Laboratory and field estimates of the rate of faecal pellet production by Antarctic krill,Euphausia superba

The time course of faecal pellet production (egestion) was monitored in January 1985 for a population of Antarctic krill, Euphausia superba Dana, maintained in flowing seawater aquaria at Palmer Station, Antarctica. Following transfer to filtered seawater, krill produced faecal strings for roughly 40 min, after which time faecal egestion virtually ceased. Similar results were obtained for freshly-trawled krill at sea in February and March 1985. There were wide daily variations in total faecal egestion rate; mean rates varied from 0.54 to 1.66 mg dry wt h^-1 and individual rates from 0.25 to 2.35 mg h^-1 (all data corrected to a standard krill of 600 mg fresh weight). Despite these wide fluctuations in total faecal egestion, the loss of organic matter showed no significant daily variation, with a mean value of 0.13 mgh^-1. The relationship between faecal egestion rate and faecal organic content suggested that feeding rate was governed by food quality; when inorganic load was high, feeding rate increased to ensure sufficient energy intake. The data suggest that superfluous feeding does not occur in krill and that values of gut-clearance time calculated from time intervals greater than about 40 min will not be representative of previous feeding history. the rates of faecal egestion observed in this study indicate that the flux of faecal pellets from krill is substantial. They imply an energy intake in E. superba of 17 to 28% body weight per day, much higher than estimated previously for this species by summing known energy losses, but similar to estimates for other euphausiids.     

Reproductive cycles of three dominant Calanus species in Disko Bay, West Greenland

This study describes the annual reproductive cycles of the three dominant Calanus species, C. finmarchicus, C. glacialis and C. hyperboreus, in Disko Bay (West Greenland) in relation to seasonal phytoplankton development. Relative abundance of females, copepodite stage V (CV) and males, and the developmental stage of the female gonad were examined from plankton samples collected at weekly to monthly intervals from May 1996 to June 1997 with a WP2 net or a pump. During spring and summer, egg production rates were determined. Females of all three species were present year round. Maximum relative abundance was reached by C. hyperboreus females at the beginning of February, by C. glacialis in mid-February, and by C. finmarchicus in April. All three species reproduced successfully in Disko Bay. Their reproductive cycles were considerably different with respect to the timing of final gonad maturation and spawning, and hence in their relation to seasonal phytoplankton development. In all three species, early gonad development took place during winter, before living food became plentiful, suggesting that these processes were largely food independent. Final gonad maturation and spawning in C. finmarchicus was related to the phytoplankton concentration, reflecting that final gonad maturation processes are food dependent in this species. C. glacialis females matured and spawned prior to the spring bloom. Our results indicate that first internal lipid stores and later ice alga grazing supplied final gonad maturation and egg production. Maximum egg production rates of C. glacialis were found in spring and summer, when the chlorophyll a concentration was high. Mature female C. hyperboreus were found from February until mid-April, when the chlorophyll a concentration was still low. In this species, reproductive activity was decoupled from phytoplankton development, and final maturation processes and spawning were solely fuelled by internal energy stores.     

Reproductive cycle of Zidona dufresnei (Caenogastropoda: Volutidae) from the southwestern Atlantic Ocean

Gonadal development of Zidona dufresnei (Donovan, 1823) (Caenogastropoda: Volutidae) was studied over a period of two consecutive years, through analysis of gonadal tissue. Individuals were sampled monthly at the Mar del Plata area, Argentina. Gonadosomatic index was estimated for males and oocyte size was used to estimate the stage of gonadal development in females. The reproductive season in this species in the sampled locality extended from October to March (austral spring-summer). During summer, a stage of advanced gonadal development and spawning predominated in the adult population. In autumn, gonads were generally under atresia; during wintertime (June-August), they underwent a period of recovery that lasted until the spring months, when gametes were released again. Synchronism between both sexes was evident. Marked periods of spawning were followed by resorption periods and then a growing phase; it was very clear that reproductive seasonality was linked to changes in bottom water temperature. These results suggest that Z. dufresnei gonads have a yearly cycle of gamete production, with two major activity peaks in September-October and January-February.     

