Algal overgrowth of alcyonacean soft corals

Colonies of the soft coral Lobophytum pauciflorum (Ehrenberg, 1834) (Coelenterata: Octocorallia: Alcyonacea: Alcyoniidae), some of which were heavily overgrown by the algae Ceranium flaccidum (Kuetzing) Ardissone and Enteromorpha sp., and other minor epizoites, were collected at Taylor Reef (17°50S; 146°35E) in the Great Barrier Reef. Overgrown colonies contained the diterpene 2-epi-sarcophytoxide as the major secondary metabolite, while conspecific colonies with clean polyparies contained two diterpenes in approximately equal amounts: 14-hydroxycembra-1,3,7,11-tetraene and 15-hydroxycembra-1,3,7,11-tetraene. By contrast, twenty conspecific pairs of overgrown and clean colonies of other alcyoniid soft corals collected from Pelorus Channel, Palm Island Group (18°34S; 146°29E), showed no chemical differences within in the pairs. Cultures of a common species of Ceramium [C. codii (Richards) Mazoyer] were incubated with different concentrations of nine soft-coral-derived diterpenes and significant algal growth inhibition was observed in many cases. It appears that terpenoids from soft coral may contribute to the lack of epizoic organisms on soft-coral polyparies.     

Chemical aspects of mass spawning in corals. I. Sperm-attractant molecules in the eggs of the scleractinian coral Montipora digitata

This paper provides the first evidence for sperm chemotaxis in the Scleractinia. Montipora digitata Dana, 1845 (Scleractinia: Coelenterata) is a hermaphroditic coral which reproduces bi-annually, releasing egg-sperm bundles during the mass spawning at Magnetic Island (19°10S; 146°52E) in late spring-early summer, and autumn each year. The buoyant egg-sperm bundles float to the surface where they break apart, releasing eggs and sperm into the ocean. Fertilisation occurs after 30 min. Unfertilized eggs were collected, washed free of sperm, and freeze-dried. The eggs were extracted with dichloromethane, fractionated by chromatography on silica gel, and the fractions assayed for their ability to attract M. digitata sperm. The active fraction was further fractionated by high-performance liquid chromatography, resulting in the isolation of three highly unsaturated fatty alcohols: (1) dodeca-2,4-diynol; (2) tetradec-13-ene-2,4-diynol; (3) (14Z)-heptadeca-14,16-diene-2,4-diynol. Of these three compounds, only Compound 1 attracted sperm of M. digitata. Synthetic Compound 1, produced from simple precursors by known reactions, possessed sperm-attracting activity comparable to the naturally derived attractant. Preliminary experiments suggest that the natural mixture of Compounds 1, 2 and 3 in the ratio 1:4:9 is more effective in attracting sperm from M. digitata than sperm from other Montipora species. Sperm attractants may act to reduce the incidence of hybridisation between different species of Montipora.     

Experimental investigation into the effects of suspended sediment on fertilisation, larval survival and settlement in a scleractinian coral

Laboratory and field experiments were used to determine whether high (≃100 mg l⁻¹), low (≃50 mg l⁻¹) and control (≃0 mg l⁻¹) levels of suspended sediment affected fertilisation, larval survival, and larval settlement in the scleractinian coral Acropora digitifera (Dana, 1846). Both high- and low-sediment treatments significantly decreased fertilisation, but post-fertilisation embryonic development was not inhibited by suspended sediments. Larval survival and larval settlement were significantly reduced in high- and low-sediment treatments. No difference was found between high- and low-sediment treatments in any of the three post-spawning processes investigated, suggesting that they are susceptible to sediment concentrations which are not exceptionally high even under natural conditions (>50 mg l⁻¹). The introduction of an additional stress in the form of high levels of suspended sediments coupled with naturally high variability in recruitment may have a considerable effect on the successful supply and settlement of coral larvae to a reef. Given that many coral communities are open reproductive systems, the consequences of disturbance events are not likely to be restricted to the impact area. Recruitment to a population may be reduced significantly in the presence of high levels of suspended sediments because of effects on larval survival and settlement. Recruitment of larvae to adjacent populations may also be affected due to a decreased fertilisation success and potential increases in mortality of larvae passing through the affected site. Received: 13 August 1998 / Accepted: 22 July 1999     

Transformation of soft coral (Coelenterata: Octocorallia) terpenes by Ovula ovum (Mollusca: Prosobranchia)

