Organ-like gonads, complex oocyte formation, and long-term spawning in Periphylla periphylla (Cnidaria, Scyphozoa, Coronatae)

An account is given about the development of the gametes of the holopelagic coronate scyphomedusa Periphylla periphylla (Péron and Lesueur 1809). The gonads of the species are complex and differ from those thus far described in Scyphozoa in having this characteristic composition of trophocytes, follicle cells, gamete-releasing pores, mucus cells, and resorption cells. Our results differ from those of previous coronate studies with respect to the contact of the oocytes with the gonad tissue throughout the duration of development. Among the Medusozoa, follicle cells have thus far been considered as an apomorphy for the Staurozoa, but their presence in the Scyphozoa casts some doubts on this assumption. From morphological structure and examination of the gonads and gametes, it can be deduced that P. periphylla has true organs and that gamete release is continuous. These results supplement our knowledge of the reproductive biology of this mesopelagic species.     

Life history and settlement preferences of the edible jellyfish Catostylus mosaicus (Scyphozoa: Rhizostomeae)

 The life history and settlement preferences of larvae of Catostylus mosaicus (Scyphozoa: Rhizostomeae) (Quoy and Gaimard, 1824) were investigated in New South Wales, Australia, over a 2 mo period beginning in November 1998. The life history consisted of an alternation between a sexual, medusoid stage and an asexual, polypoid stage, and was similar to that described for other rhizostomes. Planula larvae were brooded by the adults. Approximately 4 d after collection, larvae settled on a variety of substrata including wood, sandstone, shell, seagrass and glass, and metamorphosed into four-tentacled polyps. The number of tentacles increased and polyps strobilated when they had between 12 and 20 tentacles. Strobilation occurred within 15 d of settlement, but only polyps that settled on the concave surfaces of the shells strobilated. Both monodisk and polydisk strobilation was observed. Ephyrae were raised for one month and were observed developing oral arms. Polyps reproduced asexually via the formation of podocysts, by production of buds, and by partial fission. Received: 30 April 1999 / Accepted: 27 August 1999     

Asexual repooduction in Glossobalanus crozieri (Ptychoderidae,Enteropneusta, Hemichordata)

Several aspects of asexual reproduction in Glossobalanus crozieri (a species first found in Bermuda, but occurring also on the coast of Brazil) have been observed. The results are compared with those by earlier authors on other species of Enteropneusta. A revision of asexual reproduction in the Enteropneusta (Hemichordata) is made. Developmental rates and regeneration were studied in branchiogenital individuals, hepatic individuals and regenerands. The habitat where these forms are found is described. Possible meaning of gonad maturation and sexual reproduction in relation to asexual reproduction in different species is discussed. The need for further studies on the variation of forms which reproduce both sexually and asexually is emphasized, with a view to taxonomic problems, based both upon the proportions of the different body regions and on the number of branchial pores. The present findings indicate that, when compared to the results reported for other species, asexual reproduction in all ptychoderids (Balanoglossus, Glossobalanus and Ptychodera) follows the same general pattern. The phenomenon of asexual reproduction in this group is also confirmed to be worldwide.     

Black spots and female association preferences in a sexual/asexual mating complex (Poecilia, Poeciliidae, Teleostei)

We investigated whether female association preferences for males are influenced by black spot disease (BSD), a parasite induced change of the host phenotype. We compared three different species of fish: a gynogenetic hybrid species, Poecilia formosa (amazon molly) and two sexual species (Poecilia latipinna and Poecilia mexicana), which were involved in the natural hybridisation leading to the amazon molly. Contrary to their sexual relatives, asexual amazon mollies significantly avoided images of males infected with black spot disease. We propose that amazon molly females have direct fitness benefits from choosing healthy males. The adaptive significance of the preference for BSD-uninfected males in the asexual amazon molly is yet unclear but may involve avoidance of predation or parasite infection as well as increased sperm availability.     