Reproduction of the commercial sea cucumber Holothuria scabra (Echinodermata: Holothuroidea) in the Solomon Islands

Over a 3-year period (1996-1998), reproduction of the commercial sea cucumber Holothuria scabra (Jaeger, 1833) was investigated in the Solomon Islands to determine the spawning pattern and whether gametogenesis is continuous or seasonal. The gonad consisted of a single cohort of tubules that developed uniformly. Macroscopic examination of the gonads revealed that mature gametes were present throughout the year. Individuals with gonads at different stages of maturity were present in most samples. Partly spawned gonads were prevalent in females, whereas mature gonads were prevalent in males. The time at which the peak gonad index was recorded differed among years. Although gametogenesis was continuous, with a potential for prolonged gamete release, a period of enhanced spawning occurred during the dry season, from September to December. Maximum gonad indices were reached prior to and during this period of enhanced spawning. Histology revealed that gametogenesis reinitiated in partly spawned gonads, resulting in the presence of gametes at different stages of development in the gonad. The uniform growth of gonad tubules indicated that H. scabra does not conform to the progressive tubule recruitment model described for holothurian oogenesis. Continuous reproduction in H. scabra and prolonged availability of mature gametes would facilitate use of this species for aquaculture.     

Growth dynamics and mortality of the intertidal encrusting sponge Halichondria okadai (Demospongiae, Halichondrida)

A periodical survey of the growth of an intertidal encrusting sponge Halichondria okadai was conducted on a rocky intertidal shore on the Izu Peninsula, southern Japan. The area covered by each of 16 marked sponges was monitored, and the fusion and fission of specimens were recorded from June 1995 to June 1998. During the investigation, 15 of the original specimens managed to survive accompanying fusions or fissions and only one colony completely disappeared. Ten fusions and 23 fissions were observed in total. The maximum death rate during a semilunar period was 16.7% and cumulative mortality throughout the study period was 46.1%, indicating that the mortality of H. okadai is lower than that of other species previously reported. There was no particular seasonal pattern in the occurrences of fusion, fission and death of sponges. However, fissions were frequently observed in the latter half of the study period and were sometimes followed by fusions or death of specimens. Positive growth rates were often observed during warmer months, and specimens showed little growth or regression in winter. Seasonal patterns in surface area were thought to be related to seasonality in reproductive activity.     

Seasonal changes in biochemical composition of the bivalve Lima hians from the Clyde Sea Area

Seasonal changes in the tissue weight and biochemical composition of the bivalve Lima hians (Gmelin) from a station in the Skelmorlie Bank, Clyde Sea Area, were studied during 1969–1971. The composition of an animal of standard size was calculated for each sampling date to give information on seasonal changes independent of shell growth. Increases in tissue weight took place between April and July as the gonad developed and reserves were built up. Between July and March tissue weight fell, at first rapidly as a result of spawning in the population, and later more slowly as reserves were utilized. Differentiation of the gonad began during the winter, at the expense of stored reserves, but was greatly accelerated during the spring due to increased food from the plankton. The lipid and nitrogen content of a standard animal rose to a pre-spawning maximum in July and then fell. Carbohydrate content also rose during this period, but continued to rise to reach maximum values in September. Carbohydrate and nitrogen both declined during the winter, but there was little indication of any net utilization of lipid during the winter months. The tissues of the ripe female contained more lipid than the male, and the tissues of the ripe male contained more nitrogen than those of the female. The total calorific content of the tissues of the male was greater than that of the female.     

Effects of the 1996 and 1998 positive sea-surface temperature anomalies on corals,coral diseases and fish in the Arabian Gulf (Dubai,UAE)

