The effects of Hurricane Allen on Back Reef populations of Discovery Bay,Jamaica

Sixteen months after Hurricane Allen, an assessment of the condition (living vs. dead and encrusted) and volume of staghorn coral, Acropora cervicomis Lamarck, patches within the East Back Reef of Discovery Bay, Jamaica was made. Data generated by this assessment were compared with similar data collected in 1975–1976 prior to the storm. Densities of two urchins, Diadema antillarum Philippi and Echinometra viridis A. Agassiz, and the threespot damselfish, Eupomacentrus planifrons Cuvier, within the coral patches were also measured.Although staghorn coral patches were significantly smaller (? 65%) in mean volume in 1981 compared to 1975–1976, 22% of the patches were unchanged since 1976 or had increased in volume and only 9% were reduced to piles of rubble. Diadema and threespot densities were significantly higher than in 1976. Mortality of damselfish and larger Diadema appeared to have been reduced. Coral patches with both damselfish and Diadema present exhibited a high proportion of living coral tissue, while those patches dominated by either damselfish or Diadema were overgrazed with < 5% of the substrata covered by living coral. Similarly, the fore reef exhibited high urchin and low damselfish densities, possibly contributing to its low proportion of living coral.     

Interference behavior and ecology of threespot damselfish (Eupomacentrus planifrons)

Summary Interference behavior by threespot damselfish directed at the sea urchins, Diadema antillarum and Echinometra viridis, differed in levels of aggression and discrimination. Higher aggression towards and recognition of Diadema by threespots was correlated with the distribution of Diadema along edges of coral patches. Lower aggression levels combined with lower recognition levels of Echinometra were correlated with a distribution of Echinometra closer to damselfish algal lawns. Differences in behavior of the urchins stimulated differing levels of aggression by the damselfish directly affecting the distribution of the urchins in the back-reef environment.This is contribution number 167 from the Discovery Bay Marine Laboratory of the University of the West Indies, Jamaica     

Mass mortality of Diadema antillarum on the Caribbean coast of Panama

The ecologically important sea urchin Diadema antillarum suffered mass mortalities in 1983, first noted in Panama and then reported from the rest of the Caribbean. We documented the effects of this mortality at two localities on the Atlantic coast of Panama, Punta Galeta and the San Blas Archipelago. At Punta Galeta, affected by the mortality in January 1983, the numbers of D. antillarum changed from an estimated 14,000 per ha in June 1982 to 0.5 per ha in May 1983; by February 1984 they had increased to 38 per ha. In the San Blas, where mass mortality started in April 1983, the number of D. antillarum in permanent quadrats on 8 reefs was reduced by an average of 94.2%. The average reduction in population density measured in transects on nine reefs was 98.9%. Data taken in permanent quadrats on four reefs in 1978, 1979 and 1980 indicate that population fluctuations of D. antillarum are normally much smaller, justifying the labeling of the 1983 event as mass mortality. Size structure of the San Blas populations was also affected; mean test diameter of D. antillarum on four reefs was reduced from 48.6 mm to 25.0 mm. Other echinoids (Echinometra viridis, E. lucunter, Lytechinus variegatus, L. williamsi, Eucidaris tribuloides, Tripneustes ventricosus, Clypeaster rosaceus and Echinoneus cyclostomus) suffered no ill effects at either Galeta or the San Blas; their population densities remained stable or increased. Density determinations of Diadema mexicanum at the island of Taboguilla on the Pacific side of Panama indicate that Diadema mass mortality did not extend to the eastern Pacific. Sea surface temperatures, tidal levels, rainfall and salinity showed no abnormal fluctuations during the time of D. antillarum mass mortality at Galeta, suggesting that mortality was not due to physical stress. The wide geographical spread and species-specificity of the mortality suggest a water-borne pathogen as the most likely causative agent. Recovery of D. antillarum populations is likely to be slow because there are few, if any, unaffected populations in the Caribbean to contribute larvae for the recolonization of depleted areas. The absence of D. antillarum will probably be reflected by changes in the algal, coral and echinoid communities, and by altered patterns of bioerosion.     

