A molecular phylogenetic analysis of reproductive trait evolution in the soft coral genus Alcyonium

The soft coral genus Alcyonium is among the most reproductively diverse invertebrate taxa known: The genus includes species that vary both in mode of reproduction (including broadcast spawners, internal brooders, and external brooders) and sexual expression (gonochores, hermaphrodites, and a unisexual parthenogen). Such diversity offers a unique opportunity to examine associations between reproductive and morphological traits in a phylogenetic context. We used an approximately 900-bp sequence of the nuclear ribosomal gene complex spanning the internal transcribed spacer (ITS) regions to construct a molecular phylogeny for 14 European and North American species of Alcyonium onto which we mapped the known distribution of reproductive and morphological traits. The phylogeny suggests that hermaphroditism or parthenogenesis has evolved independently at least twice in this genus, and always in internally brooding species. Broadcast spawning and external brooding only occur in species with large colony size, whereas all species with small colony size brood their larvae internally. Internal brooding and small size appear to be ancestral in this genus; if this is the case, an association between broadcast spawning and large colony size has evolved independently in at least two clades. This tendency of small adults to brood their larvae while large adults broadcast spawn them into the plankton has been observed in a variety of solitary invertebrate taxa, but to date has not been documented in any other colonial invertebrates. Moreoever, it has been suggested that organisms with a colonial growth form should not experience the allometric constraints on brood space that have been proposed to explain the association between adult size and mode of reproduction in solitary organisms. Unlike many other colonial groups, however, module (polyp) size is strongly correlated with colony size in Alcyonium, and constraints on brooding may be imposed by module, rather than colony, allometry. The very close genetic relationship (< 1% sequence divergence) and shared polymorphisms among A. digitatum (a large, gonochoric broadcast spawner), A. siderium, and A. sp. A (intermediate-sized and small hermaphroditic, internal brooders) suggest that evolutionary transitions between broadcast spawning and brooding and between gonochorism and hermaphroditism can occur easily and rapidly in this group.     

Correlated evolution of sex and reproductive mode in corals (Anthozoa: Scleractinia)

Sexuality and reproductive mode are two fundamental life-history traits that exhibit largely unexplained macroevolutionary patterns among the major groups of multicellular organisms. For example, the cnidarian class Anthozoa (corals and anemones) is mainly comprised of gonochoric (separate sex) brooders or spawners, while one order, Scleractinia (skeleton-forming corals), appears to be mostly hermaphroditic spawners. Here, using the most complete phylogeny of scleractinians, we reconstruct how evolutionary transitions between sexual systems (gonochorism versus hermaphrodism) and reproductive modes (brooding versus spawning) have generated large-scale taxonomic patterns in these characters. Hermaphrodites have independently evolved in three large, distantly related lineages consisting of mostly reef-building species. Reproductive mode in corals has evolved at twice the rate of sexuality, while the evolution of sexuality has been heavily biased: gonochorism is over 100 times more likely to be lost than gained, and can only be acquired by brooders. This circuitous evolutionary pathway accounts for the prevalence of hermaphroditic spawners among reef-forming scleractinians, despite their ancient gonochoric heritage.     

Simultaneous hermaphroditism; cost and benefit

Current theories explain simultaneous hermaphroditism by the advantage it gives to organisms which are widely dispersed or sluggish, resulting in a low frequency of reproductive contacts. It is difficult to see why hermaphroditism is not more widespread unless there is some counterbalancing disadvantage.It is suggested that hermaphrodites suffer an energetic cost because they maintain two reproductive systems and a cost due to the reduced number or viability of offspring which may result from accidental self-fertilization. These costs will result in a disadvantage to hermaphroditism (compared to gonochorism) when reproductive contacts are frequent. However, even in widely dispersed or sluggish organisms behavioural mechanisms may exist which increase the frequency of reproductive contacts, favouring gonochorism instead of hermaphroditism.It is argued that externally fertilizing species should as a rule be gonochoric and that species which brood their young may often be hermaphroditic. Hermaphroditism in species which form permanent male/female pairs in the breeding season could result in more zygotes being produced. However, where parental care of the young is important, it is suggested that gonochorism and sexual dimorphism may result in more progeny being reared.     

