Comparative study of the chemistry and structure of gorgonian and antipatharian coral skeletons

The comparative morphology of the axial skeleton was studied in representatives of 5 reef-dwelling gorgonian families and 2 antipatharian genera by electron and light microscopy. Comparative axial histochemistry, amino acid composition and the physico-chemical character of the protein were also studied. Evidence obtained indicates that collagen is a widespread and prominent structural feature of the gorgonian axial skeleton. Aromatic crosslinkage (sclerotization) also appears to be of widespread, if not universal occurrence in these animals. Sclerotization is equally characteristic of the black coral skeletons studied, but this material does not contain collagen. Its unusual composition, structure, and reactivity is discussed.     

Population structure in the coral Pavona cactus: clonal genotypes show little phenotypic plasticity

Electrophoretic data were used to examine the relationship between genotype and growth form, and to assess the contribution of asexual reproduction to recruitment within six local populations of the agaricid coral Pavona cactus from the central and northern Great Barrier Reef. The data revealed the presence of highly replicated clonal genotypes in the five densest populations. In three cases, samples of 50 to 60 colonies collected from 25 m² areas within the Eclipse Island and Pandora Reef populations each consisted of colonies with only two distinct 4-locus genotypes. More intensive sampling of the population at Eclipse Island showed that colonies with the same 4-locus genotypes were separated by distances of up to 93 m. In contrast, the population at Watson's Bay (Lizard Island) consisted of a few widely scattered and genetically distinct colonies. The samples collected from each population contained a range of growth forms including, in some cases, the entire morphological range described for this species. A strong association of genotype and growth form was detected in samples from all populations, with the exception of Watson's Bay where no such comparison was possible. Nevertheless, some genotypes were represented by more than one growth form and this could reflect the effects of limited phenotypic plasticity. The effects of asexual reproduction reduced the value of these data as a test of the genetic connectedness of the six populations studied. The genetic distance between samples was not simply correlated with geographic distance. This may reflect either the true structure of the breeding population(s) or the effects of asexual reproduction on estimates of allelic frequencies.Contribution No. 47 from the Ecology and Genetics Group of the University of Wollongong     

The gorgonian coral Acabaria biserialis: life history of a successful colonizer of artificial substrata

The azooxanthellate coral Acabaria biserialis Kükenthal, 1908 (Octocorallia: Alcyonacea) is highly abundant on the vertical underwater structures of the oil jetties at Eilat (Red Sea), but it is rare on the adjacent natural reefs. To understand its success on such artificial substrata we examined various aspects of its life history and population dynamics. A. biserialis is a gonochoric brooder. The percentage of colonies on the artificial substrata bearing gonads ranged from none (September 1994 and 1995) to 100% (January 1994, 1995 and April 1994). Mature oocytes and sperm sacs reached rather small maximum diameters of 240 and 160 μm, respectively, probably dictated by the small gastrovascular cavities. A continuous release of planulae was observed in the laboratory from March to July 1995. Planulation occurred during various lunar phases, and both by day and night. Recruitment of A. biserialis on PVC plates attached to the jetties coincided with the breeding period inferred from the laboratory findings. Recruits reached a maximum height of 3 cm within 3 months, thus exhibiting a remarkably rapid growth rate compared with other gorgonians. This high colony growth rate may compensate for the lack of clonal propagation in the studied A. biserialis population. The complex substrata of the jetties provide the conditions required for successful colonization, which include upside-down orientation, an adequate light regime and exposure to flow. The findings of our study may provide a useful contribution to the design of artificial reefs aimed at attracting rapid colonization by A. biserialis colonies. Received: 30 December 1998 / Accepted: 19 August 1999     

Spatial autocorrelation analysis of small-scale genetic structure in a clonal soft coral with limited larval dispersal

The philopatric larval dispesal and small effective population sizes characteristic of many clonal species should promote the development of significant small-scale genetic structure within populations as a result of isolation-by-distance. We used spatial autocorrelation statistics to detect genetic structure, arising from both clonal reproduction and philopatric dispersal of sexual propagules, for five allozyme loci within populations of the soft coral Alcyonium sp. In a population on Tatoosh Island, Washington, USA, sampled in 1991/1992, we found significant positive spatial autocorrelation at all loci among individuals separated by <40 cm, reflecting the presence of significant smallscale genetic structure due to associations among clonemates. For 4 of 5 loci, however, we detected no significant spatial autocorrelation among the different clones within this population over distances of 1 to 40 m. Analysis of soft-coral populations from six additional, topographically diverse sites in the north-east Pacific also did not reveal significant spatial autocorrelation among clones at any loci. This general lack of spatial autocorrelation of genotypes among clones suggests that significant small-scale genetic structure has not arisen in populations of Alcyonium sp. as a consequence of isolation-by-distance.     

