Depth-related shift in life history strategies of a brooding and broadcasting deep-sea asteroid

Combining field and laboratory work, this study investigated the reproductive cycle, aggregative behavior, spawning periodicity, development and early growth of the sea star Henricia lisa living at bathyal depths off eastern Canada. Marked differences were found between individuals from ~1,300 and ~600 m deep. The former had a male biased sex ratio and an aperiodic reproductive cycle, whereas the latter displayed an equal sex ratio and a biannual breeding pattern. Furthermore, the maximum size was larger and female fecundity roughly five times higher in shallower compared to deeper populations. In the tanks, aggregative behavior was recorded twice a year during the summer and winter breeding periods. The onset of aggregations and spawning coincided with a temperature of 3–4°C. Males spawned first and females typically responded inside 30–60 min. Between 12 and 20 eggs were retained to be brooded under the arched arms of the female, whereas the remainder were broadcasted and developed without parental care. The fertilized eggs underwent a first cleavage after 12 h, reached the brachiolaria stage in 1 month, became juveniles within 3–4 months and reached ~ 4 mm in diameter after 14–17 months of growth. The embryos and juveniles developed at the same rate whether brooded or not, and development of winter cohorts was typically slower due to lower prevailing temperatures. This study of H. lisa provides the first evidence of lecithotrophy in a seasonally breeding deep-sea echinoderm and of brooding in a deep-sea asteroid.     

Depth-related shift in life history strategies of a brooding and broadcasting deep-sea asteroid

Combining field and laboratory work, this study investigated the reproductive cycle, aggregative behavior, spawning periodicity, development and early growth of the sea star Henricia lisa living at bathyal depths off eastern Canada. Marked differences were found between individuals from ~1,300 and ~600 m deep. The former had a male biased sex ratio and an aperiodic reproductive cycle, whereas the latter displayed an equal sex ratio and a biannual breeding pattern. Furthermore, the maximum size was larger and female fecundity roughly five times higher in shallower compared to deeper populations. In the tanks, aggregative behavior was recorded twice a year during the summer and winter breeding periods. The onset of aggregations and spawning coincided with a temperature of 3–4°C. Males spawned first and females typically responded inside 30–60 min. Between 12 and 20 eggs were retained to be brooded under the arched arms of the female, whereas the remainder were broadcasted and developed without parental care. The fertilized eggs underwent a first cleavage after 12 h, reached the brachiolaria stage in 1 month, became juveniles within 3–4 months and reached ~ 4 mm in diameter after 14–17 months of growth. The embryos and juveniles developed at the same rate whether brooded or not, and development of winter cohorts was typically slower due to lower prevailing temperatures. This study of H. lisa provides the first evidence of lecithotrophy in a seasonally breeding deep-sea echinoderm and of brooding in a deep-sea asteroid.     

Growth and reproduction in the Antarctic brooding bivalve Adacnarca nitens (Philobryidae) from the Ross Sea

We present information on the reproductive biology, population structure, and growth of the brooding Antarctic bivalve Adacnarca nitens Pelseneer 1903, from the Ross Sea, Antarctica. Individuals ranging from 0.85 to 6.00 mm were found attached to a hydrozoan colony. This species shows low fecundity and large egg size, common to other brooding species. The minimum size at which oogenesis was detected was 2.3 mm and the minimum size at which brooding was evident was 3.9 mm. Embryos of a full range of developmental stages were brooded simultaneously in females. The population showed a log–normal distribution and results suggest non-periodic reproduction with continuous embryonic development. The reproductive traits of A. nitens are discussed in the context of circum-Antarctic species distribution and limitations to dispersal in brooding benthic invertebrates.     