Inferring the ancestral sexuality and reproductive condition in sponges (Porifera)

Considerable diversity abounds among sponges with respect to reproductive and developmental biology. Their ancestral sexual mode (gonochorism vs. hermaphroditism) and reproductive condition (oviparity vs. viviparity) however remain unclear, and these traits appear to have undergone correlated evolution in the phylum. To infer ancestral traits and investigate this putative correlation, we used DNA sequence data from two loci (18S ribosomal RNA and cytochrome c oxidase subunit I) to explore the phylogenetic relationships of 62 sponges whose reproductive traits have been previously documented. Although the inferred tree topologies, using the limited data available, favoured paraphyly of sponges, we also investigated ancestral character‐state reconstruction on a phylogeny with constrained sponge monophyly. Both parsimony‐ and likelihood‐based ancestral state reconstructions indicate that viviparity (brooding) was the likely reproductive mode of the ancestral sponge. Hermaphroditism is favoured over gonochorism as the sexual condition of the sponge ancestor under parsimony, but the reconstruction is ambiguous under likelihood, rendering the ancestry of sexuality unresolved in our study. These results are insensitive to the constraint of sponge monophyly when tracing the reproductive characters using parsimony methods. However, the maximum likelihood analysis of the monophyletic hypothetical tree rendered gonochorism as ancestral for the phylum. A test of trait correlation unambiguously favours the concerted evolution of sexuality and reproductive mode in sponges (hermaphroditism/viviparity, gonochorism/oviparity). Although testing ecological hypotheses for the pattern of sponge reproduction is beyond the scope of our analyses, we postulate that certain physiological constrains might be key causes for the correlation of reproductive characters.     

The evolution of mating systems in a group of brooding sea anemones (Epiactis)

Summary

The sea anemone genus Epiactis provides an unusually good opportunity to study the evolution of brooding and mating systems. The four Epiactis species on the Pacific coast of North America all brood their offspring up to the juvenile stage, but each has a different combination of internal vs. external brooding and gonochory vs. simultaneous or gynodioecious hermaphroditism. Two of the four species (E. prolifera and E. lisbethae) were indistinguishable with allozymes (20 loci), but could be differentiated using multilocus DNA fingerprinting. Phylogenetic analyses of the allozyme data by distance and parsimony methods using three outgroups suggest that the four nominal Epiactis species are polyphyletic, with the two internal brooders evolving independently of the two external brooders. This topology does not allow inferences about the evolutionary order of hermaphroditism, dioecy and gynodioecy. Separate sexes and obligate outcrossing are of ten believed to be ancestral, with hermaphroditism and the potential for self-fertilization being favored in taxa where restricted dispersal promotes inbreeding. Previous studies of population genetic structure in these Epiactis species is consistent with this hypothesis, as even the cross-fertile species were highly inbred.     

Reproductive ecology of Caribbean reef corals

The last decade has seen a resurgence of interest in the processes of sexual reproduction by scleractinian reef corals. Earlier investigations had focused fortuitously on brooding (planulating) species, which resulted in the general misconception that brooding was the main form of larval development of reef corals. More recent work on Indo-Pacific species has shown broadcast spawning and short annual reproductive periods to predominate. This report presents the reproductive patterns of eleven Caribbean coral species and attempts to explain the adaptive features and selective pressures that have led to the evolution of the four reproductive patterns described to date: (a) hermaphroditic broadcasters; (b) gonochoric broadcasters; (c) hermaphroditic broadcasters; (b) gonochoric brooders. Both (a) and (b) correlate with large colony size and short annual spawning periods; and (c) and (d) correlate with small colony size, multiple planulating cycles per year, and occupation of unstable habitats. Selection for outcrossing between long-lived individuals is proposed as the reason for gonochorism and for synchronous spawning of hermaphroditic broadcasters, and also for the large amount of sperm produced by hermaphroditic brooders. Selection for high rates of local recruitment is proposed as the force behind the evolution of brooding by species inhabiting unstable habitats and suffering high rates of adult mortality.     