Linking local to global properties in branching modular networks: gorgonian coral colonies

Branching growth is present both in plants and animals, either marine or terrestrial. Although cellular and other modular levels of organization in plants and animals have evolved through different molecular and physiological mechanisms, several aspects of their branching modular system and morphology are similar. We studied vessel organization and colony integration, in order to comprehend underlying relationships between different structural components in a gorgonian coral network. The theoretical formalism was validated in the gorgonian coral Eunicea mammosa (Plexauridae, Octocorallia) in Belize. As in vascular plants, these colonial animals create a complex network of connections among modular branches integrated in stem canals downstream toward the base. A new formalism is proposed for describing gorgonian branching. A global property of a colony is for instance the size of its base or its weight whereas a local property is the size of branch in a particular place of the colony. However, a global property is not the simple addition of local modular properties, as the case of stem canals in the colony base. Theoretically, the process of branching is tightly intertwined with the internal network organization. The colony network centralization is driven by a linear relationship between the total number of branches and the stem canals at the base of the colony. If stem canals play important roles in the transport of nutrients throughout the colony and the biomechanical support from the base up to the tips, we can assume that there is an underlying association between the number of stem canals at the base and the number of for example, terminal branches. These associations may provide new findings that extend our understanding of the functional organization of tree-like networks in octocorals and their vascular systems. The idea that the external components of a tree-like plant network are directly correlated and connected down to the main trunk seems to be analogous in an animal system.     

DNA fingerprints of orange roughy,Hoplostethus atlanticus: a population comparison

Genetic variability among Hoplostethus atlanticus collected from two spawning grounds east and west of New Zealand was examined using DNA fingerprints as revealed by hybridization with three clonal probes: 33.15, M13 and 3HVR. The 33.15 and 3HVR fingerprints revealed a complex pattern of restriction fragments, apparently refecting a multi-locus system of highly variable minisatellite alleles similar to the pattern of alleles reported in other vertebrates. The M13 fingerprints revealed a distinct pattern of restriction fragments of high molecular weight, reflecting a single-locus system that overlapped with the family of minisatellite alleles observed in 33.15 fingerprints. In a sample of 12 orange roughy collected on a single regional spawning site, the average percent similarity of 33.15 fingerprints was 21.15% (SD=17.75), the average percent similarity of 3HVR fingerprints was 14.32% (SD=14.45) and the inferred average allelic frequency of the M13 single-locus system was 0.071. A comparison of 33.15 and M13 fingerprints from two distant spawning sites ground New Zealand revealed no obvious regional differences. The variability of orange roughy fingerprints was so great, however, that regional comparisons could not be considered conclusive indicators of genetic identity. Our results provide a preliminary assessment of the power and pitfalls of using DNA-level markers for the population analysis of marine fish.     

Genetic variation and clonal structure in the scleractinian coral Pocillopora damicornis in the Ryukyu Archipelago, southern Japan

Samples of the scleractinian coral Pocillopora damicornis were collected from six sites located around four islands in the Ryukyu Archipelago, southern Japan, and subjected to allozyme electrophoresis. Seven polymorphic loci were examined for their allelic patterns. The ratio of observed to expected genotypic diversity (0.30 < G _o :G _e  < 0.64), the ratio of the observed number of genotypes to the number of individuals (0.47 < N _g :N _i  < 0.75), and deviations from Hardy–Weinberg equilibrium indicated that asexual reproduction plays a major role in the maintenance of established populations. However, populations were not completely dominated by a single or a few clones, and most clones were represented by only a few individual samples. The high frequency of typhoons in the region suggests that, in P. damicornis, fragmentation caused through occasional exposure to powerful waves is a major mode of asexual reproduction, but asexual production of planulae may also be contributing to the maintenance of populations. A significant genetic differentiation (F ST) was found between the six populations examined (0.027 < F ST < 0.092, average F ST = 0.056). The moderate gene flow is discussed according to characteristics of the larval stage of the species, and to circulation patterns in the region. Received: 7 August 1998 / Accepted: 18 May 1999     

Hidden genetic diversity in a key model species of coral

A total of 43 colonies of the scleractinian coral Pocillopora damicornis from lagoonal and reef slope sites in the western Indian Ocean (WIO) region were genetically characterised at one nuclear and two mitochondrial sequence markers and six microsatellite loci. Both mitochondrial and microsatellite data support the existence of two reciprocally monophyletic clusters (F- and NF-types) and provide evidence of the existence of two cryptic species of P. damicornis on reefs in WIO region and put current morphological delineation and geographical boundaries of P. damicornis and Pocillopora molokensis into question. The results add to ongoing studies on the phylogeny and phylogeography within the genus Pocillopora, which all point towards a range of unresolved morphological and molecular species boundaries. Nuclear phylogenies derived from the present and previously published sequences show evidence for incomplete lineage sorting and/or introgressive hybridisation between Pocillopora morphospecies. However, the two WIO types largely remain in separate clusters, further supporting the theory that these represent two different species.     