Echinoid grazing as a structuring force in coral communities: Whole reef manipulations

To examine the effects of echinoid grazing on shallow water coral community structure, the entire Diadema antillarum Philippi population (> 3000 individuals) was eliminated from a patch reef in Discovery Bay, Jamaica W.I. in January 1974. All remaining regular echinoids, notably including Echinometra viridis A. Agassiz (> 7000 individuals), were removed from one-half of the same reef 6 months later.Where Echinometra was present, its grazing was highly patchy, even at densities of up to 50/m². On the other hand, the effects of grazing by Diadema were more uniformly distributed at equivalent or lower densities.Percent-cover of adult corals was enhanced in the presence of Echinometra viridis, however, it was greatly reduced in the absence of all echinoid grazing as a result of massive overgrowth by algae. Success of coral recruitment (density of coral spat) increased in the absence of Diadema due to the alleviation of biological disturbance imposed by this urchin's feeding activities. The presence of Echinometra viridis, even at the high densities naturally present on the reef, did not deter coral settlement.Coral diversity was highest in the presence of all echinoids, intermediate in the presence of E. viridis (without Diadema), and lowest in the absence of all echinoids. This was due to the disproportionately successful recruitment and resultant high relative abundance of Agaricia at low echinoid densities. Favia, Millepora, Helioseris and Acropora all exhibited universally low recruitment levels. Predation by Diadema and Echinometra is responsible for damaging larger coral colonies, particularly Porites.As echinoid species composition and density was altered, the species composition of benthic algae and associated invertebrates shifted. In this way, the array of species competing for space with coral spat shifted as did their competitive relationships. Major epifaunal competitors included spirorbid polychaetes, various foraminifera (particularly Gypsina), and sponges. Algal competitors included filamentous chlorophytes, corallines (e.g. Jania), and numerous other red algae; the species composition of algal competitors shifted through time. Competitive success in corals varied as echinoid density changed and was species-specific with respect to coral genus. Interspecific competition between coral spat (i.e., interspecific aggression via extracoelenteric digestion) was negligible under all treatments.The presence of Echinometra viridis in high densities (in the absence of Diadema) created conditions where growth and fusion in Agaricia spat were optimized. Under these conditions, fusion allows a young coral colony to attain a large enough size to survive damage incurred from grazing or from competitive overgrowth, decreasing the probability of whole-colony mortality during the early stages of development.     

Predation and the Control of the Sea Urchin Echinometra viridisand Fleshy Algae in the Patch Reefs of Glovers Reef, Belize

The massive reduction in sea urchin Diadema antillarum populations since the mid-1980s has been associated with large increases in the abundance of fleshy algae on many Caribbean reefs despite the availability of other sea urchin and finfish grazers. This study examined the ecology of a grazer living sympatrically with D. antillarum, the common and abundant sea urchin Echinometra viridis. I examined the role that finfish and invertebrate predators play in controlling the distribution of E. viridis as well as the ability of this sea urchin to control exposed fleshy algae on the patch reefs of the Glovers Reef Atoll lagoon. I found that the major predators of this sea urchin were Calamus bajonado (jolthead porgy), Balistes vetula and Canthidermis sufflamen (queen and ocean triggerfish), Lachnolaimus maximus (hogfish), and a gastropod, probably Cassis madagascariensis. The abundance of E. viridis is constrained by predation, which restricts E. viridis to cryptic locations, such as crevices. Sea urchins bit a smaller percentage of experimental algal assays than finfish. Finfish herbivory was associated positively with patch reef topographic complexity. Unexpectedly, E. viridis abundance was positively correlated with fleshy algal abundance, but negatively correlated with the frequency of finfish bites. Predators restrict E. viridis to crevices and therefore reduce their influence on exposed fleshy algae, even at moderately high population densities (up to 10 per square meter). Since net benthic primary production of coral reefs is most strongly associated with herbivory on exposed surfaces, it would appear that E. viridis is unable to maintain the same production as reefs dominated by D. antillarum. Received 5 November 1998; accepted 2 June 1999.     