The influence of season and the tidal cycle on division of labour by the radula during feeding in the estuarine brooding gastropod <Emphasis Type="Italic">Crepipatella dilatata</Emphasis> (Calyptraeidae)

The brooding gastropod Crepipatella dilatata can feed by scraping the substrate with the radula and by suspension-feeding, which also requires use of the radula. There is a “division of labour” for the radula among three discrete tasks associated with feeding: (1) removing mucous balls from the food pouch; (2) transferring the mucous cord from the neck channel to the mouth (both components of suspension-feeding); (3) scraping the substrate. We hypothesised that the proportion of time used for each feeding activity varies according to environmental conditions. Total radular activity in females was greatest at high tide and in summer. The rate of radular extrusion for ingesting the mucous cord varied seasonally and between brooding and non-brooding females. Non-brooding females exhibited higher rates of radular extrusion for ingesting the mucous cord and for scraping the substrate than did brooders. In females, radular activity in removing the mucous ball from the food pouch was strongly influenced by the tidal cycle during winter, reaching minimum values at low tide. Differences were recorded in substrate scraping among seasons and within tidal cycles, and among males, brooding females and non-brooding females. Brooding females displayed less rasping than non-brooders, since the area available for grazing was restricted by the egg mass. Throughout the year, including low salinity periods, males allocated a greater proportion of total radular activity to rasping than to removing the mucous ball or ingesting the mucous cord. The feeding behaviour of both males and females is modulated by salinity, but the principal determinants of radular activity are the mode of reproduction (brooding in females) and, in males, motility.     

Phylogenetic analysis of the Sphaeriidae (Mollusca: Bivalvia) based on partial mitochondrial 16s rDNA gene sequences

Abstract. We have constructed molecular phylogenetic trees for members of the Sphaeriidae in order to test proposed generic level relationships, and to reconstruct the evolutionary pattern of parental care, in this exclusively freshwater family of heterodont bivalves. An ∼480 nucleotide fragment of the mitochondrial large ribosomal subunit (16s rDNA) was sequenced for 4 corbiculid outgroups in addition to 19 sphaeriid ingroup taxa. Ingroup species were obtained from North and South America, Europe, and Australasia and included representatives of the main sphaeriid genera. Our analyses support four primary conclusions: 1) the Sphaeriinae are robustly monophyletic with respect to Eupera platensis ; 2) the genus Pisidium is paraphyletic and P. sterkianum is sister to the 17 other sphaeriine taxa in our dataset; 3) synchronous brooding is the ancestral reproductive pattern in the Sphaeriinae; 4) the sequential brooders form a clade in which Musculium taxa are monophyletic and nested among lineages of Sphueriunz. Our gene trees reveal an evolutionary progression in parental care complexity from the relatively simple pattern in the Euperinae, to the origin of brood sacs and of extraoogonial embryonic nutrition in the common ancestor of the Sphaeriinae, and ultimately to the development of sequential brooding in Sphaerium/Musculium taxa.     

Sperm competition in fish: `bourgeois' males and parasitic spawning

Fish exhibit an enormous variety of reproductive patterns. There is external and internal fertilization, simultaneous and sequential hermaphroditism as well as gonochorism, and an extremely widespread occurrence of parasitic reproductive behaviour among males. In most fish species there is a great size range of reproductive males, setting the stage for divergent, intraspecific reproductive patterns and an unparalleled concentration of alternative male reproductive phenotypes. Recent theoretical, empirical and comparative evidence suggests that adaptations to sperm competition in fish might be responsible for some of the most intriguing examples of reproductive design known.     

Molecular phylogeny of the model annelid Ophryotrocha.

