Temporal succession and spatial segregation of clupeoid larvae in the coastal waters off the Tanshui River Estuary, northern Taiwan

In the coastal waters off the Tanshui River Estuary in northern Taiwan, eight species of clupeoid larvae were observed. They exhibited a distinct temporal succession in association with seasonal temperature changing. The time of peak abundance of Etrumeus teres was in January, Engraulis japonicus in late April, Sardinella spp. in mid-May, Dussumieria elopsoides in early June, Thryssa dussumieri in late June, Stolephorus insularis in mid-September, Encrasicholina heteroloba in early October and E. punctifer in mid-November. The time intervals of the temporal succession of the fishes were approximately 15 to 25 d in the spring/summer and 25 to 35 d in the autumn/winter fishing seasons. Also, they showed spatial segregation by distributing in areas with different water depths: Sardinella spp. at a water depth of 10 to 20 m, T. dussumieri at less than 10 m, E. heteroloba at 20 to 50 m and E. punctifer at 10 to 40 m. The larvae of these sympatric clupeoid species segregated their nursing periods and areas apparently to reduce competition for habitat and thus to maximize the utilization of resources. Received: 3 March 1997 / Accepted: 19 March 1997     

Functional response of sport divers to lobsters with application to fisheries management.

Fishery managers must understand the dynamics of fishers and their prey to successfully predict the outcome of management actions. We measured the impact of a two-day exclusively recreational fishery on Caribbean spiny lobster in the Florida Keys, USA, over large spatial scales (>100 km) and multiple years and used a theoretical, predator-prey functional response approach to identify whether or not sport diver catch rates were density-independent (type I) or density-dependent (type II or III functional response), and if catch rates were saturated (i.e., reached an asymptote) at relatively high lobster densities. We then describe how this predator-prey framework can be applied to fisheries management for spiny lobster and other species. In the lower Keys, divers exhibited a type-I functional response, whereby they removed a constant and relatively high proportion of lobsters (0.74-0.84) across all pre-fishing-season lobster densities. Diver fishing effort increased in a linear manner with lobster prey densities, as would be expected with a type-I functional response, and was an order of magnitude lower in the upper Keys than lower Keys. There were numerous instances in the upper Keys where the density of lobsters actually increased from before to after the fishing season, suggesting some type of "spill-in effect" from surrounding diver-disturbed areas. With the exception of isolated reefs in the upper Keys, the proportion of lobsters removed by divers was density independent (type-I functional response) and never reached saturation at natural lobster densities. Thus, recreational divers have a relatively simple predatory response to spiny lobster, whereby catch rates increase linearly with lobster density such that catch is a reliable indicator of abundance. Although diver predation is extremely high (approximately 80%), diver predation pressure is not expected to increase proportionally with a decline in lobster density (i.e., a depensatory response), which could exacerbate local extinction. Furthermore, management actions that reduce diver effort should have a concomitant and desired reduction in catch. The recreational diver-lobster predator-prey construct in this study provides a useful predictive framework to apply to both recreational and commercial fisheries, and on which to build as management actions are implemented.     

Seasonal reproductive biology of the egg-carrying calanoid copepod Pseudodiaptomus marinus in a eutrophic inlet of the Inland Sea of Japan

In situ egg production of the egg-carrying calanoid copepod Pseudodiaptomus marinus was investigated in Fukuyama Harbor, a eutrophic inlet of the Inland Sea of Japan, at 3- to 5-d intervals for a year. This species reproduced throughout the year, and the adults showed a large abundance peak in June/July and a small peak in September/October. Females usually outnumbered males, comprising 61.4% of the annual mean. The composition of ovigerous females varied from 7.9 to 100%, with an annual mean of 55.7%. Adult prosome length was consistently large throughout winter and spring, and decreased with increasing temperature in summer and fall. Egg diameter varied from 98 to 121 μm, and was negatively correlated to temperature. The seasonal variation in clutch size (range: 15.1 to 38.2 eggs) was bicyclical, with peaks in May and December. The egg production rate of breeding females was low in January to March (mean: 2.3 eggs female⁻¹ d⁻¹), while it was constantly high from mid-May to early October (mean: 12.1 eggs female⁻¹ d⁻¹). The specific egg production rate for the breeding females was highly correlated to temperature; it increased linearly from 0.03 d⁻¹ at 9 °C to 0.27 d⁻¹ at 26 °C. Compared to other co-occurring copepods, the reproductive rate of P. marinus was lowest, which is one of the reasons why this species never dominates in this inlet. Received: 11 November 1996 / Accepted: 7 December 1996     

