Seasonal cycle of sexual reproduction of the Mediterranean soft coral Alcyonium acaule (Anthozoa, Octocorallia)

Alcyonium acaule (Cnidaria, Octocorallia) is a common, hard-bottom soft coral in the northwestern Mediterranean Sea. This study describes sexual reproduction and the gamete development cycle of this soft coral. A population at 15–18 m depth in the Marine Protected Area of the Medes Islands (42º02′55″ N, 3º13′30″ E) was sampled from July 1994–August 1995. A. acaule is gonochoristic and a surface brooder, spawning once a year in early summer. The mean diameter of ripe spermatic sacs was 400 ± 91 (SD) μm, and the mean diameter of mature oocytes was 473 ± 37 (SD) μm. There were 30 spermatic sacs polyp^?1 in males and 14 oocytes polyp^?1 in females. Different phases of gametogenesis in female and male colonies were examined separately with respect to seasonal changes in bottom temperature and solar irradiance. The data suggest that the relatively constant temperatures in January–April are probably not related to oocyte maturation, but that rising temperatures in May could affect sperm maturation. Rapidly increasing solar irradiance in March may be the trigger for vitellogenesis and oocyte maturation, although the mechanism for this in anthozoans is not understood.     

Stalked tunicates Boltenia ovifera form biogenic habitat in the rocky subtidal zone of Nova Scotia

The stalked tunicate Boltenia ovifera is widely distributed in the Arctic and North Atlantic oceans on rocky substrata at 10–300 m depth, although its ecological role in benthic communities is poorly understood. The distribution and abundance of B. ovifera were recorded at 10–100 m depth in towed-video transects in November 2011 and in June, July, and November 2012 off a wave-exposed headland near Halifax, Nova Scotia. Specimens also were collected at 30 m depth using SCUBA (44°26.88′N, 63°31.59′W) in September 2012. Areas dominated by B. ovifera had densities of 10–100 individuals m^?2 on rocky substrata at 30–60 m depth. These tunicate beds often were associated with sparse kelp (Agarum clathratum) and turf-forming red algae. A generalized additive model indicated that depth, substrate type, and benthic algal type were strong predictors of tunicate abundance. Twenty-two epibiotic species were found on specimens of B. ovifera, including juvenile conspecifics. Filter-feeding macroinvertebrates, including anemones and soft corals, were more abundant in areas with B. ovifera than in areas without these tunicates. Our findings suggest that beds of B. ovifera can act as biogenic habitat to enhance local species richness in the rocky subtidal zone of Nova Scotia.     

Predation on zooplankton by the benthic anthozoans Alcyonium siderium (Alcyonacea) and Metridium senile (Actiniaria) in the New England subtidal

The Alcyonacean octocoral Alcyonium siderium Verrill and the sea anemone Metridium senile (L.), the only common perennial zooplanktivores on shallow (16-m depth) subtidal rock walls in much of northern New England, USA, are of similar heights and overlap in their habitat and microhabit distributions. The coelenteron contents of both species were sampled at four-hour intervals over a diel cycle and were compared to zooplankton available in the water at 1 to 5 cm from the rock wall, the height at which the cnidarians held their feeding tentacles. Prey in coelenterons of A. siderium were significantly smaller (means of 256 to 345 m), and those in coelenterons of M. senile were equal to or slightly greater in length (means of 415 to 1006 m) than the available zooplankton. The diets of A. siderium and M. senile differed significantly from each other and from the available zooplankton. A. siderium showed strong positive electivites for ascidian larvae and for foraminiferans, and strongly negative electivities for most crustaceans. M. senile had strong positive electivities for barnacle cyprids, ascidian larvae, and gammarid amphipods, and strong negative electivities for invertebrate eggs, foraminiferans, calanoid and harpacticoid copepods, and ostracods. Electivities may reflect tentacle avoidance or escape by motile prey as well as predator preference. Substratum-associated organisms (e.g. demersal crustaceans, larvae of benthic invertebrates) were the most common items in the diets of both species, suggesting a tight benthic food web, similar to the situation for coral reef anthrozoans which rely on reef-generated zooplankton. A. siderium ate large numbers of ascidian larvae which, as benthic adults, compete for space with A. siderium and can overgrow small colonies. Predation on the larvae of a competing species may alleviate competition by decreasing the competitor's recruitment.     