Evidence for positive density-dependent effects in recovering Diadema antillarum populations

Recovering populations may experience positive density-dependent feedbacks that contribute to population increases. Diadema antillarum, a keystone herbivore on Caribbean coral reefs, suffered a well-documented mass mortality in 1983-84. High densities of adults of this long-spined urchin could provide effective refuge from predation for juveniles under a spine canopy, as has been suggested for other urchin species. We evaluated the effect of adult density on juvenile persistence of D. antillarum experimentally, and examined size-frequency distributions of recovering local populations for evidence of positively density-dependent juvenile persistence at St. Croix, U.S. Virgin Islands. Juvenile persistence was significantly higher in high adult density treatments, and bimodal population size distributions also suggest potential positive effects of adult density on juveniles. This positive feedback could accelerate the recovery of this important coral reef grazer.     

On the dynamics of the mass mortality of Diadema antillarum in Barbados

Widespread mortality of the black sea urchin Diadema antillarum occurred in the Caribbean in 1983; beginning in Panama in January, and having its major impact at Barbados in September. Mortality on ten reefs surveyed in Barbados was 93.2%, with the highest being 99.9% and the lowest 86.9%. Mortality on each reef was independent of the pre-mortality density on the reef. Urchins with test diameters between 20 and 40 mm were more severely affected than smaller or larger urchins. Populations on reefs exposed to incoming oceanic water suffered heavier mortality than those on protected reefs. Mean size of urchins was smallest on high density reefs. This may indicate a negative effect of density on urchin growth. At post-mortality densities, urchins may grow faster and reach sexual maturity sooner.     

GAMETIC INCOMPATIBILITY BETWEEN SPECIES OF THE SEA URCHIN ECHINOMETRA ON THE TWO SIDES OF THE ISTHMUS OF PANAMA

The Pliocene rise of the Central American Isthmus has resulted in numerous “geminate pairs,” i.e., closely related species, one on each coast. Such species pairs can provide information on the evolution of isolating mechanisms in allopatry and on the relationship between genetic divergence and reproductive isolation in populations separated at a known time. The sea urchin genus Echinometra has one species, E. vanbrunti, in the eastern Pacific, and two, E. lucunter and E. viridis, in the Caribbean. E. viridis is morphologically distinct from the other two species, leading to the conclusion that E. lucunter and E. vanbrunti constitute a geminate pair. Allozyme data, on the other hand, place the speciation event of the two currently sympatric species after the rise of the Isthmus. We report fertilization experiments between the gametes of the three species performed to determine degree of reproductive isolation. Crosses between E. viridis and E. vanbrunti produce rates of fertilization almost equal to those manifested in homogamic crosses. Sperm of E. lucunter can fertilize eggs of the other two species, but few of its eggs permit fertilization by heterospecific sperm. Contrary to the predictions of the “speciation by reinforcement” hypothesis, degree of incompatibility between the allopatric E. lucunter and E. vanbrunti is higher than between the sympatric E. lucunter and E. viridis. Despite the incomplete and unidirectional nature of their gametic isolation, E. lucunter and E. viridis maintain their genetic integrities. Consideration of the likely phylogenetic relationships between the three species suggests that incompatibility of E. lucunter eggs with heterospecific sperm has evolved in the last 3.5 million years, after the rise of the Isthmus. There is no correlation between genetic divergence and strength of reproductive isolation, either within Echinometra, or among the geminate species pairs of echinoids that have been studied to date. Because recognition between echinoid gametes depends on the chemical compositions of a sperm protein and an egg glycoprotein, the appearance of gametic isolation would require only the fixation of a few mutations in each population on either side of a geographic barrier and could be independent of any other kind of genetic divergence. Thus, in animals with external fertilization, speciation need not be accompanied by major genomic reorganization.     

