Reproduction and developmental variability in the opisthobranchTenellia pallida

Pelagic and non-pelagic development occurs within a single population of the nudibranchTenellia pallida (Alder and Hancock) from South Carolina (USA). Both developmental types occurred simultaneously, under identical conditions, and without the aid of nurse eggs. Adults laid either ≈70 μm diameter eggs that hatched as pelagic (planktotrophic?) veligers in 68 to 122 h, or ≈100 μm diameter eggs that became well-developed veligers, metamorphosed within their egg capsules, and hatched as shelled juveniles in 140 to 190 h (15°C, 30‰ S). Adults that produced pelagic larvae laid more eggs per day and per egg mass and twice the daily egg volume produced by adults of the opposing type.     

Effects of gamma-aminobutyric acid on the settlement of larvae of the black chiton Katharina tunicata

Laboratory-cultured larvae of the black chiton Katharina tunicata (Wood), collected from central California in May and June of 1979 and 1980, settled preferentially on the encrusting coralline alga Lithothamnium sp. Metamorphosis, which involves the loss of the prototrochal ciliary band, occurred within a period of 2.5 h after larvae began crawling upon the alga. In response to gamma-aminobutyric acid (GABA) at a concentration of 10^-6 M, the lecithotrophic trochophore larvae ceased swimming and settled rapidly. Developmental metamorphosis did not follow settlement in the absence of encrusting coralline alga. However, when both GABA and Lithothamnium sp. were present, larval metamorphosis occurred within 2 h of settlement. Significantly different rates of settlement exist for sibling larvae of different ages; 12 d-old larvae responded more rapily and in greater numbers to treatment with GABA than 10 d-old larvae. Differences in settling rates are attributed to the onset of metamorphic competence.     

Habitat selection in the tropical polychaeteSpirobranchus giganteus

Total body weight, tube length, abdomen weight and branchial crown weight of individualSpirobranchus giganteus (Pallas) living on four different coral species on the bank reef on the west coast of Barbados were investigated in 1986. Worms onDiploria strigosa were larger in all size parameters than those onMontastrea annularis, which were larger than those onMontastrea cavernosa, which were larger than those onPorites porites. The differences between worms onM. annularis andM. cavernosa were not significant. Variation in worm size on the different corals could result from variation in mortality and/or variation in growth. Whichever the case, larger worms may have higher lifetime reproductive success. Abdomen weight is an index of gamete production inS. giganteus, and increases with increasing body weight for worms on all corals. Moreover, larger worms do not have lower abdomen weight at a given body size than smaller worms. This may imply that faster growth does not reduce gamete production at body size. The coral species on which worms are largest are those most preferred by worm larvae in the laboratory and most heavily colonised by adult worms in the field. The results suggest adaptive habitat selection by planktonic larvae ofS. giganteus.     

Reproductive isolation between intertidal and subtidal<Emphasis Type="Italic"> Scoloplos armiger</Emphasis> (Polychaeta,Orbiniidae) indicates sibling species in the North Sea

Two distinct modes of development in the common polychaete Scoloplos armiger (O. F. Müller, 1776) occur in the North Sea region: holobenthic development in egg cocoons and pelagic larvae hatching from suspended eggs. In the northern Wadden Sea near the island of Sylt, we observed that egg cocoons are produced intertidally while pelagic larvae originate from the adjacent subtidal zone. A previous genetic comparison between these subtidal and intertidal populations revealed distinct gene pools, suggesting that reproductive differences are not phenotypic but heritable. In this study, crossbreeding experiments show that intertidal and subtidal populations are reproductively isolated. Couples with males and females from different habitats had no offspring. Production of egg cocoons is determined by female origin from the intertidal zone. Pelagic larvae occurred only in couples with subtidal females and subtidal males. Intertidal males have spermatozoa with heads twice as long as those from subtidal males and a significantly shorter flagellum. We suspect that deviating sperm morphology may cause the reproductive breakdown at the fertilization stage. Juveniles hatching from cocoons have shorter anal cirri compared to juveniles that metamorphosed from pelagic larvae. We conclude there to be two sympatric sibling species in S. armiger: 'type I' in intertidal areas, which have egg cocoons, no pelagic larvae, elongated sperm heads, shortened sperm flagella and anal cirri; and a subtidal 'type S', lacking egg cocoons but with pelagic larvae, short sperm heads, long sperm flagella and anal cirri.Communicated by O. Kinne, Oldendorf/Luhe     

Sexual reproduction and settlement of the coral reef spongeChalinula sp. from the Red Sea

