Genetic structure of populations of two closely related brittle stars with contrasting sexual and asexual life histories,with observations on the genetic structure of a second asexual species

Ophiocoma pumila Lütken andOphiocomella ophiactoides (H. L. Clark) are morphologically similar brittle stars with contrasting life histories, the former obligately sexual, the latter fissiparous (capable of both sexual reproduction and asexual proliferation by binary fission). Electrophoretic analysis of five polymorphic enzymes was used to assess the genetic consequences of these differing life histories and provide a genetic perspective on the taxonomic relationship between the two species. Genotypic diversity ofOphiocoma pumila collected at Discovery Bay, Jamaica, in 1985 conformed to expectations for a sexually reproducing population. In contrast, genotypic diversity ofOphiocomella ophiactoides at this site was significantly lower than expected for a sexually reproducing population, due largely to the predominance of clonal proliferation over larval recruitment. Large variation in clonal composition over a short (50 m) distance emphasized the very localized scale of clonal mixing in this species. Allozymic data are indicative of a close sibling species relationship betweenOphiocoma pumila andOphiocomella ophiactoides which suggests that the present generic separation of the two species should be re-examined. Electrophoretic analysis was also used to examine the genetic structure of sponge- and alga-dwelling populations of a second fissiparous brittle star,Ophiactis savignyi (Müller & Troschel), which was also collected at Jamaica in 1985. Striking differences in the allelic composition of sponge- and alga-dwellingO. savignyi were observed. Genotypic diversity ofO. savignyi in sponges was very low, each sponge being dominated by a single genotype. Genotypic diversity ofO. savignyi in algae was higher, although still significantly lower than expectations for a sexually reproducing population. In the light of the highly clonal composition of fissiparous brittle-star populations, the adaptive significance of clonal growth may be related to an increase in the overall fitness of dispersed clones (genets), compared to individuals of strictly sexual counterparts, through greater genotypespecific biomass and, hence, fecundity.Contribution No. 476 of the Discovery Bay Marine Laboratory of the University of the West Indies     

Divide and broadcast: sexual reproduction in the West Indian brittle star Ophiocomella ophiactoides and its relationship to fissiparity

It is known that asexual reproduction by fission is the dominant mode of propagation in the small (disc diameter <5.0 mm) ophiocomid brittle star Ophiocomella ophiactoides (H. L. Clark). This species is, however, able to reproduce sexually as well. Sexual reproduction occurs throughout much or all of the year in a population of O. ophiactoides at Jamaica. Gonads are often present in recently split individuals, indicating that sexual and asexual reproduction could occur simultaneously. Regeneration of new gonads in the new disc half is underway by the time an individual is competent to split again. Unsexable individuals are significantly smaller than males, which in turn are significantly smaller than females. The unsexable individuals appear to be below a threshold size for gonad production. Fecundity is low, with the largest females producing a maximum of about 7 400 eggs of small (mean diameter 80 m) size. Fertilized eggs develop into planktotrophic ophioplutei which are typical of the larvae of other species of ophiocomid brittle stars in morphology, rate of development and duration of planktonic period. Egg size and larval type for O. ophiactoides are characteristic of brittle stars with the planktotrophic mode of development, but egg number is uncharacteristically low. Fissiparity in O. ophiactoides may have evolved in association with small body size, concomitant low fecundity and a high-risk planktotrophic larva as a means of ensuring reliable recruitment. Available data, although sparse, suggest that this hypothesis may be applicable to fissiparous brittle stars in general. Fission in brittle stars has a broad systematic incidence at the familial level which suggests that it has evolved recurrently. The ability both to divide and broadcast appears to be a useful combination for O. ophiactoides: it can recruit reliably at large size to occupy favourable habitats quickly, but, it can still maintain low level dispersal of larvae which could provide escape from deteriorating habitats and the potential to colonize new habitats.Contribution No. 328 of the Discovery Bay Marine Laboratory, University of the West Indies     

Sea anemones and brittle stars: unexpected predatory interactions during induced in situ oxygen crises

