availability in the environment.     

Can synchronous spawning be predicted from environmental parameters? A case study of the lugworm Arenicola marina

The annual epidemic spawning period of a Scottish population of Arenicola marina (L.) has been recorded over a period of 13 yr. This population spawns between mid-October and mid-November in a discrete spawning event over a period of 4 to 5 d. Endocrine manipulation experiments showed that spawning is induced in females only if sufficient titres of PMH (prostomial maturation hormone) are present in the prostomia. These levels are attained during the 2 to 3 wk prior to the natural spawning date. The East Sands, St. Andrews population always spawns during periods of spring tides regardless of tidal amplitude or whether they are full- or new-moon tides. Meteorological data, including sea-temperature data were collected for each year, and correlation of the environmental data with spawning time was attempted. Correlation of spawning times with weather patterns showed that mean daily air pressures were significantly higher during the spawning period than from September to November as a whole. Evidence also suggests that a reduction in sea temperature is required prior to spawning. A significant moderate negative correlation was found between May to July air temperatures and spawning date, suggesting that higher May to July temperatures may induce early spawning. Daily rainfall and wind speed were also lower during the spawning period, but not significantly so. These results indicate that air pressure (or changes therein) may act as a final spawning cue, and the advantages of this are discussed in relation to fertilization success. A model of the interplay between environmental parameters and the endocrine mechanisms controlling the induction of spawning is proposed. Higher than average summer temperatures may advance gametogenesis to bring the population into a state of maturity (full-size oocytes, well-developed sperm morulae), and may also advance spawning time. Once the population has completed gametogenesis, a drop in sea temperature is then required to trigger an increase in endocrine titres within the prostomium, without which spawning cannot be induced by prostomial injection. The population spawns on spring tides; however a lack of clement weather coinciding with the spring tide will result in population-wide spawning being aborted, as in 1996. Clement weather (high pressure, low rainfall and wind speed) in conjunction with spring tides permits spawning to proceed to completion. Received: 21 June 1999 / Accepted: 25 January 2000     

The Arctic sea butterfly<Emphasis Type="Italic"> Limacina helicina</Emphasis>: lipids and life strategy

The sea butterfly Limacina helicina was collected from May to September 2001 in Kongsfjorden, Spitsbergen, to investigate population structure and body and lipid composition with regard to life cycle and reproductive strategy. Veligers and juveniles were only found in late autumn and spring, whereas females occurred from July to September. The size of the females increased until mid-August and decreased in September. Dry and lipid mass were closely related to size; dry mass increased exponentially and lipids linearly with size. The lipid content was highest in veligers (31.5% of dry mass) and juveniles (23.6%) but low in females (<10%). Phospholipids were the dominating lipid class followed by triacylglycerols. Females, veligers, and egg ribbons, all from September, were richest in phospholipids. Juveniles contained the highest amounts of triacylglycerols, whereas females had low levels in July and the beginning of August. In mid-August, levels of triacylglycerols were variable and higher. This suggests that females were in their main spawning period and the high variability in triacylglycerols points to different stages within the spawning cycle. Enhanced amounts of free fatty acids in females from July may be related to gonad development. The 16:1(n-7) fatty acid was more dominant in spring whereas 18:4(n-3) increased in summer and autumn, which reflects a change in diet from diatom-dominated food items in spring to dinoflagellates in summer/autumn. Small amounts of long-chain monounsaturated fatty acids suggest ingestion of copepods, and the fatty acid composition of veligers feeding on particulate matter. L. helicina has a one-year life cycle with peak spawning in August and over-winters as veligers that may grow to juveniles during the winter period. They metamorphose into juveniles during spring, develop to males in early summer, and mature into females in July and August.Communicated by M. Kühl, Helsingør     

Energy investment in the annual reproduction cycle of female red porgy, <Emphasis Type="Italic">Pagrus pagrus</Emphasis> (L.)