Characteristics of the sexual reproduction and larval settlement of the haplosclerid spongeChalinula sp., which inhabits the shallow waters (1 to 6 m) of Eilat, Red Sea, were investigated from September 1985 through to November 1987. This species was found to be a simultaneous hermaphroditic brooder, hence gonochorism is not the rule in the order Haplosclerida. Brooding always takes place in special brooding chambers. While the oocytes in the brooding chambers are among the largest known in sponges (355±37µm), the spermatic cysts distributed in the choanosome are among the smallest known for this phylum (average 26±7µm).Chalinula sp. breeds throughout the year and in experiments most larvae (74%) settled within 1 to 8 h post-release, generally within 4.5 h. Metamorphosis from larval shape to a sessile sponge lasts 1 to 6 h. Thus, larvae had a short swimming period, settled fast, and metamorphosed rapidly (within 1 to 6 h). The large size of the larvae may contribute to their ability to rapidly reorganize their body shape into that of a sessile sponge. In addition, the existence of already differentiated choanocyte chambers in the larvae, facilitates fast construction of the water filtration system in the newly settled sponges. The reproductive and larval characteristics ofChalinula sp. enable the larvae to settle on any vacant space in the reef, which may explain its abundance in the Red Sea.     

Sibling species or poecilogony in the polychaete<Emphasis Type="Italic"> Scoloplos armiger</Emphasis>?

In marine invertebrates multiple modes of development, or poecilogony, may occur in a single species. However, after close examination, many of such putative cases turned out to be sibling species. A case in point may be the cosmopolitan orbiniid polychaete Scoloplos armiger, which inhabits marine shallow sediments. In addition to the well-known direct, holobenthic development from egg cocoons, pelagic larvae have also been described. Our culture experiments revealed a spatially segregated source of the two developmental modes. All females of an intertidal population produced egg cocoons and no pelagic larvae. All but 2 out of 15 females of an adjacent subtidal population produced pelagic larvae and no egg cocoons. Based on these results we performed a molecular genetic analysis (RAPD-PCR) on three intertidal and four subtidal populations in the North Sea. Selected samples from all sites were analysed also by the AFLP method. We found significantly higher genetic diversity within subtidal than within intertidal populations. This is consistent with a wider dispersal by pelagic larvae and a smaller effective population size when development is holobenthic. Total genetic divergence is not related to distance but to the intertidal/subtidal division. We suggest that S. armiger actually represents two sibling species.     

Effects of self-fertilisation and other factors on the early development of the scallop Pecten maximus

Adult Pecten maximus (L.) were dredged off north-east Anglesey, Wales, UK, during 1981. A 2×5 factorial mating was carried out involving self- and cross-fertilisation and the use of stripped spermatozoa. Assessments of yield, normality, and larval size were made at the D larva stage and larval size and mortality were measured after a fortnight's growth of the veligers. Underlying genetic variation was evident at all stages, with egg and sperm generally having a significant interactive effect. Cultures sired with stripped spermatozoa had fewer larvae, with more abnormality, grew slower and suffered higher mortality than most other cultures. Larvae from selfed cultures grew significantly slower than all other larvae. Data from past larval cultures also show that selfed larvae have a reduced growth rate. it is suggested that stripped spermatozoa may interfere with egg and sperm interaction at fertilisation, thus reducing the viability of larvae. On the other hand, the poor growth rate of selfed larvae is probably due to overall reduced heterozygosity compared to outbred larvae.     

Reproductive success of parasitized males in a marine reef fish

Parasitism is hypothesized to reduce reproductive success in heavily parasitized males because females may preferentially mate with less parasitized males (parasite-mediated sexual selection) or parasites may compromise male competitiveness. In marine systems, this hypothesis is largely unexplored. This paper provides the first confirmed record of a copepod ectoparasite (Caligus buechlerae Hewitt 1964) on the common triplefin (Forsterygion lapillum) and evaluates the hypothesis that males parasitized with C. buechlerae experience lower reproductive success than unparasitized males (as determined by the presence and area of eggs within male nests). We found that 38 % of males we surveyed were infected with at least one C. buechlerae, with a median of two individuals per infected male. About 32 % of males were defending eggs, with 62.5 % of those males infected with at least one parasite. Males of greater total length (TL) were both more likely to be infected and more likely to be defending eggs. However, when statistically accounting for the effects of TL, parasite infection had no effect on the probability of defending eggs, or the average surface area of eggs when present. Positive covariation in fish length, the presence of eggs and parasite infection observed here potentially suggest that the importance of parasitic infection on reproductive success may depend upon the strength of selection for larger male body size. Our study is one of the few studies to investigate the effects of ectoparasites on reproductive success in reef fish and also provides a quantitative measure of infection for a widespread species within New Zealand.     