During oxygen crises, benthic faunas exhibit a series of behavioural patterns that reflect the duration and severity of the event. During artificially induced oxygen deficiencies at 24 m depth in the Northern Adriatic Sea, we photographically documented predation by the sea anemones Cereus pedunculatus (Pennant, 1777) and Calliactis parasitica (Couch, 1842) on the brittle star Ophiothrix quinquemaculata (DelleChiaje, 1828). Five predatory events were recorded with four anemones during nine deployments totalling 817 h of observation. Under near-anoxic conditions, individuals of both actinians made contact with, pulled in and consumed the brittle stars. The duration of each predatory event was 1.5–7.5 h. In three of the five events, brittle star remains were regurgitated after an additional 2.0–12.5 h of digestion by the anemones. Our time-lapse sequences demonstrate that oxygen deficiency, beyond eliciting a series of specific behaviours in members of each species, also promotes previously unobserved interspecific interactions. Our results show that sea anemones are not only highly resistant to anoxia, but may also benefit by taking advantage of prey that are more vulnerable to anoxic conditions.     

Antimicrobial activity of extracts of Caribbean gorgonian corals

Extracts of 39 species of Caribbean gorgonians were tested for antimicrobial activity against 15 strains of marine bacteria. The bacteria consisted of three opportunistic pathogens, Vibrio parahaemolyticus, Leucothrix mucor, and Aerococcus viridans, and 12 strains isolated from either healthy or decayed gorgonians. Overall, only 15% (79 out of 544) of the tests resulted in antibacterial activity with 33% (13 out of 39) of the gorgonians inhibiting only one bacterial strain and 23% (9 out of 39) showing no activity. The extracts of four Pseudopterogorgia species showed relatively high levels of activity, inhibiting 43 to 86% of the bacterial strains. The potency of the active Pseudopterogorgia species was variable, however, and three additional Pseudopterogorgia species were inactive against all bacterial strains. With the exception of one sensitive strain, Vibrio species were resistant to gorgonian metabolites. Our results indicate that organic extracts of most Caribbean gorgonians do not possess potent, broad-spectrum antibacterial activity inhibitory to the growth of opportunistic marine pathogens and bacteria associated with healthy and decayed gorgonian surfaces. These findings suggest that the inhibition of bacterial growth is not the primary ecological function of gorgonian secondary metabolites and that bacteria may not be important selective agents in the evolution of gorgonian secondary chemistry.     

A new quantitative analysis of ovarian development in echinoderms: the maturity stage index

This study developed an objective quantitative method for detecting small-scale temporal or spatial differences in gametogenesis in echinoderms. The method was applied to conventional monthly samples of the planktotrophic brittle star, Ophiopholis aculeata, collected at a single site in Newfoundland (eastern Canada) at 10–15 m depth. The samples were analysed to determine gonad index, oocyte size and gonadal stage using histology. The maturity stage index (MSI) was developed to integrate a measure of brittle star size (disc diameter), oocyte size and oocyte density. The MSIs ranged from 0 to 800 and had significantly different means among the four gametogenic stages (early growth, growth, mature and spent). The MSI was more sensitive in revealing significant differences between consecutive stages than any of its individual constituents. The MSI was also applied to gametogenic data from the lecithotrophic holothuroid, Mesothuria lactea, again revealing significant differences between successive oogenic stages. This method is expected to be useful in field and experimental studies of gametogenesis in echinoderms (and possibly other taxa), where it is important to detect not just the timing of annual peaks in reproduction but small differences in reproductive status among individuals or populations (e.g. from different habitats or feeding regimes).     