Predation on the urchin Echinometra lucunter (Linnaeus) by migratory shorebirds on a tropical reef flat

Predation on urchins by migratory shorebirds was investigated during tidal exposure of the reef flat at Galeta, Panama. Avian predators, primarily ruddy turnstones Arenaria interpres (Linnaeus) reduced the patchiness of Echinometra lucunter (Linnaeus), the most abundant urchin on the reef flat. No mortality of E. lucunter occurred in quadrats protected from avian predation. No mortality occurred when urchin crowding was increased experimentally in protected areas. Stress-induced mortality could not be determined from the condition of tests alone, since turnstones were capable of removing lantern and internal organs without damage to the test. The importance of shorebird predation in reef communities during aerial exposure can be hard to detect because mortality can occur during brief periods, at night, and without damage to urchin tests.     

Shell utilization,predation pressure,and thermal stress in panamanian hermit crabs: An interoceanic comparison

Shell preference patterns of two common hermit crabs from hard bottom reef flats on the Caribbean coast of Panama are examined in relation to the predation pressures and physical stresses of their habitat. Clibanarius antillensis Stimpson lives in the high intertidal habitat and minimizes exposure to predators by seeking refuge during high tides. It prefers high-spired shells which maximize protection from thermal stress. Calcinus tibicen Herbst avoids tidal emersion and prefers low-spired shells which enhance resistance to the predators common on Caribbean reef flats.The results are compared with similar results from the tropical eastern Pacific Bay of Panama. Shell-crushing predation on Caribbean hermit crabs is suggested to differ quantitatively and qualitatively from predation on hermit crabs in the Bay of Panama. Predation on hermit crabs in the Bay of Panama is more intense and effects larger individuals than predation on Caribbean reef flat hermit crabs. In addition, shell-crushing predation on hermit crabs in the Bay of Panama is primarily from teleost fish predators (Diodon spp.), while predation on Caribbean hermit crabs is primarily by bottom-dwelling crustaceans.Differences in predation pressures and tidal regimes between the Caribbean and Pacific coasts of Panama are reflected in the shell preferences and behavior of hermit crabs from the two areas.     

DNA barcoding of echinopluteus larvae uncovers cryptic diversity in neotropical echinoids

Surveys of larval diversity consistently increase biodiversity estimates when applied to poorly documented groups of marine invertebrates such as phoronids and hemichordates. However, it remains to be seen how helpful this approach is for detecting unsampled species in well‐studied groups. Echinoids represent a large, robust, well‐studied macrofauna, with low diversity and low incidence of cryptic species, making them an ideal test case for the efficacy of larval barcoding to discover diversity in such groups. We developed a reference dataset of DNA barcodes for the shallow‐water adult echinoids from both coasts of Panama and compared them to DNA sequences obtained from larvae collected primarily on the Caribbean coast of Panama. We sequenced mitochondrial cytochrome c oxidase subunit I (COI) for 43 species of adult sea urchins to expand the number and coverage of sequences available in GenBank. Sequences were successfully obtained for COI and 16S ribosomal DNA from 272 larvae and assigned to 17 operational taxonomic units (OTUs): 4 from the Pacific coast of Panama, where larvae were not sampled as intensively, and 13 from the Caribbean coast. Of these 17 OTUs, 13 were identified from comparisons with our adult sequences and belonged to species well documented in these regions. Another larva was identified from comparisons with unpublished sequences in the Barcode of Life Database (BOLD) as belonging to Pseudoboletia, a genus scarcely known in the Caribbean and previously unreported in Panama. Three OTUs remained unidentified. Based on larval morphology, at least two of these OTUs appeared to be spatangoids, which are difficult to collect and whose presence often goes undetected in standard surveys of benthic diversity. Despite its ability to capture unanticipated diversity, larval sampling failed to collect some species that are locally common along the Caribbean coast of Panama, such as Leodia sexiesperforata, Diadema antillarum, and Clypeaster rosaceus.     

