Reproduction and growth of Holothuria atra (Echinodermata: Holothuroidea) at two contrasting sites in southern Taiwan

Reproductive periods and growth of two populations of Holothuria atra Jaeger distinctly different in body size at two sites of southern Taiwan were determined. Individuals examined in the present study were collected between march 1990 and February 1992. At Nanwan (21° 57N, 120° 45E), large individuals (351 to 1400 g wet wt) spawned from June to September. At Wanlitung (22°N, 120° 42E), a small proportion of frequently dividing individuals (<190 g wet wt) have mature gonads in May, June and September, but histological examination revealed no sign of spawning. Sexual recruits, defined as small individuals <5 g wet wt without sign of regeneration, were not found at either site during this 2-yr study. After the peak of fission at Wanlitung, 40% of the population showed signs of external regeneration. At Nanwan, small individuals transferred from Wanlitung grew from 6 g (n=6) to 166±8 g within 8 mo, and from 48±4 g (n=50) to 324±16 g within 1 yr, with a 6.8-fold biomass increase in 1 yr. At Wanlitung, the monthly average body weight of H. atra is between 33 and 62 g, apparently due to frequent fission, and the biomass increased only 2.9-fold in 1 yr. In southern Taiwan, sexual reproduction of H. atra occurs in large individuals. Asexual reproduction, occurring in small individuals, is the chief mechanism for population maintenance and increase, but it may decrease sexual reproductive potential.     

Sexual and asexual reproduction of<Emphasis Type="Italic"> Allostichaster capensis</Emphasis> (Echinodermata: Asteroidea) in Golfo Nuevo

Sexual and asexual reproduction of the fissiparous starfish Allostichaster capensis were examined for 24 months at Bahía Kaiser, Golfo Nuevo, Chubut, Argentina. Gonad indices revealed an annual reproductive cycle, with peak development occurring in late winter (August). Spawning occurred in spring (September). Pyloric caeca indices displayed a clear annual cycle and a reciprocal relationship with gonad indices, showing the storage function of pyloric caeca. The starfish population appears to be largely maintained by fission, as only two females were found. A. capensis also showed an annual cycle of fission, with the highest frequency in spring and summer (from November to January), just after spawning. Incidence of fission was correlated with environmental factors such as photoperiod, seawater temperature, and salinity.Communicated by P.W. Sammarco, Chauvin     

Genetical structure within populations of the coral Pocillopora damicornis

Genetic variation of allozymes within populations of Pocillopora damicornis from southwestern Australia was consistent with a primary role of local asexual proliferation of clones in population maintenance. Populations were composed typically of two to four multilocus genotypes accounting for 40 to 80% of individuals, with the remainder assigned to genotypes occasionally in twos or threes but more commonly singly. In the three populations where recruitment was examined genetically, 84% of all first-year recruits was assigned to clones represented in the population's resident adults. The majority of these recruits came from the most highly-replicated of the adult clones. The observed genotypic diversity was, on average, about half that calculated to occur for the same allelic frequencies in a sexually-reproducing population with free recombination. Despite the prevalence of asexual reproduction, both through planulae and fragments, the existence of a sexual mode of reproduction was inferred from the high level of variation produced by pooling populations, the existence of novel genotypes and the concordance of clonal gene frequencies at many sites with the predictions of Hardy-Weinberg equilibria.     

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

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

Population genetic structures of the fissiparous seastar Coscinasterias acutispina in the Sea of Japan

The morphological characteristics and the population genetic structures of the fissiparous seastar Coscinasterias acutispina were investigated for eight sites in the Sea of Japan in order to clarify the presence of sexual and asexual reproduction. Morphological observation based on arm length showed that fission was common at all eight sites examined, indicating the likely production of clonal individuals. A random amplified polymorphic DNA (RAPD) marker was used to detect clones arising by fission and to assess gene flow among sites. A simulation approach using RAPD data revealed the presence of clonal individuals at almost all sites, suggesting the existence of asexual reproduction. The result of phylogenetic analysis according to RAPD genotype showed no relationship between genetic and geographic distances. Considering the limited movement ability of seastar species during the adult phase, these observations suggest the existence of marked gene flow among sites, due to dispersal of planktonic larvae produced by sexual reproduction. These observations suggest that multi-locus genotypic compositions depend on the relative amounts of recruitment from sexual and asexual reproduction in each population.     

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

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

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.     