New epizooic symbioses between sponges of the genera Plakortis and Xestospongia in cryptic habitats of the Caribbean

Three new cases of sponge symbiosis between species of Plakortis and Xestospongia were found in reef caves and mesophotic reef habitats of the Caribbean. Plakortis sp. 1 from the Bahamas associates exclusively with Xestospongia deweerdtae which was originally described living freely on the deep fore-reef and caves of Jamaica. In addition, we found Plakortis sp. 2 from Puerto Rico which associates with both X. deweerdtae and a different Xestospongia sp. Sponge specimens were identified using cytochrome oxidase subunit 1, 28S rRNA and 18S rRNA gene sequence fragments, spicule analysis, and histological sections with SEM. Unlike previous sponge pairs, Xestospongia spp. not only grew as a thin veneer of tissue over the Plakortis host sponge but through the mesohyl, forming inner channels (0.1–1 cm) that may provide a benefit by facilitating more efficient water transport through the dense Plakortis tissue. Symbioses with both Plakortis spp. were documented from an early recruit stage through adulthood. Spicule measurements conducted on symbiotic versus free-living X. deweerdtae revealed significantly smaller spicule sizes for symbiotic individuals, suggesting a cost in terms of silicon availability, or a benefit in terms of a lower investment in skeleton synthesis for support. This study reveals new specialized symbiotic associations between distantly related sponge genera that likely represent an alternative strategy of adaptation for life in reef caves and mesophotic reefs.     

Effects of three years’ increase in density of the geometrid Epirrita autumnata on the change in metabolome of mountain birch trees (Betula pubescens ssp. czerepanovii)

The principal objective of the study was to characterize effects of increasing density of the autumnal moth, Epirrita autumnata (Lepidoptera, Geometridae), for 3 successive years on the change in metabolome of mountain birch trees (Betula pubescens ssp. czerepanovii). During the three study years (2000–2002), the larval density varied from low to very high (outbreak) density. Samples of leaves were collected from the same 12 trees at the same phenological stage of the trees each year. The leaves were collected from undamaged trees in the first year, from trees slightly damaged by larvae in the second year, and from trees heavily damaged by larvae in the third year. Metabolome analysis showed that the increase in density of E. autumnata larvae and degree of damage of birch trees caused multiple biochemical changes in the leaves, including increased concentrations of phenolic compounds (proanthocyanidins and hydrolysable tannins) and reduced concentrations of nutritive metabolites (monosaccharides, amino acids and some organic acids). These changes reduced the quality of leaves as food for larvae and, probably, were associated with induced chemical resistance of the birches to herbivorous insects. Additionally, the concentration of α-tocopherol was significantly higher in both slightly and heavily damaged trees. Mechanisms of changes in the metabolism of phenolic compounds and carbohydrates, and the role of α-tocopherol in their regulation are discussed.     

An experimental study of the relationship between below-ground structure and meiofaunal taxa

An experimental study was conducted in a South Carolina, USA salt marsh to examine relationships between meiofaunal-sized tube structure and densities of burrowing meiofauna. Abundance of the tube-building polychaete Manayunkia aestuarina Bourne was increased by erecting exciosure areas from June–August 1979. Non-manipulated areas of identical dimensions were staked out in adjacent locales. Densities of burrowing meiofauna and nematode trophic groups, and volumes of Spartina alterniflora root mat and tube-builders were monitored approximately every 2 wk from each experimental treatment. Volumes of tube-builders were significantly higher inside exclosure sites on all dates in July and August. Root-mat volume and total number of nematodes were not different between treatments at any time. The dominant copepod species, Stenhelia (D.) bifidia, and the nematode species, Spirinia sp., first increased in number inside manipulated sites and then decreased in abundance as tube volume increased. No general trend was apparent for nematode trophic groups when areas with increased volumes of tubebuilders were compared to unmanipulated sites. The fact that densities of S (D.) bifidia and Spirina sp. decreased only when tube-builders attained highest abundance suggests that negative interactions between tube-builders and these meiofaunal taxa may be effective only at elevated abundances.     

