A floating mid-water coral nursery as larval dispersion hub: testing an idea

The global decline in reef health has prompted the need for effective management methodologies, including the development of active restoration measures. One such approach is the ‘gardening concept’ that involves use of underwater nurseries where coral fragments are farmed before their transplantation into denuded reefs. Here we document enhanced sexual reproduction in colonies of the coral Stylophora pistillata cultured in mid-water floating nursery situated in nutrient enriched water, near the fish farms in Eilat, Red Sea. We found that after 2 years of nursery, the average number of oocytes per polyp in farmed colonies was ca. 35% higher than in corresponding naturally growing colonies. Small branches in the nursery developed gravid colonies that released equal (or more) numbers of planula larvae as compared to similar size, 5-year old naturally growing colonies. These nursery-borne planulae possessed more zooxanthellae and contained more chlorophyll per larva. While settled and metamorphosed in equal rates compared to planulae originated from reef-grown colonies, the nursery borne planulae developed faster growing young colonies. We estimate that a coral nursery could generate, during the reproductive season, tens of millions of planula larvae and therefore should be regarded as a ‘larval dispersion hub’ that can be used as a management tool for natural recruitment enhancement.     

Steps towards the evaluation of coral reef restoration by using small branch fragments

“Gardening” of denuded coral reef habitats is a novel restoration approach in which sexual and asexual recruits are used. The present study aimed at the evaluation of the potentiality for restoration use of different types of small fragments subcloned from the Red Sea coral species Stylophora pistillata. In situ short-term (24 h, ⁴⁵Ca method) and long-term (1 year, alizarin Red S vital staining) experiments revealed high variation (up to 70%) in growth rates between up-growing branches of a specific genet, and that tip ratios in dichotomous branches (n = 880) differ significantly between newly formed and older branches, further emphasizing the within-colony genetic background for spatial configuration. Small, isolated branches (<4 cm) revealed high survivorship (up to 90%, 1 year) and up to 20–30% (1 year, single- vs. dichotomous-tip branches, respectively) growth, showing that small-sized branches are suitable for restoration purposes. Results differed significantly between genets. Total length added for dichotomous-tip branches was in general at least twice that recorded for single tips of a specific genet. Restoration protocols may be applied either by sacrificing whole large colonies via pruning high numbers of small fragments or, by pruning only a few small branches from each one of many genets. An in situ “nursery period” of approximately 8 years is predicted for S. pistillata small fragments. Received: 17 August 1999 / Accepted: 15 February 2000     

Three party symbiosis: acoelomorph worms,corals and unicellular algal symbionts in Eilat (Red Sea)

Epizoic worms were found to occur on certain coral colonies from reefs off the coast of Eilat (Red Sea). We identified 14 coral species infested by acoelomorph worms at a depth range of 2–50 m. The host corals were all zooxanthellate and included both massive and branching stony corals and a soft coral. Worms from all hosts were identified as belonging to the genus Waminoa and contained two distinct algal symbionts differing in size. The smaller one was identified as Symbiodinium sp. and the larger one is presumed to belong to the genus Amphidinium. Worm-infested colonies of the soft coral, Stereonephthya cundabiluensis, lacked a mucus layer and exhibited distinct cell microvilli, a phenotype not present in colonies lacking Waminoa sp. In most cases, both cnidarian and Acoelomorph hosts displayed high specificity for genetically distinctive Symbiodinium spp. These observations show that the epizoic worms do not acquire their symbionts from the “host” coral.     

Chemical mimicry in an incipient leaf-cutting ant social parasite

Some social parasites of insect societies are known to use brute force when usurping a host colony, but most use more subtle forms of chemical cheating either by expressing as few recognition cues as possible to avoid being recognized or by producing similar recognition cues to the host to achieve positive discrimination. The former “chemical insignificance” strategy represents a more general adaptive syndrome than the latter “chemical mimicry” strategy and is expected to be characteristic of early evolutionary stages of social parasitism. We tested this hypothesis by experimentally analyzing the efficiency by which Acromyrmex echinatior leaf-cutting ants recognize intruding workers of the incipient social parasite Acromyrmex insinuator. The results were consistent with the parasite being “chemically insignificant” and not with the “chemical mimicry” hypothesis. Gas chromatography–mass spectrometry analysis of cuticular hydrocarbon profiles showed that social parasite workers produce significantly fewer hydrocarbons overall and that their typical profiles have very low amounts of hydrocarbons in the “normal” C29–C35 range but large quantities of unusually heavy C43–C45 hydrocarbons. This suggests that the C29–C35 hydrocarbons are instrumental in normal host nestmate recognition and that the C43–C45 compounds, all of which are dienes and thus more fluid than the corresponding saturated compounds, may reinforce “chemical insignificance” by blurring any remaining variation in recognition cues.     