Seasonal cycle in energy content of gonad, perivisceral fat, liver, and muscle was used to estimate the energy cost of spawning of female red porgy (Pagrus pagrus) from Buenos Aires coasts, Argentina. Spawning was observed primarily between November and January, although a low-intensity spawning was also observed at the beginning of this period. The energy required for gonad development of female red porgy (the difference between the maximum and minimum energy content of ovaries along the year) was 2,652 J g⁻¹ dry weight or 558 kJ. Female red porgy expend 22.8% of their total body energy (3,481 kJ) during the spawning period. Of this energy, relative contributions from different sources were as follows: ovaries 62.5% (496 kJ), perivisceral fat 17.8% (142 kJ), liver 10.3% (82 kJ), and muscles 9.4% (75 kJ). Female spawners lost about 7.2% (119 g) of their total body weight (1,654 g) in the spawning period. From the 7.2% of the total weight lost, female gonad represented a relative reduction of 73.8% (87.6 g), the perivisceral fat declined by 5.7% (6.8 g), the liver declined by 10.6% (12.5 g), and muscle tissue exhibited a 10% loss in total weight (11.9 g). Reproductive effort represented 7.6% of food energy, while resting metabolism accounted for almost 38% of food energy. Despite the fact that red porgy females feed throughout the entire spawning period, it is shown that between pre-spawning (September, water temperature 15°C) and mid-spawning (December, water temperature 19°C) they lost 45 and 38% of their muscle and liver lipids, respectively, and 89% of the perivisceral fat associated with the gut and body cavity. Red porgy used both their energy reserves and food energy during ovarian maturation. This asynchronous species have a strategy of a lower energy allocation and a higher dependence of food during the spawning time (late-spring and summer) for successful reproduction.     

Seasonal cycle of lysosomal enzyme activities in the mantle tissue and isolated cells from the musselMytilus edulis

InMytilus edulis L., gametogenesis takes place in the mantle at the expense of the connective storage tissue. There are two main types of storage cells: vesicular (VC) cells storing large amounts of glycogen and adipogranular (ADG) cells containing large numbers of protein granules, lipid droplets and lesser amounts of glycogen. One of the ways in which stored reserves can be mobilized for gamete formation is by controlled autophagy, in which the cellular constituents are degraded by lysosomes. Mussels were collected from the Menai Strait, North Wales, and monthly measurements made, over two years (1984–1986), of the activities of lysosomal acid hydrolases (acid phosphatase,-N-acetyl-D-glucosaminidase and-glucuronidase) and Cathepsins B and L in the mantle tissue, isolated ADG cells, low-density cells and, during spawning, in the mature oocytes of female mussels. The lysosomal proteinases, Cathepsins B, L and H, were further characterised by activation with thiol compounds and inhibition with thiol blockers and by leupeptin. Because of the low activity in the mantle tissue ofM. edulis, Cathepsin H was not assayed on a seasonal basis. There was a general increase in lysosomal enzyme activity during the winter, which can be related to increased autolysis in the storage cells and to the process of maturation in the developing oocytes. The activity of Cathepsin B was highest in the ADG and low-density cells, implying an important role in proteolysis within the ADG cells. By contrast, Cathepsin L displayed the highest activity in the mature oocytes, suggesting a major function of Cathepsin L in the development and maturation of the oocytes. Two different-glucosidase activities were measured in the monthly assays, one with a pH optimum of 4.5 (acid) and the other at pH 7.5 (neutral). Highest activities of the acid-glycosidase were found in the low-density cells, but there were no significant seasonal changes in the mantle tissue as a whole. Activities of the neutral-glucosidase were low in the ADG cells and mature oocytes, but showed high activities in the mantle tissue, with marked seasonal changes that corresponded to the mobilization of glycogen reserves in the VC cells.     

