Size-dependent reproduction in the Caribbean gorgonian Pseudoplexaura porosa

Reproductive success among clonal taxa is often portrayed as a simple function of clone size, but reproduction in Pseudoplexaura porosa (Houtuyn), a common Caribbean gorgonian, reflects a more complex relationship between size and gamete production. Tagged colonies were sampled at two reefs in the San Blas Islands, Panama during the 1995 spawning season to determine the colony size at first reproduction, size-dependent polyp fecundities, and whole colony gonad production, and to follow the gametogenic cycle. Additional data were obtained during non-spawning months in 1984 and 1985. Of 120 colonies ranging in height from 20 to 250 cm, only colonies taller than 50 cm were reproductive. An average of 4.33 oocytes polyp⁻¹, 560 to 800 μm in diameter, were released during monthly synchronous spawning events from June to September. Oocytes released each month developed from a group of intermediate size oocytes present at the end of the preceding month. Almost all oocytes >560 μm were released during each spawning event. Spermatogenesis was a shorter process than oogenesis. During each spawning month, mature spermaries developed from a group of <125 μm spermaries. Virtually all spermaries >190 μm were released each month, and most of the spermary volume in mature male polyps was generated anew each month of spawning. Among reproductive colonies, polyp fecundity increased with colony height from 50 to 200 cm. Per capita fecundity of >200 cm tall female colonies was lower than for 150 to 200 cm colonies, but whole colony fecundity of large colonies was greater due to the exponential increase in the number of polyps that occurs with increasing colony height. In male colonies, both polyp and colony reproductive output increased with colony height. The large amounts of sperm produced by large male colonies probably contributes to the high rates of in situ fertilization observed in P. porosa. Differences in fecundity as a function of colony size and sex suggest differences in some combination of the cost of reproduction and/or allocation of resources to reproduction. Received: 9 September 1998 / Accepted: 7 June 1999     

Reproductive patterns of scleractinian corals in the northern Red Sea

The majority of published accounts on scleractinian coral reproduction are from the tropical Pacific and Caribbean, with very little information known about Red Sea species. This report examines variation in reproductive mode in 24 species of hermatypic corals (belonging to seven families) in the Gulf of Eilat, Red Sea. Eighteen species are hermaphroditic broadcasters, two are hermaphroditic brooders and three are gonochoric broadcasters. In the Pocilloporidae, the gonads project into the body cavity, while in the other six families the gonads reside inside the mesenteries. The number of gonads per polyp in broadcasting species follows family or genus lines. Fecundity (eggs per polyp) increases with polyp size. Brooding species usually exhibit one or two gonads per polyp and each gonad contains only one to three oocytes. Oocyte size varies widely and does not relate to mode of reproduction. The largest oocytes (diameter = 450 μm) occur in the brooding coral Alveopora daedalea and in broadcasting species of the genus Acropora (diameter = 420 μm). Gonad morphology and gonochorism versus hermaphroditism appear to be constrained phylogenetically at the family or genus level. Lastly, this report compares the data presented for Red Sea scleractinian species with the data available on scleractinian corals from other geographical regions. Received: 2 February 1993 / Accepted: 9 March 1998     

Interannual, seasonal, local and body size variations in reproduction of the prawn Penaeus (Marsupenaeus) japonicus (Crustacea: Decapoda: Penaeidae) in the Ariake Sea and Tachibana Bay, Japan

 The reproductive biology of female Penaeus japonicus Bate was investigated in the Ariake Sea and Tachibana Bay (located outside the Ariake Sea), Japan from 1994 to 1996. Interannual, seasonal, individual female body size and spatial influences on the incidence of spawning were examined. The proportion of inseminated females, increased with increasing body size up to 170 mm body length (BL) and decreased thereafter. The minimum size at maturity was similar between years; 130 to 140 mm BL. The minimum size of individuals with developing ovaries or spermatophores differed from that of ripe females by 20 to 25 mm. Spawning occurred in the central part of the Ariake Sea and Tachibana Bay but rarely in the inner part. P. japonicus had a clear reproductive cycle. Spawning started earlier and ended later, occurring from mid-May to mid-October, in the eastern central part of and outside of the Ariake Sea compared with the western central part, where spawning occurred from mid-June to mid-September. Spawning dynamics differed across seasons, body sizes and areas but not across years. The seasonal peak in the proportion of ripe females varied with body size. In small individuals (130 to 169 mm BL), no peak was observed, whereas in large individuals (>170 mm BL) the proportion of ripe individuals peaked in June. The proportion of ripe individuals increased with increasing body size and was high outside the Ariake Sea. Factors causing the variation in spawning dynamics are discussed. Received: 30 November 1998 / Accepted: 12 October 1999     