Survey of deep-dwelling red coral (<Emphasis Type="Italic">Corallium rubrum</Emphasis>) populations at Cap de Creus (NW Mediterranean)

The distribution and population structure of the eurybathic gorgonian Corallium rubrum were studied off Cap de Creus (Costa Brava, Northwestern Mediterranean Sea). Red coral is endemic to the Mediterranean Sea and the adjacent NE Atlantic coast, where it has been over exploited for centuries. This study presents, the first quantitative data on the spatial distribution and structure of a population extending between 50 (common SCUBA limits) and 230 m depth, and compared it with shallow populations previously studied in the same area. Different remotely operated vehicles (ROV) and two methodological approaches were employed during four cruises between 2002 and 2006: 1-Extensive surveys: sea to coast transects in which red coral density and patch frequency were recorded; 2-Intensive surveys, in which parameters describing colony morphology were recorded. Most of the hard substrate between 50 and 85 m depth was inhabited by red coral colonies, showing a patch frequency of 8.3 ± 7.9 SD patches per 100 m-transect (total transect area: 34 m²), and within-patch colony densities of 16–376 colonies m⁻² (mean of 43 ± 53 colonies m⁻²). Below 120 m depth red coral was less abundant, and rather than forming dense patches as in shallow water, isolated colonies were more common. The population structure differed between sites that are easily accessible to red coral fishermen, and remote ones (both at similar depth, 60–80 m), as colonies in easily accessible locations were smaller in height and diameter, and showed a less developed branching pattern. At shallower locations (10–50 m depth) the population structure was significantly different from those at deeper locations, due to the heavy harvesting pressure they are exposed to in the shallows. Twenty-five to forty-six percentage of the deeper colonies were taller than 6 cm, while only 7–16% of the shallow water colonies exceeded 6 cm colony height. Forty-six to seventy-nine percentage of the colonies in deeper waters were large enough to be legally harvested, while only 9–20% of the shallow water colonies met the 7 mm legal basal diameter to be collected. The branching pattern was also better developed in deeper colonies, as up to 16% of the colonies showed fourth order branches, compared to less than 1% of the shallow water colonies (of which 96% consisted of only one single branch). The results thus confirm that C. rubrum populations above 50 m depth are exposed to a higher harvesting intensity than deeper populations in the same area.     

Predators selectively graze reproductive structures in a clonal marine organism

Although the fitness consequences of herbivory on terrestrial plants have been extensively studied, considerably less is known about how partial predation impacts the fitness of clonal marine organisms. The trophic role of Caribbean parrotfish on coral reefs is complex: while these fish are important herbivores, as corallivores (consumers of live coral tissue), they selectively graze specific species and colonies of reef-building corals. Though the benefits of parrotfish herbivory for reef resilience and conservation are well documented, the negative consequences of parrotfish grazing for coral reproductive fitness have not been previously determined. We examined recently grazed colonies of Montastraea annularis corals to determine whether grazing was positively associated with coral reproductive effort. We measured gonad number, egg number and size, and proportional reproductive allocation for grazed and intact coral colonies 2–5 days prior to their annual spawning time. We found that parrotfish selectively grazed coral polyps with high total reproductive effort (number of gonads), providing the first evidence that parrotfish selectively target specific tissue areas within a single coral colony. The removal of polyps with high reproductive effort has direct adverse affects on coral fitness, with additional indirect implications for colony growth and survival. We conclude that chronic grazing by parrotfishes has negative fitness consequences for reef-building corals, and by extension, reef ecosystems.     

