Sperm morphology of scleractinians from the Caribbean

This study compares the sperm morphology of 2 hermaphroditic (Diploria strigosa, Montastrea annularis) and 2 gonochoric (Siderastrea siderea, Dendrogyra cylindrus) Caribbean reef corals. The spermatozoa of the gonochoric species are morphologically more specialized than those of the hermaphrodites. The cytological differences are apparent in the formation of a Spitzenkörper and pro-acrosomal vesicles, a mitochondrial mass (Nebenkern) and a large lipid vesicle in the gonochorists studied. Harrison (1985) first demonstrated that sperm morphology is correlated with sexual reproductive pattern and systematic position in scleractinian corals, and this is supported by the results of this study. The possible functional and phylogenetic bases of sperm morphology are discussed and a brief outline is given of further studies required.     

Annual cycle of gametogenesis and spawning in a tropical zoanthid,Protopalythoa sp.

Gametogenesis and spawning have been studied since 1986 in a population of Protopalythoa sp. (Anthozoa, Zoanthidea) from Orpheus Island, Great Barrier Reef. Fertile zooids were gonochoric or hermaphroditic with oocytes and sperm vesicles intermingled in the same mesentery; colonies were single sex or mixed. Oocytes arose in macrocnemes below the actinopharynx and were first observed in July. Their mean diameter increased from < 50 µm to 160–240 µm by November, by which time they contained zooxanthellae. Female zooids each produced 800–2400 ova. No testicular vesicles were visible before October but spermatozoa were present in November and December. Spawning in each of three years occurred in November (4th to 6th nights after full moon) simultaneously with mass spawning of scleractinian corals. The buoyant eggs (along with sperm in the hermaphroditic zooids) were released in bundles. Some large oocytes and mature sperm were still present in December.     

Reproductive ecology of Caribbean reef corals

The last decade has seen a resurgence of interest in the processes of sexual reproduction by scleractinian reef corals. Earlier investigations had focused fortuitously on brooding (planulating) species, which resulted in the general misconception that brooding was the main form of larval development of reef corals. More recent work on Indo-Pacific species has shown broadcast spawning and short annual reproductive periods to predominate. This report presents the reproductive patterns of eleven Caribbean coral species and attempts to explain the adaptive features and selective pressures that have led to the evolution of the four reproductive patterns described to date: (a) hermaphroditic broadcasters; (b) gonochoric broadcasters; (c) hermaphroditic broadcasters; (b) gonochoric brooders. Both (a) and (b) correlate with large colony size and short annual spawning periods; and (c) and (d) correlate with small colony size, multiple planulating cycles per year, and occupation of unstable habitats. Selection for outcrossing between long-lived individuals is proposed as the reason for gonochorism and for synchronous spawning of hermaphroditic broadcasters, and also for the large amount of sperm produced by hermaphroditic brooders. Selection for high rates of local recruitment is proposed as the force behind the evolution of brooding by species inhabiting unstable habitats and suffering high rates of adult mortality.     

Hermaphroditic spawning by the gonochoric pillar coral Dendrogyra cylindrus

Coral Reefs - Scleractinian corals typically reproduce sexually as either gonochoric (separate male and female) or hermaphroditic (producing both eggs and sperm) colonies. The Caribbean pillar...     

An experimental analysis of the effects of light and zooplankton on coral zonation

Summary In Panamá, vertical zonation of coral species is well marked; branching corals (Pocillopora spp.) predominate in shallow (1–6 m) water while massive forms (Pavona spp.) occupy the deeper (6–10 m) areas of the reef. To test the hypothesis that this zonation represents differential resource utilization (i.e., niche partitioning of food resources), one year manipulative field experiments were conducted at two depths (1 m and 7 m below Mean Lower Sea Level) assessing the relative contribution of light and zooplankton to the nutrition of three Pacific corals: Pocillopora damicornis, Pavona clavus and Pavona gigantea. Also tested were the related hypotheses that (i) energy for the maintenance and growth of corals comes mainly from light, independent of zooplankton supply and (ii) Porter's model, in which coral morphology is a predictor of the phototrophic-heterotrophic capabilities of a particular species. That is, corals with a branching morphology and a small polyp diameter (and short tentacle length) should be primarily phototrophic while those species with larger polyps and/or a massive form should show a greater degree of heterotrophy.The comparison of caged versus control corals indicated that the branching coral, Pocillopora damicornis (polyp diameter — 1 mm) grew independent of zooplankton supply (>95) but was markedly affected by shading. Pavona clavus (massive form with intermediate size polyps — 2 mm) was also negatively affected by shading but this effect was minimized when zooplankton was present. Surprisingly, under ambient light conditions, the growth of this specics was independent of zooplankton (>95); perhaps indicating a facultative reliance on zooplankton especially in shallow water. The third species, Pavona gigan-tea (massive form with largest polyps — 3 mm) was highly dependent on both light and zooplankton and was unable to compensate for the effects of shading by zooplankton feeding. This high dependence on light was unexpected since large polyps, according to Porter's model, would predict a more heterotrophic existence.Under all treatments, in both shallow and deep water, Pocillopora was found to grow more rapidly than either massive species. Thus, this study indicates that zonation on eastern Pacific reefs cannot be explained by depth-related differences in nutritional requirements among the species present. These results do provide the first long-term, experimental field evidence that corals are largely phototrophic organisms. The validity of Porter's model attempting to correlate phototrophic-heterotrophic abilities to morphological characteristics is, at least, partially verified: species with larger tentacles do appear to utilize zooplankton to a greater extent than those species with smaller tentacles. However, the significance of corallum morphology in predicting the mode of nutrient acquisition is unclear. Rather, behavioral patterns (tentacle expansion-contraction cycles) and location on the reef may be more important factors to consider.     