Variability in the chemical defense of the sponge Chondrilla nucula against predatory reef fishes

Chondrilla nucula is a common Caribbean demosponge that grows in a range of habitats, from coral reefs to mangrove swamps. On reefs, C. nucula grows as a thinly encrusting sheet, while in mangrove habitats it surrounds submerged mangrove roots as fleshy, lobate clumps. Previous feeding experiments using predatory reef fish revealed a high degree of variability in the chemical defenses of C. nucula. The present study was undertaken to determine whether a relationship exists between habitat, growth form, and chemical defense of C. nucula. Both laboratory and field feeding-assays of crude extracts confirmed that C. nucula possesses a chemical defense with high intercolony variability, but there was no significant variation in feeding deterrency between reef and mangrove habitats at either geographic location (Bahamas and Florida). Extracts of C. nucula collected during September and October 1994 from the Bahamas were significantly more deterrent than those collected during August 1993, May 1994, and May 1995 from Florida, and extracts of these spring and summer Florida collections were more deterrent than extracts of C. nucula collected in December 1994 and February 1995 in the same locations. There was no evidence that deterrent compounds were concentrated in the surface tissues of the sponge, or that chemical defense could be induced by simulated predation. Laboratory and field assays of the fractionated crude extract revealed that feeding deterrency was confined to the most polar metabolites in the extract. Field transplants were used to determine whether predation influenced the growth form of C. nucula. Uncaged sponges transplanted from the mangrove to the reef were readily consumed by spongivorous reef fishes. Lobate mangrove sponges became thinner after being caged on the reef for 3 mo, but encrusting reef sponges did not become thicker after being caged in the mangroves for the same period of time. Reef sponges that were caged for 3 to 15 mo thickened by only a small amount (<1 mm) compared to uncaged and open-caged (i.e. in cages lacking tops) sponges. Simulated bite marks on both reef and mangrove sponges were repaired at a rapid rate (0.8 to 1.6 mm d⁻¹). Fish predation has an important impact on the distribution and abundance of C. nucula, but the thin growth form common to reef environments may be more the result of hydrodynamics than of grazing by spongivorous fishes. Received: 6 October 1997 / Accepted: 19 March 1998     

Behaviour and tolerance to hypoxia in juvenile Norway lobster (Nephrops norvegicus ) of different ages

The annual occurrence of hypoxia (<25% oxygen saturation) in the bottom waters along the Swedish west coast coincides with the postlarval settlement of Norway lobster, Nephrops norvegicus (L.). This study investigates behaviour and the experimental effects of low oxygen concentrations in juvenile N. norvegicus of different ages. All experimental individuals were reared to the juvenile (postlarval) stage in the laboratory and then given sediment as a substratum. Behavioural responses to low oxygen concentrations were tested in early and late Postlarvae 1 exposed to normoxia (>80% oxygen saturation, pO₂ > 16.7 kPa), moderate hypoxia (30% oxygen saturation, pO₂ = 6.3 kPa) and hypoxia (25% oxygen saturation, pO₂ = 5.2 kPa). The experiments were run for a maximum period of 24 h or until individuals died. Behaviour was studied using sequential video recordings of four behavioural activities: digging, walking, inactivity or flight (escape swimming up into the water column). Behaviour and mortality changed with lowered oxygen concentrations; energetically costly activities (such as walking) were reduced, and activity in general declined. In normoxia, juveniles initially walked and then burrowed, but when exposed to hypoxia they were mainly inactive with occasional outbursts of escape swimming. To increase oxygen availability the juveniles were observed to raise their bodies on stilted legs (similar to adults in hypoxic conditions), but oxygen saturations of 25% were lethal within 24 h. The results suggest that the main gas exchanges of early postlarval stages occur over the general body surface. Burrowing behaviour was tested in Postlarvae 1 and 2 of different ages held in >80% oxygen saturation for 1 wk. The difference in time taken to complete a V-shaped depression or a U-shaped burrow was measured. The results showed a strong negative relationship between postlarval age and burrowing time, but all individuals made a burrow. Juveniles were more sensitive to hypoxia than adults. Thus, the possible consequences of episodic hypoxia for the recruitment of Nephrops norvegicus and for the recolonization of severely affected areas are discussed. Received: 4 August 1996 / Accepted: 11 October 1996     