Reproductive investment reduces recuperation from exhaustive escape activity in the tropical scallop Euvola zizac

In scallops, mobilization of reserves from the adductor muscle to support maintenance and reproductive activity may impinge upon a major role of the adductor muscle, the movement of the valves during swimming and escape responses. The tropical scallop Euvola ziczac (Linnaeus 1758) invests energy reserves to different degrees during its two periods of reproduction each year. We evaluated the impact of reproductive investment on the escape responses (clapping capacity and recovery after exhaustive exercise) of E. ziczac sampled at different reproductive stages (immature, mature, spawned) during the two reproductive periods in 1997. We compared the escape response capacities with the levels of muscle energetic reserves (glycogen, proteins, and arginine phosphate) and muscle metabolic capacities [activity of the glycolytic enzymes: glycogen phosphorylase (GP), pyruvate kinase (PK), phosphofructokinase (PFK), octopine dehydrogenase (ODH), arginine kinase (AK), and the mitochondrial enzyme, citrate synthase (CS)]. Gonad size, gamete volume fraction, and levels of gonadal protein and lipid were greater for mature scallops during the first than during the second reproduction. Numbers of claps during escape responses (49–57) and levels of muscle arginine phosphate remained similar throughout the different reproductive stages in both reproductive periods. In contrast, recovery was slowed during gonadal maturation in both reproductive periods and during spawning in the first reproduction. Scallops generally took more time to regain their initial clapping capacity during the first (25–40 min) than during the second reproduction (20–30 min). Muscle glycogen decreased markedly during both gonadal maturation and spawning in both reproductions; whereas, muscle proteins decreased only in the first reproduction. The levels of most enzymes decreased during gonadal maturation in both reproductions, and also after spawning, particularly during the first reproduction. We concluded that gonadal maturation and spawning did not decrease clapping capacity of E. ziczac, but decreased its capacity to recover from exhaustive exercise most likely due to decreased levels of energetic reserves and a reduced metabolic capacity of the adductor muscle. Moreover, these effects were probably stronger during the first cycle because of the greater reproductive investment coincident with decreased food availability. Received: 28 April 2000 / Accepted: 21 September 2000     

Food and feeding dynamics of the larval Antarctic fishNototheniops larseni

The feeding dynamics of larvae of the Antarctic fishNototheniops larseni were analyzed from data collected over three years in Bransfield Strait and adjacent waters (Antarctica). Seasonal feeding was examined from 1977/1978 (November–March). The diel feeding cycle was investigated during a 96 h station established in February 1976, while food selection was analyzed using larvae and zooplankton samples collected in February 1982. Hatching occurs in early spring, and larvae fed on eggs of calanoid copepods and on cyclopoid copepods. Copepod eggs were the principal food near the pack ice, and cyclopoids in open waters. Cyclopoids were the staple food in summer. Eggs of the Antarctic krillEuphausia superba were ingested selectively and formed major portions of the larval summer diet in neritic (Joinville Island) and oceanic (Elephant Island) spawning areas ofE. superba. In the fall, copepods predominated in the diets. Most abundant and most frequently ingested prey in summer and fall wereOncaea spp. Feeding commenced at dawn and continued at least until dusk. Krill eggs were taken chiefly during morning hours and egg incidence declined during the day, suggesting that eggs were ingested soon after spawning. Prey size at the onset of feeding was estimated as 0.130 to 0.330 mm. Size-selective feeding was evident in small larvae, while in larger larvae median prey length remained constant. High feeding incidence among yolk-sac larvae in spring, high overall feeding incidence in summer, and size-selective foraging of small larvae suggested favorable feeding conditions in the 1977/1978 season. Yolk-absorption times in Antarctic fish larvae vary on a scale of weeks and may be further retarded due to early feeding. Hence, year-to-year variability of yolk incidence inN. larseni indicated variable biotic environments of early feeding larvae rather than temporal shifts of hatching periods. As hatching periods are constant between years in contrast to the variable retreat of the pack ice and subsequent onset of the production cycle in space and time, maternal yolk reserves are probably utilized to compensate for such variations.     