Ultrastructure of sperm and spermathecae in Micromaldane spp. (Polychaeta: Capitellida: Maldanidae)

Assessing the possibility that external fertilization has re-evolved requires the study of monophyletic groups that exhibit various reproductive methods. Maldanid polychaetes show a range of reproductive mechanisms, though previous studies of reproduction have hitherto been restricted to larger species with external fertilization. Micromaldane pamelae Rouse and M. nutricula Rouse are small, gonochoristic maldanids that brood directly developing larvae. Both species have sperm with elongate nuclei and an acrosome extending down each side of the anterior end of the nucleus. A true midpiece is absent; two mitochondria extend along the posterior region of the nucleus. Spermatids develop synchronously in large clusters connected by a cytophore. In M. pamelae sperm are released into the water as spermatozeugmata. These are comprised of clusters of sperm with their tails oriented to the centre and the sperm heads facing outwards. Females of M. pamelae and M. nutricula bear pairs of spermathecae ventrally (M. pamelae three pairs, between setigers 10 and 11, 11 and 12 and 12 and 13 and M. nutricula two pairs, between setigers 10 and 11 and setigers 11 and 12). The blind sacs are epidermal invaginations bound closely together. The entrance to each spermatheca may only be 1 to 2 m across with each spermatheca holding several hundred sperm. This represents the first detailed study of spermathecae in the Capitellida. The occurrence and structure of spermathecae and spermatozeugmata in other groups are discussed and compared with Micromaldane spp. Comparisons are made with non-polychaetes with the purpose of discussing functional aspects of reproductive mechanisms in marine metazoans in general. Elongate sperm nuclei are associated with sperm storage and/or large egg size. The lack of an elongate sperm midpiece may be an indicator of having to swim in water but does not contraindicate sperm storage. Spermatozeugmata may serve as an indication of sperm storage and brooding of larvae. Speculations on the phylogenetic significance of these reproductive features are limited by the fact that supposedly modified (i.e., derived) states may reflect functional/structura, constraints of small body size.     

Brooding behaviour and reproductive success in two species of free-living simultaneous hermaphrodites

Although polyclads are amongst the most structurally simple of the triploblastic metazoans, they adopt a wide range of reproductive strategies. Parental care behaviour in this group is yet to be quantified for any species. We assessed the significance of brooding behaviour to the reproductive success of two free-living marine flatworms. Echinoplana celerrima and Stylochus pygmaeus were collected from the field and placed in pairs in containers of filtered seawater where they laid batches of eggs. Both parents were then removed from half of the containers and the brooding behaviour and hatching success of eggs were quantified. There were interspecific differences in brooding behaviour. Egg masses were covered by one E. celerrima parent for 12 ± 2% of time, whereas egg masses of S. pygmaeus were covered by one or both parents simultaneously for 85 ± 8% of time. Egg batches were abandoned by both species immediately prior to the onset of hatching (10–12 days). Hatching success was generally high (~90%) and brooding did not enhance the hatching success of eggs. We assessed the significance of parental care to hatching success of E. celerrima egg masses in the presence of three potential egg predators; in the presence of other organisms. E. celerrima devoted less time to brooding; however, hatching success was not affected. The amount of time spent brooding eggs differed greatly between the two polyclad species but was not essential to their reproductive success under benign conditions. Parental care may be of adaptive value under more stressful environmental conditions commonly experienced in estuarine environments such as lowered salinity, increased hypoxia or turbidity. Covering egg batches may play an additional role of advertising sexual status and a willingness to care for eggs.     

Biologie d'un copépode des mares temporaires du littoral méditerranéen français: Eurytemora velox

