Reproductive biology of three sponge species of the genus Xestospongia (Porifera: Demospongiae: Petrosida) from the Great Barrier Reef

The reproductive development of three species of the Petrosida, Xestospongia bergquistia, X. exigua, and X. testudinaria, was monitored for four years on a fringing reef at Orpheus Island, Great Barrier Reef, Australia. All three species were oviparous and female reproductive activity began prior to males becoming active. X. bergquistia and X. testudinaria were gonochoric and broadcast eggs in spawning events that were synchronous within species. Egg development occurred over more than five months in X. bergquistia and X. testudinaria and two months in X. exigua. Spawning was during periods of warm temperature and occurred in October or November for X. bergquistia and X. testudinaria, and January or February for X. exigua. Lunar phase was implicated in timing of spawning of X. testudinaria. Diel timing of spawning in X. testudinaria and X. bergquistia was consistently a morning event.     

Host Imprinting in Anemonefishes (Pisces: Pomacentridae): Does it Dictate Spawning site Preferences?

The olfactory cues, to which some species of anemonefish embryos imprint, are secreted in the mucus on the tentacles and the oral disc of the host anemone. Close contact of the eggs of anemonefishes with the host's tentacles seems therefore important to imprinting. A corollary of this observation is that if local environmental conditions sweep tentacles in one specific direction, then the eggs will be placed leeward of the tentacles, rather than to foremost way from the tentacles. Other known factors such as egg predation can also cause a spawning site preference. No study has examined the possibility of the existence of such a preference. In this study, we addressed two questions: (1) Does spawning site preference exist in anemonefishes? (2) If yes, is it possible to relate this to the imprinting hypothesis, i.e. does local ocean currents over the host anemone have any influence on this preference? Two different coral reef areas were surveyed for anemonefish groups with eggs present: Eilat and Na'ama in the Aquaba-bight, the Red Sea (RS), and areas at Lizard Island, the northern Great Barrier Reef (GBR). We found the anemonefishes Amphiprion akindynos (GBR), A. bicinctus (RS), A. melanopus (GBR), and A. perideraion (GBR), to have a distinctive spawning site preference. We discuss the relevance of these findings to anemonefish host imprinting.     

Inducing broadcast coral spawning ex situ: Closed system mesocosm design and husbandry protocol

For many corals, the timing of broadcast spawning correlates strongly with a number of environmental signals (seasonal temperature, lunar, and diel cycles). Robust experimental studies examining the role of these putative cues in triggering spawning have been lacking until recently because it has not been possible to predictably induce spawning in fully closed artificial mesocosms. Here, we present a closed system mesocosm aquarium design that utilizes microprocessor technology to accurately replicate environmental conditions, including photoperiod, seasonal insolation, lunar cycles, and seasonal temperature from Singapore and the Great Barrier Reef (GBR), Australia. Coupled with appropriate coral husbandry, these mesocosms were successful in inducing, for the first time, broadcast coral spawning in a fully closed artificial ex situ environment. Four Acropora species (A. hyacinthus, A. tenuis, A. millepora, and A. microclados) from two geographical locations, kept for over 1 year, completed full gametogenic cycles ex situ. The percentage of colonies developing oocytes varied from ~29% for A. hyacinthus to 100% for A. millepora and A. microclados. Within the Singapore mesocosm, A. hyacinthus exhibited the closest synchronization to wild spawning, with all four gravid colonies releasing gametes in the same lunar month as wild predicted dates. Spawning within the GBR mesocosm commenced at the predicted wild spawn date but extended over a period of 3 months. Gamete release in relation to the time postsunset for A. hyacinthus, A. millepora, and A. tenuis was consistent with time windows previously described in the wild. Spawn date in relation to full moon, however, was delayed in all species, possibly as a result of external light pollution. The system described here could broaden the number of institutions on a global scale, that can access material for broadcast coral spawning research, providing opportunities for institutions distant from coral reefs to produce large numbers of coral larvae and juveniles for research purposes and reef restoration efforts.     

Summer spawning of Porites lutea from north-western Australia

Most coral species off Australia??s west coast spawn in the austral autumn (March?CApril), with a few species also spawning in the southern spring or early summer (November?CDecember). This is the reverse timing to spawning recorded off Australia??s east coast. Porites lutea, a gonochoric broadcast spawner that is common on Australia??s west coast, is shown here to spawn in the months of November or December, as it does on Australia??s east coast. Spawning occurred between 2 and 5 nights after full moon, with the majority of spawning activity on night 3. Gametes developed over three to four months with rapid development in the last two weeks before spawning. Zooxanthellae were typically observed in mature oocytes, only a week before spawning so their presence may provide a useful indicator of imminent spawning.     