Physical forces experienced by echinoid eggs in the oviduct during spawning: comparison of the geminate pair Echinometra vanbrunti and Echinometra lucunter

During spawning, eggs of echinoids are released from the gonad and traverse a narrow oviduct-gonopore complex before being released into the water surrounding the spawning female. As the eggs traverse the oviduct-gonopore complex, a velocity gradient develops within the fluid formed by the eggs. This velocity gradient imposes a shear stress on the eggs that ranges from a maximal value at the walls of the oviduct-gonopore complex and declines linearly to zero at the center of the complex. In addition to shear stress, the eggs of some echinoids also experience strain. Strain is imposed on the eggs in situations where the diameter of the egg is greater than the diameter of the oviduct-gonopore complex. Both shear stress and strain have the potential to damage echinoid eggs during spawning and are likely selective pressures acting on the physical properties of protective extracellular layers, which surround the eggs of these organisms. The amounts of shear stress and strain experienced by eggs of a geminate pair of echinoids, Echinometra vanbrunti and E. lucunter, are calculated. Calculations of shear stress based on the mean spawning rates and dimensions of the oviduct-gonopore complexes of specimens collected during this study ranged up to 30 Pa for E. vanbrunti and up to 17 Pa for E. lucunter. To obtain conservative estimates of shear stress, these calculations were repeated incorporating a spawning rate that was reduced by one order of magnitude below the mean rate measured for both species. These calculations yielded shear stresses that ranged up to 13 and 6 Pa for E. vanbrunti and E. lucunter, respectively. The diameter of the oviduct-gonopore complex of echinoids increases linearly with the diameter of the tests of the adults. Because the diameter of the oviduct-gonopore complex influences the amount of shear stress experienced by eggs of echinoids, shear stress calculations were repeated incorporating the full range of potential diameters of the oviduct-gonopore complexes from data in the literature. Using the mean spawning rates obtained from each species, the amount of shear stress experienced by eggs ranged up to 32 Pa for E. vanbrunti and up to 398 Pa for E. lucunter. When the mean spawning rates were reduced by one order of magnitude, the amount of shear stress experienced by E. vanbrunti eggs ranged up to 14 Pa and up to 140 Pa for the eggs of E. lucunter. The eggs of E. vanbrunti do not experience strain during spawning because the diameter of the its eggs is smaller than the diameter of its oviduct-gonopore complex at all stages of adult growth. In contrast, the diameter of the eggs of E. lucunter is larger than the diameter of the oviduct-gonopore complex at small adult sizes and the eggs of this species experience true strains ranging up to 1.2. These results indicate that there are substantial differences in the amounts of physical forces that the eggs of echinoids experience during spawning both within species at different stages of adult growth and between species. Different amounts of shear stress and strain experienced by eggs among echinoid species may select for differences in the mechanical properties of protective extracellular layers that surround them.     

Can one species determine the structure of the benthic community on a temperate rocky reef? The case of the long-spined sea-urchin Diadema antillarum (Echinodermata: Echinoidea) in the eastern Atlantic

We sampled 36 coastal rocky reefs throughout the overall Canarian Archipelago and consider (1) the daily macroalgal consumption of the long-spined sea urchin Diadema antillarum and (2) the daily net production of macroalgae along temperate rocky-substrates, to provide evidence that Diadema antillarum plays an important role in the structure of the shallow benthic environment of the eastern Atlantic. D. antillarum was found to be the main key-herbivore species, as it controls by its own the algal assemblages, with negligible contribution of other grazing species.     

Innate immune response in the sea urchin Echinometra lucunter (Echinodermata)

Echinometra lucunter, (Pindá) is a sea urchin encountered in the Brazilian coast and exposed to high and low temperatures related to low and high tides. Despite their great distribution and importance, few studies have been done on the biological function of their coelomocytes. Thus, Echinometra lucunter perivisceral coelomocytes were characterized under optical and transmission electron microscopy. Phagocytic amoebocytes in the perivisceral coelom were labelled by injecting ferritin, and ferritin labelled phagocytic amoebocytes were found in the peristomial connective tissue after injecting India ink into the tissue, indicating the amoebocytes ability to respond to an inflammatory stimulus. Results showed that the phagocytic amoebocytes were the main inflammatory cells found in the innate immune response of E. lucunter. While other works have recorded these phenomena in sea urchins found in moderate and constant temperature, this study reports on these same phenomena in a tropical sea urchin under great variation of temperature, thus providing new data to inflammatory studies in invertebrate pathology.     