The faecal pellets from specimens of the prosobranch mollusc Ovula ovum found feeding on the soft coral Sarcophyton sp. at Eclipse Island, Palm Island Group (18°46S; 146°33E) in November 1980 were analysed. The only terpene present in the faeces, 7,8-deoxysarcophytoxide, differed from the major constituent of the soft coral, sarcophytoxide, suggesting that the latter had been transformed into the former within the cowrie. This transformation is not trivial, and could not be produced simply by acid catalysis. Subsequent analysis of tissues dissected from different regions of O. ovum indicates that the transformation is probably effected by enzymes in the digestive diverticula stomach region of the prosobranch. The transformed compound is significantly less toxic to the mosquito fish Gambusia affinis Baird and Girard than the ingested compound.     

Movement in soft corals: An interaction between Nephthea brassica (Coelenterata: Octocorallia) and Acropora hyacinthus (Coelenterata: Scleractinia)

An interaction between the alcyonarian octocoral Nephthea brassica Kukenthal, 1904 and the scleractinian Acropora hyacinthus Dana was photographed and the assemblage collected at Rib Reef (18°15S; 146°45E), Central Region of the Great Barrier Reef, in September 1980. The interaction, which did not involve allelopathy, was characterised by extensive deposition of undifferentiated aragonite by the scleractinian in the vicinity of the soft coral, which moved across the surface of the acroporid. The basal attachment region of the nephtheid showed structural features which permitted movement. The net effect of this interaction was a series of divided trails across the surface of the acroporid plate evidencing the movement through growth and asexual division of the parent colony. These trails were susceptible to secondary colonisation by epifauna and algae which may reduce the fitness of the acroporid. This movement appears to afford N. brassica a distinct advantage in competing for space with acroporid corals.     

Fitting fertilisation kinetics models for free-spawning marine invertebrates

 To determine how fertilisation varied with sperm concentration for two species of scallop, Chlamys (Equichlamys) bifrons (Lamarck) and C. asperrima (Lamarck), we performed a simple series of sperm dilution experiments, and measured egg size and sperm swimming speeds. C. bifrons eggs were much larger (average diam=116.5 μm), and sperm swimming speeds faster (209.8 μm s⁻¹), than C. asperrima (71.2 μm, 166.0 μm s⁻¹). In both species, maximum fertilisation occurred at an ambient sperm concentration of around 100 sperm μl⁻¹; the maximum proportion of eggs fertilised was less than 0.70 in the C. bifrons experiments, but nearer 1.0 with C. asperrima. At high sperm concentrations (>100 sperm μl⁻¹), fertilisation decreased (presumably due to polyspermy) with increasing sperm concentration, but decreased more rapidly in C. bifrons than C. asperrima. A polyspermy-adjusted fertilisation kinetics model could be fitted to the experimental data, but unique parameter estimates could not be determined. Received: 7 October 1999 / Accepted: 8 July 2000     

Evolution of egg size and fertilisation efficiency in sea stars: large eggs are not fertilised more readily than small eggs in the genus<Emphasis Type="Italic"> Patiriella</Emphasis> (Echinodermata: Asteroidea)

Fertilisation kinetics theory suggests that, when sperm are limiting, the larger eggs of broadcast-spawning marine organisms ought to be fertilised more frequently than smaller eggs, because they provide a bigger target for searching sperm. Whilst this effect has been demonstrated within species, it is not known if this pattern holds among species. We tested whether a large difference in egg size between congeneric seastars with contrasting planktotrophic and lecithotrophic modes of development results in differences in the likelihood of eggs being fertilised in sperm-limiting situations. Measurement of egg sizes and sperm swimming speeds led to the prediction that the sperm–egg collision rate constant for Patiriella calcar (420-µm-diameter egg) should be nine times greater than for P. regularis (140-µm-diameter egg). Although the eggs of P. calcar should be fertilised at greater rates in low sperm concentrations, they were not. When gametes were allowed to mix for 10 s, the hypothesis that P. calcar eggs required less sperm than P. regularis to ensure 50% of eggs were fertilised was rejected. When gametes were mixed for 5 min, P. regularis eggs were more frequently fertilised, but the difference was not statistically significant. We conclude there must be a difference between these species in the likelihood that when a sperm finds a conspecific egg it can successfully fertilise. This apparent uncoupling of egg size and likelihood of fertilisation suggests that fertilisation is not a major constraint on the evolution of egg size in these seastars.Communicated by G.F. Humphrey, Sydney     