Seasonality and lunar periodicity in the reproduction of Pocilloporid corals

Reproductive seasonality and lunar periodicity of planula release were investigated for the three brooding coralsPocillopora damicornis, Seriatopora hystrix, andStylophora pistillata at Heron Island in the southern Great Barrier Reef. Branch fragments collected from undisturbed colonies in the field were used to determine when planulae were present for all three species, and direct observations of planula release were made on colonies ofPocillopora kept in aquaria. All three species displayed marked seasonal variation in reproductive output, with nearly all reproductive activity occurring over the summer months.Pocillopora exhibited distinct lunar periodicity in planulation, with planula release occurring around three quarter moon, and no planulae being present in samples collected between new and full moons.Seriatopora also displayed lunar periodicity of planula release, although it was not as distinct as forPocillopora, whileStylophora did not show any lunar periodicity.     

Congruent patterns of connectivity can inform management for broadcast spawning corals on the Great Barrier Reef

Connectivity underpins the persistence and recovery of marine ecosystems. The Great Barrier Reef (GBR) is the world's largest coral reef ecosystem and managed by an extensive network of no‐take zones; however, information about connectivity was not available to optimize the network's configuration. We use multivariate analyses, Bayesian clustering algorithms and assignment tests of the largest population genetic data set for any organism on the GBR to date (Acropora tenuis, >2500 colonies; >50 reefs, genotyped for ten microsatellite loci) to demonstrate highly congruent patterns of connectivity between this common broadcast spawning reef‐building coral and its congener Acropora millepora (~950 colonies; 20 reefs, genotyped for 12 microsatellite loci). For both species, there is a genetic divide at around 19°S latitude, most probably reflecting allopatric differentiation during the Pleistocene. GBR reefs north of 19°S are essentially panmictic whereas southern reefs are genetically distinct with higher levels of genetic diversity and population structure, most notably genetic subdivision between inshore and offshore reefs south of 19°S. These broadly congruent patterns of higher genetic diversities found on southern GBR reefs most likely represent the accumulation of alleles via the southward flowing East Australia Current. In addition, signatures of genetic admixture between the Coral Sea and outer‐shelf reefs in the northern, central and southern GBR provide evidence of recent gene flow. Our connectivity results are consistent with predictions from recently published larval dispersal models for broadcast spawning corals on the GBR, thereby providing robust connectivity information about the dominant reef‐building genus Acropora for coral reef managers.     

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.     

Does simultaneous and sympatric reproduction between two native spined loaches lead to reproductive interference and local extinction?

Reproductive interference occurs when fitness of females of one species decreases because of misdirected courtship behaviors by males of another species. Reproductive interference strongly excludes either species in the interaction but it does not persist for long on an evolutionary time scale. Therefore, reproductive interference between a pair of co-evolved native species is difficult to observe because exclusion ends the interaction even if even the potential for reproductive interference still exists. However, anthropogenic environmental changes can bring about secondary contact between two species and trigger reproductive interference, leading to local species extinction. We demonstrated this phenomenon with two native spined loaches: Cobitis magnostriata (Cmag) and C. minamorii oumiensis (Cmio). In 2015 and 2016, we investigated the spatiotemporal dynamics of adults and juveniles of both species to estimate the effects of the relative abundance of adults on reproductive success. Additionally, we surveyed the distribution of juveniles and investigated the proportion of gravid females which failed to reproduce at the end of the spawning season. In 2016, Cmio juveniles were fewer than in 2015, although the abundances of Cmio adults were similarly low in both years and the Cmag adults were more abundant in 2016 than in 2015. The juveniles of the two species exhibited similar spatiotemporal dynamics. At the end of the spawning season, Cmag males were abundant and most Cmio females failed to reproduce. These results suggest that the two species share spawning habitats and seasons and that Cmag males exert reproductive interference on Cmio females. This report is the first of a study suggesting that anthropogenic environmental changes triggered reproductive interference between native species in the field.     