Population structure and feeding deterrence in three shallow-water antarctic soft corals

Alcyonium paessleri and Clavularia frankliniana are numerically abundant soft corals in the nearshore (12 to 33 m depth) benthic communities of eastern McMurdo Sound. They are much less abundant in western McMurdo Sound where a third species, Gersemia antarctica, co-occurs in low numbers. The body tissues of these three species are comprised mainly of organic material (53 to 70% dry wt), which is primarily dervied from NaOH-soluble protein and refractory material. The energetic contents of the whole-body tissues of A. paessleri, C. frankliniana and G. antarctica are 15.9, 17.3, and 14.5 kJ g^-1 dry wt, respectively. The mean biomass per individual is 1.81, 0.008, and 45 g dry wt for each respective species. Based on population densities of 7.3, 1337.3, and 0.04 soft corals m^-2 for A. paessleri, C. frankliniana and G. antarctica, respectively, the population energetic densities are estimated to be 210.1, 185.1, and 26.1 kJ m^-2. Despite the relatively rich energetic content of the tissue and apparent vulnerability to predators, very little predation occurs on these soft corals. Two potential predators, the antarctic sea stars Perknaster fuscus and Odontaster validus, exhibited significant chemotactic defensive tube-foot retractions to hexane, chloroform, methanol, and aqueous methanol extracts of each soft coral. In addition, wholebody tissue of each soft coral was rejected by the demersal fish Pseudotrematomus bernacchii and the cryopelagic fish Pagothenia borchgrevinki. In contrast, whole soft-coral tissues sequentially extracted in four increasingly polar solvents were readily ingested by these antarctic fishes, indicating that sclerites do not play a significant role in deterring predators. Our results indicate that these antarctic soft corals contain bioactive compounds which deter common predatory seastars and fishes.     

Long-term trend in substratum occupation by a clonal, carbonate bryozoan in a temperate rocky reef in times of thermal anomalies

Studies over time provide opportunities to detect variations in the spatial and temporal patterns of clonal organisms and measure changes on their population dynamics related to extreme events. We assessed population dynamics for a bryozoan species dominating a subtidal rocky reef at Tino Island, in the eastern Ligurian Sea (NW Mediterranean). Using 9 years of annual photosurveys (1997–2005), rapid decline in Pentapora fascialis colony cover was shown at 11 and 22 m depths following the anomalous warming events in 1999 and 2003. An 86 % reduction in live colony portion was found after the 1999 warming event (2.3 °C higher than normal), with larger colonies being most affected. Effects from the 2003 event were delayed, and gradual cover decline occurred during the following 2 years. At the “Shallow” photostations, none of the larger colonies (>1,000 cm²) survived after the first cover decline. Availability of new substrate after the 1999 disturbance resulted in enhanced recovery through new colony production. At the “Deep” photostations, the population structure did not change over the duration of the monitoring period showing the same monomodal structure and same dominant size class (50–500 cm²). In the 4 years following the first cover decline, the deeper population regained colony cover to levels similar to pre-disturbance level, showing a good resilience. This 9-year monitoring analysis provided the temporal resolution needed to detect changes occurring in the P. fascialis population and will contribute to the assessment of long-term changes on benthic populations suffering during recent decades from dramatic increases in extreme events.     

Bionomische Studien an mediterranen Anthozoen: die Anthozoenfauna des Strombolicchio (Äolische Inseln)