Chemical resistance of gorgonian corals against fungal infections

The frequency and impact of diseases affecting corals throughout the Caribbean have been increasing but little is known about the factors promoting the emergence and outbreak of disease. A disease caused by a fungal pathogen [Aspergillus sydowii (Thom et Church)] which affects Caribbean sea fan corals provided an opportunity to examine the efficacy of coral crude extracts in disease resistance. Minimum inhibitory concentration (MIC) assays showed that of the 20 common gorgonian species in the Florida Keys, extracts from 15 species had MICs < 15 mg ml⁻¹ against A. sydowii pathogenic to sea fans. Extracts from several species in two gorgonian genera (Pseudoplexaura and Pseudopterogorgia) were among the most active, with MICs < 10 mg ml⁻¹. Gorgonia ventalina L., one of two sea fan species known to be hosts to A. sydowii in the field, had an MIC < 10 mg ml⁻¹, suggesting that complete disease resistance requires more active extracts. For the antifungal compounds to be effective in situ, they must also occur in sufficiently high concentrations in living coral tissue. For example, Pseudopterogorgia americana (Gmelin) had comparatively potent extracts but did not have sufficient concentrations in the tissue to be effective. Conversely, Plexaura homomalla Esper extracts were less potent but occurred in high enough concentrations in the tissue to be effective against A. sydowii. When potency and extract concentration are considered together (i.e. potency × concentration), several other gorgonian corals emerge as likely hosts to A. sydowii. Crude extracts from the most active gorgonian species were also effective against two geographic variants of A. sydowii pathogenic to sea fans, a non-pathogenic terrestrial strain of A. sydowii, and three strains of A. flavus Link known to be human, plant, and insect pathogens (MIC range, 7.5 to > 15 mg ml⁻¹). Although the potency in these assays did not attain a clinically significant level, the potency is comparable to a known antifungal agent, hygromycin B, which had an MIC ≤7.5 mg ml⁻¹ in our assays, highlighting the potential of these gorgonian corals for bioprospecting. Received: 29 May 1999 / Accepted: 22 May 2000     

Spatial variation of growth and gonadal developments of Maurolicus muelleri in the Norwegian Sea and in a Norwegian fjord

The epibenthic megafauna of the high-Arctic Northeast Greenland shelf was investigated by means of seafloor photography and Agassiz trawl catches. At 54 stations in water depths between 40 and 770 m, sequences of color slides, each depicting about 1 m² of the seafloor, were obtained along photographic transects of about 100 to 600 m length. The photographs were quantitatively analyzed for abundance of epibenthic organisms identified by comparison with specimens collected from trawl catches. Megabenthic biomass was estimated by multiplying density values with averge body mass figures. For five dominant brittle star species, the population oxygen uptake and, thus, organic carbon mineralization potential were approximated by applying individual respiration rates of average-sized specimens to density figures. Multivariate analyses of the megabenthic species distribution revealed a distinct depth zonation. Shallow shelf banks (<150 m), characterized by coarse sediments, many stones and boulders as well as negative bottom water temperatures, housed a rich epifauna (30 to 340 ind m^–2, 1.8 to 10.5 g AFDW m^–2), strongly dominated (80 to 98% by numbers) by the brittle stars Ophiocten sericeum and Ophiura robusta. The oxygen uptake by brittle stars ranged from 0.4 to 95 mol O₂ m^–2 h^–1 (i.e., assuming a respiratory quotient of 0.8, an organic carbon mineralization of 0.1 to 21.9 mg C m^–2 d^–1). At the bank flanks sloping to the shelf troughs (100 to 580 m), finer sediments prevailed, stones were rare, and bottom water temperatures were positive due to the inflow of Atlantic water. Compared to bank sites, total epibenthic abundances as well as carbon mineralization by brittle stars were roughly ten times and total biomass about four times smaller. In deep shelf depressions as well as at the continental slope (200 to 770 m), stones were completely lacking, and sediments very fine. Epibenthic standing stock and carbon mineralization were one to two orders of magnitude lower than on the banks. The estimation of brittle star oxygen uptake indicates that a considerable portion of the organic carbon produced in the polynya and partitioned to the benthos may be remineralized by epibenthic bank assemblages.     