Two positive sea-surface temperature anomalies occurred in the Arabian Gulf in short sequence. Between May and August 1996 and 1998, sea-surface temperatures in the southern Arabian Gulf were elevated by 2°C above average. The consequences for coral fauna, coral diseases and coral regeneration were studied in Dubai (United Arab Emirates) between Jebel Ali and Ras Hasyan. In 1996, coral death was widespread, affecting primarily the genus Acropora. In Acropora-dominated areas, live coral cover was reduced from 90% to about 26% in 1996, while in 1998 only a reduction from 26% to 22% of the remaining coral cover occurred. In the study area, all six Acropora species suffered total mortality in 1996, thus the coral fauna was reduced from 34 species to 27. The nearest areas with surviving Acropora were 30 km to the east (Deira) and 20 km to the west (Al Jazira). Massive coral species suffered negligible mortality, and slowly increased in space cover. The Acropora overkill turned 7.9 km² (19.7% of total coral-covered area) of previously lush coral gardens into a dead framework that was increasingly bioeroded. Acropora recruitment only started in 1998, average recruit size in 1999 was 7Dž cm, and recruits were rare. Prior to the mass mortality event, coral diseases were common and seasonal (14LJ% of corals, mainly Acropora, affected in summer, in winter 7Lj%, mainly massives), after the mortality event seasonality was lost and infection remained below winter levels (6LJ%, only massives infected). In fish, overall species richness decreased from 95 to 64 species in point counts, but frequency only decreased in one species (Pseudochromis persicus). Guild structure changed inasmuch as herbivores and planktivores increased, and invertivores decreased, although differences were not statistically significant. The most abundant family, both prior to and after the coral mass mortality, was Lutjanidae. It appears that even though much of the coral was dead, the maintenance of structural complexity allowed the fish assemblage to avoid a similar catastrophic change to that experienced by the coral assemblage.     

Seasonal changes in biochemical composition of the bivalve Chlamys septemradiata from the Clyde Sea Area

Seasonal changes in the tissue weight and biochemical composition of the pectinid bivalve Chlamys septemradiata (Müller) from a station in the Cumbrae Deep, Clyde Sea Area (UK), were studied during 1969–1971. Separate analyses were made of three divisions of the tissues: the adductor muscle, the gonad, and the remaining tissues referred to as mantle. The composition of each of these divisions, and of the total tissues of an animal of standard size was calculated for each sampling date to give information on seasonal changes independent of shell growth. Increase in gonad weight took place mainly between March and May, although there was a slow increase during the winter months (November-March). Spawning in July and August resulted in a fall in gonad weight to very low levels in September and October. The female gonad contained a greater proportion of lipid than the male, and the male a greater proportion of nitrogen than the female, although the total calorific content was similar in both sexes. The adductor muscle showed a clear seasonal cycle of tissue weight and composition, and it was demonstrated that the adductor muscle tissues are a site for storage of reserves. The weight of the adductor muscle was maximal in October-November, and declined through the winter to reach minimum values in March. There was a steady increase during the summer. Lipid, carbohydrate and protein all contributed to the decrease in winter, in part contributing to gonad differentiation but mainly meeting the bivalve's metabolic maintenance requirements. The remaining tissues displayed some irregular fluctuations, but in general the changes showed an upward trend in weight until November followed by a general downward trend to March. The changes are compared with those for other bivalves from the same area.     

Larval survivorship, competency periods and settlement of two brooding corals, Heliopora coerulea and Pocillopora damicornis

Larval dispersal and recruitment are important in determining adult coral distribution; however, few studies have been made of coral larval dispersal. This study examined the larval behavior, survivorship competency periods and settlement of two brooding corals, Heliopora coerulea and Pocillopora damicornis, in relation to different potential larval dispersal patterns. We also examined the lipid content of H. coerulea as a means of flotation and a source of energy. Planulae of H. coerulea were on average 3.7 mm in length, lacked zooxanthellae, and were mostly benthic, probably because of restricted movement and low lipid content (54% by dry weight). Planulae of P. damicornis were on average 1.0 mm in length, had zooxanthellae and swam actively. The competency period of H. coerulea was shorter (30 days) than that of P. damicornis (100 days). Forty percent of H. coerulea planulae crawled onto the substrata within 1 h of release, and 47% settled within 6 h. By contrast, fewer than 10% of P. damicornis planulae crawled onto the substrata within the first hour and 25% settled within 6 h of release. The planulae of H. coerulea may have a narrower dispersal range than those of P. damicornis, settling and recruiting near parent colonies. Thus, brooding corals exhibit variations in larval dispersal patterns, which are characterized by their position in the water column and competency periods.     