Strombolicchio is a rocky islet situated about 1.6 km northeast of the isle of Stromboli. The islet consists of three steep walls having different light and surface conditions. The zonation on the northern wall differs essentially from the zonation found in caves because of the greater turbulence. On smooth rocks near the water surface, there is only one actinian species which resists turbulence, Actinia equina. The best turbulence resisting anthozoan species are the scleractinians (Astroides calycularis Balanophyllia verrucaria). They are followed by stolonial growing forms (Paralcyonium elegans, Cornularia cornucopiae). All other actinians settle in crevices and small holes. The turbulence prevents Parazoanthus axinellae and Corynactis viridis from settling on the smooth rock near the surface. Therefore, these species occur on Strombolicchio only at comparatively deep levels. As a result of being forced to settle in crevices, space competition occurs among the various actinian species. Cereus pedunculatus, Aiptasia mutabilis, and Telmatactis forskalii co-exist, while Anemonia sulcata lives isolated in few crevices only. Contrary to other localities, A. sulcata is not the most abundant actinian species on Strombolicchio. Most frequent are the viviparous forms of Cereus pedunculatus and Actinia equina mediterranea. This mode of reproduction is rather rare in these species in the Mediterranean Sea, but common in the Atlantic Ocean. It is supposed that the viviparous varieties of Strombolicchio are geographical isolates of Atlantic populations. The influence of light on the colour of some anthozoans has also been studied. Zooxanthellae were found to be much more abundant in Aiptasia mutabilis in darkness than in bright surface waters. Gerardia savaglia proved to be quite abundant in deeper caves and on overhanging walls; it is more sciaphilic than Paramuricea clavata. Paralcyonium elegans occurs near the surface as well as at different depth levels; it has been observed in differing growth forms, corresponding to different patterns of turbulence. Alcyonium brioniense is considered to be a growth form of Alcyonium acaule.     

Evidence for benthic primary production support of an apex predator–dominated coral reef food web

Five hundred and ninety-nine primary producers and consumers in the Papahānaumokuākea Marine National Monument (PMNM) (22°N–30°N, 160°W–180°W) were sampled for carbon and nitrogen stable isotope composition to elucidate trophic relationships in a relatively unimpacted, apex predator–dominated coral reef ecosystem. A one-isotope (δ¹³C), two-source (phytoplankton and benthic primary production) mixing model provided evidence for an average minimum benthic primary production contribution of 65 % to consumer production. Primary producer δ¹⁵N values ranged from ?1.6 to 8.0 ‰ with an average (2.1 ‰) consistent with a prevalence of N₂ fixation. Consumer group δ¹⁵N means ranged from 6.6 ‰ (herbivore) to 12.1 ‰ (Galeocerdo cuvier), and differences between consumer group δ¹⁵N values suggest an average trophic enrichment factor of 1.8 ‰ Δ¹⁵N. Based on relative δ¹⁵N values, the larger G. cuvier may feed at a trophic position above other apex predators. The results provide baseline data for investigating the trophic ecology of healthy coral reef ecosystems.     

Early signs of recovery of Acropora palmata in St. John,US Virgin Islands

Since the 1980s, diseases have caused significant declines in the population of the threatened Caribbean coral Acropora palmata. Yet it is largely unknown whether the population densities have recovered from these declines and whether there have been any recent shifts in size-frequency distributions toward large colonies. It is also unknown whether colony size influences the risk of disease infection, the most common stressor affecting this species. To address these unknowns, we examined A. palmata colonies at ten sites around St. John, US Virgin Islands, in 2004 and 2010. The prevalence of white-pox disease was highly variable among sites, ranging from 0 to 53 %, and this disease preferentially targeted large colonies. We found that colony density did not significantly change over the 6-year period, although six out of ten sites showed higher densities through time. The size-frequency distributions of coral colonies at all sites were positively skewed in both 2004 and 2010, however, most sites showed a temporal shift toward more large-sized colonies. This increase in large-sized colonies occurred despite the presence of white-pox disease, a severe bleaching event, and several storms. This study provides evidence of slow recovery of the A. palmata population around St. John despite the persistence of several stressors.     

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.     