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     

Reproduction in Anthelia glauca (Octocorallia: Xeniidae). I. Gametogenesis and larval brooding

Anthelia glauca Lamarck, 1816 is a gonochoric, external-brooding soft coral found in KwaZulu-Natal. It is reproductively active in the summer months. The development of gametes produced in late summer is arrested in winter. Several stages of gametes are found at the base of the polyps, and female polyps produce several cycles of larvae over an extended breeding period of 4 to 5 months. Larvae are brooded in a unique pharyngeal brooding pouch not yet described in other coral species. The brood pouch consists of an expansion of the pharynx with constrictions proximal and distal to the embryos and larvae. Our data suggest that egg transfer and fertilization occur at full moon and the mature larvae are released after new moon. Zooxanthellae are absent in A. glauca oocytes, but zooxanthella infestation commences at the immature larval stage. Received: 15 July 1997 / Accepted: 12 March 1998     

Evidence of amictic reproduction in the brittle-star Ophiomyxa brevirima

Ophiomyxa brevirima is a direct-developing species of brittle-star found around the coast of New Zealand. The mode of reproduction of this ophiuroid was studied using isozyme electrophoresis of mother and brooded offspring. Multi-locus assessment of the genotypes of 35 adult females and their brooded offspring was undertaken. A minimum of 45% of broods assessed showed evidence of amictic reproduction, with all offspring in these broods possessing genotypes that were identical to the genotype of their heterozygous mother. In most other broods there was evidence of segregation. These observations are interpreted as evidence of both sexual and asexual reproduction in this ophiuroid species. Received: 19 November 1996 / Accepted: 28 January 1997     

Size–frequency dynamics of NE Pacific abyssal ophiuroids (Echinodermata: Ophiuroidea)

The 17-year time-series study at Station M in the NE Pacific has provided one of the longest datasets on deep-sea ophiuroids to date. Station M is an abyssal site characterized by low topographical relief and seasonal and interannual variation in surface-derived food inputs. From 1989 to 2005, over 31,000 ophiuroid specimens were collected. Size–frequency distributions of the four dominant species, Ophiura bathybia, Amphilepis patens, Amphiura carchara and Ophiacantha cosmica, were examined for recruitment and the role of surface-derived food supplies on body size distributions. Juveniles were collected in sediment traps and used to investigate settlement patterns and seasonality. Trawl samples showed no indication of seasonal changes in recruitment to larger size classes; however, there was evidence of seasonal settling of juveniles. Interannual differences in median disk diameters and size distributions of trawl-collected adults are greater than those at the seasonal scale. Three of the four species, O. bathybia, A. patens and O. cosmica, had co-varying monthly median disk diameters, suggesting they may have a similar factor(s) controlling their growth and abundance. Interannual differences in monthly size distributions were generally greater than those between seasons. Cross-correlations between the particulate organic carbon (POC) flux (food supply) and size distribution indices for O. bathybia, A. patens and O. cosmica all were significant indicating that increases in food supply were followed by increases in the proportion of smaller size classes after approximately 17–22 months. These findings suggest that food inputs are indeed an important factor influencing deep-sea ophiuroid populations on interannual time scales, more generally supporting the long-hypothesized connection between food availability and population size structure in the deep sea. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The genetic mating system of a sex-role-reversed pipefish (Syngnathus typhle): a molecular inquiry

In the pipefish Syngnathus typhle as in other species of Syngnathidae, developing embryos are reared on the male's ventral surface. Although much laboratory research has been directed toward understanding sexual selection in this sex-role-reversed species, few studies have addressed the mating behavior of S. typhle in the wild, and none has capitalized upon the power of molecular genetic assays. Here we present the first direct assessment of the genetic mating system of S. typhle in nature. Novel microsatellite loci were cloned and characterized from this species, and employed to assay entire broods from 30 pregnant, field-captured males. Genetic analysis of 1340 embryos revealed that 1–6 females (mean = 3.1) contributed to each brooded clutch, the highest rate of multiple maternity yet documented in any pipefish. Evidence of multiple mating by females was also detected. Thus, this population of S. typhle displays a polygynandrous mating system, a finding consistent with previous field and laboratory observations. Our results, considered together with similar studies of other syngnathid species, provide preliminary support for the hypothesis that the genetic mating system is related to the evolution of sexual dimorphism in the fish family Syngnathidae. Received: 19 January 1999 / Received in revised form: 15 April 1999 / Accepted: 9 May 1999     