Nucleotide sequencing of highly repetitive DNA from seven species in the coral genusAcropora (Cnidaria: Scleractinia) implies a division contrary to morphological criteria

Nucleotide sequences have been determined for 31 homologous 118 base-pair highly repeated DNA sequences from seven species ofAcropora. A matrix was constructed from the sequence data and subjected to phylogenetic analysis using heuristic search routines in the PAUP (phylogenetic analysis using parsimony) program, Version 3.0L. These analyses confirm a close relationship between two species of one subgeneric group (A. pulchra andA. millepora), but identify a division in a group of six species which is contrary to taxonomic groupings based on morphological criteria.     

Indices for detecting differences in species composition: some simplifications of RDA and CCA

We provide algebraic simplifications for the redundancy analysis (RDA) eigenvalue and the canonical correspondence analysis (CCA) eigenvalue in the special case of permanent plots sampled twice. The indices for RDA and CCA are interrelated and are intuitively interpretable. These simplifications also apply to simple split-plot designs and to a balanced design with two independent samples.     

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     

Resident mysids: community structure,abundance and small-scale distributions in a coral reef lagoon

Seasonal and diel variations in community structure and abundance of coral-reef lagoon mysids were examined at Davies Reef in the central region of the Great Barrier Reef (GBR) between June 1980 and May 1981. Twenty-five mysid species belonging to three subfamilies of the family Mysidae were captured during the study, with six new records for the GBR. The epibenthic mysid community differed from that in the overlying water, was faunistically uniform, but formed characteristic seasonal and diel groupings. The dominant epibenthic species were Erythrops sp., Anisomysis pelewensis, Doxomysis littoralis, A. laticauda, Prionomysis stenolepis, A. lamellicauda, and A. australis, five of which formed schools. Total mysid abundances ranged between 110 and 790 m^-3 with peak abundance in October. Schooling species occurred at local densities of up to 500 000 m^-3. Mysids were absent from shallow and midwater depths during the day, but were distributed throughout all depths at night with peak abundances in mid-water and deep layers. The dominant species in the water column at night were Pseudanchialina inermis, A. laticauda and Gastrosaccus indicus, in descending order of abundance. Lagoonal mysids contribute little to the food of sessile reef planktivores, as all but three species remain concentrated near or on the lagoon floor both day and night. The contribution of resident lagoon mysids to reef trophodynamics is probably through remineralization of lagoon detritus. Given the vast reef areas comprised of sandy lagoons, the large populations and relatively large size of lagoon mysids, this trophodynamic role may be of considerable importance.A.I.M.S. Contribution No. 477     

Stability and structure of a fish community on a coral patch reef in Hawaii

Two complete collections of the fishes residing on an isolated coral patch reef ( 1500 m²) at Oahu, Hawaii, were made 11 years apart. Of the 112 species of fishes in both collections combined, only 40% were in common, but these made up more than 85% of the wet biomass in each collection. The two assemblages of fishes were similar in trophic structure and standing crop. Many coral reef fish communities are dominated by carnivorous forms. In the present study, planktivorous fishes were the most important trophic group in the community; this was related to abundant zooplankton resources. Following the second collection in 1977, recolonization by fishes was followed for 1 year. Recolonization proceeded rapidly and was primarily by juvenile fishes well beyond larval metamorphosis. Within 6 months of the second collection, the trophic structure had been re-established. The MacArthur-Wilson model of insular colonization described the recolonization process and predicted an equilibrium situation in less than 2 years. The recolonization data suggested that chance factors may explain the colonization process on a small scale, but a relatively deterministic pattern emerged when considering the entire reef. Thus, at the community level the fishes are a persistent and predictable entity.     

Mitochondrial DNA sequence variation in deep-sea bamboo coral (Keratoisidinae) species in the southwest and northwest Pacific Ocean

The Keratoisidinae are a poorly known, but phenotypically diverse group of deepwater corals. Recent developments in deepwater trawling in the southwest Pacific have provided many more specimens of bamboo corals. Two sub-regions of the mitochondrial genome were sequenced to test genetic relationships among specimens collected over a wide geographical range (27–50° S): a sub-region of the large-subunit rRNA (16S rRNA), characterized by a highly variable insertion/deletion (INDEL#2) region; and a non-coding region between COII and COI. Based on DNA haplotypes, 14 species of Keratoisidinae were recognized among 88 specimens from deep water in the southwest Pacific Ocean. The common haplotypes also appeared in specimens collected in the northwest Pacific Ocean and may indicate that some bamboo coral species are widespread in the Pacific, or that the mitochondrial markers are insensitive to recent speciation events. Many specimens were taken from flat bottom areas and, contrary to assumptions, the bamboo corals are not endemic to seamounts. The closure of some seamounts to trawling will protect bamboo corals from extinction, but not from local depletion.Communicated by M.S. Johnson, Crawley     