Fecundity of red coral <Emphasis Type="Italic">Corallium rubrum</Emphasis> (L.) populations inhabiting in contrasting environmental conditions in the NW Mediterranean

In this study, we examined the variability and potential patterns of fecundity in the precious Mediterranean red coral Corallium rubrum (L. 1758). A total of 12 populations were selected from the NW Mediterranean Sea. We used a hierarchical sampling design to explore fecundity patterns associated with different environmental conditions found in different cave zones (entrance vs. interior), depths (15–22 vs. 39–42 m), and geographic locations (Côte Bleue vs. Calanques). Overall, 240 apical tips from colonies (10 male + 10 female colonies per population) were analyzed. Fecundity ranged between 1.0 ± 0.7 and 3.2 ± 2.3 mature gonads per polyp in female colonies and between 2.5 ± 1.6 and 6.9 ± 2.5 mature gonads per polyp in male colonies. The fecundity of red coral varied significantly for populations dwelling in different cave zones and geographic areas but not for the examined depths. Our results contribute to the knowledge of red coral fecundity in populations not yet studied in the NW Mediterranean and elucidate significant variability in fecundity within different environmental conditions. The information on coral fecundity can contribute to the development of management and conservation plans for red coral populations.     

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.     

Mathematical modelling for conservation and management of gorgonians corals: youngs and olds,could they coexist?

Gorgonian corals are long-lived, slow-growing marine species dominating Mediterranean rocky bottoms. Endowed with complex morphologies they give a structure to the whole community, moreover, being efficient suspension feeders, they play a key role in plankton-benthos energy flow and CO₂${\text{CO}}_2$ storage. Thus, the structure and the development of benthic, hard bottom communities are linked to gorgonian survival. The red coral Corallium rubrum (L. 1758) is a precious gorgonian endemic to the Mediterranean Sea. Harvested and traded world-wide since ancient times red coral is a clear example of overexploited marine resource. This species is structured into self-seeding, genetically differentiated populations, some of which, living in the shallower part of the species bathymetric distribution, was recently affected by anomalous mortality events linked to global climate change. The co-occurrence of overharvesting and mass mortality could dramatically affect such populations. Demographic population models, widely applied by conservation biologists to check population viability and to project population trends over time are fundamental to foster survival of such populations matching harvesting to population growth rates. Therefore we set out a dynamic model of a genetically differentiated red coral population living in shallow waters. This population is characterised by small/young, crowded colonies and high recruitment rate. On the basis of the size–age structure determined for this population, a static life-history table, in which survival and reproduction coefficients of the different size–age classes were reported, has been set out. Demographic data were included in a non-linear, discrete, age-structured dynamic model, based on a Leslie-Lewis transition matrix. Our field data indicate that the recruits-to-larvae ratio is actually density-dependent. Such dependence, positive for low and negative for high density values, was included into the model and the effect of colonies of different size–age classes on recruits-to-larvae ratio was considered to be proportional to the number of polyps they have. We applied such model to simulate the trends of the studied population under different increases of survival and life-span. As some populations of gorgonians actually show the dominance of sparse, big/old colonies and low recruitment rate, while others are characterised by crowded, small/young colonies and high recruitment rate, we simulated the shift from the former to the latter structure increasing survival and life-span. Our results suggest that a dramatic mortality increase of bigger–older colonies (due, in the case of red coral to overfishing) could have determined the population structure we found.     

Size,spatial, and bathymetrical distribution of the ascidian Halocynthia papillosa in Mediterranean coastal bottoms: benthic–pelagic coupling implications

Benthic suspension feeders are abundant in Mediterranean coastal environments, though most of them are threatened due to overexploitation, climate change impacts, and unregulated diving or fishing practices. Little is known about most of the coastal communities in terms of large-scale distribution and realistic benthic–pelagic coupling implications, which are keys to understand and manage those threatened ecosystems. The active suspension feeder Halocynthia papillosa (one of the most common ascidian species of the Mediterranean Sea) was selected as a model organism to help to understand the ecological role in benthic–pelagic coupling processes and its importance as a carbon sink (an essential ecosystem service). The spatial and bathymetrical distribution of this organism has been studied using remotely operated vehicle video transects. The species was distributed throughout the study area, with a maximum density of 4 specimens m^?2. The highest abundances and the biggest sizes were observed on the range of 20–50 m depth. The role as carbon and nitrogen sink of this suspension feeder has been quantified coupling distribution data with existing in situ studies of feeding and respiration. Along the 1.24 ha of the study area, H. papillosa yearly ingested 519.4 g C and 31.4 g N and retained 20.2 g C. As long as the physiological data are known, this new methodology could be very useful in assessing bentho–pelagic links and the capacity of being C and N sinks of a wide range of species. This new approach may be essential for the future management of benthic communities.     