Complex reproductive cycle inEucheilota paradoxica (Hydrozoa: Leptomedusae): Medusae,polyps and frustules produced from medusa stage

Eucheilota paradoxica Mayer, 1900 is a small leptomedusa unknown in the Mediterranean Sea until 1977. Since then, it has become endemic and occurs in great abundance in autumn in the bay of Villefranche-sur-mer. Since no sexual stages have ever been collected, either from the Atlantic Ocean or from the Mediterranean Sea, we decided to investigate the still unknown life cycle of this species. Specimens were obtained in the bay of Villefranche-sur-mer from plankton hauls between 0 and 50 m depth during October and November 1986. At temperatures >18°C, the medusa reproduces intensively by asexual medusal budding (one young medusa liberated per two days). This asexual reproduction explains the rapid proliferation of the species and the origin of the large population every autumn. In cultures maintained at temperatures <18°C, two other reproduction modes, not yet reported in a leptomedusa, were observed: (1) At 16° to 17°C, frustules bud along the canals; these frustules are dormant, giving rise to a small polyp when the temperature increases once more above 18°C. (2) At temperatures close to 15°C each medusa, in place of the gonads, differentiate into one or two polyps; these polyps develop progressively, concomitant with gradual regression of the mother medusa.     

Mode of reproduction,recruitment, and genetic subdivision in the brooding sponge<Emphasis Type="Italic"> Haliclona</Emphasis> sp.

Sponges display a variety of reproductive strategies that have the potential to influence population genetic structure. Histological examination of ten reproductive individuals of the Western Australian sponge Haliclona sp. showed that this species broods embryonic larvae that are potentially limited in dispersal capabilities. Because sponges have the potential to propagate in a number of modes, allozyme electrophoresis was used to assess the relative importance of asexual and sexual reproduction to recruitment, and to quantify genetic subdivision over different spatial scales. Tissue samples from 227 sponges were collected from reefs within two areas 400 km apart: Hamelin Bay and Rottnest Island. Contrary to expectations for highly clonal populations, genotypic diversity within sites was high, no linkage disequilibrium was found, and there was no evidence of genotypic clustering within reefs. There was no genetic evidence that asexual reproduction is important for the maintenance of populations. Genetic comparisons were consistent with mixing of sexually produced recruits within reefs, on a scale up to a few hundred metres, but significant genetic subdivision between reefs (F_ST=0.069 at Hamelin Bay, 0.130 at Rottnest Island) indicated that water gaps of several hundred metres are effective at preventing dispersal. Subdivision between the two areas, separated by 400 km, was moderately greater (F_ST=0.142) than within, but the same alleles were predominant in the two areas. These genetic patterns are consistent with limited dispersal capabilities of brooded larvae.Communicated by G.F. Humphrey, Sydney     

Life cycle of <Emphasis Type="Italic">Carybdea marsupialis</Emphasis> Linnaeus, 1758 (Cubozoa,Carybdeidae) reveals metamorphosis to be a modified strobilation

Two types of metamorphosis occur in the life cycle of Carybdea marsupialis Linnaeus 1758 (Cubozoa, Carybdeidae). In addition to the metamorphosis described several times over, during which the entire polyp transforms into one medusa, we investigated a second type of metamorphosis which leaves a regenerative remnant. This mode of medusa formation seems to be derived from the strobilation known from Scyphozoa. Both types are described and the implications these results have on the idea about the origin of the class Cubozoa, are discussed.     

Effect of temperature,salinity and food level on sexual and asexual reproduction in Brachionus plicatilis (Rotifera)

The reproductive response of sexual and asexual female Brachionus plicatilis (Muller) was examined over temperatures ranging from 20° to 40°C, salinities from 5 to 40 S, and food levels from 0.25 to 20 g Chlorella vulgaris dry-weight per ml. Reduced food levels, as well as temperature and salinity extremes, reduced reproduction of both sexual and asexual females, but did so differentially. Reproduction by sexual females was reduced to a greater extent at environmental extremes than asexual females. The broad, flat reproductive response curve of asexual females extended beyond the limits of the narrower, more sharply peaked curve of sexual females. Thus zones of exclusively asexual reproduction exist at environmental extremes where sexual reproduction is physiologically restricted. These results are corroborated by a comparison of the lifetime fecundity of individual sexual and asexual females over a 20°C temperature range. No differences in lifetime fecundity occurred between sexual and asexual females at 18° and 28°C. At 38°C, however, asexual female fecundity reached its highest level, while sexual female fecundity declined 15%. The appearance of sexual females in rotifer populations in the result of both inducible and repressible factors.     