Distribution and abundance of ctenophores and their zooplankton food in the Black Sea. II. Mnemiopsis leidyi

The distribution of Mnemiopsis leidyi Agassiz, 1865 in the Black Sea was determined using plankton samples collected above the anoxic zone (maximum depth 200 m) in the summer, winter, and spring from 1991 to 1995. Distribution was patchy. Average biomasses of 15 to 500 g m⁻² were measured, and abundances varied from 10 to 180 ind m⁻². Biomass and abundance peaked in winter, and there was a secondary peak in the summer. The distribution of M. leidyi was correlated with hydrographic features in the Black Sea with higher concentrations in anticyclonic gyres. The centers of the two main cyclonic gyres generally had a low biomass of M. leidyi. From July 1992 to March 1995, the populations were largely offshore. M. leidyi were confined to the upper part of the mixed layer both day and night. Some individuals displayed a negative taxis to daylight and were concentrated below the thermocline at night. Smaller M. leidyi (1.5 to 2 cm) were present in the winter, and individuals reached maximum size in the summer. Although reproduction was continuous throughout the year, there were two distinct peaks: the larger peak in the summer and the smaller peak in the winter. Microscopic analysis of stomach contents showed that copepods and molluscs form their main diet. Received: 1 November 1997 / Accepted: 30 August 1999     

Diurnal and lunar patterns of larval recruitment of Brachyura into a mangrove estuary system in Ranong Province, Thailand

The present study followed the temporal recruitment pattern of brachyuran larvae in a mangrove tidal creek on the Andaman Sea coast of Ranong Province, Thailand, based on the assumption that the processes governing recruitment are important for the overall population dynamics of mangrove brachyuran crabs. Plankton net samples were taken on five occasions: on two new moon spring tides, one waxing moon neap tide, one full moon spring tide and one waning moon neap tide during October and November 1997. In addition collectors for larval crab megalopae were employed every 3 days through one dry season and one wet season (March–October 1998). Both the plankton net samples and collector samples revealed four major brachyuran groups in three families: Ocypodidae, Grapsidae and Portunidae. The grapsid group was further separated into two morphotypes which were identified as Metaplax and sesarmid species. Identified group mean numbers per cubic metre were ocypodids 3.0, sesarmids 0.8 and Metaplax 0.5, while portunid megalopae were very scarce (≪0.1 m⁻³). Further analysis of plankton net samples showed that when considering the parameters date, depth, current direction and the diel cycle, Metaplax and ocypodids distribute differently in the tidal and lunar cycle. Metaplax recruitment dominates on flood tides and on bottom layers, followed by middle and surface layers. Conversely, ocypodid abundance varied significantly with date only. Notably recruitment was not dependent on the diel cycle for either group. The collector samples of megalopae showed that recruitment of ocypodids, Metaplax and sesarmids occurred on full and new moon spring tides, while portunid megalopae preferred to settle on full moon spring tides. Since tidal currents were related to the lunar cycle megalopa groups are also cross-correlated with tidal amplitude, except for the portunid group. It is concluded that megalopae recruit in a similar manner to what has been found in other regions of the world, except that the abundance of ocypodids and Metaplax is not influenced by the diel cycle. Received: 14 February 2000 / Accepted: 24 November 2000     

Distribution and abundance of moon jellyfish (Aurelia aurita) and its zooplankton food in the Black Sea