Comparative biometry of Eunicella singularis (Gorgonian) sclerites at East Mediterranean Sea (North Aegean Sea, Greece)

The two forms of Eunicella singularis sclerites, spindles and clubs, were examined in two sites in East Mediterranean Sea, Greece. The comparison of the sclerites size was based on the hydrodynamic profile of the sites (the site of Arethoussa was exposed and the Phidonissi site was sheltered), on the sites bathymetry (AR 5–8 m, AR 9–13 m, PH 5–10 m, PH 11–13 m) and on the position occupied by the sclerites on the gorgonian colonies (top, middle and base). With the usage of image system analysis the following microfeatures were measured or calculated: projection, length, width, elongation factor, compactness factor, length × width and length/width. The present study confirms the hypothesis that spindles are involved in the recovering of the gorgonian colonies and clubs contribute to the flexion capacity of the colonies. The spindles of the shallow E. singularis colonies (AR 5–8 m) were significantly bigger (length, width) compared with the others, in order to assure the best recovery of the colonies after their flexion, induced by higher water current velocity in shallow water. The clubs occupy the intern part of the mesoglea and their size (length, width) increased in colonies from the shallow exposed site in order to facilitate the gorgonian flexion. The present study demonstrates the gorgonian adaptation (in a microscopic scale) to the environmental pressure of hydrodynamic conditions.     

Development of a new standardised method for sustainable monitoring of the vulnerable pink sea fan Eunicella verrucosa

The aim of this survey was to test a standardised method to follow the demographic evolution of a dense aggregative ‘forest’ population of the temperate gorgonian Eunicella verrucosa (Octocorallia, Gorgoniidae) using in situ photographic recordings. Distribution, density, growth and demographic evolution of the colonies was compared along two parallel transects. Computer treatment allowed the estimation of the total branch fan surface area, and the individual growth of tagged colonies was determined by measuring the increase in this surface area, using consecutive photographs taken at two-year intervals. To integrate the potential bias of branch overlapping, we proposed a correction factor between the in situ photographic surface area and the surface area of the gorgonian calculated from ex situ photographic surface area with all branches deployed. The surface-frequency distribution of colonies was converted to estimated-age-frequency distribution using an estimated growth curve based on the net growth rate. The technique used revealed significant differences in population structure and the dynamics of gorgonian colonies, as the two transects appeared to be influenced by different environmental conditions. The recruitment also seemed to vary according to year and transect, resulting in different densities. Our work showed clear results in characterising the variations of gorgonian demographic evolution at a small spatial scale; thus, it is assumed that this method could be a sustainable tool for coastal environmental managers.     

Population genetic structure of North American <Emphasis Type="Italic">Ophiactis</Emphasis> spp. brittle stars possessing hemoglobin

With the discovery of previously unreported populations of hemoglobin-possessing Ophiactis from the Texas coast in the Gulf of Mexico, an investigation into its population structure, including populations of O. simplex from the Pacific coast of California and O. rubropoda from the Atlantic coast of Florida, was undertaken using DNA sequence data from the mitochondrial COI gene. The reconstructed haplotype network suggests that California populations contain the ancestral source of mtDNA variation, and there is no evidence of recent introductions into Texas. Population genetic analyses reveal the California, Florida, and Texas Ophiactis populations to each be significantly differentiated from one another. Sequence divergence among the three areas is shallower than would be predicted given biogeographic history. Texas and Florida populations are equally genetically diverged from California populations as they are to one another, despite the greater potential for gene flow between these areas. The genetic distinctiveness of the Texas populations and the concordance of this pattern with phylogeographic patterns in other brittle star systems indicate an isolated and independent evolutionary history and we hypothesize that the three geographic regions included in this study each serve as hypotheses of population-level lineages that remain to be tested with independent sources of data.     

Possible control mechanism of cell motility in the gorgonian Eunicella cavolinii

In a previous study it was demonstrated that a lectin controls cell-cell interaction in the gorgonian Eunicella cavolinii (Koch) as a negative modulator. Now we describe the procedure to purify this lectin to homogeneity; its molecular weight is 23 400. The homologous proteoglycans were identified as positive modulators of cell-cell (and/or cell substrate) interaction. The purified single proteoglycan aggregates were 1200±700 nm long and the distance between the attachment points of the proteoglycan subunits was about 45 nm. The glycosaminoglycan residues of the gorgonian proteoglycans were identified as hyaluronic acid (35.5%), heparan sulfate (47.9%) and dermatan sulfate (14.1%). Binding studies with immobilized homologous proteoglycan preparations revealed that gorgonian cells attach to this substratum presumably via its glycosaminoglycans. Quantitative determinations of these two modulators of cell-cell recognition in the three regions of individual colonies revealed concentration gradients these concentration differences reflect a lower cell motility in the polyp-bearing branches and a higher motility of the cells in the polyp-free stem and in the basis, is discussed.     