Importance of cysts in the population dynamics of the red tide flagellate Heterosigma akashiwo (Raphidophyceae)

To elucidate roles of cysts in occurrences of Heterosigma akashiwo blooms, cyst dynamics were studied in northern Hiroshima Bay, the Seto Inland Sea of Japan, where H. akashiwo regularly forms red tide in June. Monthly measurements of seasonal changes in the densities of vegetative cells of H. akashiwo and their germinable cysts in surface sediments (top 1-cm layer) were made for 2 years at three stations. Vegetative cells of H. akashiwo could be detected from April through December throughout the water column, and the existence of vegetative cells was confirmed in surface waters even in winter after incubation of sampled seawater in culture medium. Germinable cysts, enumerated by the extinction dilution method, existed in sediments in all seasons, even before and after the seasonal bloom. The effects of incubation temperature on the germination of natural cysts of H. akashiwo in sediments were examined. Germination was not observed at 5 °C, was low at 10 °C, while it increased at 15 °C, and maintained a high level to 25 °C. The bottom water temperature reached 15 °C (suitable for the germination of cysts) and the surface about 18 °C or more (suitable for the growth of vegetative cells) 2 to 3 weeks before the blooms. The dark survival of H. akashiwo cysts was tested, and it was found that the cysts were viable for at least 650 d at 11 °C, and for 165 d at 25 °C, indicating a significant role of cysts in the survival during winter and summer seasons. The cysts presumably also play an important role in seeding primary populations into water columns when the bottom water reaches a suitable temperature (around 15 °C); thereafter the populations develop with great annual regularity to bloom in June. These results suggest that initiation of H. akashiwo red tides in the Seto Inland Sea could be triggered by bottom water temperature. Received: 3 July 1998 / Accepted: 12 January 1999     

Sexual reproduction in the coral Pocillopora verrucosa (Cnidaria: Scleractinia) in KwaZulu-Natal, South Africa

The mode of reproduction in Pocillopora verrucosa Ellis and Solander, 1786 varies between geographically isolated regions. This scleractinian coral is common along the KwaZulu-Natal coast, and its reproductive mode and period of reproduction were assessed using histological preparations. The study was undertaken from 1992 to 1994 and showed that P. verrucosa is a simultaneous hermaphrodite and broadcast spawner in KwaZulu-Natal. Gametogenesis occurs from October to January, with the gametes maturing simultaneously so that spawning can be synchronised at new moon in January (mid-summer). Zooxanthellae are present in the mature oocytes. Received: 10 September 1997 / Accepted: 6 July 1998     

Sexuality and cyst formation of the spring-bloom dinoflagellate Scrippsiella hangoei in the coastal northern Baltic Sea

The temporal sequence and the magnitude of the sexual reproduction and subsequent cyst deposition of the common spring-bloom dinoflagellate Scrippsiella hangoei (Schiller) Larsen was studied during spring 1996 on the SW coast of Finland, Baltic Sea. The abundances of the different size of fractions of S. hangoei (14 to 18 μm, 18 to 22 μm and >22 μm) were monitored in the water column, and the deposition of resting cysts was measured using moored sediment traps. Cyst sedimentation rates were measured throughout the seasonal cycle in order to estimate cyst resuspension rates for the quantitative assessment of the fraction of population undergoing encystment. The onset of sexual reproduction, indicated by a significant increase of the small cells (14 to 18 μm) representing gametes, occurred in a nutrient replete environment well before the exponential growth phase and peak abundances of vegetative cells. Gamete formation was followed by high abundances of large cells (>22 μm) representing planozygotes, and subsequent sedimentation of resting cysts. Approximately 60% of the asexually growing bloom population was estimated to form planozygotes, suggesting that encystment was an important factor in bloom termination and possibly plays a role in the regulation of the magnitude of the bloom. Finally encystment accounted for 40% of the entire S. hangoei population, resulting in a considerable loss of the bloom population and an input of the vernal phytoplankton biomass to the benthos. Received: 11 December 1998 / Accepted: 8 April 1999     