The reproduction of Eurytemora velox, a brackish copepod from temporary lakes of the south of France, was studied in winter and spring 1978, under various temperature and salinity conditions, using Chlorella sp. and Amphidinium sp. as food. Maximum numbers of successive eggs sacs (9), eggs per sac (39.3) and total egg production per female (311) were recorded for the witner generation, only 4.8, 34.8 and 109, respectively, for the spring generation. In contrast, the number of eggs per female per day was highest (11.3) in the spring generation, which displayed a more rapid reproductive cycle. Despite strong individual variations in the rhythm of egg sac production and in abundance of eggs per sac, egg production was generally higher during the first third of adult life, attaining a maximum after production of the second or the third egg sac. The continuous presence of the males was necessary to assure complete fertilization of eggs throughout the whole life of adult females. Hatching rate displayed high individual variability, in particular for the spring generation, which had lower average hatching rates (between 0 and 26%, depending on salinity or temperature) than the winter generation (14 to 64%). These differences may be related to the ability of E. velox to produce resting eggs during spring, allowing the species to maintain itself in a temporary water milieu. Temperature significantly affected longevity and daily egg production of females; presence or absence of males did not affect these parameters. An increase in salinity from 20 to 30%. reduced longevity, number of egg sacs, and daily egg production in the winter generation, but not in the spring generation. The specific daily production of females during their adult life was calculated from the egg production:biomass ratio of females, in carbon units. In the winter generation, this ratio increased between 10° and 15°C compared to ratios between 15° and 20°C; the opposite was observed for the spring generation. The seasonal differences in the effects of temperature and salinity on reproduction could indicate an adaptation mechanism to the strongly thermal and haline seasonal fluctuations which characterize the habitat of Ex. velox (brackish waters, drying-up in summer). Larval mortality was high, except at 20%. S for the spring generation. The sex-ratio of the offspring was unaffected by variations in breeding conditions. Hatching time and development time of larvae could be described by two Blehrádek equations displaying close b and () coefficients. We calculated the energy balance of adult females from data obtained in a previous study on feeding and respiration in E. velox, and this is discussed in context with the egg production results. Net growth efficiency varied with algal concentration according to an asymptotic curve, reaching a maximum of 0.43 with Tetraselmis maculata as food or 0.53 with Amphidinium sp. Actual egg production rate obtained in the present study was in good agreement with that calculated by the difference between assimilated food and respiration expenses.     

Brood or broadcast? The adaptive significance of different reproductive strategies in the two intertidal sea stars Leptasterias hexactis and Pisaster ochraceus

This paper considers the adaptive significance of two different reproductive methods in two co-occurring, competing sea stars. The smaller (3 to 8 g mean wet weight) Leptasterias hexactis broods relatively few, large young in the winter, while the large (300 to 650 g mean wet weight) Pisaster ochraceus broadcasts relatively many, small eggs each spring. L. hexactis matures at a small size (2 g wet weight) in about 2 years, and P. ochraceus matures at a larger size (70 to 90 g wet weight) in about 5 years (Menge, 1974). As in many broadcasting asteroids, gonad and storage organ indices of P. ochraceus are inversely related over time, and maximum storage-organ index correlates with the summer feeding maximum (Mauzey, 1966). In contrast, both organ indices of L. hexactis and feeding increase and are positively correlated until early autumn, when feeding activity begins to decline. At this time the male gonad index continues to rise, and the storage-organ index drops. In contrast, both organ indices of females rise. Spawning occurs from November to January. Thereafter storage-organ indices decline in females, presumably because females draw upon energy reserves while brooding; storage-organ indices rise in males, presumably because males do not brood and can feed if food is available. The primary cause for the differences between annual reproductive cycles of P. ochraceus and L. hexactis is suggested to be patterns of food availability for the released young (planktonic food for the broadcasted young of P. ochraceus and benthic prey for the brooded young of L. hexactis. Estimates of pre-maturity survival and post-maturity longevity indicate that the probability of survival per individual of young P. ochraceus is vastly lower than that of L. hexactis. However, once mature, P. ochraceus has a much longer expected lifespan. Brooding is suggested to be a coadaptive consequence of competition-induced small size. Assuming planktonic mortality rates in this environment are roughly constant across broadcasting species, I suggest that a small broadcasting species could not produce enough offspring in its expected lifespan to replace itself. This hypothesis is partly supported by some simple simulations. Broadcasting is suggested to permit rapid location and utilization of spatially and temporally unpredictable, but highly desirable, resources by allowing rapid and widespread dispersal. Brooders presumably cannot disperse rapidly and must rely on more reliable, but perhaps less desirable, resources. Factors affecting reproductive patterns in marine invertebrates include (1) food availability for both adults and offspring, (2) planktonic mortality rates, (3) interactions between species and latitudinal changes in these factors, and (4) various physical factors. This paper suggests that competition and predation can have an important effect on the evolution of reproductive methods, a possibility heretofore largely ignored. Although several similar examples of co-occurring species' pairs which differ in reproductive method and size are available, the role of adult interactions is unknown in these examples.     

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.     