Reproductive cycle and gametogenesis in the black abalone Haliotis cracheroidii (Gastropoda: Prosobranchiata)

Seasonal changes in gonald size and stages of gametogenesis in the black abalone Haliotis cracheroidii were related to changes in environmental parameters. H. cracheroidii showed an annual reproductive cycle terminating in a synchronized spawning in late summer. Gametogenesis was initiated immediately after spawning. Gametes were present in the gonad through the winter months. Gametogenesis was initiated a second time in the spring months. Maximal gonad growth (to a gonad index of 20%) occurred during summer months prior to spawning. Changes in gonad size andperiods of initiation of gametogenesis revealed no apparent correlation with changes in seasonal water temperature. Changes in gonad size showed no apparent relation to change in day length. Total polysaccharide levels in foot tissue changed seasonally, indicating that food availability is probably not a factor in directly regulating gonad growth. Gonad index data for the chiton Katharina tunicata (collected over a 10 year period) showed no apparent correlation to seasonal change in water temperature.     

Reproductive cycle of the sea urchin Holopneustes purpurascens (Temnopleuridae: Echinodermata)

This study documents the annual reproductive cycle of the Temnopleurid echinoid Holopneustes purpurascens near Sydney. H. purpurascens reached sexual maturity between 11 and 26 mm diameter and was gonochoristic. The sequence of oogenesis was similar to previously described echinoids. However, the proportion of eggs at each stage showed no temporal pattern in abundance, suggesting that vitellogenesis is continuous throughout the year. Spermatogenesis was also similar to other echinoids and showed a significant increase in the proportion of partially spawned testes in spring, followed by a similar increase in the proportion of partially spawned testes in late spring and summer. Gonad indices showed a peak in weight from late winter to mid-spring (9.10ǃ.47 for females; 6.28ǂ.43 for males; mean-SE), then a decline for both sexes. Although spawning was variable over time, there was a peak in the proportion of spawning from late winter to early summer that correlated with the observed peaks in gonad indices and gametogenic cycles. The demographic implications of this reproductive pattern are that new recruits can enter the population through synchronous spawning during the breeding period, and smaller haphazard spawning events at other times of the year.     

Sexual and asexual reproduction of <Emphasis Type="Italic">Coscinasterias</Emphasis><Emphasis Type="Italic">tenuispina</Emphasis> (Echinodermata: Asteroidea) from Rio de Janeiro,Brazil

An intertidal population of the fissiparous sea star Coscinasterias tenuispina (Lamarck, 1816) was sampled over a 1 year period at Ponta de Itaipu, Niterói, Rio de Janeiro. Gonadal indices revealed an annual reproductive cycle. Spawning occurred in the late winter and early spring between August and November. Pyloric caecal indices did not show a clear annual cycle or a reciprocal relationship with gonadal indices. The population appears to be sustained only by fission, as only males were found. Fission occurred throughout the year, but was most frequent in the winter (July, August and September). Incidence of fission was correlated with the occurrence of the extremely low tides during daytime that exposes many specimens to air. Austral winter tides are low by day and high by night, while the austral summer tides are low by night and high by day. The most extreme annual deviation in seawater temperature is about 5°C. Larval recruitment seems to be negligible, since very small individuals were not found.     

Mate-guarding constrains feeding activity but not energetic status of wild male long-tailed macaques (Macaca fascicularis)

Mate-guarding is an important determinant of male reproductive success in a number of species. Little is known however about the constraints of this behaviour, e.g. the associated energetic costs. We investigated these costs in long-tailed macaques where alpha males mate guard females to a lesser extent than predicted by the priority of access model. The study was carried out during two mating periods on three wild groups living in the Gunung Leuser National Park, Indonesia. We combined behavioural observations on males’ locomotion and feeding activity, GPS records of distance travelled and non-invasive measurements of urinary C-peptide (UCP), a physiological indicator of male energetic status. Mate-guarding led to a decrease in feeding time and fruit consumption suggesting a reduced intake of energy. At the same time, vertical locomotion was reduced, which potentially saved energy. These findings, together with the fact that we did not find an effect of mate-guarding on UCP levels, suggest that energy intake and expenditure was balanced during mate-guarding in our study males. Mate-guarding thus seems to not be energetically costly under all circumstances. Given that in strictly seasonal rhesus macaques, high-ranking males lose physical condition over the mating period, we hypothesise that the energetic costs of mate-guarding vary inter-specifically depending on the degree of seasonality and that males of non-strictly seasonal species might be better adapted to maintain balanced energetic condition year-round. Finally, our results illustrate the importance of combining behavioural assessments of both energy intake and expenditure with physiological measures when investigating energetic costs of behavioural strategies.     