<Emphasis Type="Italic">Symbiodinium</Emphasis> sp. associations in the gorgonian<Emphasis Type="Italic"> Pseudopterogorgia elisabethae</Emphasis> in the Bahamas: high levels of genetic variability and population structure in symbiotic dinoflagellates

Little is known concerning the fine-scale diversity, population structure, and biogeography for Symbiodinium spp. populations inhabiting particular invertebrate species, including the gorgonian corals, which are prevalent members of reef communities in the Gulf of Mexico, the Caribbean, and the western Atlantic. This study examined the Symbiodinium sp. clade B symbionts hosted by the Caribbean gorgonian Pseudopterogorgia elisabethae (Bayer). A total of 575 colonies of P. elisabethae were sampled in 1995 and 1998–2000 from 12 populations lying along an ~450 km transect in the Bahamas and their Symbiodinium sp. clade B symbionts genotyped at two polymorphic dinucleotide microsatellite loci. Twenty-three unique, two-locus genotypes were identified in association with these P. elisabethae colonies. Most colonies hosted only a single Symbiodinium sp. clade B genotype; however, in some instances ( n=25), two genotypes were harbored simultaneously. For 10 of the 12 populations, 66–100% of the P. elisabethae colonies hosted the same symbiont genotype. Added to this, in 9 of the 12 populations, a Symbiodinium sp. clade B genotype was either unique to a population or found infrequently in other populations. This distribution of Symbiodinium sp. clade B genotypes resulted in statistically significant ( P<0.05 or <0.001) differentiation in 62 of 66 pairwise comparisons of P. elisabethae populations. Tests of linkage disequilibrium suggested that a combination of clonal propagation of the haploid phase and recombination is responsible for maintaining these distinct Symbiodinium sp. clade B populations.     

Phylogeography of the Antarctic planktotrophic brittle star Ophionotus victoriae reveals genetic structure inconsistent with early life history

In the marine environment, connectivity is influenced by physical oceanography as well as life history and behavioral traits, which in combination with historical events such as geology, physical oceanography, and climate, determine population structure. The Antarctic brittle star Ophionotus victoriae develops via a feeding planktonic larval stage, and therefore has potential for long-distance dispersal throughout its Antarctic/subantarctic range. To evaluate this hypothesis, phylogeography of this ecologically dominant species was elucidated by sequence analysis of two mtDNA genes from individuals collected throughout the Antarctic Peninsula and from two subantarctic islands. Counter to expectations of genetic homogeneity, mtDNA data revealed substantial levels of genetic differentiation as well as diversity. Although there were some genetically homogeneous populations, such as those throughout Bransfield Strait, we found O. victoriae to have significant population structure throughout much of the Antarctic Peninsula, with evidence of potential cryptic speciation between the western and eastern Antarctic Peninsula. Furthermore, Antarctic Peninsula populations were genetically distinct from subantarctic island populations. The low levels of connectivity implied for O. victoriae contrast with those found for many other Antarctic benthic taxa, and suggest a complex interplay between oceanography, recent climate history, and larval ecology.     

Chemical mediation of invertebrate defensive behaviors and the ability to distinguish between foraging and inactive predators

When the predatory sea star Pycnopodia helianthoides was placed upstream, the sea urchin Strongylocentrotus purpuratus responded defensively by extending and opening its globiferous pedicellariae. No pedicellaria response was given in control seawater or when the sea star was downstream. The snail Tegula funebralis responded by moving up vertical surfaces when Pycnopodia helianthoides or when Pisaster ochraceus were placed upstream. When these sea stars were introduced downstream, the snail's response was not significantly different from that in control seawater. Water collected from an aquarium containing a single sea star was sufficient to trigger the response of S. purpuratus and T. funebralis; the physical presence of the sea star was not essential. This indicated that a chemical stimulus was involved, and the lack of responses when sea stars were downstream argued strongly against the possible additional involvement of visual or vibrational stimuli. S. purpuratus gave stronger pedicellaria responses to water flowing over an active Pycnopodia helianthoides than to water flowing over the same sea star when it was inactive. The significance of the ability to distinguish between actively foraging and inactive predators is discussed, and a mechanism is proposed to explain differences in the amount of stimulatory chemicals released by active and inactive sea stars.     