Use of the fluorochrome calcein as an in situ growth marker in the brown mussel Perna perna

This study assesses the potential of the fluorochrome calcein for use as a growth marker in bivalve shell growth studies. Calcein solutions were administered in situ to the brown mussel Perna perna (Linneaus), both by injection and immersion, and the effect of calcein concentrations on fluorescent mark deposition and mussel mortality was investigated in the laboratory. Field investigations showed that, 1 month after administration, calcein injection (125 mg l^−l) into the mantle cavity produced superior results to the immersion treatments (150 and 500 mg l^−l). Both methods resulted in fluorescent mark incorporation at the growing edge, but during immersion general calcein deposition associated with endolith activity resulted in fluorescence that made identification of a distinct datum point difficult. In contrast, the injection method produced clearly defined growth marks, which were easily distinguished from autofluorescence and persisted without visible degradation for a minimum of 9 months. Shell growth rates estimated using the fluorescent mark as a datum point were similar to those from earlier studies using different methods. Laboratory investigations revealed that at␣calcein concentrations of 80 mg l^−l and above, 100% of juvenile (20 to 30 mm) and adult mussels (60 to 70 mm) retained a visible growth mark, while at concentrations >160 mg l^−l all marks were bright and clearly defined. No mussel mortality was exhibited at any time, even at calcein concentrations of 640 mg l^−l, eight times higher than those required for mark deposition. These results suggest that, compared to traditional methods of bivalve growth determination, the use of fluorochromes presents a relatively inexpensive, non-invasive and rapid alternative. When using calcein as a growth marker, problems associated with some other fluorochromes (e.g. inconsistent mark incorporation, high post-treatment mortality) were not exhibited. Received: 14 July 1998 / Accepted: 23 October 1998     

Sexual reproduction of the scyphomedusa Aurelia aurita in relation to temperature and variable food supply

The effects of food availability and temperature on sexual maturation and female reproductive output of the scyphomedusa Aurelia aurita was examined in two populations from the contrasting environments of Southampton Water and Horsea Lake, England. Trends in oogenesis and subsequent reproductive output differed markedly between the two populations. In Southampton Water, the onset of sexual maturation occurred earliest in the larger medusae, but eventually all females became ripe, the smallest being 45 mm bell diameter (BD). The decrease in minimum size at maturity was correlated with increasing temperature. In A. aurita from Horsea Lake, size at maturity varied on a seasonal basis, with the smallest ripe female being only 19 to 20 mm BD. There were spring and autumn periods of sexual maturation in this population. During the autumn period, it is likely that food limitation was playing a more critical role in determining medusa size, with decreasing temperature indirectly affecting A. aurita by limiting primary and secondary production. In similar-sized ripe medusae, fecundity was greater in Southampton Water, but the planula larvae produced were significantly smaller than those in Horsea Lake. It is suggested that in Horsea Lake, the quality of the larvae are greater in terms of biochemical content to ensure survival of the few gametes produced (i.e. K-strategy). Comparison of the reproductive effort of the two A. aurita populations revealed that medusae from Southampton Water, which experience greater food availability, are able to direct more energy to reproduction than Horsea Lake medusae. In the latter, A. aurita medusae appear to partition the available food resources into either somatic growth (and therefore increased future fecundity) when food is abundant, or reproductive growth when food is scarce. Received: 24 June 1997 / Accepted: 23 March 1998     

Early life history traits of the gold-eye rockfish, Sebastes thompsoni, in relation to successful utilization of drifting seaweed