Mutualist-induced morphological changes enhance growth and survival of corals

Species interactions can induce morphological changes in organisms that affect their subsequent growth and survival. In Moorea, French Polynesia, epibiotic gammaridean amphipods induce the formation of long, branch-like coral “fingers” on otherwise flat, encrusting, or plating Montipora coral colonies. The fingers form as corals encrust tubes built by the amphipods and lead to significant changes in colony morphology. This study examines the costs and benefits of this association to the amphipods and corals and demonstrates that the interaction is a mutualism. Amphipods gain protection from predators by living within corals, and corals benefit by enhanced growth and survival. Benefits to the coral arise through direct effects due to the amphipods’ presence as well as through benefits derived from the altered colony morphology. This study demonstrates that induced morphological plasticity can be a mechanism for facilitation, adding to our knowledge of the roles mutualism, and phenotypic plasticity play in ecology.     

Effects of spatial variability and colony size on the reproductive output and gonadal development cycle of the Mediterranean red coral (Corallium rubrum L.)

Red coral (Corallium rubrum, L. 1758) is an over-exploited Mediterranean gorgonian. The gonadal development cycle of this gorgonian is examined at the Costa Brava (NW Mediterranean) taking into account for the first time colony size, depth and spatial horizontal variability. This study compares the gonad development and fertility in two colony size classes (colonies <6-cm height, and >10-cm height, both at 40–45-m depth), and two populations at different depths (16–18-m depth, and 40–45-m depth, both consisting of <6-cm high colonies) in a 15-month period. The fertility of seven size classes (<2 cm to >12 cm high colonies, in 2 cm intervals) was examined in the deep population, where large colonies were present. Furthermore, reproductive output was compared in 6 populations (distributed along more than 70-km coastline) one month before spawning (June). Red coral was found to be dioecious and gonochoric with a sex ratio of 1:1, which differs from other NW Mediterranean populations. On the other hand, fertility of different size classes indicates that small colonies of 2-cm height already produce gonads, which is in line with previous studies. Female and male polyp fertility and sperm sac size increase significantly with colony size [sperm sac diameter: 476±144 μm (mean±SD) and 305±150 μm in the >10-cm and <6-cm height colonies, respectively), whereas no significant effect on oocyte diameter was found (oocyte diameter: 373.7±18.7 μm). Depth staggered spawning, that is, an earlier release of gonads in the shallow populations, was observed in summer 2003, coinciding with the highest temperature gradient between shallow and deep water during the study period. Colonies of <6-cm height were significantly less fertile than colonies >12 cm, thus the recommendation of this study is that a minimum height should be incorporated into fishing regulations. The six studied populations at the Costa Brava showed a comparable reproductive potential, which demonstrates little variability within the homogenous population structure and range of size classes (due to overharvesting) found at the Costa Brava. The study of reproductive output is an important tool for ecosystem management, and this work recommends basing specific exploitation laws for distinctive populations on colony size, which is found to have a larger effect on reproductive potential than mesoscale variability. An erratum to this article can be found at     

The importance of life stage to population connectivity in whiting (Merlangius merlangus) from the northern European shelf

Juvenile movements are rarely considered in studies of connectivity. This study investigates movements in juvenile and adult life stages of an ecologically important temperate piscivore, the whiting (Merlangius merlangus). Adult movement was assessed from tag-recapture data collected between 1958 and 1980 around the coast of Scotland. Displacement distances ranged from 53 to 123 km. Adults rarely dispersed offshore in the North Sea, and exchange with the west of Scotland was negligible. Connectivity between nursery and spawning grounds was investigated using otolith microchemistry. Five juvenile nursery areas were discriminated with an accuracy of 75–85%. Comparing isotopic signatures of adults and juveniles indicated that several nursery areas contributed to each spawning aggregation. The lack of evidence for return migrations suggested an opportunistic and non-philopatric recruitment strategy within a single population unit. Differences in the proportion contribution of juveniles to adult aggregations also indicated that the “value” of nursery areas differed and the Scottish west coast could be viewed as a net source of recruits to the North Sea. Given the extent of juvenile dispersal, this study highlights the fact that all stages of the life cycle should be considered in studies of connectivity.     