Changes in the biochemical composition of the pyloric caeca of female seastars,Asterias rubens,during their annual reproductive cycle

The biochemical composition of the pyloric caeca of female seastars (Asterias rubens) was studied throughout the annual reproductive cycle. The pyloric caeca index is high during the pseudo-rest stage of the ovaries and decreases gradually during ovarian growth. The pyloric caeca dry weight varies between 20 and 38% of the fresh weight. Lipids, and less prominently carbohydrates (both glycogen and other reducing sugars), seem to constitute the primary nutrient reserves for gonad growth. Proteins are also available to meet ovarian requirements during vitellogenesis. Glycogen levels are high in late autumn and winter and low at spawning time and in summer, whereas levels of the other reducing sugars are high in post-spawning months and subsequently gradually decrease during gonad development. The total lipid level is extremely variable (averaging 75 mg g^-1 organ fresh weight), peak levels of over 200 mg g^-1 being found in some individuals in summer and at spawning time. The free amino acid level is constant at about 20 mg g^-1. The protein level is rather high throughout the annual cycle, and displays only minor changes. The observed patterns of decrease in major constituents suggest a rapid release of nutrient reserves from the pyloric caeca (deposited during the summer) at the onset of vitellogenesis. Moreover, the needs of the ovaries during further development seem to be so tremendous that during the winter months the digested food may be transported to the ovaries without prior deposition in the pyloric caeca. The changes observed in the pyloric caeca are discussed and compared with those occurring simultaneously in the ovaries.     

Growth and spawning of Urechis caupo (Echiura) in Bodega Harbor,California

Growth and spawning of the large, infaunal echiuran worm Urechis caupo Fisher and MacGinitie were studied at Bodega Harbor on the coast of central California, USA, from 1978 through 1981. In situ growth rates of marked worms were negatively related to initial size. Short-term, summer growth rates (volmo^–1) of small worms (<80 ml) were greater than longer-term growth rates measured over several seasons (asesonal). Size-frequency distributions of worms sampled from two sites also suggested a seasonal growth pattern with relatively fast spring-summer growth and slower winter growth. However, larger worms sometimes lost volume during in situ growth experiments, and the loss was most pronounced during short-term, summer growth periods. It is suggested that energy used in burrow construction may have contributed to volume loss during short-term growth experiments. In contrast, longer-term, aseasonal growth rates were nearly always positive, and indicated that reproductive size (about 56 ml) could be reached within about 1.5 yr of recruitment, and a large size (about 158 ml) could be reached within about 6 yr. A seasonal pattern of spawning was observed during three consecutive years, as indicated by ripeness indices (storage organ dry weight ÷ body wall dry weight). At least two spawning episodes occurred annually: ripe gametes that accumulated in the storage organs during the summer and fall were spawned during the winter; gametes that accumulated during late winter and early spring were spawned during the spring or early summer. Worms were spawned-out by mid-summer.     

Reproduction and population dynamics of the fissiparous brittle star Ophiactis savignyi (Echinodermata: Ophiuroidea)

Reproductive cycle, asexual reproduction, and population dynamics of the fissiparous brittle star, Ophiactis savignyi, which inhabits the exhalant passages of the sponge Haliclona sp. were examined monthly from February 1991 to January 1992 at Wanlitung, southern Taiwan (22°N; 120°E). Mature gametes were found from March to December, but release was mainly in May and June. Sexual recruits were found from May to December, with the highest frequency (14.1% of population) in June. Sexual recruits composed 2.4% of the 1-yr sample. Sex ratio of male to female was 24:1. Fission occurred throughout the year, although the frequency of recently split individuals was lower from January to June (6 to 31%), and higher from July to December (42 to 52%). The occurrence of fission was highest after spawning. Regenerating individuals composed 48.2% of the 1-yr sample. Population density fluctuated greatly during summer due to recruitment by fission and mortality or dispersal due to the stressful environmental conditions. Both sexual and asexual reproduction of O. savignyi were successful at this site.     

Food quality and season affect gene expression of the mucosal lectin MeML and particle sorting in the blue mussel Mytilus edulis

The blue mussel, Mytilus edulis, is able to use biochemical cues to sort food particles. Recently, a mucosal lectin (MeML) identified in pallial organs of M. edulis was suggested to play a role in the capture and sorting of food particles. The current study presents data obtained in 2010 showing that sorting efficiencies and MeML gene expression in M. edulis increased when mussels were either fed poor quality food or starved. In addition, results suggest that particle capture and degree of selection increased during the spawning period, although the quality of food delivered to mussels before the experiments seemed to have a stronger impact on particle-sorting efficiency. Overall, the impact of both endogenous (physiological status) and exogenous (food quality) factors were shown to affect the expression of MeML in mussel feeding organs, and the food sorting abilities.     