Chemical consequences of relocation of the soft coral Lobophytum compactum and its placement in contact with the red alga Plocamium hamatum

Seven colonies of Lobophytum compactum Tixier-Durivault, 1956, which produce isolobophytolide as the major secondary metabolite, were selected from a fringing reef in the Pelorus Channel, Palm Island Group (18°34S; 146°29E), North Queensland, Australia. In September 1991, they were sectioned to afford two portions which were relocated to a grid, and a significant part of the parent colony which was left in place. The aim of the experiment was to determine the effect of relocation and contact with a toxic alga on the secondary metabolite content of a soft coral. A significant increase in the concentration of isolobophytolide was observed for all relocated colonies (n=14, p=0.001) compared to the non-relocated control colonies. This decreased after 2 mo, and was not significantly different from that of the non-relocated control colonies (n=14, p=0.881). After 1 mo, Plocamium hamatum J. Agardh plants were placed in direct contact with 50% of the relocated colonies. All soft-coral colonies in contact with the alga (n=7), showed tissue necrosis on the parts in direct contact with the alga after a further 2 wk. Tissues of the relocated control colonies (n=7), and those portions of treated colonies which were not in direct contact with the alga, were not affected. The parts of the colonies in contact with the alga showed a significant decrease in lipid content over time (n=7, p=0.001) and also a decrease in the concentration of the diterpene isolobophytolide (n=7, p=0.001). The effects of P. hamatum on the soft coral were essentially restricted to contract necrosis; chemical variations in the affected tissue were the outcomes of this necrosis. These results indicate that stress due to relocation is a more important factor in the variation of isolobophytolide levels in the soft coral L. compactum than is contact with the alga P. hamatum.     

Male characteristics, sperm traits, and reproductive success in winter-spawning Celtic Sea Atlantic herring, Clupea harengus

The relationship between sperm characteristics and reproductive success was examined in male herring, Clupea harengus L. Males were categorised as being first-time or repeat spawners on the basis of their age; they were also grouped according to whether their sperm were immediately active and exhibited forward motion on contact with seawater (FM) or had little or only vibratory motion (VM). Unlike the Pacific herring C. pallasii Valencienes, Atlantic herring sperm is usually motile on contact with seawater. The age, weight and gonadosomatic index (testes mass as a percentage of somatic mass = GSI) were measured and used as characteristics for individual fish. Sperm traits measured were (1) adenosine triphosphate (ATP) concentration, (2) sperm count, (3) duration of sperm motility. Reproductive success for each male was estimated from the fertilisation rate and from the length of larvae at hatching. Fertilisation rates for all fish were generally >80%. The ATP concentration of non-activated spermatozoa was negatively correlated with fertilisation rate. Among repeat spawners, fish with higher GSIs produced larvae that were larger at hatching. Although VM sperm fertilised eggs at rates equivalent to fertilisation by FM sperm, the larvae produced by VM sperm were significantly smaller at hatching. Larval length tended to increase in parallel with the duration of sperm motility, but the relationship was not significant in these tests. The results did not indicate any age or size pattern to spawning readiness in male herring. Sperm that are not yet ready to be shed are not fully motile on contact with seawater, but are still capable of fertilising eggs that hatch successfully. There is likely to be a progression of males which come into spawning readiness within a spawning shoal; therefore it is possible that paternal influences would result in a progressive decrease in larval size over the spawning period in winter-spawning Celtic Sea herring. Received: 22 November 1997 / Accepted: 8 June 1998     

Sequential polyandry affords post-mating sexual selection in the mouths of cichlid females

Females mating with multiple males may obtain direct benefits such as nuptial gifts or paternal care or indirect (i.e. genetic) benefits resulting in higher-quality offspring. While direct benefits are easily identified, it is difficult to determine indirect benefits, and it is hence largely unclear how they are obtained. This is particularly true in species with external fertilisation, where females seem to have little control over fertilisation. In cichlids, most maternal mouthbrooders show sequential multiple mating, where females visit several males for egg deposition. Genetic data revealed that multiple paternity of eggs and young in the mouth of females is common, but behavioural data of female spawning decisions are missing. Here, we test four hypotheses to explain female multiple mating in the maternally mouthbrooding cichlid, Ophthalmotilapia ventralis: (1) fertilisation insurance, (2) genetic bet-hedging, (3) female choice and (4) ‘sperm shopping’ (i.e. induction of sperm competition resulting in sexually selected sperm). Detailed observations of spawning behaviour in the field combined with histological analyses of the male reproductive organs suggest that fertilisation insurance, genetic bet-hedging and pre-mating female choice are unlikely to explain the sequential female multiple mating in O. ventralis. Instead, cryptic female choice by sperm shopping, i.e. post-mating sexual selection, is most compatible with our data and might be the major ultimate cause of multiple mating in females of this species and of mouthbrooding cichlids with maternal care in general. Our study provides new insight into ultimate causes of sequential polyandry in species with external fertilisation, as hitherto post-mating sexual selection by cryptic female choice has been assumed to be incompatible with external fertilisation mechanisms except by components of the ovarian fluid.     