The gold-eye rockfish Sebastes thompsoni lives in association with drifting seaweed during the early developmental period. Early life history traits of the gold-eye rockfish were studied by means of otolith analysis of larvae and juveniles associated and unassociated with drifting seaweed in the waters off southern Sanriku, Tohoku, northern Japan, from 1993 to 1995.␣Daily growth increments and the existence of an extrusion check were confirmed by both a series of tetracycline marking experiments and an analysis of the relation between the number of increments and the number of days after extrusion. Larvae and juveniles associated with drifting seaweed occurred from late May to late July, with a peak in late June, showing the same seasonal fluctuation in abundance as drifting seaweed. Larvae seemed to associate with drifting seaweed after aggregation to surface slick areas. Body size at the commencement and the end of the seaweed-associated period was estimated at 18 and 50 mm total length, respectively. Growth rates were low until the beginning of the association with seaweed; thereafter they became high but declined towards the end of seaweed-associated life. Larvae and juveniles born later grew faster than those born earlier. This tendency was detected both before and during the association with drifting seaweed. Although the season of abundant drifting seaweed was about 2␣months shorter, and occurred 2 months later, than the extrusion season of the gold-eye rockfish from early February to early June, this seemed to be compensated by the growth variation dependent on birth month. Early life history traits of the gold-eye rockfish are discussed in relation to the successful utilization of drifting seaweed. Received: 3 August 1997 / Accepted: 24 March 1998     

Reproductive patterns in the Antarctic brachiopod Liothyrella uva

The reproductive cycle of the Antarctic articulate brachiopod Liothyrella uva (Broderip, 1833) is described from monthly samples collected between September 1985 and July 1987 from a population at Signy Island, Antarctica. Spermatogenesis and oogenesis are described for the first time in this species. Surface dried-tissue masses for a standard individual (41 mm shell length) were calculated for the digestive diverticula, gut, lophophore and gonad from monthly dissections of 15 brachiopods. Seasonal patterns, with summer peaks, were observed in the digestive diverticula and gut. The lophophore and gonad masses did not exhibit seasonal trends. Females showed a sharp decrease in proportion of large oocytes between October and November in 1986, suggesting spawning during this period. There was no similar decrease during the same period in 1985. Mean percent spermatozoa measurements revealed a large increase in November 1985 and a rapid decline in December 1985. This suggested a large spawning event for males in 1985 which was not repeated the following year. These data indicate large inter-annual differences in reproductive activity as well as differences between males and females. They also suggest the possibility of sperm storage by females. Brood characteristics were also highly variable. The smallest brooding female was 31.5 mm in length. Some females brooded more than one developmental stage simultaneously, and variation in brood size (numbers of embryos or larvae held in the lophophore) and brood composition between individuals was high. All samples collected throughout the 2 yr period contained some females with broods. The data suggest that the reproductive strategy of L. uva is highly plastic, and that there may be three reproductive periodicities on seasonal, annual and inter-annual time scales. Received: 16 May 1997 / Accepted: 2 March 1998     

Metallothionein-like metalloproteins in the Baltic clam Macoma balthica : seasonal variations and induction upon metal exposure

Metallothionein-like proteins (MTLs) have been measured by differential pulse polarography every two months during a period of two and half years in the marine bivalve Macoma balthica from two locations of the Western Scheldt Estuary. The MTL concentrations (0.85 to 7.81 mg g⁻¹ dry wt) are comparable to most values found for other marine invertebrates. These concentrations vary significantly with the seasons (higher in winter, lower in summer), which is mainly due to body weight fluctuations. Strong correlations exist between MTL and metal levels. Upon short-term exposure to a mixture of Cd, Cu and Zn, there are on average significant MTL increases, but important seasonal variations have been observed: in winter, the clams are more sensitive to metals, uptake more Cd and Cu and induce more MTL than during the warmer months. Received: 16 May 1997 / Accepted: 6 June 1997     

Effects of food type and ration on reproductive maturation and growth of the sea urchin Strongylocentrotus droebachiensis