Spatial variation in the effects of size and age on reproductive dynamics of common coral trout Plectropomus leopardus

The effects of size and age on reproductive dynamics of common coral trout Plectropomus leopardus populations were compared between coral reefs open or closed (no‐take marine reserves) to fishing and among four geographic regions of the Great Barrier Reef (GBR), Australia. The specific reproductive metrics investigated were the sex ratio, the proportion of vitellogenic females and the spawning fraction of local populations. Sex ratios became increasingly male biased with length and age, as expected for a protogyne, but were more male biased in southern regions of the GBR (Mackay and Storm Cay) than in northern regions (Lizard Island and Townsville) across all lengths and ages. The proportion of vitellogenic females also increased with length and age. Female P. leopardus were capable of daily spawning during the spawning season, but on average spawned every 4·3 days. Mature females spawned most frequently on Townsville reserve reefs (every 2·3 days) and Lizard Island fished reefs (every 3·2 days). Females on Mackay reefs open to fishing showed no evidence of spawning over 4 years of sampling, while females on reserve reefs spawned only once every 2–3 months. No effect of length on spawning frequency was detected. Spawning frequency increased with age on Lizard Island fished reefs, declined with age on Storm Cay fished reefs, and declined with age on reserve reefs in all regions. It is hypothesized that the variation in P. leopardus sex ratios and spawning frequency among GBR regions is primarily driven by water temperature, while no‐take management zones influence spawning frequency depending on the region in which the reserve is located. Male bias and lack of spawning activity on southern GBR, where densities of adult P. leopardus are highest, suggest that recruits may be supplied from central or northern GBR. Significant regional variation in reproductive traits suggests that a regional approach to management of P. leopardus is appropriate and highlights the need for considering spatial variation in reproduction where reserves are used as fishery or conservation management tools.     

Captive rearing for Chinook salmon (Oncorhynchus tshawytscha) and Atlantic salmon (Salmo salar): the Idaho and Maine experiences

Captive rearing is a conservation strategy where juveniles are collected from the natural environment, reared to maturity in a hatchery environment, and then released back into the natural environment at maturity for volitional spawning. This strategy has been used to produce adult outplants for stock enhancement where natural escapement is poor or capture of adults is difficult. In both Idaho (Chinook salmon, Oncorhynchus tshawytscha) and Maine (Atlantic salmon, Salmo salar), captive rearing programs have been initiated as an experimental strategy to prevent cohort collapse and conserve genetic integrity of select depressed populations. In this paper, we provide an overview of these programs and describe some of the methods used to evaluate the effectiveness of this approach. Behaviors such as habitat selection, courting, and spawn timing were monitored. Data collected for both programs indicate that the captive fish display similar behaviors as their wild conspecifics in terms of habitat selection and spawning, although there were some differences in spawn timing. Evaluations of egg and fry production also indicate that captive-reared adults are successfully spawning and producing offspring. Each program is still waiting on final evaluations of reproductive success through genetic analyses of returning adults, but results so far indicate that this could be an additional captive propagation strategy for depressed populations.     

Scallop Size does not Predict Amount or Rate of Induced Sperm Release

Within populations of broadcast spawning marine invertebrates such as scallops, larger animals typically have larger gonads. Presumably, this means those larger males have more sperm to release than small males. However, there has never been a direct test of whether larger males actually release more sperm, at a higher rate, during spawning. To address this, we compared the allometry of induced sperm release with that of reproductive investment (gonad weight) in ripe males of 2 species of scallops, Chlamys bifrons and Chlamys asperrima. We did not find that larger scallops released more sperm or released it faster than small scallops, and were able to reject the hypothesis that instantaneous sperm release was related to body size in the same way as gonad weight. Consequently, we speculate that if larger broadcast spawning males do release more sperm, they may do so by spawning on more occasions within a reproductive season.     

Male spawning success and female choice in the mottled triplefin,Forsterygion varium (Pisces: Tripterygiidae)

A detailed analysis was made of the variation in spawning success among male Forsterygion (э Tripterygion) varium. The aim was to assess whether females were mating selectively and, if so, to examine the basis of that choice. Eggs were laid on small rocks within the males' territories. Females did not distribute eggs randomly among males. Some males guarded as many as 23000 eggs in one breeding season, while others experienced little or no breeding success. Male spawning success was positively correlated with both their size and some characteristics of their territories, namely the amount of spawning substratum and the presence of a large boulder adjacent to the territory. Experimental manipulations indicated that female choice of males was based on these features. Addition of small rocks into the territories of previously unsuccessful males resulted in a significant increase in spawning success. Conversely, spawning success of the most successful males decreased when small rocks were removed from their territories. When successful males were removed, each territory was occupied by small males; however, there was no significant change in the spawning success of these males. This experiment suggested that, although territory quality was important, female mating decisions could also be influenced by male characteristics such as body size.     

