Life history and viability of a long-lived marine invertebrate: the octocoral Paramuricea clavata

The red gorgonian Paramuricea clavata is a long-lived, slow-growing sessile invertebrate of ecological and conservation importance in the northwestern Mediterranean Sea. We develop a series of size-based matrix models for two Paramuricea clavata populations. These models were used to estimate basic life history traits for this species and to evaluate the viability of the red gorgonian populations we studied. As for many other slow-growing species, sensitivity and elasticity analysis demonstrate that gorgonian population growth is far more sensitive to changes in survival rates than to growth, shrinkage, or reproductive rates. The slow growth and low mortality of red gorgonians results in low damping ratios, indicating slow convergence to stable size structures (at least 50 years). The stable distributions predicted by the model did not differ from the observed ones. However, our simulations point out the fragility of this species, showing both populations in decline and high risk of extinction over moderate time horizons. These declines appear to be related to a recent increase in anthropogenic disturbances. Relative to their life span, the values of recruitment elasticity for Paramuricea clavata are lower than those reported for other marine organisms but are similar to those reported for some long-lived plants. These values and the delayed age of sexual maturity, in combination with the longevity of the species, show a clear fecundity/mortality trade-off. Full demographic studies of sessile marine species are quite scarce but can provide insight into population dynamics and life history patterns for these difficult and under-studied species. While our work shows clear results for the red gorgonian, the variability in some of our estimates suggest that future work should include data collection over longer temporal and spatial scales to better understand the long-term effects of natural and anthropogenic disturbances on red gorgonian populations.     

DNA fingerprints of a gorgonian coral: a method for detecting clonal structure in a vegetative species

Clonal reproduction, a common life history strategy among sessile marine invertebrates, can lead to high local abundances of one to a few genotypes in a population. Analysis of the clonal structure of such populations can provide insight into the ecological and evolutionary history of the population, but requires markers that can identify individual genets. Forensic and demographic studies have demonstrated that DNA fingerprinting can provide markers that are unique for an individual genotype. We have generated DNA fingerprints for over 70 colonies of the clonal gorgonian, Plexaura A (Plexaura sp. A) collected from June 1990 through July 1991 in the San Blas Islands, Panama. DNA fingerprints within a singic individual were identical and fingerprinting resolved multiple genotypes within and among reefs. On one reef in the San Blas Islands, Panama, 59% of the colonies sampled were of one genotype and this genotype was not found on any other sampled reefs. A previous study using tissue grafts identified 13 putative clones on these reefs, while DNA fingerprints of the same colonies differentiated 17 genotypes. The present study demonstrates the utility of DNA fingerprinting for distinguishing clones and for identifying clonal structure of marine invertebrate populations.     

Cytogenetic characterization of the greater amberjack, Seriola dumerili (Pisces: Carangidae), by different staining techniques and fluorescence in situ hybridization

A cytogenetic analysis by several staining techniques was carried out on specimens of Seriola dumerili from the Tyrrhenian coast of Sicily. The karyotype was found to be composed of 48 chromosomes in all the specimens examined. All chromosomes are subtelo- and acrocentrics, except for one pair of small-sized submetacentric chromosomes. After Ag-staining, nucleolus organizer regions (NORs) can be observed on the short arms of a medium-sized subtelocentric chromosome pair, and after chromomycin A₃-staining a bright fluorescent signal is evident at the same sites. The 18S rDNA mapping by fluorescence in situ hybridization confirms this unique NOR location. The constitutive heterochromatin is mainly restricted to centromeres and the fluorochrome-staining by chromomycin A₃ and 4′,6-diamidino-2-phenylindole revealed a uniform base composition along the chromosomal arms; therefore stock/population-specific marker chromosomes cannot be identified. Such cytogenetic features, however, rule out the presence of morphologically differentiated sex-chromosomes in the greater amberjack. Received: 16 January 1997 / Accepted: 31 January 1997     

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.     