Ecological aspects of ophiuroids from the phytal of S. W. Atlantic Ocean warm waters

Nine species of ophiuroids — Amphipholis januarii, Axiognathus squamatus, Hemipholis clongata, Ophiocnida scabriuscula, Ophiactis lymani, Ophiactis savignyi, Ophionereis reticulata, Ophioderma appressum and Ophiothrix angulata —were collected in various habitats from the mid-littoral and upper infralittoral on the northern coast of the State of São Paulo, Brazil. The abundance of 6 species occurring in 28 species of algae was investigated. A total of 220 samples was collected at monthly intervals from April, 1969 to May, 1970. The distributional pattern of the populations, as determined by Mobisita's index, shows aggregation of the ophiuroids. No ophiuroid species was limited to one algal species. Each species prefers certain algal species and/or a different level on the shore. The most abundant species on algae was Ophiactis lymani, with a maximum density of 1,510 individuals/100 ml of algae. Four species of ophiuroids occurred inside sponges. The most abundant species in sponges was O. savignyi, with a maximum density of 1,892 individuals/100 g of dry sponge. Two species of ophiuroids occurred inside the bryozoan Schizoporella unicornis. Feeding observations have shown Axiognathus squamatus to capture its food by mucous nets. Decapod crustaceans fed on ophiuroids in aquaria. The copepod Thespesiopsyllus sp. was found in the body cavity of the 2 Ophiactis species. Simultaneous sexual and asexual reproduction were abserved in O. savignyi.     

Rates of fission, somatic growth and gonadal development of a fissiparous sea star, Allostichaster insignis, in New Zealand

Seasonal changes in population structure and incidence of fission were measured in intertidal and subtidal populations of Allostichaster insignis, a fissiparous sea star. Population size structure was stable over the course of the 1-year study. Sea stars in the subtidal zone attained greater maximum size (mean arm length, R = 35 mm) than those in the intertidal population (20 mm). Fission rates were greatest among small individuals (R < 20 mm). The frequency of fission ranged from 5 to 32% with peaks in early austral summer in the intertidal zone, and in autumn and winter in the subtidal zone. Sexual reproduction occurred in early spring in sea stars larger than 12 mm. The populations were heavily biased toward males. In the laboratory, A. insignis of three size classes (small, R = 9–13 mm; medium, 19–21 mm; and large, 29–31 mm) were fed mussels ad libitum or starved (not fed macroscopic food) for ∼1 year in a 3 × 2 factorial experiment. Small and medium-sized sea stars divided throughout the experiment and the ramets of most individuals regenerated sufficiently to divide again after 6–9 months. Unfed sea stars did not undergo fission (with one exception), had a higher mortality rate, and did not grow. Small, fed sea stars grew significantly faster than medium-sized or large individuals. At the end of the experiment, the pyloric caeca index (a measure of nutritional condition) was greater in fed than in unfed animals. Gonads (only testes were observed) developed in medium-sized and large, fed sea stars. Our field and laboratory results indicate that asexual reproduction in A. insignis predominantly occurs in small, well-nourished individuals. Ramets grow gradually through repeated fission and regeneration to a size (mean length of regenerating arms, R _r ∼ 20 mm) at which they begin to switch to sexual reproduction as the dominant reproductive mode.     

Genetic structure of populations of two closely related brittle stars with contrasting sexual and asexual life histories,with observations on the genetic structure of a second asexual species

Ophiocoma pumila Lütken andOphiocomella ophiactoides (H. L. Clark) are morphologically similar brittle stars with contrasting life histories, the former obligately sexual, the latter fissiparous (capable of both sexual reproduction and asexual proliferation by binary fission). Electrophoretic analysis of five polymorphic enzymes was used to assess the genetic consequences of these differing life histories and provide a genetic perspective on the taxonomic relationship between the two species. Genotypic diversity ofOphiocoma pumila collected at Discovery Bay, Jamaica, in 1985 conformed to expectations for a sexually reproducing population. In contrast, genotypic diversity ofOphiocomella ophiactoides at this site was significantly lower than expected for a sexually reproducing population, due largely to the predominance of clonal proliferation over larval recruitment. Large variation in clonal composition over a short (50 m) distance emphasized the very localized scale of clonal mixing in this species. Allozymic data are indicative of a close sibling species relationship betweenOphiocoma pumila andOphiocomella ophiactoides which suggests that the present generic separation of the two species should be re-examined. Electrophoretic analysis was also used to examine the genetic structure of sponge- and alga-dwelling populations of a second fissiparous brittle star,Ophiactis savignyi (Müller & Troschel), which was also collected at Jamaica in 1985. Striking differences in the allelic composition of sponge- and alga-dwellingO. savignyi were observed. Genotypic diversity ofO. savignyi in sponges was very low, each sponge being dominated by a single genotype. Genotypic diversity ofO. savignyi in algae was higher, although still significantly lower than expectations for a sexually reproducing population. In the light of the highly clonal composition of fissiparous brittle-star populations, the adaptive significance of clonal growth may be related to an increase in the overall fitness of dispersed clones (genets), compared to individuals of strictly sexual counterparts, through greater genotypespecific biomass and, hence, fecundity.Contribution No. 476 of the Discovery Bay Marine Laboratory of the University of the West Indies     

Genetic structure of fissiparous populations of Holothuria (Halodeima) atra on the Great Barrier Reef