Reproduction and recruitment of scleractinian corals in a high-latitude coral community, Amakusa, southwestern Japan

Reproduction and recruitment in high-latitude coral populations in Japan have been little studied. A comprehensive study of the reproduction and early life history was conducted on nine common scleractinian coral species in Amakusa, southwestern Japan (32°N) from 2001 to 2003 including; (1) fecundity (the proportion of colonies with mature eggs), (2) timing and synchrony of spawning, (3) initial larval settlement pattern, (4) recruitment, (5) post-settlement mortality. The fecundity was high (76.7–100%) in six of seven species examined in 2002 and 2003. Annual spawning of the seven species occurred from mid July to August in 2001–2003, when seawater temperature was at the annual maximum. Spawning was highly synchronised among conspecific colonies and species in 2002 and 2003, with five species spawning five to nine nights after the full moon and another two spawning around the new moon. Temporal patterns of larval settlement of three spawning species during the first 10 days after spawning were similar to those of other spawning species from low latitudes. The number of scleractinian recruits on settlement plates, deployed from July to October (the major recruitment period at the study site), was low (2 recruits/m²) for the three consecutive years. Post-settlement mortality of 1–1.5 month old spat of five species ranged between 88 and 100% over 3–10 months in the field, similar to the values reported for both high and low latitude species (>94–99%). Among the key stages examined, the low recruitment rate may be the most important step in limiting successful reproduction and recruitment of these high-latitude scleractinian populations. The low recruitment rate may be attributable to (1) the reduced influx of larval supply from other coral populations, which are smaller and more isolated at high-latitudes and (2) the longer precompetent larval phase of broadcast-spawning corals which results in an increased chance of larvae being dispersed away from parent populations.     

Do corals select zooxanthellae by alternative discharge?

Loss of zooxanthellae (dinoflagellate Symbiodinium) from corals will sometimes lead to mass mortality of corals. To detect and quantify Symbiodinium released from corals, we developed a zooxanthellae “trap” and a quantitative PCR (qPCR) system with Symbiodinium clades A–F-specific primer sets. The trap was attached to a branch or the surface of several wild stony corals, and the water samples within the traps, including released Symbiodinium, were subjected to qPCR. All tested corals released clade C Symbiodinium at estimates of ~5,900 cells h⁻¹ cm⁻² of coral surface. Although all tested Pocillopora eydouxi harboured both clades C and D, some of these colonies released only clade C or released a lesser amount of clade D than that in the tissues. Our Symbiodinium quantification system revealed that wild hermatypic corals constantly release Symbiodinium to the environment. Our result suggests that some corals may discharge certain clades of Symbiodinium alternatively.     

δ<Superscript>13</Superscript>C and δ<Superscript>15</Superscript>N values in reef corals <Emphasis Type="Italic">Porites lutea</Emphasis> and <Emphasis Type="Italic">P. cylindrica</Emphasis> and in their epilithic and endolithic algae

In summer 1998, shallow water corals at Sesoko Island, Japan (26°38′N, 127°52′E) were damaged by bleaching. In August 2003, partially damaged colonies of the massive Porites lutea and the branching P. cylindrica were collected at depths of 1.0–2.5 m. The species composition of epilithic algal communities on dead skeletal surfaces of the colonies (‘red turfs’, ‘green turfs’, ‘red crusts’) and the endolithic algae (living in coral skeletons) growing close to and away from living coral polyps was determined. Carbon and nitrogen stable isotope values of organic matter (δ¹³C and δ¹⁵N) from all six of these biological entities were determined. There were no significant differences in the isotope composition of coral tissues of the two corals, with P. lutea having δ¹³C of −15.3 to −9.6‰ and δ¹⁵N of 4.7–6.1‰ and P. cylindrica having similar values. Polyps in both species living close to an interface with epilithic algae had similar isotope values to polyps distant from such an interface. Despite differences in the relative abundance of the algal species in red turfs and crusts, their δ¹³C and δ¹⁵N values were not significantly different from each other (−18.2 to −13.9, −20.6 to −16.2, 1.1–4.3, and 3.3 to 4.9‰, respectively). The green algal turf had significantly higher δ¹³C values (−14.9 to −9.3‰) than that of red turfs and crusts but similar δ¹⁵N (1.2–4.1‰) to the red algae. The data do not suggest that adjoining associations of epilithic algae and coral polyps exchange carbon- and nitrogen-containing metabolites to a significant extent. The endolithic algae in the coral skeletons had δ¹³C values of −14.8 to −12.3‰ and δ¹⁵N of 4.0–5.4‰. Thus they did not differ significantly from the coral polyps in their carbon and nitrogen isotope values. The similarity in carbon isotope values between the coral polyps and endolithic algae may be attributed to a common source of CO₂ for zooxanthellae and endolithic algae, namely, from respiration by the coral host. While it is difficult to fully interpret similarity in the nitrogen isotope composition of coral tissue and of green endolithic algae and the difference in δ¹⁵N between green epilithic and endolithic algae, the data are consistent with nitrogen-containing metabolites from the scleractinian coral serving as a significant source of nitrogen for the endolithic algae.     