Karyotypic characterization of the great sturgeon, Huso huso, by multiple staining techniques and fluorescent in situ hybridization

A karyotype analysis by several staining techniques was carried out on the great sturgeon, Huso huso (Linnaeus, 1758). The karyotype (2n = 118 ± 2) was composed of 42 pairs of meta-/submetacentric chromosomes and 17 pairs of acrocentrics/microchromosomes. Constitutive heterochromatin was mainly located at the centromeric regions of the acrocentric chromosomes. The biarmed chromosomes showed weak C-bands. Fluorescent staining with GC-specific chromomycin A₃ showed clearly recognizable fluorescent regions, whereas a more uniform base composition was revealed by the AT-specific 4,6-diamidino-2 phenylindole. After Ag-staining, nucleolar organizer regions could be observed on the short arms of two medium-sized submetacentrics and on two acrocentrics. Digoxigenated 28S and 5S rDNA probes, prepared from Acipenser naccarii DNA and hybridized to metaphase chromosomes, showed signals on six and two chromosomes, respectively. The telomeric sequence (TTAGGG) _n detected by FISH was located at both ends of each chromosome. Results are discussed in relation to karyotype organization and evolution in sturgeons. Received: 1 April 1998 / Accepted: 21 July 1998     

Growth and bleaching of the coral Oculina patagonica under different environmental conditions in the western Mediterranean Sea

The success that the putative alien species Oculina patagonica is able to survive under different environmental conditions may be benefiting its establishment and spreading along the Mediterranean Basin. Our objectives were to determine the response of this species, in terms of growth and bleaching, under different environmental conditions. Field data on colony growth and bleaching were obtained for a period of 18 months (from June 2010 to December 2011), in the Alicante Harbour (38°20′11″N, 00°29′11″W) and the Marine Protected Area of Tabarca (38°09′59″N, 00°28′56″W). Additionally, data on sedimentation rates, chlorophyll a concentration and organic matter were also collected. Moreover, the role of light over growth and bleaching of the coral was also studied with a field experiment. Our results showed that growth rates were similar among localities (eutrophic and oligotrophic environments), decreasing with increasing perimeter of the colony. Growth rates were at a minimum during cold months (13 °C) and enhanced during warm months until a threshold temperature; thereafter, bleaching was observed (>28 °C), being adverse for coral growth. In addition, light attenuation could act such as local stress, increasing the coral bleaching with the increase in seawater temperature. Our findings confirmed that O. patagonica has a broad tolerance to seawater temperature, irradiance and trophic water conditions, in addition to its ability to thrive through bleaching events, mainly in eutrophic environments, probably related to food availability.     

Chromosomal mapping of major ribosomal rRNA genes in the hard clam (<Emphasis Type="Italic">Mercenaria mercenaria</Emphasis>) using fluorescence in situ hybridization

Karyotype and chromosomal location of the major ribosomal RNA genes were studied in the hard clam (Mercenaria mercenaria Linnaeus) using fluorescence in situ hybridization (FISH). Metaphase chromosomes were obtained from early embryos. Internal transcribed spacers (ITS) between major RNA genes were amplified and used as FISH probes. The probes were labeled with digoxigenin-11-dUTP by polymerase chain reaction and detected with fluorescein-labeled anti-digoxigenin antibodies. FISH with the ITS probes produced two to four signals per nucleus or metaphase. M. mercenaria had a haploid number of 19 chromosomes with a karyotype of seven metacentric, four metacentric or submetacentric, seven submetacentric, and one submetacentric or subtelocentric chromosomes (7M + 4M/SM + 7SM + 1SM/ST). Two ITS loci were observed: one located near the centromere on the long arm of Chromosome 10 and the other at the telomere of the short arm of Chromosome 12. FISH signals on Chromosome 10 are strong and consistent, while signals on Chromosome 12 are variable. This study provides the first karyotype and chromosomal assignment of the major RNA genes in M. mercenaria. Similar studies in a wide range of species are needed to understand the role of chromosomal changes in bivalve evolution.     