<Emphasis Type="Italic">Diadema antillarum</Emphasis> 17 years after mass mortality: is recovery beginning on St. Croix?

Diadema antillarum was once ubiquitous in the Caribbean, but mass mortality in 1983–84 reduced its numbers by >97%. We measured Diadema abundance on back reefs and patch reefs that have been well studied for >25 years. From June 2000 to June 2001, populations on back reefs have increased >100% (June 2001 mean densities 0.004–0.368/m²), while patch reef populations increased >350% (June 2001 densities 0.236–0.516/m²). Populations are dominated by small urchins, suggesting high recent recruitment. Increased Diadema densities appear to be affecting macroalgae abundance. The general spatio-temporal pattern of recovery around St. Croix seems to be following that of the die-off, suggesting that the same oceanographic features that spread Diadema's pathogen are now carrying urchin larvae.     

Large-scale associations between macroalgal cover and grazer biomass on mid-depth reefs in the Caribbean

Since the 1970s, macroalgae have become considerably more abundant on many Caribbean reefs and overfishing of grazing fishes has been implicated as a contributory factor. We explored relationships between algal cover and grazers (biomass of herbivorous fishes and abundance of the sea-urchin Diadema antillarum) on mid-depth reefs (12-15 m) in 19 areas at seven locations in Jamaica, Barbados, Belize, Grand Cayman and Cuba, between April 1997 and April 1998. Diadema antillarum density was never >0.01 m^-2, while herbivorous fish biomass (acanthurids and scarids ⁸ cm total length) varied from 2-5 g m^-2 in Jamaica to 17.1 g m^-2 in Barbados, and was strongly correlated, negatively with macroalgal cover and positively with 'cropped' substratum (sum of 'bare', turf and crustose-coralline substrata) cover. However, overfishing of herbivorous fishes alone cannot explain the widespread abundance of macroalgae, as even on lightly fished reefs, macroalgal cover was mostly >20%. Herbivorous fish populations on those reefs were apparently only able to maintain approximately 40-60% of reef substratum in cropped states, but due to low space-occupation by coral and other invertebrates, 70-90% of substratum was available to algae. The abundance of macroalgae on lightly fished reefs may therefore be a symptom of low coral cover in combination with the continuing absence of Diadema antillarum.     

What makes a species common? No evidence of density-dependent recruitment or mortality of the sea urchin Diadema antillarum after the 1983–1984 mass mortality

A potential consequence of individuals compensating for density-dependent processes is that rare or infrequent events can produce profound and long-term shifts in species abundance. In 1983–1984 a mass mortality event reduced the numbers of the abundant sea urchin Diadema antillarum by 95–99 % throughout the Caribbean and western Atlantic. Following this event, the abundance of macroalgae increased and the few surviving D. antillarum responded by increasing in body size and fecundity. These initial observations suggested that populations of D. antillarum could recover rapidly following release from food limitation. In contrast, published studies of field manipulations indicate that this species had traits making it resistant to density-dependent effects on offspring production and adult mortality; this evidence raises the possibility that density-independent processes might keep populations at a diminished level. Decadal-scale (1983–2011) monitoring of recruitment, mortality, population density and size structure of D. antillarum from St John, US Virgin Islands, indicates that population density has remained relatively stable and more than an order of magnitude lower than that before the mortality event of 1983–1984. We detected no evidence of density-dependent mortality or recruitment since this mortality event. In this location, model estimates of equilibrium population density, assuming density-independent processes and based on parameters generated over the first decade following the mortality event, accurately predict the low population density 20 years later (2011). We find no evidence to support the notion that this historically dominant species will rebound from this temporally brief, but spatially widespread, perturbation.     