Rafting of reef corals and other organisms at Kwajalein Atoll

Studies conducted at Kwajalein Atoll (9°N; 168°E) in early 1988 reveal that marine organisms are commonly rafted into the area on drift pumice, drift wood and other flotsam. Coralskeletons on pumice provide the most useful quantitative data because they persist after rafted motile organisms have departed and rafted sessile forms have decomposed or been scavenged. The estimated minimum number of pumice fragments carrying corals into Kwajalein Atoll during its geological history is on the order of 10⁹, with a more realistic estimate in excess of 10¹¹. The estimated number of coral colonies rafted into the atoll would probably have to be increased several-fold if rafting on floating organic materials such as wood, charcoal, nuts and seeds could be determined. In the present study, a typical sample of beached pumice from Kwajalein Atoll contained 10³ coral colonies per m³ of bulk pumice fragments. Major pumice rafting episodes frequently result from volcanic eruptions. Estimates based on the observed K wajalein coral fouling rate and literature values for drift pumice production suggest that 10⁵ colonies could be rafted through the tropics by the smallest reported pumice-producing events and up to 10¹² colonies for large events. In the present study, most of the rafted corals that were recovered belong to species of the genus Pocillopora, but speccics of Porites and Millepora were also collected. Analysis of surface-current data and reported drift patterns for pumice, logs, drift bottles and wrecks suggests that movement of rafted corals and drifting larvae is predominantly from peripheral areas of low coral-species diversity into centers of high coral diversity. Coral diversity centers might be viewed as areas of coral species accumulation rather than centers of coral species origin.     

Phylogenetic analyses of potentially free-living <Emphasis Type="Italic">Symbiodinium</Emphasis> spp. isolated from coral reef sand in Okinawa,Japan

The existence of “free-living” Symbiodinium that can form symbioses with hosts is implied by the presence of hosts that produce Symbiodinium-free gametes and expulsion and/or expelled symbiotic algae from host. However, it is still unclear if potentially symbiotic Symbiodinium are found “free-living” in the coral reef environment. Sixteen Symbiodinium strains were established from samples taken from three sampling locations of coral reef sand in Okinawa, Japan. Phylogenetic analyses of the partial large subunit ribosomal DNA (28S-rDNA) and the internal transcribed spacer of ribosomal DNA (ITS-rDNA) conclusively showed that all 16 isolates belonged to Symbiodinium clade A sensu Rowan and Powers (1991). The lack of other Symbiodinium clades besides clade A in this study may be due to other clades not being readily culturable under culture conditions used here. The new isolates could be phylogenetically divided into four groups, though no sequences were identical to previously reported Symbiodinium. Two of the four groups were closely related to symbiotic Symbiodinium clade A isolated from a variety of host species. One isolate group formed a highly supported monophyly with Symbiodinium types that have previously been characterized as “free-living”. The remaining isolate group, although within clade A, was quite divergent from other clade A Symbiodinium. These results indicate that novel diversity of free-living Symbiodinium exists in coral sand.     

Phase-shift in coral reef communities in the Florida Keys National Marine Sanctuary (FKNMS), USA