Modelling the dynamics of coral reef macroalgae using a Bayesian belief network approach

Macroalgae are a major benthic component of coral reefs and their dynamics influence the resilience of coral reefs to disturbance. However, the relative importance of physical and ecological processes in driving macroalgal dynamics is poorly understood. Here we develop a Bayesian belief network (BBN) model to integrate many of these processes and predict the growth of coral reef macroalgae. Bayesian belief networks use probabilistic relationships rather than deterministic rules to quantify the cause and effect assumptions. The model was developed using both new empirical data and quantified relationships elicited from previous studies. We demonstrate the efficacy of the BBN to predict the dynamics of a common Caribbean macroalgal genus Dictyota. Predictions of the model have an average accuracy of 55% (implying that 55% of the predicted categories of Dictyota cover were assigned to the correct class). Sensitivity analysis suggested that macroalgal dynamics were primarily driven by top–down processes of grazing rather than bottom–up nutrification. BBNs provide a useful framework for modelling complex systems, identifying gaps in our scientific understanding and communicating the complexities of the associated uncertainties in an explicit manner to stakeholders. We anticipate that accuracies will improve as new data are added to the model.     

Microbial associations in sponges. III. Ultrastructure of the in situ associations in coral reef sponges

Symbiotic cyanobacteria are associated with marine sponges in three ways: the majority are free-living in the mesohyl; large aggregates occur in cyanocytes (specialized, vacuolated archeocytes); and few are present in digestive vacuoles. The cyanobacteria in Jaspis stellifera and Neofibularia irata are morphologically similar to those described in Mediterranean sponges, whereas those in Pericharax heteroraphis are different. The freeliving bacterial populations are morphologically similar, although the number of bacteria varies between the species. The fourth sponge Ircinia wistarii contains a mixed bacterial population unlike those in the other sponges. Sponge digestion of microbial associates is rare and not considered to contribute significant nutrients.     

Size and spatial structure in deep versus shallow populations of the Mediterranean gorgonian <Emphasis Type="Italic">Eunicella singularis</Emphasis> (Cap de Creus,northwestern Mediterranean Sea)

In the Western Mediterranean Sea, the gorgonian Eunicella singularis (Esper, 1794) is found at high densities on sublittoral bottoms at depths from 10 to 70 m. Shallow colonies have symbiotic zooxanthellae that deeper colonies lack. While knowledge of the ecology of the shallow populations has increased during the last decades, there is almost no information on the ecology of the deep sublittoral populations. In October and November 2004 at Cap de Creus (42°19′12″ N; 03°19′34″ E), an analysis of video transects made by a remotely operated vehicle showed that shallow populations (10–25 m depth) were dominated by small, non-reproductive colonies, while deep sublittoral populations (50–67 m depth) were dominated by medium-sized colonies. Average and maximum colony heights were greater in the deeper populations, with these deeper populations also forming larger patch sizes and more extensive regions of continuous substrate coverage. These results suggest that shallow habitats are suitable for E. singularis, as shown by the high recruitment rate, but perturbations may limit or delay the development of these populations into a mature stage. This contrasts with the deep sublittoral habitats where higher environmental stability may allow the development of mature populations dominated by larger, sexually mature colonies.     

Genetic structure of a scleractinian coral, Mycedium elephantotus, in Taiwan

The genetic structure of Mycedium elephantotus (Pallas, 1766) populations from three regions around Taiwan was examined using allozyme electrophoresis. Eight loci were scored from seven enzyme systems. Seven loci were polymorphic under the 95% criterion. The high ratio of observed to expected genotypic diversities at the collection sites (G _O:G _E=0.8 to 1.0) indicate that M. elephantotus propagates predominantly by sexual reproduction. Allele frequencies of M. elephantotus differed significantly among regions (D=0.024 to 0.256, F _ST=0.032 to 0.218, p < 0.001), while populations among collection sites within each region were homogeneous (D=0.000 to 0.015, F _ST=0.010 to 0.022, p > 0.05). Genetic differentiation between populations from southern Taiwan and the Penghu Islands is greater than that between southern and northern Taiwan populations, although the former two regions are much closer geographically. The UPGMA dendrogram based on Nei's unbiased genetic distance showed a clear subdivision of populations into two groupings, northern Taiwan/Penghu Islands and southern Taiwan. A higher level of gene flow was found between M. elephantotus populations in northern Taiwan and the Penghu Islands (N _m=7.56) than that between populations in southern Taiwan and other regions (N _m=0.90 to 1.72). The pattern of genetic subdivision among regions is consistent with the pattern of ocean currents, indicating that genetic differentiation is likely driven by surface circulation vicariance. Received: 2 December 1997 / Accepted: 15 September 1998     