Annelids of the genus Ophryotrocha are small opportunistic worms commonly found in polluted and nutrient-rich habitats such as harbors. Within this small group of about 40 described taxa a large variety of reproductive strategies are found, ranging from gonochoristic broadcast spawners to sequential hermaphroditic brooders. Many of the species have a short generation time and are easily maintained as laboratory cultures. Thus they have become a popular system for exploring a variety of biological questions including developmental genetics, ethology, and sexual selection. Despite considerable behavioral, reproductive, and karyological studies, a phylogenetic framework is lacking because most taxa are morphologically similar. In this study we use 16S mitochondrial gene sequence data to infer the phylogeny of Ophryotrocha strains commonly used in the laboratory. The resulting mtDNA topologies are generally well resolved and support a genetic split between hermaphroditic and gonochoristic species. Although the ancestral state could not be unambiguously identified, a change in reproductive strategy (i.e., hermaphroditism and gonochorism) occurred once within Ophryotrocha. Additionally, we show that sequential hermaphroditism evolved from a simultaneous hermaphroditic ancestor, and that characters previously used in phylogenetic reconstruction (i.e., jaw morphology and shape of egg mass) are homoplasic within the group.     

Broadcasting fables: Is external fertilization really primitive? Sex,size, and larvae in sabellid polychaetes

Traditionally, broadcast spawning and planktonic larvae have been considered the plesiomorphic ‘ground plan’ for the Polychaeta and other metazoan groups. To assess whether this reproductive mode is in fact ‘primitive’, the study of monophyletic groups with various reproductive modes should be informative. A large range of body sizes would allow testing the ideas that aspects of reproductive mode may be functionally constrained. The family Sabellidac is one such group, with sexual reproductive modes ranging from broadcast spawning to intratubular brooding to ovovivi-parity, and a body size range over more than five orders of magnitude. Sabellids have previously been the subject of detailed cladistic analyses (Fitzhugh 1989, 1991); here we introduce several new characters based on morphology of reproductive structures. Larval development in four brooding sabellid species is also described with the aim of introducing new characters for future systematic analyses. Our cladistic analysis of sabellid genera suggests that gonochorism and brooding of direct-developing larvae are plesiomorphic in the Sabellidae, with external fertilization and swimming larvae limited to apomorphie clades in the subfamily Sabellinae. The presence of sperm with elongate heads may be correlated with the presence of intratubular brooding, though an adequate causal explanation for this relationship can not yet be presented. The concept that ‘modified’ sperm must be derived from ‘primitive’ sperm is shown to be false, with ‘modified’ sperm being plesiomorphic for the Sabellidae, from which ‘primitive’ sperm is derived in apomorphic Sabellinae. All sabellids have lecithotrophic development and appear to be phylogenetically constrained in this regard. Data gathered on body size and reproductive variables in the Sabellidac suggests the following (when phylogenetic effects are not controlled): (1) egg number and total egg volume are significantly correlated with body size, with small animals having fewer, larger eggs than large animals; (2) individual egg volume is not correlated with body size; (3) reproductive mode is significantly correlated with body size; intratubular brooders tend to be small-bodied, whereas broadcast spawners are large. However when the effect of body size is controlled for, then (4) egg number, egg volume and total egg volume all vary significantly with reproductive mode. Broadcast spawners expel a large number of small eggs for a high total egg volurne. Intratubular brooders have a few relatively large eggs for a small total egg volume. When statistics arc performed using phylogenetically independent contrasts there is a significant correlation between total egg volume and body size but not for egg number and body size. The effect of non-independence (due to phylogeny) of our data needs to be more fully controlled in future analyses but methods of incorporating continuous data into cladistic analyses should also be investigated. We show that some predictions can be made about reproductive mode based on body size but ad hoc patterns of reproductive character-state transformation should not be made independent of empirical hypotheses of phylogenetic relationship. Further studies of this kind throughout the Annelida are needed to determine the plesiomorphic reproductive mode for the phylum.     