Characterizing the Florida Keys National Marine Sanctuary (FKNMS), USA, has gained much attention over the past several decades because of apparent changes in the benthic community structure over space and time representative of patterns occurring in the Caribbean region. We used a 5-year dataset (1996–2000) of macroalgal and sponge cover and water quality measurements as predictor variables of hard coral community structure in the FKNMS. The 16 water quality variables were summarized into 4 groups by principal component analysis (PCA). Hierarchical agglomerative cluster analysis of the mean and standard deviation (SD) of the principal component scores of water quality variables separated the reef sites into two main groups (and five sub-groups), referred to as reefs of similar influence (RSI). The main groups corresponded with their geographical locations within the Florida Keys: the reefs in the Upper and Middle Keys being homogeneous and collectively, having lower water quality scores relative to reefs in the Lower Keys. Canonical correspondence analysis (CCA) between hard coral cover and key predictor variables (i.e., water quality, macroalgal cover and sponge cover) also separated the reefs in the Lower Keys from reefs in the Upper–Middle Keys, consistent with results of the cluster analysis, which categorized reefs based on RSI. These results suggest that the prevailing gradient of predictor variables may have influenced the structuring of coral reef communities at a spatial scale larger than the individual reef. Furthermore, it is conceivable that these predictor variables exerted influence for a long time rather than being a recent event. Results also revealed a pattern showing reduction in hard coral cover and species richness, and subsequent proliferation of macroalgae and sponges during the study period. Our analyses of the Florida Keys present a pattern that is consistent with the characteristics of a reef that has undergone a “phase-shift,” a phenomenon that is widely reported in the Caribbean region.     

Coral mortality following extreme low tides and high solar radiation

Extreme tidal events are one of the most predictable natural disturbances in marine benthic habitats and are important determinants of zonation patterns in intertidal benthic communities. On coral reefs, spring low tides are recurrent disturbances, but are rarely reported to cause mass mortality. However, in years when extremely low tides coincide with high noon irradiances, they have the potential to cause widespread damage. Here, we report on such an event on a fringing coral reef in the central Great Barrier Reef (Australia) in September 2005. Visual surveys of colony mortality and bleaching status of more than 13,000 corals at 14 reef sites indicated that most coral taxa at wave-protected sites were severely affected by the event. Between 40 and 75% of colonies in the major coral taxa (Acropora, Porites, Faviidae, Mussidae and Pocilloporidae) were either bleached or suffered partial mortality. In contrast, corals at wave-exposed sites were largely unaffected (<1% of the corals were bleached), as periodic washing by waves prevented desiccation. Surveys along a 1–9 m depth gradient indicated that high coral mortality was confined to the tidal zone. However, 20–30% of faviid colonies were bleached throughout the depth range, suggesting that the increase in benthic irradiances during extreme low tides caused light stress in deeper water. Analyses of an 8-year dataset of tidal records for the area indicated that the combination of extended periods of aerial exposure and high irradiances occurs during May–September in most years, but that the event in September 2005 was the most severe. We argue that extreme low-tide, high-irradiance events are important structuring forces of intertidal coral reef communities, and can be as damaging as thermal stress events. Importantly, they occur at a time of year when risks from thermal stress, cyclones and monsoon-associated river run-off are minimal.     

Bacterial colonization and endocytosis on the gill of a new limpet species from a hydrothermal vent

Calcification rates in different fragments along branches of the hermatypic coral Stylophora pistillata were tested in the laboratory using a new technique, the optic glassfiber method. By this method, the tested colony remains constantly in dark conditions while a narrow beam of light, transferred through the optic fiber, illuminates a small distinct point of coral tissue (on a branch tip or base). The selected illuminated portion of the branch serves as the experimental fragment, while all the other parts of the same colony serve as the dark controls. The results indicate that significantly more calcium is incorporated in the tip fragments than in the bases, both in light and in dark conditions (4.1 to 13.2 times more). Illumination of the tips or the bases did not stimulate or enhance the calcification rates of these fragments. Thus, in all colonies tested, the calcification rates of the illuminated fragments were not significantly different from the average rates of other similar, non-illuminated fragments of the same colony. It is suggested that light does not directly enhance calcification in hermatypic corals, but rather, that light enhances O₂ production, which consequently stimulates coral metabolism. Our preliminary results indicate that calcification rates recorded in aerated dark experiments are significantly higher than calcification rates of non-aerated dark controls.     