Multi-year, seasonal genotypic surveys of coral-algal symbioses reveal prevalent stability or post-bleaching reversion

This report documents the extent to which coral colonies show fluctuations in their associations with different endosymbiotic dinoflagellates. The genetic identity of Symbiodinium from six coral species [Acropora palmata (Lamarck), A. cervicornis (Lamarck), Siderastrea siderea (Ellis and Solander), Montastrea faveolata (Ellis and Solander), M. annularis (Ellis and Solander), and M. franksi (Gregory)] was examined seasonally over five years (1998 and 2000–2004) in the Bahamas and Florida Keys at shallow (1 to 4 m) fore-reef/patch reef sites and at deeper fore-reef (12–15 m) locations. Symbionts were identified genetically using denaturing gradient gel electrophoresis (DGGE) fingerprinting of the internal transcribed spacer region 2 (ITS2) of ribosomal RNA gene loci. Repetitive sampling from most labeled colonies from the Bahamas and the Florida Keys showed little to no change in their dominant symbiont. In contrast, certain colonies of M. annularis and M. franksi from the Florida Keys exhibited shifts in their associations attributed to recovery from the stresses of the 1997–1998 El Niño southern oscillation (ENSO) event. Over several years, a putatively stress-tolerant clade D type of Symbiodinium was progressively replaced in these colonies by symbionts typically found in M. annularis and M. franksi in Florida and at other Caribbean locations. Greater environmental fluctuations in Florida may explain the observed changes among some of the symbioses. Furthermore, symbiotic associations were more heterogeneous at shallow sites, relative to deep sites. The exposure to greater environmental variability near the surface may explain the higher symbiont diversity found within and between host colonies.     

Seasonality in reproduction of the deep-water pennatulacean coral Anthoptilum grandiflorum

The deep-sea pennatulacean coral Anthoptilum grandiflorum exhibits a cosmopolitan distribution and was recently determined to serve as habitat for other invertebrates and fish larvae in the northwest Atlantic. Colonies collected at bathyal depths between 2006 and 2010 in eastern Canada were analysed to determine their fecundity and characterize spatial and temporal trends in their reproductive cycle. Anthoptilum grandiflorum is a gonochoric broadcast spawner with a sex ratio that does not differ significantly from equality (although one hermaphrodite colony was observed). In male colonies, all the spermatocysts synthesized become mature over the annual cycle, while only ~21 % of the initial production of oocytes reaches maturity in female colonies. Female potential fecundity based on mature oocytes just before spawning was on average 13 oocytes polyp^?1; male potential fecundity was 48 spermatocysts polyp^?1. The spawning period of A. grandiflorum differs between geographic regions, from April (in southern Newfoundland) to July (in Labrador), closely following regional spring phytoplankton blooms after accounting for the deposition of planktic detritus. Release of oocytes by a live colony held in the laboratory was recorded in April 2011, coincident with field data for similar latitudes. Seawater temperatures at the time of spawning were around 3.6–4.8 °C in all regions and in the laboratory. Early stages of gametogenesis were detected in colonies collected shortly after the spawning season, and early and late growth stages occurred successively until December. Mature colonies were observed between April and July (depending on latitude). The diameter of mature oocytes (~1,100 μm maximum diameter) is consistent with lecithotrophic larval development.     

Transplantation of juvenile corals: a new approach for enhancing colonization of artificial reefs