Growth of the staghorn coral Acropora formosa at Houtman Abrolhos, Western Australia

The growth (extension rate, number of radial branches, skeletal mass, branch diameter) of the␣staghorn coral Acropora formosa (Dana, 1846) was examined at four sites on the Beacon Island platform at Houtman Abrolhos, in subtropical Western Australia (28°S). Sites were at depths of 7 to 11 m, with variable exposure to weather and swell conditions. Two sites on the western reef slope were partly exposed to the oceanic swell, and two sites in the lagoon were largely protected from wave action. Linear extension rate between 1994 and 1995 varied significantly between sites, with greater linear extension at the more protected lagoonal sites. However, accumulation of skeletal mass per branch and number of newly initiated radial branches did not vary significantly between the sites. Carbonate was deposited in similar amounts, but either as porous, rapidly extending branches, or as denser branches which extended more slowly. Branch extension rate over 11.5 mo ranged from a mean of 50.3 mm (range=13 to 93 mm) at a reef slope site to a mean of 76.0 mm (range=31 to 115 mm) at a sheltered lagoonal site. Mean extension rates were almost twice that previously reported for this species in Houtman Abrolhos (37 to 43 mm yr⁻¹) from a shallower site where environmental conditions were apparently sub-optimal. Growth was within the range reported for A. formosa from tropical sites, which is consistent with the relatively high calcification and reef-accretion rates recorded for Houtman Abrolhos in geological and metabolic studies. The role of reduced coral growth-rate in limiting coral reef formation at high latitudes remains equivocal. Received: 19 November 1997 / Accepted: 5 May 1998     

Effects of temperature on photosynthesis and respiration in hermatypic corals

Photosynthesis and respiration rates of the reef corals Pocillopora damicornis (Linn.), Montipora verrucosa (Lamarck), Porites compressa Dana and Fungia scutaria Lamarck were measured under controlled temperatures. Results indicate that coral metabolism is closely adapted to ambient temperature conditions. Tropical corals measured at Enewetak, Marshall Islands, showed greater primary production compared to maintenance requirements at elevated temperatures than did subtropical varieties of the same species in Hawaii. Photosynthesis: respiration (P:R) ratios were significantly and negatively related with temperature between 18° and 31°C for all Hawaiian corals, whereas at Enewetak this ratio generally showed a curvilinear relationship for this temperature range. Extrapolations of P:R regressions on temperatures to a value of 2.0 (estimated as a minimum required for long-term functional autotrophy) coincide for Hawaiian specimens with published upper lethal temperatures. Extrapolation of P:R regressions for Enewetak specimens at temperatures above 25°C suggests lethal temperatures for these corals to be 2 to 5 C° higher than for Hawaiian corals, in good agreement with recent experimental findings. Interspecific differences in P:R temperature regressions for Hawaiian corals correlating with upper lethal temperature tolerances are described.Contribution No. 505 of the Hawaii Institute of Marine Biology.     

Laying-order effects on sperm numbers and on paternity: comparing three passerine birds with different life histories

In birds, the number of sperm trapped between the perivitelline membranes around the ovum is an estimate of sperm numbers present at the time and place of fertilisation in the female reproductive tract. Sperm numbers may vary among species and between eggs in a clutch and can provide information about sperm utilisation and mechanisms of sperm competition. Here, we describe patterns of variation in sperm numbers through the egg-laying sequence in three passerines in which extra-pair paternity is common, but copulation behaviour differs. Sperm numbers showed no systematic change across the laying sequence in blue tits (Cyanistes caeruleus), but decreased significantly with laying order in bluethroat (Luscinia svecica) and tree swallow (Tachycineta bicolor) clutches. This is consistent with observations that blue tits regularly copulate throughout the laying sequence, while bluethroats stop mate guarding and tree swallows reduce their copulation frequency once the first egg is laid. Nevertheless, cases of a sudden increase in sperm numbers in clutches of bluethroats and tree swallows suggest that successful inseminations also occurred after laying started. In blue tits and bluethroats, sperm numbers were not higher on extra-pair sired eggs than on eggs sired by the social male, suggesting that extra-pair copulations are not timed to the period of peak fertility for each egg. More extra-pair offspring originated from eggs laid early in the sequence in blue tits, while there was no systematic bias in bluethroats. Our results suggest that copulations during the laying sequence are predominantly performed by within-pair males in our study species.     