Ecological roles of natural products from the marine sponge Geodia corticostylifera

In the Brazilian coast, high numbers of the small brittle star Ophiactis savignyi usually live associated with the sponge Geodia corticostylifera (Demospongiae, Geodidae), but not with other sympatric sponge species. In order to check whether this association was related only with the physical shelter provided by the sponge body or was chemically mediated, the crude organic extract of G. corticostylifera was added to sponge mimics made of phytagel and spongin skeleton. Control and treated mimics were simultaneously offered to previously sponge-associated O. savignyi in both static seawater and flow-through laboratory experiments. Ophiuroids were allowed to move towards the preferred mimic. The defensive properties of the sponge extract against fish predation and fouling were also evaluated. Chemotaxis assays showed that symbiotic ophiuroids were able to chemically recognize its host sponge, moving significantly more towards mimics containing G. corticostylifera extract. Chemical deterrence assays showed that the natural concentration of the extract of this sponge was also able to inhibit generalist fish predation on field experiments and the attachment of the common mussel Perna perna in laboratory assays. These results indicate that the crude extract of G. corticostylifera plays multiple functions in the marine environment, presumably being responsible for a closer association of this sponge with O. savignyi, providing protection for this ophiuroid and inhibition of epibionts on itself.     

On the ecology of a North Adriatic benthic community: Distribution,standing crop and composition of the macrobenthos

The boundaries as well as the macro-epibenthic biomass distribution and composition of a North Adriatic benthic community, dominated by the brittle star Ophiothrix quinquemaculata (D.Ch.), the sponges Reniera ssp., and the ascidians Microcosmus ssp. were determined. Transects totalling more than 80 km (i.e., a recorded area of approximately 150 000 m²) were taken by means of a combined TV- and photocamera sled. The observations, TV-recordings, and photos, together with 392 diver-collected quantitative samples were evaluated. The biomass values were used to establish isobenths. Within the community, the mean biomass, measured as wet weight, amounted to 370 (±73) g/m² with maxima of more than 1000 g/m². 64% of the biomass was due to the designating group Ophiothrix-Reniera-Micrcosmus, 87.5% were represented by filter- and suspension-feeding species alone. The mean biomass in the peripheral areas was evaluated at 166 (±62) g/m². Biomass distribution and composition is examined, and the ecological function and meaning of the observed patterns is discussed.This investigation was carried out within the frame of a joint programme between the Lehrkanzel für Meeresbiologie, University of Vienna, and the Marine Biological Station, Portoroz, University of Ljubljana.     

Cryopreservation of the dinoflagellate symbiont of the octocoral <Emphasis Type="Italic">Pseudopterogorgia elisabethae</Emphasis>

In this paper we describe a cryopreservation protocol followed by the culture of Symbiodinium sp. isolated from the Caribbean gorgonian Pseudopterogorgia elisabethae as a potential renewable source of the dinoflagellate symbiont. Four different freezing protocols were designed: a controlled cooling device designed to cool at 1°C/min, a three-step protocol (−20°C for 2 h, −70°C for 2 h, liquid nitrogen-LN₂), a two-step protocol (−70°C for 2 h, LN₂), and a one-step protocol (LN₂). All cells were stored in LN₂ after cryopreservation. The cryoprotective agents (CPA) used were ethanol (EtOH) and methanol (MeOH) at 10 and 20%, and seawater (FSW) was used as a control. Viability measurements using cell counts showed that all cryopreservation protocols were relatively successful, and no trends were observed regarding freezing protocol or CPA used. After 19 weeks in culture the viability of samples which had high biomass was determined by the fluorescent assay CellTiter Blue™. The most viable cultures were those cryopreserved by a two-step protocol using 20% MeOH or 20% EtOH as a CPA. A genetic examination of the DNA of these samples using Symbiodinium-specific PCR primers confirmed that the composition of the culture had not changed. For the first time, we report that Symbiodinium sp. isolated from a gorgonian can be cryopreserved and subsequently cultured successfully. Lory Z. Santiago-Vázquez and Nealie C. Newberger contributed equally to this publication.     