Bud formation and metamorphosis inCassiopea andromeda (Cnidaria: Scyphozoa): A developmental and ultrastructural study

Asexual reproduction by formation of swimming buds which metamorphose directly into polyps plays a most important role in the propagation ofCassiopea andromeda (Cnidaria: Scyphozoa). (C. andromeda polyps, originally supplied by the Löbbecke Museum and Aquarium Düsseldorf, FRG, were cultured in our laboratories.) We have defined five budding stages and investigated epithelial recruitment and dynamics during bud formation using intracellular vital stains. The region of cell recruitment was found to encircle the budding site asymmetrically. The aboral side contributing considerably less to the developing bud than the oral and lateral sides. Furthermore, it was found that the epithelial flow involved in bud formation is part of a permanent apico-basal displacement of ectodermal cells. Light and electronmicroscopic investigations revealed that no drastic changes occur at the cellular level, neither in the ectoderm nor in the endoderm which both participate in bud formation. Scanning and transmissionelectron microscopic investigations of the swimming bud revealed that the ectoderm is composed of three, and the endoderm of two, cell types. Nerve elements have been detected near the mesoglea between both ecto- and endodermal cells. Amoebocytes are regularly found either at the basis of epidermal cells or within the mesoglea, whereas symbionts are located in the endoderm. Buds induced to metamorphose by a bacterial-inducing factor were used to investigate morphological and ultrastructural changes occurring during metamorphosis and scyphistoma morphogenesis. Metamorphosis starts with the settling of a bud, followed by the formation of the pedal disk in which desmocytes, as typical cnidarian adhesive structures, are differentiated. Metamorphosis is completed with the formation of the mouth and tentacles. Whereas metamorphosis of cnidarian planulae implies considerable changes at the cellular level, tissue remodeling processes prevail in bud metamorphosis ofC. andromeda.     

Asexual viviparity and population genetics of Actinia tenebrosa

The intertidal anemone Actinia tenebrosa is viviparous. An electrophoretic study of 3 polymorphic enzymes in Western Australian populations has confirmed genetic identity of adults and their brood young, indicating asexual reproduction. The population effects of this clonal reproduction are seen as linkage disequilibrium and departures from Hardy-Weinberg equilibrium within populations, and large differences between populations. The data also suggest occasional sexual reproduction, and the likelihood of a mixture of reproductive modes in this species.     

Reproductive strategies of Mycalecontarenii (Porifera: Demospongiae)

The sexual and asexual phases of the reproductive cycle of the sponge Mycalecontarenii (Martens, 1824) were studied from samples collected over a period of 2 years (June 1994 to May 1996) in a Mediterranean coastal basin (Porto Cesareo, southwestern Apulia). The species is viviparous; year round it shows a discontinuous oocyte production. No males were found, possibly owing to the very short period of spermatogenesis. M.␣contarenii produces asexual buds during autumn and winter, and this process seems to be related to changes in water temperature. Even though, at present, we cannot evaluate the incidence of larval recruitment, we postulate that asexual reproduction in M.contarenii plays a crucial role in maintaining the sponge population. Received: 8 July 1997 / Accepted: 5 November 1997     

Ovarian morphology and oogenesis in Aurelia aurita (Scyphozoa: Semaeostomae): ultrastructural evidence of heterosynthetic yolk formation in a primitive metazoan

In Aurelia aurita, the ovaries arise as horseshoe-shaped evaginations of the gastrodermis in the floor of four interradial gastric pouches. Germ-cell islands arise within endodermally-derived gastrodermal cells. Oocytes grow and gradually bulge into the mesoglea while maintaining physical contact throughout vitellogenesis with specialized cells called trophocytes. Ultrastructural changes suggest that these cells transport yolk precursors from the coelenteron to the oocytes in a manner similar to that reported for the trophonema cells of anthozoan ovaries. Vitellogenesis involves both the autosynthetic activity of the oocyte organelles (Golgi complex and rough endoplasmic reticulum) and the heterosynthetic incorporation of precursors through endocytotic processes involving both coated pits and vesicles and smooth-surfaced tubules. Ultrastructural data suggest that different types or classes of yolk precursors enter the oocyte through the trophocytes and via the surrounding mesoglea. To our knowledge, this is the most primitive animal in which this type of yolk synthesis has been described. The trophocyte-oocyte relationship in oocytes of A. aurita is reminiscent of the trophonema-oocyte relationship in anthozoans and supports the belief that the Anthozoa and Scyphozoa share a close phylogenetic relationship.     