The distribution of moon jellyfish (Aurelia aurita Linnaeus, 1758) in the Black Sea was determined from plankton samples collected above the anoxic zone (maximum depth 200 m) in the summer, winter and spring during 1991–1995. Distribution was patchy. Average biomass ranged from 98 to 380 g m⁻², and abundance varied from 2 to 14 individuals m⁻². Biomass and abundance peaked in late spring and summer. The distribution was correlated with hydrographic features in the Black Sea, with higher concentrations occurring at the peripheries of anticyclonic eddies. Centers of the two main cyclonic gyres generally had a low biomass of A. aurita. From July 1992 to March 1995, the populations were largely concentrated in offshore regions. A. aurita were confined to the upper part of the mixed layer. Smaller A. aurita (≤1 cm) were present in early spring (March), and individuals reached maximum size in the summer. Release of the epyhrae occurred in spring on the northwestern shelf of the sea when the seawater temperature was 11–12 °C. Microscopic analysis of stomach contents showed that copepods and mollusks form their main diet. Received: 3 September 2000 / Accepted: 29 September 2000     

The reproductive biology of a new species of sea cucumber, Holothuria (Mertensiothuria) arenacava in a Kenyan marine protected area: the possible role of light and temperature on gametogenesis and spawning

The sea cucumber Holothuria arenacava was discovered in the Mombasa marine reserve in 1997 and described by Samyn et al. (2001). The reproductive biology of this holothurian was investigated in order to (1) characterize the reproductive pattern, (2) examine the relationship among environmental parameters including temperature, light and lunar period, and (3) examine the relationship between the reproductive pattern and feeding of this new species. The gonad index method and microscopic examination of gonads was used to analyze samples collected for a period of 13 months. H. arenacava displayed an annual reproductive cycle with gametogenesis commencing in July during the south-east monsoons, when temperature and light intensity are lowest along the Kenyan coast. Gonad growth peaked in February–March at the end of the north-east monsoons when temperatures and light reach their annual maxima along the Kenyan coast. The higher correlation between light intensity and gonad growth (r=93) than temperature (r=0.71), coupled with the fact that temperatures continued to drop for a month after gametogenesis had already commenced, suggests that light intensity and not temperature is the cue for the onset of gametogenesis in this species. Spawning was synchronized between females and males and occurred during a short period between March and May (inter-monsoonal period) when both temperature and light intensity decrease along the Kenyan coast. Male and female gonad indices showed significant variation with lunar day and no lunar periodicity was observed in this sea cucumber. The sex ratio of the population of H. arenacava was skewed towards significantly more females than males, and females were significantly larger and had larger gonads and gonad indices than males. These life history strategies including spawning during a short discrete period, more and larger females that have larger gonads (i.e., typically more fecund), and spawning just prior to the peak in phytoplankton concentrations, a time that is probably more favorable for larval development, may serve to increase the reproductive success of this sea cucumber.     

Annual cycles of phytoplankton community-structure and bloom dynamics in the Neuse River Estuary, North Carolina

The spatiotemporal distributions of major phytoplankton taxa were quantified to estimate the relative contribution of different microalgal groups to biomass and bloom dynamics in the eutrophic Neuse River Estuary, North Carolina, USA. Biweekly water samples and ambient physical and chemical data were examined at sites along a salinity gradient from January 1994 through December 1996. Chemosystematic photopigments (chlorophylls and carotenoids) were identified and quantified using high-performance liquid chromatography (HPLC). A recently-developed factor-analysis procedure (CHEMTAX) was used to partition the algal group-specific chlorophyll a (chl a) concentrations based on photopigment concentrations. Results were spatially and temporally integrated to determine the ecosystem-level dynamics of phytoplankton community-constituents. Seasonal patterns of phytoplankton community-composition changes were observed over the 3 yr. Dinoflagellates reached maximum abundance in the late winter to early spring (January to March), followed by a spring diatom bloom (May to July). Cyanobacteria were more prevalent during summer months and made a large contribution to phytoplankton biomass, possibly in response to nutrient-enriched freshwater discharge. Cryptomonad blooms were not associated with a particular season, and varied from year to year. Chlorophyte abundance was low, but occasional blooms occurred during spring and summer. Over the 3 yr period, the total contribution of each algal group, in terms of chl a, was evenly balanced, with each contributing nearly 20% of the total chl a. Cryptomonad, chlorophyte, and cyanobacterial dynamics did not exhibit regular seasonal bloom patterns. High dissolved inorganic-nitrogen loading during the summer months promoted major blooms of cryptomonads, chlorophytes, and cyanobacteria. Received: 12 September 1997 / Accepted: 12 December 1997     