Seasonal cycle of gonadal development and serum levels of vitellogenin of the red porgy, Pagrus pagrus (Teleostei: Sparidae)

Histological examination of gonads of female and male red porgy, Pagrus pagrus, reared in captivity, was carried out in order to describe the main gonadal changes related to gametogenesis and the seasonal changes of environmental factors. The gonadosomatic index (GSI) and the concentration of serum vitellogenin (VTG) were also determined. The frequency distribution of gonad development stages and the GSI and vitellogenin concentration during the annual cycle indicated the separation of the female and male reproductive cycles into three main periods. The autumn period when gametogenesis begins (October-November), the period of exogenous vitellogenesis (December-March) or spermiation (December-March), and the spawning season (March-May). The spawning period coincided with an increase in temperature (15-19°C) and daylength. Serum levels of vitellogenin rose significantly in January, reached a peak at the beginning of the spawning period (March, 405.5 µg ml^-1) and remained high until the end of the spawning period (May).     

Seasonal variation in ecophysiological patterns in macroalgae from an Arctic fjord. I. Sensitivity of photosynthesis to ultraviolet radiation

Marine macroalgae inhabiting Arctic coastal ecosystems are exposed to pronounced seasonal variations in the radiation regime, including harmful UVB radiation. This study presents the first data on the seasonal changes in the sensitivity of macroalgal photosynthesis towards UV exposure by comparing under-ice, clear-water and turbid-water conditions characteristic for late winter, spring and summer. Various brown (Laminaria saccharina, L. digitata, L. solidungula, Saccorhiza dermatodea, Desmarestia aculeata), red (Palmaria palmata, Devaleraea ramentacea) and one green macroalgal species (Monostroma aff. arcticum) were collected at the same water depth throughout the seasons in the Kongsfjord (Spitsbergen, Svalbard, Norway). Maximum quantum yield (F_v/F_m) and maximum photosynthetic electron transport rates (ETR_max) were determined immediately after collection, after 2 h exposure to artificial UV radiation and after 18 h recovery in dim white light. Photosynthesis of the studied species showed different responses depending on their morpho-functional and physiological characteristics, their life strategies, phenology and depth distribution. Within the genus Laminaria, maximum quantum yield of adult specimens of the deep-water species L. solidungula was most strongly UV sensitive. Adult L. saccharina exhibited a lower UV sensitivity than a 6-month-old specimen. Inhibition of photosynthesis after UV exposure remained at the same level throughout the study period, both in adult L. saccharina and S. dermatodea. However, adult specimens of L. saccharina collected in May showed partial recovery only, whereas photosynthesis of specimens from both species collected later recovered fully. D. aculeata exhibited a remarkable decrease of UV sensitivity during the study period. Photosynthesis of specimens collected under the ice was strongly inhibited by UV, but the degree of inhibition decreased during spring and summer. Concomitantly ETR_max values were low after UV exposure in specimens collected in June, but increased later in the season. P. palmata exhibited a relatively flexible response. Photosynthesis in specimens collected under the ice in June or in turbid water in July/August was relatively strongly inhibited; specimens collected during sunny periods and in clear water in spring showed a much lower degree of photoinhibition after UV exposure. The seasonal pattern of low/high ETR_max values in spring/summer is probably a characteristic of the life strategy of this species. The UV sensitivity of D. ramentacea exhibited a similar seasonal pattern. In M. aff. arcticum, UV sensitivity increased and ETR_max values decreased during the study period, reflecting the life strategy of this annual late winter/spring species. The physiological basis for the seasonal changes in UV sensitivity of photosynthesis is presented in a companion paper (this issue).     