We investigated the effects of food quality and quantity on reproductive maturation and growth of juveniles of Strongylocentrotus droebachiensis (Müller) in a 22 month laboratory experiment in which we fed sea urchins four diets: (1) kelp (Laminaria spp.) for 6 d wk⁻¹ and mussel (Mytilus spp.) flesh for 1 d wk⁻¹ (KM); (2) kelp for 7 d wk⁻¹ (high ration, KH); (3) kelp for 1 d wk⁻¹ (low ration, KL) and (4) no food other than encrusting coralline algae (NF). At their first and second opportunity for reproduction, all sea urchins in the KM and KH treatments, and most in the KL treatment were reproductively mature, whereas all sea urchins in the NF treatment remained immature. Gonad index differed significantly among all fed treatments at first and second reproduction, and was highest in the KM and lowest in the KL treatment. Gonad index was similar in both sexes at first reproduction, but it was higher in females than in males at second reproduction. Diet had little or no effect on the relative abundance of spermatocytes, spermatozoa, or nutritive phagocytes in testes at first and second reproduction. In ovaries, nutritive phagocytes were significantly more abundant in females in the KM and KH treatments than in the KL treatment at first reproduction, and significantly more abundant in unfed (NF) than fed (KM, KH, KL) females at second reproduction. Mean oocyte size was similar in all fed females at first reproduction, but significantly larger in fed than unfed females at second reproduction. Mean ovum size was similar in all fed females in both reproductive periods. Increase in test diameter was greatest in the KM treatment and smallest in the KL treatment; sea urchins in the NF treatment decreased slightly in size. Survival was 95 to 100% in all fed sea urchins but significantly lower in unfed ones. The feeding rate on kelp was significantly greater in the KL than the KM and KH treatments. In the KM treatment, the feeding rate on kelp increased significantly over a 6 d period after mussel flesh was provided. Our results demonstrate that a diet of high food quality and quantity accelerates reproductive maturation and growth rate, and enhances gonad production and survival in juvenile and young adult S. droebachiensis. These findings contribute to our understanding of the reproductive ecology of S. droebachiensis in habitats with differing food supplies (e.g., kelp beds and barrens). Our results also can be used to improve aquacultural practices for sea urchins. Received: 3 June 1998 / Accepted: 2 February 1999     

Life cycle of the copepod Calanus hyperboreus in the Greenland Sea

The seasonal ontogenetic migration of the Arctic copepod Calanus hyperboreus was described from surface-to-bottom hauls in the central Greenland Sea Gyre (GSG) and in the Westspitsbergen Current (WSC). All stages except females spent the winter below 500 m in the GSG and below 1000 m in the WSC. Seasonal ascent begins in April, and descent in July. For the C.␣hyperboreus population an active downward transport of 8.1 g m⁻² dry weight during 8 months of overwintering was estimated, similar to flux rates of particulate matter in sediment traps. Seasonal distribution of biomass was determined from weight measurements of single stages. Annual means varied from 4.0 to 9.2 g m⁻² in two different years in the GSG and were 1.1 in 1 year in the WSC. The life cycle in the Greenland Sea was reconstructed from field data on stage composition, vertical distribution, reproduction, and moult cycle phase from tooth development of CV. Laboratory experiments were conducted on the moulting of CIV and CV in fall. A 3-year (males) and 3- to 4-year (females) life cycle is proposed for the GSG and 2 to 3 years for the WSC. However, the small number of young larvae and the incomplete spring ascent by older copepodites observed in the WSC cast doubt on the reproductive success in the WSC. A suite of physiological strategies and adaptations performed by the developmental stages support survival of this species in harsh environments. Received: 25 January 1997 / Accepted: 11 February 1997     

Bathymetric patterns of genetic variation in a deep-sea protobranch bivalve, Deminucula atacellana

The origin of the deep-sea benthic fauna is poorly understood and represents an enormous gap in our understanding of basic evolutionary phenomena. One obstacle to studying evolutionary patterns in the deep sea has been the technical difficulty of measuring genetic variation in species that are typically minute, rare, and must be recovered from extreme depths. We used molecular genetic techniques to quantify variation in the 16S rRNA mitochondrial gene within and among populations of the common protobranch bivalve Deminucula atacellana (Schenck, 1939). We analyzed 89 individuals from nine samples collected in the 1960s along a depth gradient from 1100 to 3800 m in the western North Atlantic. Genetic variability within populations is much lower than between populations, and peak haplotype numbers occur near the center of its depth distribution. Continental slope (<2500 m) and rise (>2500 m) populations were genetically distinct despite the lack of any obvious topographic or oceanographic features that would impede gene flow. These findings indicate that the deep-sea macrofauna can have strong population structure over small (134 km) spatial scales, similar to that observed in shallow-water and terrestrial organisms. This surprisingly high biodiversity at the genetic level affords the potential for adaptation and evolutionary diversification, the ultimate historical causes of high species diversity in the deep-sea benthos. Received: 24 July 1997 / Accepted: 26 January 1998     