Permeability barrier in the mantle epithelium lining the testis in the apple-snail Pomacea canaliculata (Gastropoda: Ampullariidae)

Intercellular junctions are studied in the epithelium lining the testis of the freshwater snail Pomacea canaliculata by conventional staining and lanthanum tracer techniques. The junctional complex consists of belt desmosomes and septate junctions. Septate junctions are of the pleated-sheet type and they are constantly associated with mitochondria. Gap and tight junctions appear to be absent. These septate junctions seem to be the structural correlate of an epithelial permeability barrier that separate the testis from the extrapallial space where the shell elements are deposited. These junctions may contribute to a functional barrier in the male gonad of Pomacea canaliculata. The results indicate that freshwater prosobranchs have junctional structures very close to those found in other molluscs.     

MOLECULAR PHYLOGENETIC ANALYSIS OF LIFE-HISTORY EVOLUTION IN ASTERINID STARFISH

We analyzed phylogenetic relationships among 12 nominal species of starfish in the genera Patiriella and Asterina (Order Valvatida, Family Asterinidae), based on complete sequences for a mitochondrial protein coding gene (cytochrome oxidase subunit I) and five mitochondrial transfer RNA genes (alanine, leucine, asparagine, glutamine, and proline) (1923 bp total). The resulting phylogeny was used to test a series of hypotheses about the evolution of life-history traits. (1) A complex, feeding, planktonic larva is probably ancestral for these starfish, but this is not the most parsimonious reconstruction of ancestral larval states. (2) The feeding larval form was lost at least four times among these species, and three of these losses occurred among members of a single clade. (3) Small adult size evolved before both cases of hermaphroditism and viviparous brooding, but viviparity was not always preceded by an intermediate form of external brooding. (4) An ordered transformation series from feeding planktonic development to viviparous brooding has been predicted for starfish, but we could not find an example of this transformation series. (5) Viviparity evolved recently (< 2 Mya). (6) Both species selection and transformation of lineages may have contributed to the accumulation of species with nonfeeding development among these starfish. (7) Neither Asterina nor Patiriella are monophyletic genera. Larval forms and life-history traits of these starfish have evolved freely under no obvious constraints, contrary to the widely assumed evolutionary conservatism of early development.     

Hermaphroditism in tardigrades

Evidence is presented to show the existence of hermaphroditism in tardigrades, a phenomenon hitherto unknown for this phylum: specimens were obtained containing male and female germ cells in maturation in the same gonad. Hermaphroditic animals have been found in a few species of Isohypsibius; in many species of other genera and also of Isohypsibius, there is gonochorism.     

Hermaphroditism as a reproductive strategy for metazoans; some correlated benefits

Abstract

The concept of hermaphroditism is examined, the terminology is reviewed, and several new terms are proposed: EUHERMAPHRODITISM for the occurrence together of eggs and sperm in the individual that produced them; TEMPORARY HERMAPHRODITISM for the few instances where euhermaphroditism is a brief transitional phase between sexes in protandry and protogyny, and opsiautogamy to designate the condition where sperm saved from an early male phase are used to fertilise eggs produced in a later female phase. An animal that functions successively in the different sex roles is considered to be quite distinct from one that functions as both male and female simultaneously. Further, the circumstances in which these different sexual modes are appropriate and the advantages that accrue from them are quite different. For this reason protandry, protogyny, and consecutive sexuality are not considered to be part of hermaphroditism, but rather aspects of labile gonochorism.

A major idea advanced in this paper is that hermaphroditism is of primary importance as a reproductive strategy of very small metazoans. In such animals, whose eggs are so disproportionately large that very few can be produced, hermaphroditism ensures that through having the maximal number of females the maximal number of eggs are produced and fertilised economically. For very small hermaphrodites the mate-finding difficulties associated with small size and limited motility are reduced by converting each meeting into a potential mating. Further, in hermaphroditism not only is the frequency of mating maximised. but through sperm storage the effects of matings last for a long time. and through multiple reciprocal copulations genetically diverse progenies are produced. The effective size of the gene pool is maximised, and the risk of genetic drift in sex ratio in small populations is avoided. Very small animals do not have the energetic resources to produce large numbers of large, yolky eggs, nor do they have the space in which to store them before release. Thus tiny hermaphrodites tend to produce a few eggs at a time, but over extended or continuous breeding seasons. The effects of their reduced fecundity may be further offset by precise methods of fertilisation, brooding, and commonly the suppression of pelagic phases, thus minimising losses from larval predation and overdispersal.