Reproductive cycle of two commercial species of sea cucumber (Echinodermata: Holothuroidea) from Caribbean Panama

The reproductive status of the holothuroid species Isostichopus badionotus (Selenka, 1867) and Holothuria mexicana (Ludwig, 1875) was studied over 16 months in Bocas del Toro (Panama), from November 1999 to February 2001. Sexual reproduction was evaluated by the gonad index method, and by histology of gonad development. In addition, population structure was assessed based on sex ratio, minimum reproductive size, and length and weight distributions of males and females. The sex ratio in both species was 1:1, with a unimodal population distribution composed mainly of mature individuals. The minimum reproductive length and weight were 13-20 cm and 150 g, respectively, for both species, although reproductive individuals 10 cm in length were also found. A consistently higher gonad index was observed in H. mexicana, due to a high proportion of mature females and males and high gonad indices in most monthly samples. Gametogenesis and spawning patterns seemed to occur throughout the year, with periods of enhanced activity. Two periods of maximum reproductive activity were tentatively identified: July-November for I. badionotus and February-July for H. mexicana, but neither species had a single, sharply defined annual spawning event. Further work on these exploited holothuroids should examine the relationships between reproduction and environmental factors and between reproductive status and recruitment.     

Reproductive development of the barnacle Chthamalus malayensis in Hong Kong: implications for the life-history patterns of barnacles on seasonal, tropical shores

Hong Kong, lying just below the Tropic of Cancer (22°17′N, 114°09′E), experiences a strongly seasonal environment, with a cool almost temperate winter and a hot, tropical, summer. Histological sectioning of the gonads of the high-shore barnacle, Chthamalus malayensis Pilsbry, showed a seasonal trend in the development of its reproductive organs. Four stages of female gonad development were identified according to the cell types present: post-spawning, resting, growth and mature stages. The female gonad was mature from April to November, which was related to seawater temperatures, and entered a resting phase from December to March. Although the male gonad showed a seasonal developmental trend and reached maximum maturity in summer, the seminal vesicles were full of spermatozoa and functional throughout the year. The reproductive season of this species is therefore solely dependant upon the maturity of the female gonad. The estimated maximum number of broods per year was up to 10 and the maximum number of eggs produced per brood can reach 3,000 eggs. The minimum size for female gonad maturity was 6 mm rostro-carinal diameter (RCD) at which size, the barnacles were ~6-month old. Sperm production occurred at a smaller size (2 mm=2-month old). Compared with Chthamalus montagui and Chthamalus stellatus from temperate regions, C. malayensis produced a greater number of broods per year, had a longer reproductive period and faster gonad development. Chthamaloid barnacles in tropical regions may, therefore, invest more energy per year in reproduction during their life span. Contrary to the seasonal gonad developmental pattern of C. malayensis in the present study, however, C. malayensis in Singapore (which experiences only slight seasonal variation) had mature female and male gonads throughout the year, further supporting the strong role of climatic conditions effecting the reproductive biology of barnacles.     

Winter mortality, growth, and behavior of young-of-the-year of four coastal fishes in New Jersey (USA) waters

Winter mortality has been hypothesized to select for large body size in young-of-the-year (YOY) fishes, yet substantiation of winter mortality and its cause(s) are available for few estuarine or marine species. We examined seasonal length distributions of wild populations of four common marine species, black sea bass (Centropristis striata), tautog (Tautoga onitis), cunner (Tautogolabrus adspersus), and smallmouth flounder (Etropus microstomus), and mortality (i.e., frequency of death), growth, and behavior of their YOY in the laboratory at ambient winter temperatures (mean 7°C, range 2-13°C) during a 135-day period (December 1992 through mid-April 1993) to establish potential causes of their mortality in the field. Young-of-the-year black sea bass experienced 100% mortality when water temperatures decreased to 2-3°C in February, emphasizing the importance of winter emigration from estuaries in this southern species. The low mortality of two labrid species, YOY tautog (14%) and YOY cunner (3%), was consistent with their northern distribution and year-round occurrence in estuarine and nearshore coastal waters. Laboratory mortality of YOY smallmouth flounder (33%) was higher for small (<35 mm total length) fish, suggesting that this small species may experience high winter mortality in estuaries and nearshore coastal waters. Seasonal differences in fish length result potentially from several mechanisms (e.g., mortality and/or migration) that are difficult to assess, but our laboratory experiments suggest that seasonal temperature changes cause size-specific mortality of YOY smallmouth flounder and offshore migration of YOY black sea bass.     