Reef coral reproduction in the Abrolhos Reef Complex, Brazil: the endemic genus Mussismilia

The current contribution deals with the reproductive biology of a genus endemic to Brazil –Mussismilia Ortmann, 1890 – including all three species of the genus: M. braziliensis (Verrill, 1868), M. hartti (Verrill, 1868), and M. hispida (Verrill, 1902), which occur sympatrically in the studied area, the Abrolhos Reef Complex, Brazil. Sexuality patterns, modes of reproduction, synchrony and spawning periods are reported, and were determined by histological examination of material. All three species started to develop female and male gametes over different periods in the same breeding season. The three species are probably broadcast spawners, since no embryos or planulae were observed in any species at any given time of the year. Each reproductive cycle lasted approximately 11 months. Oogenesis and spermatogenesis started in different periods, with spermaries appearing in approximately the eighth month of ovary development and lasting about 3 months. Reproductive cycles were annual. Spawning probably occurred in consecutive months in each species. In M. braziliensis, spawning presumably happened between March and the middle of May in 1996 and 1997. Evidence suggested that spawning events of M. hispida took place between the end of April and mid-June. M. hartti may have spawned between September and November. The data presented here suggested that all studied species have at least one exclusive spawning period, asynchronically with the others. A possible exception may be the simultaneous (or close) spawnings of M. braziliensis and M. hispida in May. It is suggested that asynchrony in spawning periods among species may reduce the chance of hybridization, gamete waste and the competition for settlement surfaces. The occurrence of extended spawning periods for each species may also reduce the risks of reproductive failure, due to temporary adverse conditions. Received: 8 December 1998 / Accepted: 15 July 1999     

Population biology of Dyopedos monacanthus (Crustacea: Amphipoda) on estuarine soft-bottoms: importance of extended parental care and pelagic movements

The suspension-feeding amphipod Dyopedos monacanthus (Metzger, 1875) is a common epibenthic amphipod that lives on self-constructed “mud whips” (built from filamentous algae, detritus and sediment particles) in estuaries of the northern North Atlantic Ocean. The population biology of D. monacanthus at a shallow subtidal site in the Damariscotta River Estuary (Maine, USA) was examined between July 1995 and July 1997. The resident population at the study site was dominated by adult females during most months of the year. High percentages of subadults were found in late summer/early fall. Often, between 10 and 20% of the adult females were paired with males, and the percentage of ovigerous females varied between 40 and 100%, indicating continuous reproduction. The percentage of parental females varied between 40 and 80% during most months, but dropped to levels below 20% during summer/early fall. The average size of amphipods on their own mud whips was ∼4 mm during the summer/early fall, after which it increased continuously to >7.0 mm in March or April, and then dropped again. In March and April, the average number of eggs and juveniles female⁻¹ was ∼100 eggs and 55 juveniles, while during the summer/early fall the average number of eggs female⁻¹ was <20 and that of juveniles female⁻¹ was <10. Many juveniles grew to large sizes (>1.4 mm) on their mothers' whips in winter/early spring but not in the summer/fall. The average number of amphipods at the study site was low in late summer/early fall (<50 individuals m⁻²), increased steadily during the winter, and reached peak densities of >3000 individuals m⁻² in April 1996 (>1600 individuals m⁻² in May 1997), after which densities decreased again. The decrease of the D.␣monacanthus population at the study site coincided with a strong increase of amphipods found pelagic in the water column. This behavioural shift occurred when temperatures increased and benthic predators became more abundant and active on shallow soft-bottoms, suggesting that D. monacanthus at the study site is strongly affected by predation. The effects are direct (by predation on amphipods) and indirect (by reducing duration of extended parental care and enhancing pelagic movements). Both extended parental care and pelagic movements are important behavioural traits of D.␣monacanthus (and other marine amphipods), and significantly affect its population dynamics. Received: 18 January 1998 / Accepted: 27 May 1998     

Production and biomass accumulation of periphytic diatoms growing on glass slides during a 1-year cycle in a subtropical estuarine environment (Bay of Paranaguá, southern Brazil)