Fertilisation success in the fly Dryomyza anilis (Dryomyzidae): effects of male size and the mating situation

Male fertilisation success in relation to male size and the mating situation (ordinary pair formation with a single, nonvirgin female vs. take overs) was examined in the fly Dryomyza anilis. In ordinary matings, large males achieved higher fertilisation success than small ones when they were the second to mate with the female. Take overs differ from ordinary pair formation in that the second male experiences intensified sperm competition. This is because in take overs the female is not able to discharge any of the sperm inseminated by the first male as she usually does before a new mating. Compared with ordinary matings, take overs reduced the fertilisation success of the second male by 8–10%, whereas that of the first male was 7–14% higher in take overs. Even though the intruder was always larger than the paired male his superior fertilisation success did not compensate for the effect of the sperm already present in the female. In D. anilis, males can increase their fertilisation success by tapping the female's external genitalia with their claspers or having several copulation bouts per mating. Thus, in a take over, the intruder could respond to the intensified sperm competition by performing more tapping sequences per copulation bout or more copulation bouts per mating. In matings observed in the wild, males performed more tapping sequences after a take over than after pair formation with a single female, although the difference was not significant. The results show that there are differences in fertilisation success between males of different size. In addition, different mating situations can result in considerable variation in the fertilisation success of an individual male. Higher fertilisation success for the first male after a take over may be significant, in particular, for the reproductive success of small males, which frequently lose their females to large males.     

Subitaneous, diapause, and delayed-hatching eggs of planktonic copepods from the northern Gulf of Mexico: morphology and hatching success

Experiments were conducted to examine the morphology and hatching success of eggs, either spawned by freshly caught planktonic copepods or recovered from bottom sediments in the northern Gulf of Mexico. Collections were made between August 1992 and September 1995. Eggs of nine species were described and these differed in their diameter, color and surface attributes. Three types of eggs were distinguished: subitaneous, diapause, and delayed-hatching. Three species, Labidocera aestiva Wheeler, Acartia tonsa Dana, and Centropages velificatus (Oliveira) produced only subitaneous eggs. Hatching success varied greatly among these species. Two species, Labidocera mirabilis Fleminger and Centropages hamatus (Lilljeborg) produced diapause eggs and subitaneous eggs. The length of the refractory phase of the diapause eggs differed greatly both within and between these two species. A third type of dormant egg, delayed-hatching, was recognized in Labidocera scotti Giesbrecht and Pontella meadi Wheeler. The existence of delayed-hatching eggs may be an adaptive response of subtropical species to less seasonal fluctuation. Based upon morphological characteristics of the eggs and rearing of nauplii to an identifiable stage, benthic dormant eggs of eight species of calanoid copepods were also identified. Received: 9 September 1996 / Accepted: 1 October 1996     

Seasonal and sex-specific variations in levels of photo-protecting mycosporine-like amino acids (MAAs) in soft corals

This study focusses on the nature and extent of variation in mycosporine-like amino acids (MAAs) in relation to annual cycles in solar radiation, seawater temperature, and reproduction in reef-flat populations of two soft coral species. The results show MAA tissue concentrations in shallow water colonies of Lobophytum compactum and Sinularia flexibilis to be significantly correlated to annual cycles in solar radiation (P<0.0006 and P<0.0005, respectively) and seawater temperature (P<0.0006 and P<0.0004, respectively). Evidence of seasonal cycles in MAA levels in the tissues of shallow-reef invertebrates positively correlating with annual cycles in solar radiation and temperature suggests that they are an integral component of the soft corals' biochemical defence system against high irradiance and/or temperature stress and thus bleaching. This is further corroborated by the higher production of MAAs in females than males prior to spawning (up to 67% and 56% for L. compactum and S. flexibilis, respectively), presumably to provide a high level of protection against irradiance stress for progeny.     

Copepod recruitment and food composition: do diatoms affect hatching success?