The effect of micrograzers on algal community structure in a coral reef microcosm

The effect of amphipod grazing on algal community structure was studied within a 75 l refuge tank connected to a 6500 l closed-system, coral reef microcosm. When amphipods (Ampithoe ramondi) were absent or present in low numbers, a high biomass of mostly filamentous algal species resulted, including Bryopsis hypnoides, Centroceras clavulatum, Ceramium flaccidum, Derbesia vaucheriaeformis, Enteromorpha prolifera, Giffordia rallsiae, and Polysiphonia havanensis. These microalgae disappeared when amphipod density increase beyond approximately 1 individual cm^-2 of tank surface. The macroalga Hypnea spinella germinated in the system in association with amphipod tube sites. H. spinella plants remained rare until filamentous species were eliminated by amphipod grazing. Feeding trials confirmed that H. spinella was protected from grazing by its size rather than a chemical defense strategy. The H. spinella community we observed is similar to the flora described on algal ridges where physical conditions exclude fish grazing. We suggest that amphipods and similar micrograzers are responsible for the algal community structure of these ridges. Caging experiments may be subject to similar effects from increased amphipod grazing on the algae. Introduction of fish that are amphipod predators into the refuge tank caused an increase in algal species diversity but total H. spinella growth rates fell from 25 g dry wt month^-1 to less than 8 g dry wt month^-1. We describe amphipod behavior in relation to changes in population density and food supply, and we stress the potential for increasing the productivity of commercial seaweeds through maintenance of appropriate amphipod species in mariculture facilities.     

Size and structure of echinoderm populations associated with different coexisting coral species at Aldabra Atoll,Seychelles

The associated echinoderm fauna of 125 living heads of massive microatolls of Porites spp., branched heads of Porites nigrescens Dana and small, densely packed heads of the hydrocoral Millepora exacea Forskål of a shallow sublittoral lagoon flat at Aldabra Atoll, Seychelles was examined. The weight and number of echinoderms increased with coral size in all cases. The microatolls were the largest coral heads and sheltered the most diverse, underlying echinoderm cryptofauna comprising the largest bodied species, regardless of class or family. The infauna in heads of P. nigrescens was less diverse and smaller bodied than the echinoderms under microatolls. Only with P. nigrescens did species number increase with increasing coral size. Heads of M. exacea were the smallest and provided a habitat of compact interstitial spaces for the most depauperate, smallest bodied infauna which included juveniles of larger species. Ophiuroids dominated the coral-associated echinoderm faunas whereas larger bodied classes, such as echinoids and holothuroids, were less well represented. Large bodied ophiuroids such as certain ophiodermatids occurred only under microatolls, whereas the usually small bodied ophiotrichids dominated in the heads of M. exacea. The generally ubiquitous ophiocomids were well represented in, or under all corals. Heads of M. exacea could be nursery areas for recently metamorphosed juvenile ophiuroids and echinoids which, as they grow, seek shelter in corals with larger interstitial spaces such as P. nigrescens. Significant interspecific ophiuroid associations were related to space availability for larger species under microatolls and in heads of P. nigrescens. The possibility of microhabitat partitioning in corals among some ophiuroids is suggested.     

Habitat biodiversity as a determinant of fish community structure on coral reefs

Increased habitat diversity is often predicted to promote the diversity of animal communities because a greater variety of habitats increases the opportunities for species to specialize on different resources and coexist. Although positive correlations between the diversities of habitat and associated animals are often observed, the underlying mechanisms are only now starting to emerge, and none have been tested specifically in the marine environment. Scleractinian corals constitute the primary habitat-forming organisms on coral reefs and, as such, play an important role in structuring associated reef fish communities. Using the same field experimental design in two geographic localities differing in regional fish species composition, we tested the effects of coral species richness and composition on the diversity, abundance, and structure of the local fish community. Richness of coral species overall had a positive effect on fish species richness but had no effect on total fish abundance or evenness. At both localities, certain individual coral species supported similar levels of fish diversity and abundance as the high coral richness treatments, suggesting that particular coral species are disproportionately important in promoting high local fish diversity. Furthermore, in both localities, different microhabitats (coral species) supported very different fish communities, indicating that most reef fish species distinguish habitat at the level of coral species. Fish communities colonizing treatments of higher coral species richness represented a combination of those inhabiting the constituent coral species. These findings suggest that mechanisms underlying habitat-animal interaction in the terrestrial environment also apply to marine systems and highlight the importance of coral diversity to local fish diversity. The loss of particular key coral species is likely to have a disproportionate impact on the biodiversity of associated fish communities.