Ultrastructure of the ovary and oogenesis in the jellyfish Linuche unguiculata and Stomolophus meleagris,with a review of ovarian structure in the Scyphozoa

Ovarian structure and oogenesis has been examined in six scyphozoan species including the semaeostome Diplumularis antarctica Maas, 1908 (collected in 1987 in McMurdo Sound, Antarctic), the rhizostomes Cassiopea xamachana Bigelow, 1892 (collected in Belize in 1988), and Stomolophus meleagris L. Agassiz, 1862 (collected in Ft. Pierce Inlet in 1988), and the coronates Periphylla periphylla (Peron and Lesueur, 1810), Nausithoe atlantica Broch, 1914 (both collected in the Bahamas in 1988), and Linuche unguiculata (Schwartz, 1788) (collected in Nassau Harbor, Bahama Islands in 1989). Based on these findings and information on five other scyphozoan species from additional literature sources, at least two fundamentally different types of ovaries exist in the Scyphozoa. In semaeosotome and rhizostome species, oocytes develop in close association with specialized gastrodermal cells called trophocytes which may serve a nutritive function. However, coronate species lack trophocytes and oocytes develop freely in the mesoglea. The ovaries of S. meleagris and L. unguiculata are used as models to represent the ultrastructural events occurring during oogenesis in species having trophocytes and those lacking them, respectively. In both L. unguiculata and S. meleagris, the ovaries arise as evaginations of the gastrodermis in the floor of interradial pouches. Germ cells appear to originate from endodermally-derived gastrodermal cells and migrate into the mesoglea prior to vitellogenesis. In L. unguiculata, the oocytes develop freely within the mesoglea throughout vitellogenesis, while in S. meleagris each oocyte maintains contact with specialized gastrodermal cells called trophocytes. In the vitellogenic oocytes of both species, numerous invaginations of the oolemma result in the formation of intraooplasmic channels throughout the ooplasm. These channels are intimately associated with cisternae of rough endoplasmic reticulum and may play some role in yolk precursor uptake by substantially increasing the surface area of the oocyte. Vitellogenesis is similar in both species and involves the autosynthetic activity of the Golge complex and rough endoplasmic reticulum, and the heterosynthetic incorporation of yolk precursors through receptor-mediated endocytosis. However, in the oocytes of S. meleagris, the trophocytes probably play a role in the transfer of nutrients from the gastrovascular cavity to the oocyte. The present study suggests that scyphozoans were among the first metazoans to develop ovarian accessory cells during their reproductive evolution. The trophocyte-oocyte association observed in some scyphozoans is similar to but structurally less complex than the trophonema-oocyte association described from anthozoans. Scyphozoan ovarian morphology helps support the view that the Scyphozoa share a closer phylogenetic relationship with the Anthozoa than with the Hydrozoa.     

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     

Genetic study of the extent and consequences of sexual and asexual reproduction in the deep-sea epizoic anemones Amphianthus inornata and Kadosactis commensalis (Cnidaria: Anthozoa)