Plasticity of fertilization rates under varying temperature in the broadcast spawning mussel,Mytilus galloprovincialis

Oceans are a huge sink for the increased heat associated with anthropogenic climate change, and it is vital to understand the heat tolerance of marine organisms at all life stages to accurately predict species’ responses. In broadcast spawning marine invertebrates, reproduction is a vulnerable process in which sperm and eggs are released directly into the open water. Gametes are then exposed to fluctuating environmental conditions that may impact their fertilizing capacity. Using the broadcast spawning Mediterranean mussel, Mytilus galloprovincialis, as a model species, we performed blocks of factorial mating crosses to assess the variance in fertilization rates among individuals under both ambient and elevated temperatures. Overall, we found a small, but significant decline in fertilization rates with elevated temperatures. However, there was substantial plasticity in responses, with particular mussels having increased fertilization under elevated temperatures, although the majority showed decreased fertilization rates. Our results suggest possible future reproductive costs to ocean warming in M. galloprovincialis, although it is also possible that genetic variation for thermal sensitivity may allow for adaptation to changing environmental conditions.     

Phenology of sexual reproduction in the common coral reef sponge,Carteriospongia foliascens

Understanding processes that contribute to population maintenance is critical to the management and conservation of species. Despite this, very little is currently known about the reproductive biology of Great Barrier Reef (GBR) sponge species. Here, we established reproductive parameters including mode of sexuality and development, seasonality, sex ratios, gametogenesis, reproductive output, and size at sexual maturity for the common phototrophic intertidal sponge, Carteriospongia foliascens, in the central GBR over two reproductive cycles. A population sexual productivity index (PoSPi) integrating key reproductive parameters was formulated to compare population larval supply over time. This study shows that C. foliascens is reproductive all year round, gonochoric and viviparous, with larvae developing asynchronously throughout the mesohyl. The influence of environmental parameters relevant to C. foliascens reproduction [i.e., sea surface temperature (SST), photoperiod, and rainfall] was also examined, and SST was found to have the most significant effect on phenology. C. foliascens reproduction exhibited annual mono-cyclic patterns closely resembling SST fluctuations. Reproductive output was depressed at low SST (<23 °C) and increased at temperatures above 23 °C. Peak sperm release occurred at temperatures above 25 °C, while peak larval release occurred during the annual temperature maxima (>28 °C). A twofold increase in maximum larval production (PoSPi) in C. foliascens was observed in the second reproductive cycle, following a depressed PoSPi in the first cycle. This reduction in PoSPi in the first reproductive cycle was associated with elevated SST and rainfall, coinciding with one of the strongest La Niña events on record.     

Timing of final oocyte maturation in Acropora and merulinid corals

Final oocyte maturation is a gametogenic process that occurs at the end of oogenesis, which includes germinal vesicle migration (GVM) and germinal vesicle breakdown (GVBD). This process is essential for oocytes to become competent for fertilization and occurs just before spawning. Timing of GVM and GVBD has been extensively studied to understand reproductive timing in various marine organisms. For corals, however, such information remains scarce. Here, we examined the timing of GVM and GVBD in broadcast-spawning corals: two Acropora and six merulinid species, from around the full moon until spawning day(s), at Lyudao, Taiwan in 2019 and 2020. The proportion of oocytes that had completed GVM around the full moon varied among species (14–64%), while the proportion increased to > 80% by 6–7 d after the full moon when many sampled colonies spawned. By contrast, although data are limited, the timing of GVBD indicated a more uniform pattern among species, and the onset of GVBD in the majority of oocytes occurred within 3–10 h before spawning. As GVM and GVBD are prerequisites for fertilization, and probably the spawning itself, the interspecific variation in the timing of GVM likely reflects an interspecies variation in spawning timing. This hypothesis may partially explain the different spawning timing (days) among coral species observed at the study location. Further research is needed to test such a hypothesis, given the limitation of our study with regard to the number of observations, annual variation, and examined coral taxa.