Seasonality of asexual reproduction in Holothuria (Halodeima) atra, H. (H.) edulis and Stichopus chloronotus (Holothuroidea: Aspidochirotida) on the Great Barrier Reef

Asexual reproduction by fission was monitored for 18 mo in populations of Holothuria (Halodeima) atra, H. (H.) edulis and Stichopus chloronotus on three nearshore fringing reefs and one midshelf reef in the Great Barrier Reef. Fission in S. chloronotus occurred exclusively between March and October, with a peak value of 31% recently divided individuals in one population in July. H. atra showed a similar pattern, with maxima of between 16 and 26% from May to July. In H. edulis, asexual reproduction occurred only between March and July, with a maximum of 17% recently divided individuals in March. Fission rates of H. atra and S. chloronotus in winter were significantly higher than in all other seasons. For H. atra, at least 76% of all individuals at Fantome Island were estimated to undergo fission per year, whereas only 9% undergo fission on the midshelf reef. Highest annual fission rates (43%) for S. chloronotus were found in a dense population on Great Palm Island. The lower-density midshelf reef population exhibited comparatively lower annual fission rates (19%). About 24% of H. edulis undergo fission each year. Annual fission rate and population density were positively correlated in the four populations of S. chloronotus and H. atra studied. Received: 21 March 1997 / Accepted: 26 March 1997     

Temporal variation of recruits as a basis of ephemeral genetic heterogeneity in the western rock lobster Panulirus cygnus

Allozymes were examined in quantitative lunar monthly collections of larval recruits of the western rock lobster Panulirus cygnus George over three recruitment seasons at two sites nearly 350 km apart in Western Australia. At Alkimos, the southern site, recruitment occurs in a relatively narrow peak early in the spring, whereas at the northern Houtman Abrolhos Islands, recruitment extends into the summer months. In the 1995/1996 recruitment season, the frequency of the GPI ^* 100 allele increased from early to late in the season, but the frequencies were indistinguishable at the two sites in each monthly collection. The combination of this temporal variation in allelic frequencies with the contrasting patterns of recruitment at the Abrolhos Islands and Alkimos resulted in genetically different cohorts at the two sites. This pattern was ephemeral, as it was not repeated in the subsequent two years. Thus, ephemeral genetic patchiness in P. cygnus can be generated by the locally-specific genetic mix of recruits obtained from a common larval pool. This mechanism is the probable explanation of previously observed temporal and possible spatial genetic variation in adult P. cygnus, and highlights the importance of studying recruitment in order to understand the genetic structure of marine species. Received: 22 February 1999 / Accepted: 8 June 1999     

Distribution and abundance of ctenophores, and their zooplankton food in the Black Sea. I. Pleurobrachia pileus

The distribution of Pleurobrachia pileus Müller, 1776 in the Black Sea was determined using plankton samples collected above the anoxic zone (maximum of 200 m) in the winter, spring, and summer of 1991 to 1995. The summer samples were collected in 1991 to 1993 (for a previous) and are included in this paper for comparative purposes. High concentrations of P. pileus were found at the northern edges of anticyclonic eddies along the southern coastal regions. The biomass and abundance of P. pileus increased from winter through spring to a peak in summer. The highest mean wet weight during a sampling period was 250 g m⁻², while the maximum wet weight was 1429 g m⁻². P. pileus was mostly found in a layer extending from the lower parts of the thermocline down to the anoxic zone, where the temperature was <8 °C. The vertical distribution of P. pileus biomass had two clear maxima at night: an upper maximum at 20 to 40 m was less pronounced than the lower maximum at 90 to 120 m depth. Mean body length of P. pileus did not exceed 12 mm. Smaller individuals (9 to 10 mm length) occurred in winter. P. pileus had two length classes in early spring (March 1995) and late summer (August 1993), indicating the presence of both newly hatched and larger individuals. Overall, the stomach contents of P. pileus consisted mainly of Copepoda (90%), Cladocera (1%), Mollusca (1%), fish eggs and larvae (1%), and other taxa (7%). The preferred food of P. pileus (frequency of occurrence) was: Calanus euxinus (39%), Pseudocalanus elongatus (30%), Acartia clausi (28%), Oithona similis (2%), and Paracalanus parvus (1%). The endoparasite Hysterothylacium aduncum was commonly found in P. pileus. Abundances of Mnemiopsis leidyi and P. pileus were either negatively correlated (r = −0.5 to −0.7) or positively correlated at a low significance level (r = 0.25 to 0.3) with abundance of A. clausi in different months of the year. Aurelia aurita abundance was correlated mainly with the abundance of C. euxinus from June 1991 to March/April 1995. Over the same period the abundance of P. pileus was significantly correlated with the abundance of P. elongatus, an important prey species. Received: 1 November 1997 / Accepted: 30 August 1999     