Polymorphism in soft coral larvae revealed by amplified fragment-length polymorphism (AFLP) markers

The dioecious Red Sea soft coral Parerythropodium fulvum fulvum breeds its nonsymbiotic planula larvae on the surface of female colonies for less than a week. After completing their development, larvae crawl and settle near maternal colonies. Here we study the genetic polymorphism of developing larvae by the use of amplified fragment-length polymorphism markers. Four reproductive colonies from shallow water populations (two from a dense population and two from a less densely populated area 100 m away) were chosen, and ten larvae were randomly collected from each colony. DNA was analyzed by using three different primer combinations producing 61, 63, 63 polymorphic markers, respectively. All larvae exhibited different banding patterns from one another, illustrating the prominent role of sexual reproduction for the production of larvae. Nei's mean genetic distances for all 12 possible pair-wise combinations for larval origins revealed, in most cases, that sister larvae are genetically closer than larvae from different colonies and that larvae may be grouped into three statistical clusters in accordance with colony origin and population studied. The usefulness of molecular methodologies in coral population genetics is discussed. Received: 26 March 1999 / Accepted: 19 October 1999     

Reproductive ecology of the azooxanthellate coral <Emphasis Type="Italic">Tubastraea coccinea</Emphasis> in the Equatorial Eastern Pacific: Part V. Dendrophylliidae

The reproductive ecology of Tubastraea coccinea Lesson, an azooxanthellate tropical scleractinian coral, was studied over various periods from 1985 to 2006 at four principal eastern Pacific locations in Costa Rica, Panamá, and the Galápagos Islands (Ecuador). This small (polyp diameter 0.8–1.0 cm), relatively cryptic species produced ova and planulae year round, including colonies with as few as 2–10 polyps. Of 424 colonies examined histologically, 13.7% contained both ova and sperm. Mature ova varied in diameter from ∼300 to 800 μm and the time from spawning and fertilization of oocytes to release of brooded planulae was about 6 weeks. Planulae were 0.5–1.5 mm long and they settled and metamorphosed on a variety of substrates after 1–3 days. Spermaries, though more difficult to distinguish in histological sections, were present throughout the year. Spent spermaries were never observed in sections, but several colonies in Panamá and the Galápagos Islands released sperm from night one to night five after full moon, indicating the potential for cross-fertilization among colonies. Planula release was observed at Uva Island (Panamá) in March, May, June, and July, and in general planula presence was higher at warm ocean temperatures at all sites, whether or not the sites were influenced by seasonal upwelling. Annual fecundity estimates for T. coccinea are comparable with other high fecundity brooding species, including the zooxanthellate Porites panamensis, with which it co-occurs in Panamá. Tubastraea coccinea is widely distributed in the tropical Indo-Pacific and has colonized substrates in the western Atlantic. In addition to the reproductive characteristics described in the present study, other features of the biology of T. coccinea, such as an ability to withstand conditions that produce bleaching and mortality in zooxanthellate species, may account for its widespread, low-latitude distribution in multiple oceans. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users. An erratum to this article can be found at     

Population structure, growth and recruitment of the euryalinid brittle-star Astrobrachion constrictum (Echinodermata: Ophiuroidea) in Doubtful Sound, Fiordland, New Zealand

The euryalinid brittle-star (snake star) Astrobrachion constrictum (Farquhar) lives coiled around the branches of black coral (Antipathes fiordensis) colonies. Twenty-two vertical transects, 10 m wide by 30 m deep, were swum in Doubtful Sound over a 2.5 yr period from 1993 to 1995. Numbers, disc diameters and colour morphotypes of brittle-stars inhabiting coral colonies were recorded. 36.3% of the coral colonies >200 mm tall (n = 292) hosted ≥1 Astrobrachion constrictum (range 0 to 12). Overall, the population was patchily distributed on the available coral habitat. The dark red colour morphotype of A. constrictum was most common (87%, n = 279) followed by the yellow, striped, and then spotted varieties. The population was comprised mainly of large (≥10 mm disc diam) individuals, and juveniles were rarely encountered, indicating low rates of recruitment or a high mortality of recruits. Disc-diameter data gathered from this and previous studies indicated that growth in A. constrictum is initially rapid, with individuals reaching a disc diameter of 15 mm in ≃2.5 yr; growth decreases with age, as in other deep-sea ophiuroids. Growth rate within years, however, was not constant, with faster growth in the spring/summer. Maximum size for A. constrictum is reached in ≃8 yr at ˜23 mm disc diam. Anecdotal evidence indicates that A. constrictum may not be confined solely to black coral colonies. Received: 25 September 1996 / Accepted: 16 October 1996