Costs of extended brood protection in the Antarctic sea star, Neosmilaster georgianus (Echinodermata: Asteroidea)

Changes in size, morphology, and biochemical composition in adults and embryos of a brooding sea star, Neosmilaster georgianus (Sladen), were studied in a population adjacent to Anvers Island, Antarctic Peninsula (64°46′S; 64°04′W) during the austral spring, 1991. Five morphological stages of development were designated in 24 broods, and for each the weight and biochemical composition of the brooding adults and their embryos were determined. Between Stage 1 and 2, the dry weight (dw) and organic weight (ow) of the embryo did not change. From Stage 2 to 3, the dw and ow increased significantly by 10%. Stage 2 and 3 embryos were in clusters of a few (2 to 10) to as many as 40 individuals. In the smaller clusters, individual embryos were attached by tissue cords to another, sometimes atrophied, brood member. In the larger clusters, they were attached to a central mass of tissue containing remnants of embryos. We interpret these interactions as a form of cannibalism which may account for the weight gains between Stage 2 and 3. During Stages 4 and 5, as juvenile form was approached, the dw and ash weight of the young increased significantly and the ow decreased significantly. The calculated energy content for the juvenile (Stage 5) was not significantly different from the energy content of the earliest undifferentiated stage (Stage 1), an indication that most of the organic matter in the egg is the primary contribution to the large juvenile. In brooding females, pyloric caeca indices declined by 52% from Stage 1 to Stage 5 and pyloric energy stores declined by 63% due to proportionately equivalent declines in protein and lipid. The ovary index was low and increased only slightly during brood protection, while the size of the largest oocytes remained approximately 23% that of ova. Energy stores in the pyloric caeca of brooding N. georgianus thus become depleted over a long period of incubation during which the adults apparently starve. This may delay oocyte development and ultimately limit the frequency of individual reproduction. Received: 15 September 1998 / Accepted: 9 March 1999     

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     

Gill morphogenesis in the oyster Ostrea chilensis

Despite the importance of the gills in the acquisition of food by suspension-feeding bivalve mollusks, there is almost no information on gill organogenesis. By means of a series of stereoscan electron micrographs, this paper describes gill development in the Chilean oyster, Ostrea chilensis, from the brooded larval stages to 1-month-old spat. A single gill rudiment was observed on each side of the mantle at a shell length of 320 μm, and the rudiments increased in number and size until the end of the brooding period. During metamorphosis the gill filaments increased in number from 5 or 6 to between 7 and 9. The loss of the velum and the absence of functional gill filaments during metamorphosis are consistent with previous observations of weight loss during this critical period of the life history, because the newly settled juvenile lacks the ability to remove particles from suspension. The end of metamorphosis (100% of spat with dissoconch edge) was reached 36 h after larval settlement, when the gill filaments began to grow cilia, which increased in density and differentiated as the spat developed and acquired the capability of suspension-feeding, accounting for the increase in body weight previously recorded during this stage. The larval rudiments gave rise to the inner demibranchs. The outer demibranchs were observed 10 days after settlement, located between the inner demibranch and the mantle. In 1-month-old spat, the gill did not show differentiation between primary and secondary filaments, indicating that the heterorhabdic condition characteristic of adult oysters had yet to be attained. Received: 11 December 1998 / Accepted: 21 August 2000     

Mycosporine-like amino acids prevent UVB-induced abnormalities during early development of the green sea urchin Strongylocentrotus droebachiensis