It is also argued that hermaphroditism may have been advantageous because of features often associated with it. Copulation with protected sperm transfer may have been especially important, not only because of economies in gamete production and use, but also in the exploitation of new habitats. In freshwater hermanhrodites copulatory or spermatophore exchange habits protect sperm from osmotic death: in endoparasites copulation protects sperm from digestive enzymes and osmotic stress in the host’s gut, and from antibodies and phagocytosis in the bloodstream.     

Gonochorism vs. hermaphroditism: relationship between life history and fitness in three species of Ophryotrocha (Polychaeta: Dorvilleidae) with different forms of sexuality

1. The relationships between life history, fitness and sexuality, together with their ecological and evolutionary significance, has been analysed comparing the main life-history traits and demography in three closely related species belonging to the genus Ophryotrocha. The species are: the gonochoristic O. labronica, the simultaneous hermaphrodite O. diadema and the protandrous hermaphrodite O. puerilis. 2. Survivorship and reproductive data were collected weekly and were used to construct life tables and population projection matrices for each species and compare life-history characteristics. Elasticity, life-table response and decomposition analyses were performed to examine the relative contribution of fecundity and survivorship to differences in lamda between species. 3. The gonochoristic and hermaphroditic species differ in all the main life-history parameters and also in demographic characteristics. In particular the value of lamda, used commonly to express fitness, is markedly higher in the gonochoristic species while in terms of fitness simultaneous and sequential hermaphroditism are very similar. In the genus Ophryotrocha gonochorism currently represents the most widespread condition, being characteristic of the majority of the known species in the genus. 4. Given the demographic advantage ensured by gonochorism, it remains be understood why some species have retained simultaneous hermaphroditism and one has evolved a sequential type hermaphroditism; the most probable hypothesis is correlated with the density of the species in natural habitats.     

Genetic data,reproduction season and reproductive strategy data support the existence of biological species in Ophioderma longicauda

Cryptic species are numerous in the marine environment. The brittle star Ophioderma longicauda is composed of six mitochondrial lineages, encompassing brooders, which form a monophyletic group, and broadcasters, from which the brooders are derived. To clarify the species limits within O. longicauda, we compared the reproductive status of the sympatric lineages L1 and L3 (defined after sequencing a portion of the mitochondrial gene COI) during the month of May in Greece. In addition, we genotyped a nuclear marker, intron i51. Each L3 female was brooding, whereas all L1 specimens displayed full gonads, suggesting temporal pre-zygotic isolation between brooders and broadcasters. Statistical differences were found among lineages in morphology and bathymetric distribution. Finally, the intron i51 was polymorphic in L1 (60 individuals), but monomorphic in L3 (109 individuals), confirming the absence of gene flow between brooders and broadcasters. In conclusion, the broadcasting lineage L1 and the brooding lineage L3 are different biological species.     

Genotypic diversity and gene flow in brooding and spawning corals along the Great Barrier Reef, Australia

