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.     

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.     

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.     

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.     

Spermatozoan ultrastructure of scleractinian corals from the eastern Pacific: Pocilloporidae and Agariciidae

Spermatozoa ofPocillopora damicornis, Pocillopora elegans (Astrocoeniina, Pocilloporidae) andPavona gigantea (Fungiina, Agariciidae) from the eastern Pacific (Isla del Caño, Costa Rica) were examined using transmission electron microscopy. The hermaphroditic pocilloporidsP. damicornis andP. elegans are spermiomorphologically very similar to hermaphroditic acroporids, being characterized by bullet-shaped nuclei and elongated mitochondria. Such traits have not been found in other families. Thus, the suborder Astrocoeniina, including pocilloporids and acroporids, can clearly be distinguished from other scleractinian suborders. This separation underlines the isolated position of the Astrocoeniina within the order Scleractinia following the evolutionary scheme of Wells. A conical sperm type, known from gonochoric species (sexes separate) from all families except Acroporidae and Pocilloporidae, was found in the gonochoric agariciidPavona gigantea. This supports previous findings that gonochoric corals share a unique and common sperm structure regardless of which family they belong to. However, no gonochoric Astrocoeniina have ever been examined. Hence, the question whether the sperm type common to gonochorists is also represented in Astrocoeniina, which would undermine the ultrastructural distinction of Astrocoeniina and other suborders seen among hermaphrodites, as well as the systematic value of sperm structure within scleractinian corals, remains open.     

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.     

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.     

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.     

Hermaphroditism in molluscs

The functional significance of hermaphroditism is, as yet, not well understood. This study attempts to examine the relevance of some theories on this topic to reality, by reviewing the literature on hermaphroditism among molluscs. The study reveals that about 40% of the 5600 mollusc genera are either simultaneous or sequential hermaphrodites. Hermaphroditism occurs in 100% of the Solenogastres, 99% of the Opisthobranchia, 100% of the Pulmonata, 3% of the Prosobranchia and 9% of the Bivalvia. Sequential hermaphroditism, whatever its theoretical advantages over gonochorism, is not common among molluscs. Simultaneous hermaphroditism is common among the Euthyneura. Over 80% of the hermaphroditic genera belong to the Euthyneura (about 2000 recent genera), and over 99% of the Euthyneura are hermaphrodites. The almost ubiquitous occurrence of hermaphroditism throughout the Euthyneurans, coupled with this group's very widespread dispersal in marine, terrestrial and freshwater habitats (and corresponding exposure to a very wide range of predators, competitors or parasites); and also its wide range of population densities, suggests that in this group hermaphroditism is not an adaptive response to selective forces of the environment and it may well be a phylogenetic constraint. A few stylommatophorans have broken this constraint by possessing genitalia in which the male system is absent (aphally) or extremely reduced (hemiphally). They reproduce by self-fertilization. In theory this is advantageous in that these species save the cost of developing two sets of reproductive apparatus. In reality there is no record of aphally occurring throughout an entire species, as one may expect if this saving of cost were so overwhelmingly advantageous. There is also no record of stylommatophoran individuals developing only the male system, as one may expect if there was any evolutionary trend from hermaphroditism towards gonochorism. Simultaneous hermaphroditism offers a (theoretical) option to selling. Selfing by self-fertilization is widespread among freshwater basommatophorans, amphibious stylommatophorans and freshwater bivalves. Selfing by parthenogenesis is widespread among freshwater prosobranchs. Altogether, of 48 genera which self, 60% are dwellers of freshwater. Certain freshwater habitats may perhaps be less stable than most marine or terrestrial ones are. Beyond the Euthyneura, hermaphroditism is common among parasitic, deep sea and Cnidaria-eating molluscs. Perhaps these three associations should be slightly rearranged and combined into a single generalization: that hermaphroditism (beyond the Euthyneura) is widespread among molluscs that live in a close and permanent intimacy with live marine invertebrates, whether as parasites, commensals or predators. Gonochorism but with dwarf males is common among the Eulimoidea, which are parasites, and Galeommatoidea, which are commensals. This situation of gonochorism, in which there is a very close association between members of the two sexes, is functionally a hermaphroditic type. Animals that can manipulate the physiology and endocrinology of their hosts may also have the tendency and ability to manipulate the size of their own males. It may well be that the female not only determines the size of the male but also his sex, in which case dwarf males should be considered as a case of environmental sex determination. The theoretical possibility that hermaphroditism may evolve in brooding animals was examined in gastropods. Among the (predominantly gonochoric) prosobranchs brooding is recorded in 85 genera (4%), only two of which are hermaphrodites. Among the (predominantly hermaphrodite) heterobranchs it is recorded in only 95 odd genera (4%). When all gastropod genera are considered together, the frequency of hermaphroditism among the brooders is only very slightly and insignificantly higher than in the non-brooders. When only the prosobranchs are considered, the frequency of hermaphroditism among the brooders is actually lower than among non-brooders. The theory that hermaphroditism evolves in brooding animals thus appears not to be relevant to the vast majority of gastropods. Brooding is widespread among freshwater prosobranchs (48% of the brooding genera). In this habitat, brooding may perhaps be an adaptation to protect the developing embryo against osmotic stresses.     