Experiments were performed to determine how ultraviolet radiation (UVR) in the environmentally relevant range affects development of the sea urchin Strongylocentrotus droebachiensis (Müller) and whether mycosporine-like amino acids (MAAs), present in the early life stages, reduce UV-induced damage. Eggs, embryos, and larvae contained five MAAs having absorption maxima ranging from 320 to 334 nm. Eggs contained principally shinorine and porphyra-334, which absorb maximally at 334 nm and half-maximally at 312 and 348 nm, spanning much of the environmental range of biologically effective UVR. Concentrations of MAAs remained constant in unirradiated embryos through the gastrula stage, but decreased significantly in two-armed pluteus larvae. Daily exposure to combined photosynthetically active radiation (PAR, 400–700 nm) and UVR did not affect the concentration of MAAs in these embryos up to the two-armed pluteus stage. Prism larvae of sea urchins and the sand dollar Echinarachnius parma (Lamarck) did not accumulate shinorine from the surrounding seawater. Daily exposure of embryos to UVA (320–400 nm) and UVB (295–320 nm) radiation in the presence of PAR induced delays and abnormalities during development, and removing UVB eliminated this effect. Abnormalities in embryos included thickening of the blastoderm wall, filling of the blastocoel by abnormal cells, exogastrulation, and formation of abnormal spicules. The percentage of embryos that developed normally was lower in batches of embryos exposed to PAR + UVA + UVB, except in embryos from urchins maintained on MAA-rich diets. In all cases, the percentage of PAR + UVA + UVB-exposed embryos that developed normally was positively related to the concentration of MAAs in eggs from which the embryos developed. Thus, the MAAs found in S. droebachiensis embryos protect them against UVB-induced abnormalities during their development to at least the four-armed pluteus larval stage. Received: 8 May 2000 / Accepted: 29 September 2000     

Population structure, growth and recruitment of the euryalinid brittle-star Astrobrachion constrictum (Echinodermata: Ophiuroidea) in Doubtful Sound, Fiordland, New Zealand

The euryalinid brittle-star (snake star) Astrobrachion constrictum (Farquhar) lives coiled around the branches of black coral (Antipathes fiordensis) colonies. Twenty-two vertical transects, 10 m wide by 30 m deep, were swum in Doubtful Sound over a 2.5 yr period from 1993 to 1995. Numbers, disc diameters and colour morphotypes of brittle-stars inhabiting coral colonies were recorded. 36.3% of the coral colonies >200 mm tall (n = 292) hosted ≥1 Astrobrachion constrictum (range 0 to 12). Overall, the population was patchily distributed on the available coral habitat. The dark red colour morphotype of A. constrictum was most common (87%, n = 279) followed by the yellow, striped, and then spotted varieties. The population was comprised mainly of large (≥10 mm disc diam) individuals, and juveniles were rarely encountered, indicating low rates of recruitment or a high mortality of recruits. Disc-diameter data gathered from this and previous studies indicated that growth in A. constrictum is initially rapid, with individuals reaching a disc diameter of 15 mm in ≃2.5 yr; growth decreases with age, as in other deep-sea ophiuroids. Growth rate within years, however, was not constant, with faster growth in the spring/summer. Maximum size for A. constrictum is reached in ≃8 yr at ˜23 mm disc diam. Anecdotal evidence indicates that A. constrictum may not be confined solely to black coral colonies. Received: 25 September 1996 / Accepted: 16 October 1996     

Biology of the deep-sea octopus Bathypolypus sponsalis (Cephalopoda: Octopodidae) from the western Mediterranean Sea