Pink-line syndrome, a physiological crisis in the scleractinian coral Porites lutea

Coral diseases are one of the major factors that alter coral cover and their diversity. We have earlier reported the “Pink-line syndrome” (PLS) in the scleractinian coral Porites lutea wherein a colored band appears between the dead and healthy tissue of a colony. About 20% of the P. lutea colonies were affected in Kavaratti of the Lakshadweep Islands in the Arabian Sea during April 1996 and the incidence increased fourfold within the next 4 years. Fungi were associated in both PLS-affected and healthy specimens, whereas the cyanobacterium Phormidium valderianum occurred exclusively in the PLS-affected specimens. There was an increased expression of a 29 kDa protein without any significant increase in total protein content in the PLS-affected colonies. A reduced number of zooxanthellae and an increase in zooxanthellae size, mitotic index, and chl a concentrations were some of the characteristics of the PLS-affected colonies. PLS induction experiments conducted using selected fungi and the cyanobacterium P. valderianum isolated from the affected colonies and abiotic factors, such as CO₂ enrichment and the effect of cyanobacterial photosynthesis inhibition, indicated that the CO₂ build-up around the host tissue caused the pink coloration. We hypothesize that these physiological changes disturb the mutualism between the zooxanthellae and the host. When the symbiosis is disturbed by the external CO₂, the host loses control over the zooxanthellae, causing their uncontrolled division. This process may lead to a break in photosynthate transfer to the host, thereby resulting in starvation and finally leading to partial mortality. We further hypothesize that these degenerative processes are triggered by the CO₂ produced by P. valderianum through its carbon concentration mechanism. In this context, any opportunistic cyanobacteria or other agents having potential to interfere with the physiology of the host or the symbiont can cause such a physiological disorder. The mechanism of PLS formation is an early warning to protect corals as the increasing atmospheric CO₂ could induce PLS-like physiological disorder in corals.     

Demographic parameters of two populations of red coral (Corallium rubrum L. 1758) in the North Western Mediterranean

The demographic and reproductive structure of populations represents the main data set needed for conservation and management plans. Mediterranean red coral Corallium rubrum has been exploited for 2,000 years, but only recently management plans have been solicited by the international community. We examined and compared the demographic features of two red coral shallow populations located in distinct geographic locations: Portofino (Italy) and Cap de Creus (Spain). Adults and juveniles density, growth rates, population size and age structure, fecundity and fertility were examined. Juveniles were the dominant class (33 %) in both populations. The analysis of the gamete content of 653 colonies revealed that the populations have balanced sex ratios and similar fertility and polyp fecundity. The average annual growth rate, determined on 119 colonies by annual growth rings count, was similar in both populations (0.24 mm year^?1), decreasing with colony age. Maximum life span of 99 % of the colonies was 60 and 40 years at Portofino and Cap de Creus, respectively. Minimum harvestable size (7 mm basal diameter) was reached in 30–35 years, and the percentage of colonies above it was 6.7 % at Portofino and 2.1 % at Cap de Creus, where juvenile and adult colony densities were significantly lower and the percentage of commercial-sized colonies reduced by 25 % in just a few years. Notwithstanding similar growth and fecundity, the two populations showed different densities and size/age structures suggesting local factors, together with different fishing pressures, have to be taken into account in the management plans for this species.     