Reproduction and distribution of the invasive European fanworm Sabella spallanzanii (Polychaeta: Sabellidae) in Port Phillip Bay, Victoria, Australia

 The European fanworm Sabella spallanzanii (Gmelin, 1791) was recently introduced to Port Phillip Bay and is now a conspicuous component of most benthic communities. Reproduction of the worm was investigated in a population at Queenscliff over a 2 yr period (October 1995 to October 1997) using gonadal histology. The worms are dioecious (sex ratio 1:1, n=250), and attained sexual maturity at ∼50 mm body length. Reproductive periodicity followed a distinct annual cycle, and spawning proceeded through an extended autumn/winter period. Spawning was broadly synchronous between sexes, and coincided with falling seawater temperatures and shorter day-lengths. The females were highly fecund, and >50 000 eggs were probably shed from large females (>300 mm body length) during the annual spawning period. Breeding cycles of S. spallanzanii in Port Phillip Bay are ∼6 mo out of phase with endemic populations located at similar latitudes in the northern hemisphere. The spread of S. spallanzanii within Port Phillip Bay has been monitored by divers on an annual basis since 1994. The most recent dive survey (1998) indicates that S. spallanzanii has extended its range through out the entire 2000 km² embayment, and has invaded most subtidal habitats. Quantitative estimates of S. spallanzanii abundances were highest on pier pylons (12.5 individuals m⁻², 0.5 to 7 m depths). On sediments, estimates were highest at shallow sites (0.3 m⁻², 7 m depth), but numbers declined significantly with depth (0.1 m⁻², 17 to 22 m depth). Mean worm lengths and biomass were, by contrast, significantly higher at intermediate depths (12 to 17 m) than in shallower (7 m) or deeper (22 m) locations. S. spallanzanii demonstrates a clear preference for growth in sheltered, nutrient-enriched waters, so it may not spread from Port Phillip Bay into the adjacent oceanic waters of Bass Strait; however, in view of S. spallanzanii's current high abundance, fecundity and extended spawning periodicity, there is a high risk of future range expansions, mediated by shipping, into other temperate-water ports. Received: 17 November 1998 / Accepted: 6 January 2000     

Sink or swim? Copepod population maintenance in the Columbia River estuarine turbidity-maxima region

Maintenance of estuarine zooplankton populations in large river-dominated estuaries with short residence times has been an intriguing subject of investigation. During three different hydrological seasons, autumn 1990, summer 1991, and spring 1992, we intensively sampled zooplankton populations in the estuarine turbidity maxima (ETM) region of the Columbia River estuary of Oregon and Washington, USA. One of the principal objectives was to investigate retention mechanisms of the predominant zooplankton species, the harpacticoid copepod Coullana canadensis and the epibenthic calanoid copepod Eurytemora affinis. In the ETM, C. canadensis densities mirrored those of turbidity gradients and were almost always greater at the river bed, while E. affinis densities were greater higher in the water column during the flood and lower in the water column during the ebb. Cross-correlation and time-series analyses determined that C. canadensis densities were highly positively correlated with turbidity and that most of the variability was explained by the lunisolar diurnal (K₁) and principal lunar (M₂) tidal components occurring once every 23.93  h and once every 12.42 h, respectively. This indicates that C. canadensis populations are most probably maintained in the estuary through the same near-bottom circulation features that trap and concentrate particles in the ETM. In contrast, densities of the more motile species E. affinis were highly correlated with negative velocities, or ebb tide, and most of the variability in population densities could be explained by the principal lunar tidal component; therefore, we hypothesize that this species is probably vertically migrating on a tidal cycle into different flow layers to avoid population losses out of the estuary. Received: 7 April 1997 / Accepted: 22 April 1997     