Allozyme variation at five polymorphic loci was surveyed in a total of 311 individuals of the sea cucumber Holothuria (Halodeima) atra (Jäger, 1833) collected from two nearshore and two midshelf populations in the Great Barrier Reef in November 1996. Strong deviations in genotype frequencies from those expected under Hardy–Weinberg equilibrium, particularly a large number of heterozygote excesses, confirmed the occurrence of asexual reproduction. Females and males differed significantly in genotype frequencies as a result of differences in either the amount of fission in each sex or in the mortality of asexually produced recruits. The estimated maximum sexual input (number of sexually produced individuals: sample size = N*:N_i) to the two nearshore reefs (38 to 67%) was low relative to that to the two midshelf reefs (74 to 87%). The three ratios and G _o :G _e , N _go :N _i , N*:N _i , (where G _o = observed genotypic diversity, G _e = expected genotypic diversity, N _go = number of genotypes) considered to be indicators of the extent of asexual reproduction, showed a consistent trend in the degree of asexual reproduction similar to that derived from the number of regenerating individuals observed in the populations for which data were available. F-statistic analyses of clonal gene frequencies demonstrated that all populations received sexual recruits from the same gene pool. There was evidence of restricted sexual recruitment to the Fantome population, suggesting that asexual reproduction was dominant only in areas where sexual recruitment was limited by other factors.     

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     

Demography of the intertidal brown alga Pelvetia fastigiata in Southern California,USA

The demography of Pelvetia fastigiata was studied from 1973 through 1977 in the intertidal zone near La Jolla, California, USA. Populations were usually small and relatively stable as new recruits replaced older individuals that were lost; however, in 1977 high recruitment substantially increased total abundances. The survivorship of recruits was similar regardless of the season or year of recruitment. Mortality rates decreased with age, and the mean life expectancy continued to increase for at least 1 yr. Losses of juveniles were high; 50% lived 80 d and only 9% survived to the age of first reproduction (about 1.5 yr). Plants present at the start of the study had higher survivorship rates than new recruits, 19 and 7%, respectively, living more than 3 yr. Individuals within aggregations of large plants lived longer than most dispersed individuals, and aggregations remained intact throughout the study in spite of low recruitment. Established individuals had low mortality rates and populations probably relied on these persistent individuals for reproduction.     

Fissiparity and population genetics of Coscinasterias calamaria

The effect of asexual reproduction on the population genetics of the fissiparous seastar Coscinasterias calamaria was examined at Rottnest Island and the adjacent mainland, Western Australia, in 1985. Field samples and laboratory observations on growth rate and regeneration showed that fission is common at all 14 sites examined. Electrophoretic analysis of six polymorphic enzymes revealed low genotypic diversity and strong genetic disequilibrium at each site, confirming the highly clonal structure of local populations. Striking variation in clonal composition over distances as short as 50 m emphasizes the very localized subdivision of these populations. In the combined sample from all sites, however, the expected range of multilocus genotypes was found in the proportions expected from random mixing of sexually produced larvae, confirming that clonal diversity results from sexual reproduction. Local genotypic diversity was not correlated with fission, emphasizing the difficulty of determining the short-term roles of asexual and larval recruitment in the maintenance of populations. Subtidal populations, however, appear to have both lower incidence of fission and lower larval recruitment than do intertidal populations.     

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     

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.     

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     

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.     

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     

Divide or broadcast: Interrelation of asexual and sexual reproduction in a population of the fissiparous hermaphroditic seastar Nepanthia belcheri (Asteroidea: Asterinidae)

Asexual and sexual reproduction were studied in an intertidal population of Nepanthia belcheri (Perrier) at Townsville, Queensland, Australia, by regular sampling over a year (March 1976-March 1977) and by histological analysis of gonads. Fission reached a peak in early winter (April–June), when about 45% of the population showed evidence of recent fission. Propensity for fission was unrelated to longest arm length. Seven-armed seastars predominated in the population and these underwent fission in two stages to produce one 3-armed fragment and two 2-armed fragments. Fission planes were not related to numbers or positions of madreporites. Hermaphroditism was a normal sexual condition in the population. Almost all gonads contained oocytes, but some gonads functioned as ovaries (without spermatogenic tissue) while others functioned as testes. Seastars with mature ovaries were significantly larger than those with mature testes, indicating protandry, as in other hermaphroditic asteroids. However, fission complicates the pattern of gonad development by causing regression or retardation of gonads and by apparently having a masculinizing effect, so that ovaries may change to testes in fission products. There was a period of sexual reproduction in early summer (October–November). This followed the period of intense fission and regeneration, and a population change from predominately functional females to males. Consequently there was an extreme imbalance against mature females at sexual reproduction, further reducing potential fecundity. Thus, sexual reproduction was very subordinate to fission as the means of recruitment. The 450 m diam eggs probably give rise to pelagic lecithotrophic development and, if this is the case, N. belcheri retains the advantage of complementing reliable recruitment from fission with a dispersive phase. The combination of fission and hermaphroditism is particularly advantageous for a very sparse dispersal of larvae, as a functionally dioecious population may develop from one larva settling in a new locality.