 Coral reefs in the northern Gulf of Eilat are exposed to continuous man-made disturbances, resulting in decreased coral coverage and reduced recruitment at the Nature Reserve of Eilat. The construction of artificial reefs on sandy bottoms is a possible option to decrease diving pressure on natural reefs. In the present study we tested this hypothesis by submerging an experimental artificial reef anchored to the bottom at 18 m depth and floated vertically 3 m below water surface. The reef was composed of PVC plates, attached both vertically and horizontally along a wire. Propagules of two coral species, the stony coral Stylophora pistillata and the soft coral Dendronephthya hemprichi, were transplanted to this artificial reef. Planulae of S. pistillata were obtained during the breeding season, seeded in petri dishes in the laboratory and after 2 wk the dishes were transferred to the experimental artificial reef. Automized fragments of D. hemprichi which had previously settled on 10 × 10 cm PVC plates were transplanted onto the experimental artificial reef. The survivorship of the transplanted D. hemprichi colonies was significantly higher on the lower sides of shallower plates. Survivorship of S. pistillata colonies increased with depth when located on the vertical plates, or on the upper sides of the horizontal plates. The highest survivorship of this coral was on the vertical plates and on the upper sides of the horizontal plates, while very low survivorship was recorded on the lower sides. The results indicate that vertical artificial surfaces offer the optimal biotic and abiotic conditions for the survival of the two examined corals. The vertical plates are characterized by low sed imentation rates, low coverage of turf-algae, minimal grazing by sea urchins and absence of the competitor tunicate Didemnum sp. In addition, the vertical orientation of the experimental plates reduces shading and offers the required light intensity for zooxanthellate corals such as S. pistillata. Only a few studies to date have tried to implement artificial reefs in a coral reef environment. The results of the present study indicate the potential of enhancing recruitment of corals by transplantation of juvenile recruits onto appropriate artificial structures. Maximal survivorship of these recruits is dependent upon the structural features of the artificial reef, which should offer optimal conditions. Received: 25 May 1996 / Accepted: 15 July 1996     

Feeding biology and ecological impact of an introduced nudibranch,Tritonia plebeia,New England,USA

The nudibranchTritonia plebeia (Johnston) was first observed in New England in 1983, on vertical rock walls at 7 m depth off Nahant, Massachusetts. This northern European species preys exclusively on the soft coralAlcyonium digitatum (Linneaus) in its natural habitat. At Nahant, it preyed primarily on the closely relatedAlcyonium siderium Verrill. Laboratory studies indicated that it could locate its prey by distance chemoreception and by visual orientation towards tall dark surfaces which could help it find the vertical walls, overhangs, and boulder sides where the soft corals occur. Field studies showed thatT. plebeia fed primarily on colony bases, causing extensive damage and whole colony mortality. The most important endemic predator onA. siderium, Coryphella verrucosa (Sars), preyed preferentially on hydroids, but would graze polyps off the top portions ofA. sederium colonies, causing little permanent damage to the colony, during the winter months when hydroids were scarce. AlthoughC. verrucosa occasionally behaved agonistically towardT. plebeia, there was no indication in laboratory or field studies that either nudibranch had an effect on the other's foraging through interference competition. Extensive predation byT. plebeia caused the disappearance ofA. siderium at two sites (Outer and Inner Shag Rocks) and a sharp reduction at a third site (Inner East Point). The higher mortalities at the Shag Rocks sites most likely occurred because of a simultaneous urchin (Strogylocentrotus droebachiensis) population expansion. As space among aggregates ofA. siderium opened up due toT. plebeia predation, urchins were able to forage on the vertical walls and scrape off remaining colonies. At a fourth site, Halfway Rock, whereT. plebeia were seldom present,A. siderium colonies also suffered high mortalities. This increae in mortality began nearly a year before urchin populations increased, and during a summer of abnormally high water temperatures at Halfway Rock. The high temperatures, followed by urchin predation on remaining colonies could account for the disappearance of allA. siderium colonies at this site.T. plebeia disappeared at all sites by summer 1986 andA. siderium populations have since stabilized, but community-level changes at all sites whereA. siderium were removed have persisted.     