     

The reproductive biology and early life ecology of a common Caribbean brain coral, <Emphasis Type="Italic">Diploria labyrinthiformis</Emphasis> (Scleractinia: Faviinae)

Despite the fact that most of the severe demographic bottlenecks in coral populations occur during their earliest life stages, information on the reproductive biology and early life history traits of many coral species is limited and often inferred from adult traits only. This study reports on several atypical aspects of the reproductive biology and early life ecology of the grooved brain coral, Diploria labyrinthiformis (Linnaeus, 1758), a conspicuous reef-building species on Caribbean reefs. The timing of gamete release of D. labyrinthiformis was monitored in Curaçao over eight consecutive months, and embryogenesis, planulae behavior, and settlement rates were observed and quantified. We further studied growth and symbiont acquisition in juvenile D. labyrinthiformis for 3.5 yr and compared settler survival under ambient and nutrient-enriched conditions in situ. Notably, D. labyrinthiformis reproduced during daylight hours in six consecutive monthly spawning events between May and September 2013, with a peak in June. This is the largest number of reproductive events per year ever observed in a broadcast-spawning Caribbean coral species. In settlement experiments, D. labyrinthiformis planulae swam to the bottom of culture containers 13 h after spawning and rapidly settled when provided with settlement cues (42% within 14 h). After 5 months, the survival and growth rates of settled juveniles were 3.7 and 1.9 times higher, respectively, for settlers that acquired zooxanthellae within 1 month after settlement, compared to those that acquired symbionts later on. Nutrient enrichment increased settler survival fourfold, but only for settlers that had acquired symbionts within 1 month after settlement. With at least six reproductive events per year, a short planktonic larval phase, high settlement rates, and a positive response to nutrient enrichment, the broadcast-spawning species D. labyrinthiformis displays a range of reproductive and early life-history traits that are more often associated with brooding coral species, illustrating that classical divisions of coral species by reproductive mode alone do not always reflect the true biology and ecology of their earliest life stages.     

Sexual reproduction of Acropora reef corals at Moorea, French Polynesia

Little information is available on reproductive processes among corals in isolated central Pacific reef regions, including French Polynesia. This study examined the timing and mode of sexual reproduction for Acropora reef corals at Moorea. Spawning was observed and/or inferred in 110 Acropora colonies, representing 12 species, following full moon periods in September through November 2002. Gamete release was observed and inferred in four species of Acropora between 9 and 13 nights after the full moon (nAFM) in September 2002. Twelve Acropora spp. spawned gametes between 5 and 10 nAFM in October 2002, with six species spawning 7 nAFM and four species spawning 9 nAFM. In November 2002, spawning of egg and sperm bundles was observed and inferred in 27 colonies of Acropora austera, 6 nAFM. These are the first detailed records of spawning by Acropora corals in French Polynesia.     

Female nuptial signal advertises spawning readiness in a population of <Emphasis Type="Italic">Halichoeres margaritaceus</Emphasis> on the Great Barrier Reef,Australia

Females of some polygynous fish species display female nuptial signals (FNSs), advertising spawning readiness to potential mates. Halichoeres margaritaceus, a polygynous coral reef fish, reportedly display red belly color and a unique bobbing behavior as FNSs advertising spawning readiness. In this study, I examined whether FNSs were observed in an H. margaritaceus population in Australia. Females in this study displayed red belly color as an FNS, although at a lower intensity than that reported in a Japanese population. Contrary to observations made in the Japanese population, bobbing behavior performed by females in this study showed no temporal trend as spawning approached, indicating bobbing behavior did not function as an FNS in the Australian population. The role of local selective pressures on FNS intensity and lifetime reproductive benefits H. margaritaceus may potentially gain in utilizing FNSs are briefly discussed.     

Biannual Spawning and Temporal Reproductive Isolation in Acropora Corals

Coral spawning on the oceanic reef systems of north-western Australia was recently discovered during autumn and spring, but the degree to which species and particularly colonies participated in one or both of these spawnings was unknown. At the largest of the oceanic reef systems, the participation by colonies in the two discrete spawning events was investigated over three years in 13 species of Acropora corals (n = 1,855 colonies). Seven species spawned during both seasons; five only in autumn and one only in spring. The majority of tagged colonies (n = 218) spawned once a year in the same season, but five colonies from three species spawned during spring and autumn during a single year. Reproductive seasonality was not influenced by spatial variation in habitat conditions, or by Symbiodinium partners in the biannual spawner Acropora tenuis. Colonies of A. tenuis spawning during different seasons separated into two distinct yet cryptic groups, in a bayesian clustering analysis based on multiple microsatellite markers. These groups were associated with a major genetic divergence (G”_ST = 0.469), despite evidence of mixed ancestry in a small proportion of individuals. Our results confirm that temporal reproductive isolation is a common feature of Acropora populations at Scott Reef and indicate that spawning season is a genetically determined trait in at least A. tenuis. This reproductive isolation may be punctuated occasionally by interbreeding between genetic groups following favourable environmental conditions, when autumn spawners undergo a second annual gametogenic cycle and spawn during spring.     

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.