The anemone Amphianthus inornata is found at bathyal depths living on colonies of the gorgonian Acanella arbuscula. Previous studies of the morphology and reproductive stage of this anemone, during different times of the year, have indicated that it reproduces sexually on a seasonal basis. A small proportion of the study population were also reported to be undergoing asexual reproduction by fission. The anemone Kadosactis commensalis is also bathyal, but lives mainly on the holothurian Paroriza prouhoi. Previous morphological studies have indicated that K. commensalis is a protandrous hermaphrodite that exhibits non-seasonal sexual reproduction only. In the present study, allozyme electrophoresis was used to examine the prevalence and genetic consequences of asexual reproduction in a population of Amphianthus inornata from 2 200 m in the Rockall Trough, North Atlantic Ocean. Genetic evidence, from five randomly selected polymorphic enzyme loci, for asexual reproduction in this species was weak. Exact tests indicated that genotype frequencies did not differ significantly from those expected under Hardy–Weinberg equilibrium. F _IS (correlation of homologous alleles with reference to local population, assuming random mating) values also did not differ significantly from zero, and observed heterozygosity (H _o =0.446) and genotypic diversity (G _o =17.0387) were very similar to Hardy–Weinberg expected frequencies (H _e =0.446; G ^* _e =17.0010). Evidence suggests that the contribution of asexual reproduction to recruitment in the study population of A. inornata is low. For a single population of K. commensalis from 4 850 m on the Porcupine Abyssal Plain, North Atlantic Ocean, the hypothesis that inbreeding due to reduced occurrence of outcrossing between anemones on a single holothurian host was examined by electrophoresis of ten randomly selected enzyme loci. Single-locus genotypic frequencies were significantly different from expected frequencies for one locus P≤ 0.05, hexokinase-1 (Hex-1)]. F _IS values were significantly different from zero for two enzyme loci (Hex-1 and Hex-2, P≤ 0.01 and P≤ 0.05, respectively), and the overall observed heterozygosity was lower than the expected heterozygosity (H _o =0.125, H _e =0.140). The hypothesis of inbreeding could not be rejected by the present study, although sample size was small (N=55), leading to possible bias in tests of significance. Genetic variation in A. inornata was higher than that recorded for most eukaryotes, although interlocus sampling error for only five loci is high. High genetic variability has been found in other sea anemones, and has been related to high longevity and mixed reproductive modes. Genetic variation in K. commensalis was in the higher range of that found in other eukaryotes, and is not unusual for anemones. Received: 5 August 1996 / Accepted: 11 December 1996     

Divide and broadcast: sexual reproduction in the West Indian brittle star Ophiocomella ophiactoides and its relationship to fissiparity

It is known that asexual reproduction by fission is the dominant mode of propagation in the small (disc diameter <5.0 mm) ophiocomid brittle star Ophiocomella ophiactoides (H. L. Clark). This species is, however, able to reproduce sexually as well. Sexual reproduction occurs throughout much or all of the year in a population of O. ophiactoides at Jamaica. Gonads are often present in recently split individuals, indicating that sexual and asexual reproduction could occur simultaneously. Regeneration of new gonads in the new disc half is underway by the time an individual is competent to split again. Unsexable individuals are significantly smaller than males, which in turn are significantly smaller than females. The unsexable individuals appear to be below a threshold size for gonad production. Fecundity is low, with the largest females producing a maximum of about 7 400 eggs of small (mean diameter 80 m) size. Fertilized eggs develop into planktotrophic ophioplutei which are typical of the larvae of other species of ophiocomid brittle stars in morphology, rate of development and duration of planktonic period. Egg size and larval type for O. ophiactoides are characteristic of brittle stars with the planktotrophic mode of development, but egg number is uncharacteristically low. Fissiparity in O. ophiactoides may have evolved in association with small body size, concomitant low fecundity and a high-risk planktotrophic larva as a means of ensuring reliable recruitment. Available data, although sparse, suggest that this hypothesis may be applicable to fissiparous brittle stars in general. Fission in brittle stars has a broad systematic incidence at the familial level which suggests that it has evolved recurrently. The ability both to divide and broadcast appears to be a useful combination for O. ophiactoides: it can recruit reliably at large size to occupy favourable habitats quickly, but, it can still maintain low level dispersal of larvae which could provide escape from deteriorating habitats and the potential to colonize new habitats.Contribution No. 328 of the Discovery Bay Marine Laboratory, University of the West Indies     

Sexual and asexual reproduction in two species of Tethya (Porifera: Demospongiae) from a Mediterranean coastal lagoon

The sexual and asexual phases of reproductive cycles of two sponges, Tethya citrina and T. aurantium, living sympatrically in a Mediterranean coastal lagoon (Stagnone di Marsala, NW Sicily) were studied from samples collected over an 18-mo period. Both species are oviparous and gonochoric. They have a summer, partially overlapping, period of oocyte production, although T. citrina appear to mature earlier. No males were found, possibly due to the very short period of spermatogenesis. Both species produce asexual buds during the autumn/winter months. However, they seem to follow different reproductive strategies, with T. citrina showing a significantly lower production of buds than T. aurantium; by contrast, egg production is significantly lower in the latter species. The difference in the reproductive resource allocation is consistent with data reported in the literature on the anatomy features, genetic population structure and ecological distribution.     