Change in zooxanthellae and mucocyte tissue density as an adaptive response to environmental stress by the coral, <Emphasis Type="Italic">Montastraea annularis</Emphasis>

Results from controlled in situ experimentation conducted on the leeward reef tract of Curaçao, Netherlands Antilles, indicate that the coral Montastraea annularis exhibits a complex, yet consistent, cellular response to increasing sea surface temperature (SST) and decreasing irradiance. This was determined by simultaneously quantifying and tracking the tissue density of zooxanthellae and mucocytes using a novel technique that integrates the lectin histochemical stain wheat germ agglutinin (WGA) with high-resolution (200 nm) optical epifluorescence microscopy. Coral colonies growing at 6-m water depth (WD) and an irradiance of 100.2 ± 6.5 μmol m^?2 s^?1 were treated with a shading experiment for 11 days that reduced irradiance to 34.9 ± 6.6, 72.0 ± 7.0 and 90.1 ± 4.2 μmol m^?2 s^?1, respectively. While a significant decrease in the density of both zooxanthellae and mucocytes were observed at all shade levels, the largest reduction occurred between the natural non-shaded control (44,298 ± 3,242 zooxanthellae cm^?2; 4,853 ± 346 mucocytes cm^?2) and the highest shading level (13,982 ± 1,961 zooxanthallae cm^?2; 2,544 ± 372.9 mucocytes cm^?2). Colonies were also sampled during a seasonal increase in SST of 1.5°C, where the density of zooxanthellae was significantly lower (from 54,710 ± 1,755 to 34,322 ± 2,894 cells cm^?2) and the density of mucocytes was significantly higher (from 6,100 ± 304 to 29,658 ± 3,937 cells cm^?2). These observations of coral cellular response to environmental change provide evidence to support new hypotheses for coral survival and the complex role played by mucus in feeding, microbial associations and resilience to increasing SST.     

Reef coral reproduction in the eastern Pacific: Costa Rica,Panamá, and the Galápagos Islands (Ecuador). VI. Agariciidae, <Emphasis Type="Italic">Pavona clavus</Emphasis>

The reproductive ecology of the zooxanthellate reef coral Pavona clavus was investigated at several sites in Costa Rica, Panamá, and the Galápagos Islands (Ecuador) over the period 1985–2009. Pavona clavus displayed stable gonochorism as only five hermaphrodites were found in 590 samples. At four of five locations, sex ratios were skewed toward female dominance; however, at Saboga Island (Panamá) male colonies predominated. In Panamá, sexual maturity was observed in an estimated eight-year-old female colony, and several colonies of 10–20 years of age demonstrated gametogenesis. Sexual activity was observed at all study sites, but gamete development occurred in only 14–31% of colonies sampled sporadically. Seasonality of gametogenic activity occurred predominantly during the warm/wet season, June to August, at mainland sites (Caño Island, Costa Rica, and Gulfs of Chiriquí and Panamá, Panamá). This pattern was repeated in the Galápagos Islands, but mainly from March to May when seasonally high sea temperatures and rainfall prevailed there. Histological sampling and field observations indicated that spawning was centered around the full moon, most frequently on lunar day 17, and near sunset (1,800 h). Mean fecundity (mature ova cm^?2 live tissue) estimates were significantly different for two sites and ranged from ~1,780 (Saboga Island, Gulf of Panamá, seasonally upwelling) to ~4,280 (Uva Is, Gulf of Chiriquí, non-upwelling). Assuming three annual spawning events colony^?1 (August, September, October), extrapolation of minimum and maximum fecundities yield 5,340 and 12,840 ova cm^?2 year^?1. Seasonal, lunar, and diel spawning patterns in nine zooxanthellate species at Uva Island indicate asynchronous coral community spawning.     

The effects of partial mortality on the fecundity of three common Caribbean corals

The recent intensification of human disturbances in the Caribbean has increased the prevalence of partial mortality on coral colonies. Partial mortality can change colony size by directly shrinking colonies or by splitting colonies into fragments. A reduction in colony size can also adversely affect fecundity and fitness as internal resources shift away from reproduction toward colony maintenance. This study aimed to determine whether three Caribbean coral species, Siderastrea siderea, Montastraea faveolata, and Diploria strigosa, along the reef tract in Puerto Morelos, Mexico (20^o52′N, 86^o51′W), continued to dedicate resources to reproduction when colonies were fragmented to pre-maturation size. Contrary to expectations, eggs were found in colonies that were smaller than the maturation size and had been subjected to partial mortality. The continued dedication of resources toward reproduction, even in the smallest colonies, suggests that resource trade-offs away from reproduction are not as rigid as previously suggested in stressed corals.