Production rates, chlorophyll concentrations and general composition of periphytic diatom communities growing on glass slides were studied in relation to environmental parameters during one seasonal cycle in the Bay of Paranaguá, southern Brazil. Slides were routinely submersed at 1, 2 and 3 m depth and recovered weekly for microscopic examinations, analyses of chlorophyll, cell counts and in situ photosynthetic incubations using the Winkler titration method. Water samples were also collected at surface and bottom layers for determinations of temperature, salinity, nutrients and chlorophyll in the water. The periphytic community was mainly formed by epipelic and epipsammic species, dominated by Navicula phyllepta, Cylindrotheca closterium, Navicula spp. and Amphora sp. Weekly chlorophyll a and cell accumulations on slides varied from <1–32 mg m⁻² and up to 31 × 10⁸cells m⁻², respectively. Photosynthetic rates varied from <1 to 35 mg oxygen mg chlorophyll a ⁻¹ h⁻¹, with higher values in summer. Daily production varied from 5 to 3,600 mg oxygen m⁻² day⁻¹ (<0.01–1.4 g carbon m⁻² day⁻¹). Multiple regression analysis revealed that vertical differences in light conditions and grazing pressure jointly affected the influence of temperature on the seasonal patterns of cell densities and chlorophyll concentrations according to depth. Received: 27 April 2000 / Accepted: 16 August 2000     

Reproductive biology of a deep-sea brittle star Amphiura carchara (Echinodermata: Ophiuroidea)

 New information on the brooding reproduction of Amphiura carchara, and previous studies of several deep-sea congeners, contradict the prevailing notion that direct development is exceptional among deep-sea echinoderms. Over 500 specimens of A. carchara from 2,850 to 4,100-m depths off the coast of California, USA, were studied. The species was found to be gonochoric, although most brooding species of ophiuroids are hermaphroditic. Females each brooded up to 72 embryos at a time, with up to 10 in a single bursa. The embryos of individual adults were often at different stages of development, but those brooded in a single bursa tended to be at the same stage. Thus, the species has characteristics that breach the distinctions between sequential and simultaneous brooding exhibited by shallow-water ophiuroids. The embryos developed from yolky eggs that are large for an ophiuroid, with the mean largest oocytes 0.45 mm, and ranging to 1.28 mm in diameter. Almost all late-stage embryos were positioned with their mouth and arms pressed against the wall of the bursa, possibly to facilitate the uptake of nutrients from the parent. Thus, A. carchara may be matrotrophic. The largest embryos examined had a rudimentary disk skeleton, and arms with four joints and a terminal plate. Emerging juveniles probably differ in disk diameter. The species appears to brood year round, although differences in gonad size, the incidence of brooding, and the relative numbers of early developmental stages in summer and winter samples indicated that there are seasonal trends in reproduction. Received: 14 February 2000 / Accepted: 3 July 2000     

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     

Male genital traits and mating interval affect male fertilization success in the water strider Gerris lacustris

Parker's seminal work brought attention to the possibility of postmating sexual selection by non-random fertilization success. Mechanisms for these processes are still only partly understood and there is clearly a need for more studies of intraspecific variation in sperm precedence. Here, we report results from an experimental study of the variation in fertilization success between males of the water strider Gerris lacustris. Genital morphology, male body size, and copulation duration were examined as possible correlates of paternity. The significance of guarding duration was also analysed. Only male genital morphology was correlated to fertilization success. This is one of the first studies showing a relationship between male genital traits and fertilization success, supporting the view that sexual selection may be responsible for the rapid and divergent evolution of genital structures in animals with internal fertilization. The fertilization success of last males varied considerably after double matings with a short mating interval (10 min). Last-male priority ranged from 0 to 100% and usually one of the males involved fertilized almost all the eggs. After double matings with a short mating interval, the proportion of eggs fertilized by the last male averaged 0.68 and was greater than 0.5. In contrast, the average fertilization success was biased towards the first male when the matings were more spread out over time (24 h). These results do not support earlier suggestions of a widespread last-male sperm priority in water striders. Received: 28 July 1998 / Received in revised form: 15 March 1999 / Accepted: 28 March 1999