Differences between native and invasive<Emphasis Type="Italic"> Caulerpa taxifolia</Emphasis>: a link between asexual fragmentation and abundance in invasive populations

The tropical and subtropical marine green alga Caulerpa taxifolia has invaded several temperate regions throughout the world, including southeastern Australia. In this study, I examined how invasive C. taxifolia from temperate southeastern Australia differed from native C. taxifolia from subtropical Moreton Bay, Australia, in the traits proposed as being important to its invasion success: thallus size and density, levels of asexual reproduction (fragmentation) and total biomass. Against the prediction of a large size for invasive C. taxifolia, native populations from Moreton Bay had larger stolons and fronds than invasive populations. However, invasive populations consistently had much higher densities of stolons, fronds and fragmented fronds; and a greater biomass compared to native populations. Average densities at invasive sites exceeded 4,700 stolons and 9,000 fronds/m² and were as high as 27,000 stolons and 95,000 fronds/m², which are the highest reported for C. taxifolia anywhere. Average densities of fragmented fronds at invasive sites were as high as 6,000/m² and up to 45% of all stolons at invasive sites could be directly linked to asexual recruitment via fragmented fronds. Importantly, at invasive locations there was a strong association between asexual reproduction and abundance demonstrated by positive correlations between the density of fragmented fronds and total biomass. These findings are the first to describe quantitative differences between native and invasive C. taxifolia and to demonstrate a link between the high levels of asexual reproduction and high abundance in invasive populations. Although the causes and consequences of high levels of asexual reproduction remain to be explored, this study suggests that changes in demographic and life-history traits during the invasion by C. taxifolia into temperate habitats may contribute to its success there.Communicated by M. S. Johnson, Crawley     

Allozyme electrophoresis demonstrates that the scleractinian coral Montipora digitata is two species

Allozyme electrophoresis at five loci was used to clarify the species status of two morphs of the scleractinian coral Montipora digitata (Dana, 1846), using specimens collected from three locations off the Queensland coast between October 1991 and April 1993. The two morphs, occurring sympatrically, were distinguished by one fixed gene difference at Locus LT-2 and frequency differences at three other loci, indicating reproductive isolation between them. Populations of both morphs were generally in Hardy-Weinberg equilibrium, suggesting that sexual reproduction does occur. Despite sampling designed to reduce the chance of collecting clonemates, the genetic diversity ratio (G _O:G_E) indicated that asexual reproduction also occurs. This is in agreement with the known modes of reproduction for this species. There was significant genetic differentiation between populations of one of the morphs of M. digitata. This could have been influenced by greater degrees of asexual reproduction in this morph, selection within sites, or variation in larval survival and recruitment patterns. Relatively low F _st (differentiation among populations) values found for a broadcast spawning species during this study are in accordance with the hypothesis that brooded planulae are adapted for rapid settlement, whereas planulae produced by broadcast spawners are adapted for widespread dispersal.     

Genetic variation and clonal structure in the scleractinian coral Pocillopora damicornis in the Ryukyu Archipelago, southern Japan

Samples of the scleractinian coral Pocillopora damicornis were collected from six sites located around four islands in the Ryukyu Archipelago, southern Japan, and subjected to allozyme electrophoresis. Seven polymorphic loci were examined for their allelic patterns. The ratio of observed to expected genotypic diversity (0.30 < G _o :G _e  < 0.64), the ratio of the observed number of genotypes to the number of individuals (0.47 < N _g :N _i  < 0.75), and deviations from Hardy–Weinberg equilibrium indicated that asexual reproduction plays a major role in the maintenance of established populations. However, populations were not completely dominated by a single or a few clones, and most clones were represented by only a few individual samples. The high frequency of typhoons in the region suggests that, in P. damicornis, fragmentation caused through occasional exposure to powerful waves is a major mode of asexual reproduction, but asexual production of planulae may also be contributing to the maintenance of populations. A significant genetic differentiation (F ST) was found between the six populations examined (0.027 < F ST < 0.092, average F ST = 0.056). The moderate gene flow is discussed according to characteristics of the larval stage of the species, and to circulation patterns in the region. Received: 7 August 1998 / Accepted: 18 May 1999