Several aspects of the biology of Bathypolypus sponsalis were studied from 297 individuals (115 males, 180 females and 2 indeterminates) caught in a depth range of 200–800 m depth in the western Mediterranean Sea. The paper presents data on sizes (length-weight relationships, size-frequency distributions) and reproduction (sex ratio, maturation, condition), and also analyses of the diet of B. sponsalis from samples taken throughout the year. Length-weight relationships showed that females are heavier than males at the same mantle length. Although mature individuals were found all year round, the maximum number occurred in spring and summer. Sexual maturation data revealed that males mature at smaller sizes than females. The gonadosomatic index increased with maturity in both sexes; the increase was gradual in males, but abrupt in females. The digestive gland index was used as a condition index and showed a differential behaviour with maturity; it increased gradually in females, but decreased in males. Like other octopus species, B. sponsalis appears to be an opportunistic predator, feeding on a great variety of preys. Stomach content analysis yielded a total of 19 different prey items belonging to four major groups (Crustacea, Mollusca, Ophiuroidea and Osteichthya). The first three groups were the more frequent preys, since crustaceans, molluscs and ophiuroids appeared in 76%, 49% and 30% of the stomachs, respectively. Decapoda Reptantia (among crustaceans) and cephalopods and bivalves (among molluscs) constituted the more abundant prey items. While the Decapoda Reptantia group was significantly more abundant in stomach contents of females, gastropods were taken more frequently by males. These differences in diet could reveal females as a more active predators than males. Received: 5 March 2000 / Accepted: 7 November 2000     

Life on glass houses: sponge stalk communities in the deep sea

Photographs of the deep-sea floor often show organisms attached to biogenic structures that protrude from the soft bottom. In particular, the stalks of glass sponges (hexactinellids) provide hard substrata and act as habitat islands for deep-sea fauna. The primary objectives of this study were to determine the abundance of glass sponge “stalks” at an abyssal station in the NE Pacific, to identify the fauna associated with stalks, and to compare the distribution patterns of epifaunal taxa both horizontally and vertically. Densities of stalks and large epifauna were estimated from analysis of ∼9 km of photographic transects taken in 1994–1995 at station M (34°45′N; 123°00′W; 4,100 m depth) off California, USA. At least 87% of the stalks were the spicule columns of live or dead hexactinellids in the genus Hyalonema (Gray, 1832). Stalks appeared to be distributed randomly across the sea floor (density: 0.13 stalks m⁻²). A colonial zoanthid, Epizoanthus stellaris (Hertwig, 1888), inhabited 20% of the stalks and was the most commonly observed epifaunal organism, followed by other suspension feeders that generally were situated at the top of the structures. Thirty-five stalk communities were collected in tube cores in 1994–1995 using the submersible “Alvin”. A total of 139 taxa was associated with these hard-substratum habitats (another five species were observed only in photographs). Although taxon richness was high, the species diversity of these communities was relatively low due to the dominance in percentage abundance of a foraminiferan, Cibicides lobatulus (Walker and Jakob, 1798), and a serpulid polychaete, Bathyvermilia sp. (Zibrowius, 1973). The relationship between number of taxa and surface area of the stalks yielded a slope (z-value) typical of islands with a low rate of immigration. Three-dimensional complexity created by branching epifauna on the stalks provided more surface area and a variety of cryptic microhabitats. Vertical zonation on the stalks appeared to be controlled by biological interactions among species, with solitary fauna and certain functional groups of colonial organisms restricted by sheet-like colonial organisms that appeared to be dominant space competitors. Received: 13 April 2000 / Accepted: 9 November 2000     

Embryonic,larval, and post-metamorphic development of the sea urchin <Emphasis Type="Italic">Cassidulus mitis</Emphasis> (Echinoidea; Cassiduloida): an endemic brooding species from Rio de Janeiro,Brazil

Cassiduloids are currently rare irregular echinoids with a highly conserved adult morphology. Aristotle’s lantern is present only during the post-metamorphic stage, and little is known about the early development of species in this group. Cassidulus mitis produces eggs of about 375 μm in diameter, lecithotrophic larvae with four reduced arms with skeletal fenestrated rods, cilia along the body surface, and a ciliated band on arms and lobes. Offspring is brooded among the female spines from embryo to settler’s stage. The echinopluteus larval stage is reached 6 days after fertilization, and the settler’s stage is formed at the age of 17 days. Aristotle’s lantern appears around the thirteenth day of development. The lantern is well developed and functional in settlers. It remains until at least 62 days after fertilization and can be used to acquire food from the environment. The early development of C. mitis is unusual concerning features of typical lecithotrophic larvae (such as reduced arms), but retains some features of planktotrophic larvae (such as skeletal rods and a ciliated band). Regarding egg size, early development in C. mitis seems to be transitioning from facultative lecithotrophic to typical obligate lecithotrophic pattern in echinoid larval evolution.     