Calcium and magnesium carbonate concentrations in different growth regions of gorgonians

Four of the most abundant gorgonian species from the southwestern Cape waters, Eunicella papillosa (Esper, 1797), E. alba (Esper, 1797), E. tricoronata Velimirov, 1971 and Lophogorgia flamea (Ellis and Solander, 1786) were analysed for Ca and Mg by atomic absorption spectroscopy (AAS) and ethylenediaminetetraacetate (EDTA) titration. The total mineral content in the peripheral tissues, excluding the axial skeleton, expressed as the sum of CaCO₃ and MgCO₃ of dry matter was between 65.5 and 83.5%. The mineral content varied in different growth regions and all specimens showed a higher degree of mineralization at the base than at the branch tips. The MgCO₃ concentration varied with genus and species and was between 9 and 11 mol %. The variation of the MgCO₃ concentration within different growth regions of the same species was small and generally did not exceed 0.8 mol %. From the branch to the stem, CaCO₃ and total mineral content was found to increase. The CaCO₃:MgCO₃ rations in different growth regions of all species indicated that the composition of the mesoskeleton with regard to the relative concentration of CaCO₃ and MgCO₃ is constant throughout the animal. Mineralogically, the mesoskeleton consists of high magnesian calcite as identified by X-ray diffraction. MgCO₃ concentrations determined by the peak shift method and by AAS were in fair agreement. The MgCO₃ data in gorgonian samples from the cold Atlantic Ocean and the warmer Indian Ocean show a linear relationship between water temperature and MgCO₃ concentration already demonstrated by Chave (1954). However, our data were consistently lower by 1 to 2% than expected.     

Prey defense,predator preference,and nonrandom diet: The interactions between Pycnopodia helianthoides and two species of sea urchins

Interactions between the predatory sea star Pycnopodia helianthoides (Brandt, 1835) and two of its natural prey, the sea urchins Strongylocentrotus purpuratus (Stimpson, 1857) and S. franciscanus (Agassiz, 1863), are examined with regard to predator preference, predator diet, and prey defenses. The sea star is able to detect both species of sea urchin upstream in a Y-trough, but does not consistently choose one over the other (i.e., no preference). However, when the sea star is presented with equal numbers of similar-sized specimens of the two species of sea urchin, its diet is markedly nonrandom, since S. purpuratus is eaten almost 98% of the time. The defensive responses of the two species of sea urchin differ in form and effectiveness. S. franciscanus employs its long spines as defensive weapons, pinching the rays of an attacking sea star. This defensive response is more effective than the pedicellarial response used by S. purpuratus. The nonrandom diet of the predator seems to result primarily from prey defensive responses that differ in effectiveness, rather than from an intrinsic, behavioral preference of the predator at an earlier stage in the predator/prey interaction.     

Matching spatial distributions of the sea star Echinaster sepositus and crustose coralline algae in shallow rocky Mediterranean communities

Understanding why a species is present in a particular location and the consequences of its presence is complex but necessary to identify the mechanisms that generate and maintain ecological diversity. The common sea star Echinaster sepositus can be either very abundant or non-existing in nearby localities of the western Mediterranean. Yet, the factors that shape its distribution and the impact of the sea star on natural communities remain uninvestigated. Here, we quantified multiple biotic and abiotic factors that may affect the distribution of E. sepositus and tested whether this sea star can shape the organization of the community it inhabits. Our results showed that the distribution of this sea star was highly contagious and positively correlated with the abundance and distribution of crustose coralline algae from tens of meters to tens of kilometers. Despite significant differences in community composition between localities with high or low abundance of the sea star, experimental addition of E. sepositus to natural communities failed to shift the composition of the algal community in 4 months. Overall, our results suggest that within habitat variability in the abundance of crustose coralline algae may explain the abundance of E. sepositus at multiple geographic scales, emphasizing the need to investigate small-scale processes at larger geographic scales.