Cytogenetics in the sacoglossan Oxynoe olivacea (Mollusca: Opisthobranchia): karyotype, chromosome banding and fluorescent in situ hybridization

Developing embryos and sexually mature follicles of the male portion of ovotestis proved to be a suitable material as a source of cleaving cells for advanced cytological investigations on the sacoglossan species Oxynoe olivacea Rafinesque, 1819 (Mollusca: Opisthobranchia). O. olivacea has a diploid chromosomal number of 30 made up of 15 pairs of which six are metacentric/submetacentric (M/SM), four subtelocentric (ST) and five on the borderline between SM and ST. Correspondingly, 15 bivalents occur in spermatocytes at Metaphase I. Constitutive heterochromatin is scarce and restricted to small C-bands seen in five pachytene bivalents. The use of combined silver staining and fluorescent in situ hybridization (FISH) with a Paracentrotus lividus (Echinodermata) 4.3 kilobase (kb) rDNA probe (prR14) consisting of sequences from the 3′ end of 18S rDNA to the 3′ end of 26S rDNA, revealed that nucleolus organizer regions (NORs) are situated terminally on one arm of a small metacentric pair. The telomeric (TTAGGG) _n sequence did not hybridize with termini of O. olivacea chromosomes. Received: 14 December 1999 / Accepted: 10 July 2000     

Sexual reproduction of the solitary sunset cup coral Leptopsammia pruvoti (Scleractinia, Dendrophylliidae) in the Mediterranean. 2. Quantitative aspects of the annual reproductive cycle

There are few data on the reproductive biology of coral species living in temperate zones, particularly in the Mediterranean Sea. Leptopsammia pruvoti is a solitary coral that is commonly found in sea caves and under overhangs throughout the Mediterranean basin and along European coasts from Portugal to southern England. In this paper, we describe its annual reproductive cycle in the eastern Ligurian Sea near the city of Leghorn (Tuscany, Italy). Polyps were sexually mature at 3 mm in length (maximum diameter of the oral disc), were dioecious with a sex ratio of 1:1, and brooded their larvae. The maturation of spermaries took 12 months and oocytes 24 months. The rate of gonad development increased significantly from November to January, fertilization occurred from January to April and planulation during May and June. Seasonal variations in water temperature and photoperiod may have played an important role in regulating reproductive events. The amount of energy devoted to male gametogenesis (quantified by gonad index) was significantly higher in the sex separated species L. pruvoti than in the hermaphroditic dendrophylliid Balanophyllia europaea, whose reproduction has been studied in a previous work. We hypothesize that this difference is due to the contrasting sexuality or fertilization biology of these two species (cross-fertilization in the sex separated L. pruvoti versus possibly self-fertilization in the hermaphroditic B. europaea). Greater male sexual allocation in reproductive strategies characterized by dioecism or cross-fertilization when compared to those characterized by hermaphroditism or self-fertilization is common in plant mating systems. In relation to other solitary dendrophylliids, L. pruvoti presents an ‘r’-reproductive strategy.     

In situ swimming and settlement behaviour of larvae of an Indo-Pacific coral-reef fish, the coral trout Plectropomus leopardus (Pisces: Serranidae)

Late larvae of the serranid coral trout Plectropomus leopardus (Lacepède), captured in light traps, were released during the day both in open water and adjacent to two reefs, and their behaviour was observed by divers at Lizard Island, northern Great Barrier Reef. Coral trout larvae (n = 110) were present in light-trap catches from 18 November to 3 December 1997, including new moon (30 November). The swimming speed of larvae in open water or when swimming away from reefs was significantly greater (mean 17.9 cm s⁻¹) than the speed of larvae swimming towards or over reefs (mean 7.2 cm s⁻¹). Near reefs, larvae swam at average depths of 2.7 to 4.2 m, avoiding 0 to 2 m. In open water, swimming depth varied with location: larvae >1 km east of Lizard Island swam steeply downward to >20 m in 2 to 4 min; larvae >1 km west oscillated between 2.6 and 13 m; larvae 100 to 200 m east of Lizard Island oscillated between 0.8 and 15 m. Nearly all larvae swam directionally in open water and near reefs. In open water, the average swimming direction of all larvae was towards the island, and 80% (4 of 5) swam directionally (p < 0.05, Rayleigh's test). Larvae swam directionally over the reef while looking for settlement sites. The frequency of behaviours by larvae differed between two reefs of different exposure and morphology. Depending on site, 26 to 32% of larvae released adjacent to reefs swam to open water: of these, some initially swam towards or over the reef before swimming offshore. In some cases, offshore-swimming seemed to be due to the presence of predators, but usually no obvious cause was observed. Depending on the reef, 49 to 64% of the larvae settled. Non-predatory reef residents aggressively approached 19% of settlers. Between 5 and 17% of the larvae were eaten while approaching the reef or attempting to settle, primarily by lizardfishes but also by wrasses, groupers and snappers. A higher percentage of larvae settled in the second week of our study than in the first. Average time to settlement was short (138 s ± 33 SE), but some larvae took up to 15 min to settle. Average settlement depth was 7.5 to 9.9 m, and differed between locations. No settlement took place on reef flats or at depths <4.2 m. Larvae did not appear to be selective about settlement substrate, but settled most frequently on live and dead hard coral. Late-stage larvae of coral trout are capable swimmers with considerable control over speed, depth and direction. Habitat selection, avoidance of predators and settlement seem to rely on vision. Received: 7 July 1998 / Accepted: 26 January 1999     