Halogenated metabolites in two marine polychaetes and their planktotrophic and lecithotrophic larvae

This study investigated the occurrence and ontogenetic changes of halogenated secondary metabolites in planktotrophic and lecithotrophic larvae and adults of two common, infaunal polychaetes, Streblospio benedicti (Spionidae) and Capitella sp. I (Capitellidae), with different life-history traits. S. benedicti contains at least 11 chlorinated and brominated hydrocarbons (alkyl halides) while Capitella sp. I contains 3 brominated aromatic compounds. These halogenated metabolites are potential defense compounds benefiting both larvae and adults. We hypothesized that: (1) planktotrophic larvae contain halogenated metabolites because they are not protected by adult defenses, (2) quantitative and qualitative variation of planktotrophic larval halogenated metabolites parallels that of adults, and (3) brooded lecithotrophic larvae initiate the production of halogenated metabolites only after metamorphosis. To address these hypotheses, volatile halogenated compounds from polychaete extracts were separated using capillary gas chromatography and identified and quantified using mass spectrometry with selected ion monitoring. All four life stages (pre- and post-release larvae, new recruits, adults) of both S. benedicti and Capitella sp. I contained the halogenated metabolites previously identified from adults. This is the first report of halocompounds identified and quantified in polychaete larvae. Allocation of potential defense compounds to offspring varied as a function of species, feeding type and developmental stage. Pre-release larvae of S. benedicti with planktotrophic development contained the lowest concentration of total halogenated metabolites (1.75 ± 0.65 ng mm⁻³), post-release and new recruits contained intermediate concentrations (8.29 ± 1.72 and 4.73 ± 2.63 ng mm⁻³, respectively), and planktotrophic adults contained significantly greater amounts (28.9 ± 9.7 ng mm⁻³). This pattern of increasing concentrations with increasing stage of development suggests synthesis of metabolites during development. Lecithotrophic S. benedicti post-release larvae contained the greatest concentrations of halometabolites (71.1 ± 10.6 ng mm⁻³) of all S. benedicti life stages and developmental types examined, while the amount was significantly lower in new recruits (34.0 ± 15.4 ng mm⁻³). This pattern is consistent with a previously proposed hypothesis suggesting a strategy of reducing potential autotoxicity during developmental transitions. Pre-release lecithotrophic larvae of Capitella sp. I contained the highest concentration of total halogenated metabolites (1150 ± 681 ng mm⁻³), whereas the adults contained significantly lower total amounts (126 ± 68 ng mm⁻³). All concentrations of these haloaromatics are above those known to deter predation in previously conducted laboratory and field trials. As a means of conferring higher larval survivorship, lecithotrophic females of both species examined may be expending more energy on chemical defenses than their planktotrophic counterparts by supplying their lecithotrophic embryos with more of these compounds, their precursors, or with energy for their synthesis. This strategy appears common among marine lecithotrophic larval forms. Received: 14 July 1999 / Accepted: 20 January 2000     

Reproduction of the invasive slipper limpet, Crepidula fornicata, in the Bay of Brest, France

The reproduction of Crepidula fornicata was studied in the Bay of Brest in order to characterise the first step of the reproductive cycle of this invasive species. The survey was carried out from 2000 to 2003 and different parameters were measured, namely, the percentage of the different sexual stages, the straight length of the shell and the percentage of brooding females using a survey of the embryonic development and the fecundity. The juvenile frequency increases generally from mid-June or mid-August, depending on the year. In 2001 and 2003, a first peak was observed as early as May, but it was followed by a rapid disappearance of the individuals. The sex-ratio female/male increased from 0.22 to 0.46 between 2001 and 2003. The sex change between intermediates and females took place mainly in summer and was well marked in 2001 and 2003. The survey of the embryonic development in the egg capsules brooded by the females provided an annual phenology of the laying and hatching processes. The laying period extends from February to September with three to four major periods of egg-laying per year and corresponding hatching periods about 1 month later. Each female lays two to four times per year on average. The first egg-laying concerned fewer females than subsequent ones, except in 2003, and exhibited a higher fecundity. The annual mean of the number of eggs for each stage was not significantly different, thus indicating no significant mortality rate during embryonic development. For the C. fornicata population in the Bay of Brest, several reproductive characteristics tend to highlight its invasive capacity: (1) a long reproductive period, (2) reproduction in a ‘multi-trials’ process equivalent to a spreading out of the risks and (3) a relatively high fecundity.     