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.     

Sperm morphology of scleractinians from the Caribbean

This study compares the sperm morphology of 2 hermaphroditic (Diploria strigosa, Montastrea annularis) and 2 gonochoric (Siderastrea siderea, Dendrogyra cylindrus) Caribbean reef corals. The spermatozoa of the gonochoric species are morphologically more specialized than those of the hermaphrodites. The cytological differences are apparent in the formation of a Spitzenkörper and pro-acrosomal vesicles, a mitochondrial mass (Nebenkern) and a large lipid vesicle in the gonochorists studied. Harrison (1985) first demonstrated that sperm morphology is correlated with sexual reproductive pattern and systematic position in scleractinian corals, and this is supported by the results of this study. The possible functional and phylogenetic bases of sperm morphology are discussed and a brief outline is given of further studies required.     

Analysis of the mitochondrial 12S rRNA gene supports a two-clade hypothesis of the evolutionary history of scleractinian corals

Scleractinian corals have long been assumed to be a monophyletic group characterized by the possession of an aragonite skeleton. Analyses of skeletal morphology and molecular data have shown conflicting patterns of suborder and family relationships of scleractinian corals, because molecular data suggest that the scleractinian skeleton could have evolved as many as four times. Here we describe patterns of molecular evolution in a segment of the mitochondrial (mt) 12S ribosomal RNA gene from 28 species of scleractinian corals and use this gene to infer the evolutionary history of scleractinians. We show that the sequences obtained fall into two distinct clades, defined by PCR product length. Base composition among taxa did not differ significantly when the two clades were considered separately or as a single group. Overall, transition substitutions accumulated more quickly relative to transversion substitutions within both clades. Spatial patterns of substitutions along the 12S rRNA gene and likelihood ratio tests of divergence rates both indicate that the 12S rRNA gene of each clade evolved under different constraints. Phylogenetic analyses using mt 12S rRNA gene data do not support the current view of scleractinian phylogeny based upon skeletal morphology and fossil records. Rather, the two-clade hypothesis derived from the mt 16S ribosomal gene is supported.     

Hermaphroditic spawning by the gonochoric pillar coral Dendrogyra cylindrus

Coral Reefs - Scleractinian corals typically reproduce sexually as either gonochoric (separate male and female) or hermaphroditic (producing both eggs and sperm) colonies. The Caribbean pillar...     

Contributions of growth, stasis, and reproduction to fitness in brooding and broadcast spawning marine bivalves

Reproductive modes in marine invertebrates can be generally grouped into two types: those brooding larvae and those broadcast-spawning gametes into the water. We asked if these different life-history strategies differ based on how contribution to fitness is partitioned between growth, stasis, and reproduction. To investigate this question, we used published demographic data on ten diverse species of marine bivalves. We parameterized simple matrix-population models and calculated the sums of elasticities to growth, stasis, and reproduction parameters and plotted the results on triangular axes. We also assessed whether contribution patterns were correlated with reproductive mode and tropical, temperate, or polar environments. We found that some of the broadcast spawners fell in the region of the plot with high elasticities for stasis and that some of the brooders fell in the region of the plot with higher growth and reproduction elasticities than stasis ones. However, instead of a sharp dichotomy, we found a continuum in contributions of stasis parameters with long-lived brooders and short-lived broadcast spawners in the same region of the plot. There was no clear pattern of reproductive mode associated with any particular environment, but we think these preliminary results are intriguing and that further work on comparative demography of marine invertebrates is warranted. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Sexual reproduction of Hawaiian black corals,with a review of the reproduction of antipatharians (Cnidaria: Anthozoa: Hexacorallia)