Diet composition and feeding intensity of horse mackerel,<Emphasis Type="Italic"> Trachurus trachurus</Emphasis> (Osteichthyes: Carangidae) in the eastern Adriatic

Diet composition and feeding intensity of the horse mackerel, Trachurus trachurus, collected in the eastern Adriatic Sea, were examined. Stomach contents of 1,200 specimens, taken at monthly intervals (January–December 1996), were analyzed. The stomachs contents consisted of five major prey groups: Crustacea (Euphausiacea, Mysidacea, Decapoda), Cephalopoda and Teleostei. Euphausiacea constituted the most important food resource by weight, number and frequency occurrence. Teleosts were the second most important food category, while mysids, decapods and cephalopods were occasional food. There was little seasonal variation in diet. Euphausiids were dominant prey during all seasons, and were especially abundant from spring and autumn. Feeding intensity of horse mackerel varied during the year. The lowest intensity of feeding was recorded in winter (February, March) and early spring (April) during the lower sea temperature as well as at the time of intensive spawning. Feeding activity increased upon spawning period (May and June) and was also higher during summer. Feeding intensity and diet composition changed during the diurnal cycle. The highest feeding intensity was recorded at night and during early morning hours. Euphausiids, mysids and a greater part of teleosts dominated night and morning diet, while decapods and cephalopods were most frequently in the daily and evening diet.Communicated by O. Kinne, Oldendorf/Luhe     

Reproductive dynamics of endeavour prawns, Metapenaeus endeavouri and M. ensis, in Albatross Bay, Gulf of Carpentaria, Australia

 The spawning patterns of two penaeid prawns, Metapenaeus endeavouri (Schmitt) and M. ensis (De Haan), were examined from data collected at 45 stations between March 1986 and March 1992. An index of population fecundity based on the abundance, proportion and fecundity of sexually mature females was used as a measure of spawning output of the prawn stock. The population fecundity index for M. ensis was higher than that for M. endeavouri. The monthly population fecundity index for M. endeavouri varied markedly among years, while that for M. ensis was consistent among years. Spawning of M. endeavouri occurred year-round, while that of M. ensis was concentrated mainly in spring (September to November). For M. endeavouri, a minor spawning, derived from a relatively small number of summer spawners, occurred in the 20 to 30 m offshore waters in summer. In early summer (after May), the major spawning group consisted of large females from the winter-spawning cohort, and the spawning area shifted to depths of 30 to 60 m. In winter (July), the major spawning, derived from the winter-spawning cohort, occurred at depths of 20 to 40 m. For M. ensis, the major spawning, derived from the spring-spawning cohort, was observed in depths <50 m and was concentrated particularly in inshore waters (<20 m) in spring. In autumn, the spawning output was mainly from the autumn-spawning cohort, which comprised but a small number of individuals. In winter, the major spawning group again consisted of the large females from the spring-spawning cohort, and spawning increased in the oceanic waters (>50 m). These results suggest that mature female M. endeavouri and M. ensis move offshore (>40 m) by May and July, respectively, and return to shallow waters (<35 m) in July and November, respectively. The monthly reproduction patterns of both species in the “effective spawning” area showed that the major spawning season for M. endeavouri is August to October and that for M. ensis is September to December. Received: 19 February 1999 / Accepted: 18 June 2000     

Larval fish assemblages in Independencia Bay,Pisco, Peru: temporal and spatial relationships