Recruitment pattern of Diadema antillarum in La Parguera, Puerto Rico

Recruitment success is one of the factors that may be influencing the slow population recovery of Diadema antillarum at many locations in the Caribbean. D. antillarum recruitment was measured with recruitment plates at monthly intervals from September 2005 to September 2006 at three reefs along an inshore-offshore gradient in La Parguera, Puerto Rico. A total of 275 recruits of D. antillarum were collected at the shelf-edge reef during this 13-month study. Two recruits were collected at the mid-shelf reef and no recruits were collected at the inner shelf reef. Recruitment varied among months at the shelf-edge, with the highest value (1,067 ind/m²) occurring in July 2006. Previous benthic surveys in the La Parguera region have demonstrated higher densities of adult D. antillarum at inshore reefs. The higher recruitment at the shelf-edge suggests that recruitment is not a major determinant of the spatial distribution of the adult population. Recruitment occurring in this study is indicative that sources of larvae were available upstream and larval survival was occurring.     

Larval influx of Diadema antillarum to the Florida Keys linked to passage of a Tortugas Eddy

Lack of recovery of the sea urchin Diadema antillarum in the Caribbean region following a widespread epizootic in 1983–1984 has garnered great interest due to the role of this species as a grazer of macroalgae that exclude reef-building corals. In the Florida Keys, USA, previous research suggests that reestablishment of D. antillarum is limited by low fertilization success and a lack of larval supply. However, the physical mechanisms determining larval supply in the Florida Keys remain poorly resolved. Here, we use coupled biological and physical oceanographic datasets of D. antillarum larval supply to settlement collectors, sea surface temperatures and heights, and the Okubo Weiss parameter to examine a link between influx of larvae to the middle Florida Keys and Tortugas Eddy activity in the Straits of Florida. The greatest magnitude of settlement over 3 months of measurements occurred from late May to late June 2015, coinciding with the passage and dissociation of a Tortugas Eddy. Settlement occurred on collectors only at offshore bank-barrier reef sites, consistent with a temperature signal of a passing eddy at these sites. No D. antillarum were observed by divers at sites 1 yr following settlement on the collectors. The results indicate that despite the lack of population recovery, D. antillarum larval influx can occur during intermittent oceanographic events.

     

The reproductive biology of the deep-sea asteroids Benthopecten simplex (Perrier), Pectinaster filholi Perrier,and Pontaster tenuispinus Düben & Koren (phanerozonia : benthopectinidae) from the rockall trough

Samples of three species of benthopectinid starfish were collected in a time-series from stations in the northern Rockall Trough. A detailed study was made of the reproductive biology of Benthopecten simplex (Perrier) from a fixed station at 2200 m. Studies of the gametogenic cycles of Pectinaster fllholi Perrier and Pontaster tenuispinus Düben & Koren from less regular and more scattered samplings are also presented.The oogenic cycles of the three species are similar. Growing oocytes line the ovary wall and come to lie in the lumen only when well developed. Accessory cells surrounding young oocytes become more numerous as the oocytes develop. Vitellogenesis begins when oocyte diameter reaches ≈ 250–300 μm and oocytes attain a maximum diameter of 950 μm in Benthopecten simplex, 850 μm in Pectinaster filholi, and 800 μm in Pontaster tenuispinus. Unspent oocytes are broken down by phagocytes and degenerate ovaries become filled with the breakdown products. Internal degeneration of large oocytes is not common.Size frequency data show a dominance of previtellogenic oocytes and a pattern of continuous growth of vitellogenic oocytes. There is no evidence of synchrony of growth between individuals and no suggestion of seasonal reproduction is evident from summated sample data.Spermatogenesis follows the pattern previously established for deep-sea asteroids. A mature male remains in a state of constant ripeness, ready to shed sperm at any chance encounter with a female releasing ripe eggs. There is no evidence of brooding in any of the species and from the large size and yolk content of the egg, we can infer direct lecithotrophic development demersally.