Abstract. Black corals (Cnidaria: Antipatharia) are ecologically important members of the sessile invertebrate fauna, but due to their typical deep‐water environments (>50 m), very little is known about their basic biology, including reproduction. We used histological techniques to examine the sexual reproduction of members of eight species of antipatharians collected from the Hawaiian Islands over a wide depth range (10–1327 m). Gametes of all species examined were found in association with the primary transverse mesenteries, which in some cases reached into the cavity of lateral tentacles. Specimens contained either spermatocysts or oocytes, but never both within the same individual, suggesting either a gonochoric or a sequentially hermaphroditic mode of reproduction. No developing embryos or larvae were observed in any of the samples, indicating that fertilization and larval development likely occur externally in the water column and not internally within polyps. We compared our results with previously published information on the sexual reproduction of antipatharians. Our review suggests that in antipatharians, more generally, (1) entire colonies are either female or male (although sequential hermaphroditism cannot be ruled out in most cases and has been reported previously for one species), (2) gametes are confined to the primary transverse mesenteries in most species, and (3) there is no evidence of internal fertilization. Further studies are needed to determine whether gonochorism or sequential hermaphroditism is more prevalent within the Antipatharia, whether dimorphic polyps occur within this taxonomic order, and to examine the reproductive seasonality and larval behavior of individual species.     

Contrasting reproductive strategies in three deep-sea octocorals from eastern Canada: Primnoa resedaeformis, Keratoisis ornata, and Anthomastus grandiflorus

Various aspects of reproduction were studied in three deep-sea octocorals belonging to the order Alcyonacea that co-occur at bathyal depths on the continental edge and the slope of eastern Canada. The main goals were to expand knowledge of deep-water heterotrophic corals and ascertain whether reproductive strategies could explain the known patterns of occurrence. Anthomastus grandiflorus is a gonochoric species with a female-biased sex ratio that exhibits internal fertilization and brooding of planula larvae. Conversely, Primnoa resedaeformis and Keratoisis ornata rely on broadcast spawning and external fertilization; their sexuality remains undetermined as spermatocysts were not found. In P. resedaeformis, the presence of mixed size classes of oocytes in samples from all months, depths, and locations studied suggests continuous oogenesis or overlapping development of oocyte cohorts, indicative of a gametogenic cycle spanning more than a year. No evidence of periodicity was found in this species, although it could have been masked by the striking bathymetric variation in potential relative fecundity (oocytes polyp⁻¹). The two other octocorals displayed a clear annual breeding pattern. Spawning in K. ornata and larval release in A. grandiflorus occurred in late summer and fall, respectively, possibly in response to environmental factors, as supported by shifts in the reproductive peak of A. grandiflorus across latitudes. The three species are presumed to share a nonfeeding larval mode, and data on their reproductive potential do not present any striking disparities. Published data on bycatches and video surveys in Atlantic Canada indicate that the gonochoric brooder A. grandiflorus is more widely distributed than the two free spawners, P. resedaeformis and K. ornata, which is contrary to common dispersal potential paradigms.     

Sex ratio, reproductive mode and genetic diversity in Triops cancriformis

1. Aquatic invertebrates display a wide array of alternative reproductive modes from apomixis to hermaphroditism and cyclical parthenogenesis. These have important effects on genetic diversity and population structure. Populations of the 'living fossil' Triops cancriformis display a range of sex ratios, and various reproductive modes are thought to underlie this variation. Using sex ratio information and histological analyses European populations have been inferred to be gonochoric (with separate males and females), selfing hermaphroditic and androdioecious, a rare reproductive mode in which selfing hermaphrodites coexist with variable proportions of males. In addition, some populations have been described as meiotic parthenogens.
2. Here we use population genetic analysis using microsatellite loci in populations with a range of sex ratios including a gonochoric population, and marker segregation patterns in heterozygote individuals reared in isolation, to clarify the reproductive mode in this species.
3. Our data show that populations in general have very low levels of genetic diversity. Non-gonochoric populations show lower genetic diversity, more heterozygote deficiencies, higher inbreeding coefficients and stronger linkage disequilibria than the gonochoric population. The maintenance of some heterozygosity in populations is consistent with some male influence in T. cancriformis populations, as would be expected from an androdioecious reproductive system. Results of marker segregation in eggs produced in isolation from non-gonochoric populations indicate that meiosis occurs and are consistent with two reproductive modes: selfing hermaphroditism and a type of ameiotic parthenogenesis.
4. Overall, our data indicate that androdioecy and selfing hermaphroditism are the most likely reproductive modes of non-gonochoric European Triops populations. Triops populations are strongly structured, suggesting high genetic drift and low levels of gene flow.     