Lipid biochemistry and reproductive biology in two species of Gammaridae (Crustacea: Amphipoda)

Gammarus oceanicus Segerstråle, 1947 and Echinogammarus marinus (Leach, 1815) were sampled during the breeding season from Oslofjord in 1984, and their lipid composition examined in relation to reproductive condition. In G. oceanicus, female lipid content increased as the ovary matured. Both the amount of lipid stored and the rate of accumulation were greater in spring than in winter. Spring eggs contained 12.4 g lipid, of which 63% was triacylglycerol and 27% phospholipid. Both fractions decreased steadily during embryonic development. Winter eggs contained 19.2g lipid, of which 52% was triacylglycerol and 43% phospholipid. During the early stages of embryonic development the amount of phospholipid decreased sharply, whereas that of triacylglycerol increased, suggesting that some of the fatty acid released from phospholipid was sequestered temporarily as triacylglycerol. When newly spawned, both winter and spring eggs were richer in monoenoic fatty acids than adult amphipods and these acids were the major fuel consumed during development. 6 fatty acids were utilised more slowly than 3 acids, and egg carotenoid pigment content remained constant. Female E. marinus increased in lipid content as the ovary matured. Spring eggs contained 14.7 g lipid when newly spawned and this increased to 16.6 g during the early stages of development. This increase was entirely triacylglycerol, which declined in later stages; the source of the extra lipid was unclear. Eggs contained very little phospholipid or sterol, and both of these components remained at a steady low level during development. E. marinus eggs were not significantly rich in thonoenoic acids compared with adults, and saturated, monoenoic and polyenoic acids were utilised about equally during development. Both adults and eggs were rich in 20.46, which was utilised at a slower rate than the 3 polyunsaturated acids during embryonic development; again, egg carotenoid pigment content remained constant. In both species there was a decrease in the size of the egg (and as a result, of the newly hatched juvenile), but an increase in total reproductive output (i.e., the total weight of the egg clutch) per female as the breeding season proceeded. The reproductive output of an individual female is probably related to food availability during the period of ovarian maturation, whereas the size of an individual egg is dictated largely by feeding conditions for the juveniles once they are independent of the female. The different patterns of lipid utilisation during development found in this study emphasize the flexibility of response in the reproductive biology of gammarid amphipods. It is not yet possible, however, to relate the differing patterns in a simple way either to egg size or total female reproductive output. Two outstanding problems are the source of extra triacylglycerol during the early stages of development of E. marinus and the metabolic cost of brooding eggs.     

Genetic differentiation among populations of the Mediterranean hermaphroditic brooding coral<Emphasis Type="Italic"> Balanophyllia europaea</Emphasis> (Scleractinia: Dendrophylliidae)

Spatial models of genetic structure and potential gene flow were determined for five populations of Balanophyllia europaea, a simultaneous hermaphroditic and brooding coral, endemic to the Mediterranean. Six allozyme loci indicated a genetic structure that departed markedly from Hardy–Weinberg equilibrium, with a significant lack of heterozygotes. The genetic structure observed supports the hypothesis that self-fertilisation characterises the reproductive biology of B. europaea. Populations at small spatial scales (8–40 m) are genetically connected, while those at large scales (36–1,941 km) are genetically fragmented; the genetic differentiation of the populations is not correlated to geographic separation. This spatial model of genetic structure is compatible with an inbreeding mating system. Furthermore, it is also consistent with the expected dispersal potential of swimming larvae of brooding corals, i.e. larvae that are able to produce significant gene flows only within limited spatial scales.Communicated by R. Cattaneo-Vietti, Genova     

Pelagic larval development in the brooding Antarctic brachiopod Liothyrella uva

Larvae of the Antarctic brachiopod Liothyrella uva (Broderip, 1833) were released by brooding females between 10 and 15 January 1992. They were collected and cultured for the ensuing 45 d. At release they were at the gastrula stage, and were recognisable as the usual brachiopod three-lobed larva from around 18 d after release. At 22 d post-release, larvae began to congregate under shell fragments placed in culture vessels as potential settlement substrata. However, the larvae showed no signs of settlement which, combined with the absence of many of the attributes of mature larvae seen in previous studies, was taken as evidence that these larvae had not reached fully competent stages. Previous work had shown that L. uva spawns in October, indicating that the developmental period lasts for a minimum of between 115 and 160 d. The release and subsequent development of larvae also shows that what was previously thought to be a brooding species combines a long brooding period with a free-swimming phase. If this is true of other brachiopod species, then there may need to be a recategorization of some species from brooding to combined brooding and free-swimming developmental types. Even though the development period in L. uva was likely to have been underestimated because a fully competent stage was not reached, a comparison with development rates for temperate species indicated a slowing of between 8 and 70 times. This dramatic slowing of development rate in brachiopods now joins similar data previously reported for echinoderms and nemerteans, suggesting that this is a very widespred, perhaps universal, attribute of Antarctic marine invertebrates.