The structure of the larval fish assemblages in Independencia Bay on the coast of Peru was examined using a combination of univariate and multivariate techniques. The plankton of Independencia Bay was sampled during 2000, to ascertain ichthyoplankton composition, abundance, and seasonality. These data were used to assess the function of the bay as spawning and nursery grounds and were related to the regional oceanography. In total, 16,156 fish larvae, representing 34 families, 48 genera, and 48 species were collected. Engraulidae, Normanichthyidae, Blenniidae, Gobiesocidae, Haemulidae, Labrisomidae, Pinguipedidae, and Atherinidae comprised 96.8% of the larvae captured; the remaining 3.2% included 26 families. Greatest mean larval fish densities, 319–1,381 per 100 m³, were recorded between September and November, suggesting a major spring spawning period. The most abundant fish larvae during this period were preflexion stage mote sculpins (Normanichthyidae) and newly hatched and preflexion stage anchovies (Engraulidae). A second, smaller summer peak was dominated by preflexion stage anchovies, followed by preflexion stage mote sculpins. The occurrence of high larval fish densities and the wide range of larval stages suggest that Independencia Bay is a regionally important spawning and nursery ground for marine fish. The principal component analysis showed that temperature and salinity were the dominant variables within the first two principal components, which accounted for 74.4% of the variation in environmental conditions. These conditions varied over time, station, and depth; however, interaction terms could not clearly be identified. Fitting a multinomial logistic model showed that larval fish assemblages and environmental conditions were associated in a complex way. The spring and summer ichthyoplankton abundance peaks in Independencia Bay coincided with high zooplankton standing stock and also coincided approximately with the periods of increased upwelling in the area.Communicated by O. Kinne, Oldendorf/Luhe     

Spawning patterns in the leopard grouper, <Emphasis Type="Italic">Mycteroperca rosacea</Emphasis>, in comparison with other aggregating groupers

We documented the spawning patterns of the leopard grouper, Mycteroperca rosacea, from April to June 2005 in the central Gulf of California, Mexico to draw comparisons with other aggregate-spawning groupers and provide information useful for management of their fishery. Adults formed spawning aggregations of 150 to >700 individuals at specific sites, and spawning occurred daily at these sites from late April through early June. Courtship occurred throughout the day, but spawning was restricted to the evening hours. Adults spawned in groups of 6–40 fish, and pair-spawning was not observed. The group-spawning behavior of adults and the gonosomatic indices of mature males (maximum = 7.2%) suggest that sperm competition was present. The site-specificity of leopard grouper spawning aggregations and diel spawning period were typical of most aggregating groupers, and the size and structure of these aggregations was similar to other species in the genus Mycteroperca. Leopard grouper behavior patterns were unusual in that spawning aggregations persisted for extended periods, spawning was not synchronized with the lunar cycle, and adults aggregated during non-spawning periods. The extensive duration and site-specificity of spawning aggregations and the propensity of M. rosacea to form aggregations year-round increases the vulnerability of the species to overfishing. Policies that limit harvest from these aggregations are needed to improve the management of leopard grouper fisheries in the Gulf of California.     

Factors affecting the spawning period of sardine in two highly oligotrophic Seas

Ovarian histology and weight dynamics of reproductive and somatic tissues were used to investigate seasonality of spawning and factors affecting reproductive period in two E Mediterranean sardine, Sardina pilchardus, populations (central Aegean and Ionian Seas), between September 1999 and May 2001. Despite similarities in the general spawning pattern of the two populations, i.e. protracted spawning season during the colder months of the year, reproductive period in the Ionian was shorter and lagged behind the Aegean by 2–3 months. In both Seas, reproductive period was influenced by body size as smaller females tended to have both shorter and delayed spawning season. Whereas hepatosomatic index (HSI) was consistently higher in reproductively active females during the whole spawning period, seasonality of spawning did not match variations of HSI. In both Seas fluctuations of HSI matched the spring burst of primary productivity and were followed by fluctuations in somatic condition and mesenteric fat by a 2 months’ lag. Somatic condition and reproductive activity displayed inverse seasonal patterns, implying that egg production in sardine basically relies on visceral and muscular fat deposits, accumulated during the summer growing season. Reliance of egg production on past energetic reserves suggests that sardine is a capital breeder, which is a rather efficient strategy in such oligotrophic habitats as the E Mediterranean, where winter spawning is accompanied by a particular decline in adult preying fields.