Seasonal variations in biochemical constituents during the reproductive cycle of the female dog cockle Glycymeris glycymeris

In an attempt to describe the biochemical events associated with the main stages of the annual and reproductive cycles of the female dog cockle Glycymeris glycymeris L., we studied seasonal variations in the various stages of oocyte development of the ovaries, and the glycogen, total protein and total lipid content of five body tissues – adductor muscle, foot, tunic coat, visceral mass and mantle. From November 1991 to November 1994, microscopic examination of the ovaries and measurement of the tissue concentrations of glycogen, total proteins and total lipids in these five body tissues were made monthly on ten female dog cockles originating from the sea area around Douarnenez (south Brittany, France). Morphological studies revealed that in the population investigated the annual cycle is characterised by three major periods: a first period of vitellogenesis extending from February/March to April/May and preceding a spawning in spring; a second period of vitellogenesis extending from May/June to September/October and leading to either no spawning, a single autumnal spawning event, or to two spawning events in summer and autumn; and a third period extending from October/November to February/March and characterised by a high level of oocyte lysis. In the muscular body tissues of the dog cockle, i.e. the adductor muscles, the foot and the tunic coat (the muscular envelope containing the visceral mass), the concentrations of glycogen, total proteins and total lipids underwent very similar variations during the annual cycle. During each stage of vitellogenesis, a typical glycogen–protein–lipid sequence was observed in the muscular tissues that was characterised firstly by a peak of glycogen concentration 2 to 3 mo before spawning, followed by a peak in total proteins 1 mo before spawning, and finally by a peak in lipid content just before spawning. A similar glycogen–protein–lipid sequence was also recorded in the first half of the winter period. However, these events were followed by general atresia affecting all oocytes in the gonad. Maximum energetic value of biochemical constituents in females coincided with peaks in lipid content in the visceral mass and mantle. These biochemical events occurred principally immediately before and at the end of oocyte lysis (December/January). A drop in the total energetic value, affecting mainly the visceral mass and the mantle, was recorded each year during the period January to March, coinciding with the period of shell growth in this species. Our data clearly indicate that in female G. glycymeris all muscular tissues contribute to the storage of glycogen and proteins, and suggest that glycogen may be the source of energy triggering vitellogenesis. Biochemical and microscopic observations revealed that oocyte development takes place during the first half of winter, but that these oocytes undergo atresia in December/January. The metabolites produced from oocyte lysis could contribute to somatic growth, which occurs in late winter. Received: 3 March 1997 / Accepted: 23 July 1997     

Seasonality of reproduction in the cushion star Odontaster validus at Signy Island, Antarctica

The reproductive and larval biology of the common Antarctic cushion-star Odontaster validus Koehler (Echinodermata: Asteroidea) were studied in a shallow-water population at Signy Island in the maritime Antarctic. The gonad index peaked in April/May, and injection with 1-methyladenine indicated that competence to spawn peaked in May/June. Gastrulae were first detected in May, and maximal water-column larval densities were measured in July. Reproduction was therefore highly seasonal, although not all individuals appeared to spawn each year. The pyloric caeca index increased in summer (typically November to March, but variable between years) and decreased during winter (June to October), suggesting that feeding activity was seasonal. All aspects of biology investigated showed significant interannual variability. Comparison with previous studies in the high Antarctic at McMurdo Sound indicated a similar if more pronounced seasonality at Signy, but the warmer seawater temperatures in the maritime Antarctic (particularly in late winter/early spring from September onwards) resulted in faster larval development at Signy. Received: 21 April 1997 / Accepted: 28 January 1998