Sexual systems in scleractinian corals: an unusual pattern in the reef-building species Diploastrea heliopora

The sexual system in corals refers to the spatial and temporal pattern of sexual function within an individual coral polyp, colony or population. Although information on sexual systems now exists for over 400 scleractinian species, data are still lacking for some important reef-building taxa. The vast majority of scleractinians are either simultaneous hermaphrodites or gonochoric with other sexual systems rarely occurring. Diploastrea heliopora is one of the most ubiquitous and easily recognised reef-building species in the Indo-West Pacific; however, surprisingly little is known about its reproductive biology. The aim of the present study was to examine the reproductive biology of D. heliopora colonies on chronically impacted, equatorial reefs south of Singapore. Here we show that in Singapore, D. heliopora is a broadcast spawner with predominantly gonochoric polyps. Colonies, however, contained male, female and a low proportion of cosexual polyps during the 14-month sampling period. The most plausible explanation for this is that polyps switch sexes with oogenic and spermatogenic cycles occasionally overlapping. This leads to colony level alternation of sex function within and between breeding seasons. While this sexual system is atypical for scleractinians, it supports molecular evidence that D. heliopora is phylogenetically distinct from species formerly in the family Faviidae.     

Mitochondrial and nuclear DNA variability in the living fossil Triops cancriformis (Bosc, 1801) (Crustacea, Branchiopoda, Notostraca)

The living fossil Triops cancriformis comprises bisexual (either gonochoric or hermaphroditic) and unisexual populations. Genetic surveys have recently revealed a general trend of low differentiation of 12S and 16S mitochondrial genes. We, therefore, surveyed further mitochondrial (COI gene and control region) and nuclear markers (dinucleotide microsatellites) to assess the genetic variability and to establish any relationship with the different reproductive modes found in European populations. The mitochondrial analyses confirmed the pattern of low variability. Hence, the low mitochondrial genetic variability appears as a common feature of the genus Triops. The microsatellite analysis found that Italian populations are monomorphic or exhibit little polymorphism, while other European samples display a higher degree of polymorphism and private alleles. Spanish, Austrian and Italian populations show patterns of Hardy-Weinberg disequilibrium that could be explained by the mode of reproduction, or by a higher frequency of null alleles in these populations. The low diversity and differentiation among Italian populations lead us to question the Monopolization Hypothesis. One microsatellite locus appears to be sex-linked, with heterozygotes detected only in males and hermaphrodites.     

Coral reproduction in the world��s warmest reefs: southern Persian Gulf (Dubai, United Arab Emirates)

Despite extensive research on coral reproduction from numerous geographic locations, there remains limited knowledge within the Persian Gulf. Given that corals in the Persian Gulf exist in one of the most stressful environments for reef corals, with annual variations in sea surface temperature (SST) of 12°C and maximum summer mean SSTs of 36°C, understanding coral reproductive biology in the Gulf may provide clues as to how corals may cope with global warming. In this study, we examined six locally common coral species on two shallow reef sites in Dubai, United Arab Emirates (UAE), in 2008 and 2009 to investigate the patterns of reproduction, in particular the timing and synchrony of spawning. In total, 71% colonies in April 2008 and 63% colonies in April 2009 contained mature oocytes. However, the presence of mature gametes in May indicated that spawning was potentially split between April and May in all species. These results demonstrate that coral reproduction patterns within this region are highly seasonal and that multi-species spawning synchrony is highly probable. Acropora downingi, Cyphastrea microphthalma and Platygyra daedalea were all hermaphroditic broadcast spawners with a single annual gametogenic cycle. Furthermore, fecundity and mature oocyte sizes were comparable to those in other regions. We conclude that the reproductive biology of corals in the southern Persian Gulf is similar to other regions, indicating that these species have adapted to the extreme environmental conditions in the southern Persian Gulf.