Gene flow at three spatial scales in a coral reef fish, the three-spot dascyllus, Dascyllus trimaculatus

Dispersal in coral reef fishes occurs predominantly during the larval planktonic stage of their life cycle. With relatively brief larval stages, damselfishes (Pomacentridae) are likely to exhibit limited dispersal. This study evaluates gene flow at three spatial scales in one species of coral reef damselfish, Dascyllus trimaculatus. Samples were collected at seven locations at Moorea, Society Islands, French Polynesia. Phylogenetic relationships and gene flow based on mitochondrial control region DNA sequences between these locations were evaluated (first spatial scale). Although spatial structure was not found, molecular markers showed clear temporal structure, which may be because pulses of settling larvae have distinct genetic composition. Moorea samples were then compared with individuals from a distant island (750 km), Rangiroa, Tuamotu Archipelago, French Polynesia (second spatial scale). Post-recruitment events (selection) and gene flow were probably responsible for the lack of structure observed between populations from Moorea and Rangiroa. Finally, samples from six Indo-West Pacific locations, Zanzibar, Indonesia, Japan, Christmas Island, Hawaii, and French Polynesia were compared (third spatial scale). Strong population structure was observed between Indo-West Pacific populations. Received: 26 May 2000 / Accepted: 10 October 2000     

Seasonality of coral reproduction in the Dampier Archipelago,northern Western Australia

Coral spawning in Western Australia (WA) occurs predominantly in the austral autumn in contrast to the Great Barrier Reef (GBR) on Australia’s east coast where most spawning occurs in spring. Recent work, however, suggests a second spawning period in northern WA with at least 16 Acropora spp spawning in spring or early summer. This discovery has initiated a re-examination of reproductive seasonality in northern WA, particularly on inshore reefs adjacent to large development projects, such as the site of this study in Mermaid Sound, in the Dampier Archipelago. Three locally abundant taxa, Porites spp, Pavona decussata and Turbinaria mesenterina were sampled monthly from September 2006 to May 2007 to determine sexuality, the mode of reproduction and the time of gamete maturity. All three taxa were gonochoric broadcast spawners. Porites spp. colonies were mature in November and December, P. decussata in March and April. In contrast, most colonies of T. mesenterina contained mature gametes for up to 5 months beginning in November, suggesting either individuals are releasing gametes on multiple occasions, or they retain mature gametes for more than 1 month. Field surveys to determine the reproductive status of the remaining coral assemblage were conducted prior to the full moon in October 2006 and March 2007. Only four species contained mature gametes in October 2006. In contrast, 55 species contained mature gametes in March 2007. We conclude that the major spawning season of corals on shallow-inshore reefs in the Dampier Archipelago is autumn, although taxa that spawn in spring and summer include Porites spp., Acropora spp. and possibly T. mesenterina that are numerically dominant at many of these sites. Consequently, management initiatives to limit the exposure of coral spawn to stressors associated with coastal development may be required in up to five months per year.