Sponge-inhabiting barnacles on Red Sea coral reefs

In this study eight different species of barnacles were found within nine species of sponges from the Red Sea. This brings to 11 the number of sponge-symbiotic barnacles reported from the Red Sea, two of these are new Acasta species (not described herein) and one (A. tzetlini Kolbasov) is a new record for this sea. This number is much higher than that of symbiotic barnacles found within sponges from either the N. Atlantic (2) or the Mediterranean (4). Two possible explanations for this are the presence of numerous predators in coral reefs and scarcity of available substrate for settlement. These factors can lead to high incidence of symbiotic relationships. Of the nine sponge species, only one (Suberites cf. clavatus) had previously been known to contain barnacles. Even at the family level, this is the first record of symbiotic barnacles in two out of the seven sponge families (Latrunculiidae, Theonellidae). Our present findings strengthen the apparent rule that the wider the openings in a barnacle shell, the fewer the host taxa with which it will associate, usually from one or two closely related families, and the more frequent it will associate with elastic sponges. Most Neoacasta laevigata found on Carteriospongia foliascens were located on the same side as the sponge's ostia, i.e. facing the incoming water. This adaptation allows the barnacles to catch more suspended particles from the water, provides them with more oxygen and prevents their exposure to discharged sponge waste. The highest density of barnacles observed on one face of a “leaf ” (with ostia) was 0.389 barnacles cm⁻² (one barnacle per 2.57 cm²) and on average 0.181 ± 0.68, while the average on the other side was only 0.068 ± 0.52 barnacles cm⁻². As indicated by the Morisita index, these barnacles most frequently (58%, n = 12) had a clumped spatial distribution (while the rest were randomly distributed), as is to be expected from such sessile organisms with internal fertilization via copulation. The presence of N. laevigata induced the growth of secondary perpendicular projections of its host C. foliascens. Of the N. laevigata examined, 17% brooded 324 ± 41 embryos each, of 286 ± 17 μm total length; only 5.7% (n = 123) were found to be dead. Size distribution analysis of skeletal elements from dead barnacles showed them to be significantly larger than the skeletal elements of the population of live barnacles ( p < 0.05). Received: 26 June 1998 / Accepted: 1 December 1998     

Parthenogenetic reproduction of Diaphanosoma celebensis (Crustacea: Cladocera): influence of salinity on feeding, survival, growth and neonate production

 Effect of salinity on the feeding rate and parthenogenetic reproduction of asexual females of the cladoceran Diaphanosoma celebensis Stingelin was studied. Short-term (10 h) grazing experiments were conducted using Isochrysis galbana as feed at 5, 17, 25 and 30 psu salinity. Gut pigment concentration showed a significantly higher rate of feeding at lower salinities. Survival, growth, maturity attainment and neonate production of asexual females reared in the above four test salinities indicated preference for lower salinities (5 and 17 psu). The mean size of adult females decreased from 909 to 593 μm, mean life span from 24 to 5 d, mean neonate production from 12 to 2 and mean size of neonates from 434 to 400 μm as the salinity increased from 5 to 30 psu. Salinity variations also affected the size and age of primiparous females. Resting egg formation and sexual reproduction did not occur at the tested salinities. The results indicate that D. celebensis is adapted to low saline, estuarine environments. Received: 14 January 2000 / Accepted: 24 March 2000