Composition and temporal fluctuations of inshore juvenile fish populations in the Kornati Archipelago, eastern middle Adriatic

Temporal fluctuations of species composition and abundance of fish juveniles in the National Park Kornati were studied over 12 months from January to December 1992. A total of 40 168 individuals, representing 24 families and 69 species, were collected using a 50 m long beach seine. The community was dominated numerically by a few species: Atherina hepsetus (45.90%), Sarpa salpa (21.21%), Diplodus vulgaris (10.49%), and Symphodus ocellatus (6.98%), constituting over 84.58% of the total catch. Similarity between patterns in the abundance of the 17 most common species were examined using correlation matrix-based principal component analysis. Results were compared to the abiotic factors leading to the conclusion that only a low amount of variation in the abundance field can be explained by temperature, salinity and dissolved oxygen. Received: 23 March 1997 / Accepted: 11 April 1997     

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     

Coexistence and reproductive isolation of the sympatric echinoids<Emphasis Type="Italic"> Diadema savignyi</Emphasis> Michelin and<Emphasis Type="Italic"> Diadema setosum</Emphasis> (Leske) on Kenyan coral reefs

The sympatric echinoids Diadema savignyi and D. setosum coexist in shallow reef lagoons throughout East Africa. The reproductive strategies of these echinoids were studied to investigate reproductive isolation as a possible mechanism for maintaining the coexistence of these closely related species. The annual reproductive cycle and lunar periodicity were determined by gonad index measurements, histological examination of gametogenesis, and induction of spawning with injections of KCl. The peak reproductive period of D. savignyi coincided with the north-east monsoon period (when light and temperatures are high) as gonad indices were high (>8%) beginning in February and peaked at 9.7% in May. Gonad indices subsequently rapidly decreased (by 26%) in June at the beginning of the cooler south-east monsoon period. However, the presence of sperm and ova in most months of the year indicates continuous gametogenesis with reduced reproductive effort during the cooler months. The annual cycle for D. setosum showed less of a seasonal trend as gonad indices remained above 7% throughout much of the year but tended to be highest when temperatures were lower. This is the first confirmation of continuous reproduction in these two species at the equator. The reproductive patterns of both species remained consistent over 2 years of sampling. Both species exhibited a synchronized lunar spawning periodicity during the 3 months sampled, with D. setosum spawning on lunar days 8–10 and D. savignyi spawning after the full moon (lunar days 17–18). Whereas spawning in D. savignyi was very tightly synchronized, 20% of D. setosum individuals still spawned after the peak spawning period. The coexistence of these closely related species appears to be maintained by temporal reproductive isolation during the lunar spawning period reinforced by seasonal differences in reproductive effort.Communicated by O. Kinne, Oldendorf/Luhe     

Age and growth of the mesopelagic squid Ancistrocheirus lesueurii (Oegopsida: Ancistrocheiridae) from the central-east Atlantic based on statolith microstructure

Statolith microstructure was studied in 56 Ancistrocheirus lesueurii (25 to 423 mm of mantle length, ML) caught in the central-east Atlantic. Statolith growth increments were grouped into three main growth zones, distinguished mainly by increment width. The second transition in the statolith microstructure (from Zone 2 to Zone 3) coincides with the life history shift from epipelagic and upper mesopelagic to a bathyal habitat. Second-order bands (mean 27.65 growth increments) and sub-bands (mean 13.6 growth increments) within statolith microstructure appeared to be related to the lunar cycle. Striking sexual dimorphism is reflected in the age and growth rates: males live ca. 1 yr, while females only start maturing at this age and obviously live >1.5 yr. A. lesueurii is a slow growing squid, attaining 25 to 30 mm ML at the age of 100 d. After ontogenetic migrations into bathypelagic waters at ML > 30 to 35 mm, growth rates gradually decrease to the minimum known values for squids. Based on back-calculated hatching dates, A. lesueurii hatches throughout the year with a peak between November and March. Received: 28 August 1996 / Accepted: 31 January 1997