Laboratory experiments were conducted to differentiate between factors controlling the hatching success of copepod eggs. Factors that could affect viability of eggs; viz food quality, female condition and external factors were investigated. In a series of experiments the copepod Acartia tonsa Dana was fed several different diets while egg production and hatching success were monitored. The diet was analysed for fatty acid content as an indicator of food quality. Both egg production and hatching were found to be affected by the nutritional quality of the food. Hatching was also highly dependent on female fertility. External effects were tested by exposing eggs to diatom extracts. Negative effects were only evident at high extract concentrations, but disappeared when aeration was supplied to the solution. Oxygen measurements showed that failure to hatch was due to hypoxia in the extracts. No inhibitory or toxic effects of diatom cell components on hatching could be found.     

Effect of nutrient enrichment on the complementary (secondary) metabolite composition of the soft coral Sarcophyton ehrenbergi (Cnidaria: Octocorallia: Alcyonaceae) of the Great Barrier Reef

 A long-term study of the effects of nutrient enrichment on coral reefs (ENCORE Experiment) was carried out at One Tree Island (23°30′S; 152°96′E), Great Barrier Reef, between 1992 and 1996. The experiment involved the addition of water-soluble nutrients to 12 microatolls which contained a range of organisms and were situated within the larger lagoon. Three replicates of each of three nutrient treatments (nitrogen, phosphorus, nitrogen plus phosphorus) and an untreated set of three control atolls were prepared using the 12 selected microatolls. As part of the larger ENCORE experiment, changes in the chemical composition of the alcyonacean soft coral Sarcophyton ehrenbergi Marenzeller (Octocorallia: Alcyonacea), placed in the treatment microatolls, were monitored for a 1 yr period in an attempt to detect any responses attributable to nutrient enrichment. Multivariate analyses were performed to determine whether there were any patterns of response in the different nutrient treatments. At the level of individual metabolites, there were no clear treatment effects. However, the ratio of bioactive or stress metabolites (terpenes) to energy storage metabolites (lipids), referred to as the “physiological-change indicator”, revealed effects of nutrient enrichment. Nitrogen enrichment resulted in a trend towards higher physiological-change indicators than control or phosphorus treatments in the majority of cases, while phosphorus enrichment significantly decreased the ratio relative to controls. In most cases, the physiological-change indicator increased in soft soft corals relocated into contact with the scleractinian Pocillopora damicornis. The potential of soft corals to serve as indicators of a changed nutrient regime is discussed. Received: 23 November 1998 / Accepted: 23 September 1999     

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

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

Genetic differentiation among populations of a broadcast spawning soft coral, Sinularia flexibilis, on the Great Barrier Reef

The genetic structure of 12 reef populations of the soft coral Sinularia flexibilis (Octocorallia, Alcyoniidae) was studied along the Great Barrier Reef (GBR) at a maximum separation of 1,300 km to investigate the relative importance of sexual and asexual reproduction, genetic differentiation and gene flow among these populations. S. flexibilis is a widely distributed Indo-Pacific species and a gamete broadcaster that can form large aggregations of colonies on near-shore reefs of the GBR. Up to 60 individuals per reef were collected at a minimum sampling scale of 5 m at two sites per reef, from December 1998 to February 2000. Electrophoretic analyses of nine polymorphic allozymes indicated that genotypic frequencies in most populations and loci did not differ significantly from those expected from Hardy–Weinberg predictions. Analysis of multi-locus genotypes indicated a high number of unique genotypes (N _go) relative to the number of individuals sampled (N) in each reef population (range of 0.69–0.95). The maximum number of individuals likely to have been produced sexually (N*) was similar to the number of individuals sampled (i.e. N*:N ˜ 1), suggesting that even repeated genotypes may have been produced sexually. These results demonstrated a dominant role of sexual reproduction in these populations at the scale sampled. Significant genetic differentiation between some populations indicated that gene flow is restricted between some reefs (F _ST=0.026, 95% CI= 0.011 − 0.045) and even between sites within reefs (F _ST=0.041, 95% CI=0.027 − 0.055). Nevertheless, there was no relationship between geographic separation and genetic differentiation. Analyses comparing groups of populations showed no significant differentiation on a north-south gradient in the GBR. The pattern in the number of significant differences in gene frequencies in pairwise population comparisons, however, suggested that gene flow may be more restricted among inner-shelf reef populations near to the coast than among mid/outer-shelf populations further from the coast. Received: 10 July 2000 / Accepted: 5 October 2000