Marine organisms exhibit great variation in reproductive modes, larval types, and other life-history traits that may have major evolutionary consequences. We measured local and regional patterns of genetic variation in corals along Australia's Great Barrier Reef to determine the relative contributions of sexual and asexual reproduction to recruitment and to infer levels of gene flow both locally (among adjacent sites, < 5 km apart) and regionally (among reefs separated by 500-1,200 km). We selected five common brooding species (Acropora cuneata, A. palifera, Pocillopora damicornis, Seriatopora hystrix, and Stylophora pistillata) and four broadcast spawners (Acropora hyacinthus, A. cytherea, A. millepora, and A. valida), which encompassed a wide range of larval types and potential dispersal capabilities. We found substantial genotypic diversity at local scales in six of the nine species (four brooders, two spawners). For these six, each local population displayed approximately the levels of multilocus genotypic diversity (Go) expected for outcrossed sexual reproduction (mean values of Go:Ge ranged from 0.85 to 1.02), although consistent single-locus heterozygous deficits indicate that inbreeding occurs at the scale of whole reefs. The remaining three species, the brooder S. hystrix and the spawners A. valida and A. millepora displayed significantly less multilocus genotypic diversity (Go) than was expected for outcrossed sexual reproduction (Ge) within each of several sites. Acropora valida and A. millepora showed evidence of extensive localized asexual replication: (1) a small number of multilocus (clonal) genotypes were numerically dominant within some sites (Go:Ge values were as low as 0.17 and 0.20): (2) single-locus genotype frequencies were characterized by both excesses and deficits of heterozygotes (cf. Hardy-Weinberg expectations), and (3) significant linkage disequilibria occurred. For the brooding S. hystrix Go:Ge values were also low within each of four sites (x = 0.48). However, this result most likely reflects the highly restricted dispersal of gametes or larvae, because levels of genetic variation among sites within reefs were extremely high (FSR = 0.28). For all species, we detected considerable genetic subdivision among sites within each reef (high FSR-values), and we infer that larval dispersal is surprisingly limited (i.e., Nem among sites ranging from 0.6 to 3.3 migrants per generation), even in species that have relatively long planktonic durations. Nevertheless, our estimates of allelic variation among reefs (FRT) also imply that for all four broadcast spawning species and three of the brooders, larval dispersal is sufficient to maintain moderate to high levels of gene flow along the entire Great Barrier Reef (i.e., Nem among reefs ranged from 5 to 31). In contrast, widespread populations of S. hystrix and S. pistilata (the two remaining brooders) are relatively weakly connected (Nem among reefs was 1.4 and 2.5, respectively). We conclude that most recruitment by corals is very local, particularly in brooders, but that enough propagules are widely dispersed to ensure that both broadcast spawning and brooding species form vast effectively panmictic populations on the Great Barrier Reef.     

Sexual development and sexuality in the Nassau grouper

Hermaphroditic sexual patterns are widespread among teleosts and are especially characteristic of certain perciform families. Among the most diverse groups of hermaphroditic species are the groupers, hamlets and sea basses of the family Serranidae. Like other groupers, the epinepheline serranid, Epinephelus striatus , the Nassau grouper, has long been assumed to exhibit monandric protogyny (a form of hermaphroditism in which all males derive from adult females by sex change). Histological and demographic data, however, indicate that the sexual pattern of this species is not monandric, and that, unlike other group species, the juveniles pass through a bisexual stage of gonadal development. Sexuality in the Nassau grouper is essentially gonochoristic, with potential for sex change. The combination of bisexuality and gonochorism is previously undescribed in the serranids.     

A study on snail location by Fasciola hepatica using radioisotopically labelled miracidia.

A number of common freshwater molluscs were exposed to Fasciola hepatica miracidia labelled in vivo with radioselenium. Radioactivity was rapidly incorporated in all lymnaeid species tested (L. stagnalis, L. tomentosa and L. truncatula) whereas no radioactivity could be demonstrated in non-lymnaeid pulmonate species (Aplexa hypnorum, Physa acuta, Physa fontinalis, Anisus vortex, Gyraulus albus and Planorbis planorbis) or in the prosobranchs Valvata cristata and Valvata piscinalis. However, the prosobranchs Bithynia leachii, Bithynia tentaculata and Potamopyrgus jenkinsi showed a slow but significant uptake of radioactivity which was presumably explained by uptake of labelled miracidia through a 'filter-feeding' mechanism. Sphaerium corneum, a true filter-feeder, showed a similar course of uptake of radioactivity.