Ultrastructural observations of the spermatogenesis of the hermaphroditic solitary coral Balanophyllia europaea (Anthozoa, Scleractinia)

Spermatogenesis ultrastructure was studied in a simultaneous hermaphrodite population of the solitary coral Balanophyllia europaea. In this species, spermatogenesis takes place in spermatocysts located within gametogenetic mesenteries surrounded by a bilayered boundary. Spermatogonia and spermatocytes are large flagellate cells, densely packed at the outermost edges of the spermatocyst. Spermatids and sperm are loosely distributed near the centre of the spermatocyst. The cytoplasm of spermatogonia and primary spermatocytes often contains short lengths of free axonemes, probably derived from the reabsorption of a primitive flagellum. Maturing spermatids either contain long intracytoplasmic axonemes, that may be stages of the tail synthesis, or have a flagellum. The morphological features of the sperm of this hermaphroditic scleractinian, very similar to those observed in the sperm of gonochoric taxa, support the hypothesis that the hermaphroditism of this population is an adaptive condition. Accepted: 1 October 1999     

Competitive strategies of soft corals (Coelenterata: Octocorallia). II. Variable defensive responses and susceptibility to scleractinian corals

Interactions involving competition for space between several species of alcyonacean and scleractinian corals were assessed experimentally on Britomart Reef, central region of the Great Barrier Reef, Australia. Colonies of three soft coral species, Sarcophyton ehrenbergi Marenzeller, Nephthea brassica Kukenthal, and Capnella lacertiliensis Macfayden Forskal (Coelenterata:Alcyonacea) were relocated within stands of two scleractinian corals, Parités andrewsi Vaughan (= P. cylindrica Dana) and Pavona cactus Förskal (Coelenterata:Scleractinia). Undisturbed scleractinian and relocated alcyonacean controls were also monitored.Alcyonacean corals induced necrosis of tissue in scleractinian corals. Necrosis was significantly more pronounced when colonies were in contact but was also observed in the absence of contact, implicating the presence of active allelopathic agents. Scleractinian coral species varied in their susceptibility to the ill effects of alcyonaceans, with Pontes andrewsi being more susceptible than Pavona cactus. Of the soft corals, Nephthea caused the highest degree of mortality in the two scleractinian corals examined and Sarcophyton the least. Some soft corals appear to retain their toxins while others release them, implying a combination of anti-predatory and anti-competitor roles for the secondary metabolites. Scleractinian corals were often overgrown by soft corals.Both species of scleractinian corals were found to cause approximately equal amounts of tissue necrosis in alcyonaceans. These effects were more pronounced when colonies were in direct contact. The necrotic effects among alcyonacean corals were species-specific. Alcyonaceans also overgrew scleractinian corals and secreted a protective polysaccharide layer in areas proximal to scleractinians. Secretion of this layer was stimulated differentially by the two scleractinian species and also varied in frequency of occurrence among the alcyonaceans.High levels of tissue necrosis were observed in both groups of organisms within 3 wk of initiation of the experiment. Necrosis increased with time in the scleractinian corals and decreased in the alcyonaceans. The development of a protective polysaccharide layer in the alcyonaceans increased with time.     

A new sequence data set of SSU rRNA gene for Scleractinia and its phylogenetic and ecological applications

Scleractinian corals (i.e. hard corals) play a fundamental role in building and maintaining coral reefs, one of the most diverse ecosystems on Earth. Nevertheless, their phylogenies remain largely unresolved and little is known about dispersal and survival of their planktonic larval phase. The small subunit ribosomal RNA (SSU rRNA) is a commonly used gene for DNA barcoding in several metazoans, and small variable regions of SSU rRNA are widely adopted as barcode marker to investigate marine plankton community structure worldwide. Here, we provide a large sequence data set of the complete SSU rRNA gene from 298 specimens, representing all known extant reef coral families and a total of 106 genera. The secondary structure was extremely conserved within the order with few exceptions due to insertions or deletions occurring in the variable regions. Remarkable differences in SSU rRNA length and base composition were detected between and within acroporids (Acropora, Montipora, Isopora and Alveopora) compared to other corals. The V4 and V9 regions seem to be promising barcode loci because variation at commonly used barcode primer binding sites was extremely low, while their levels of divergence allowed families and genera to be distinguished. A time‐calibrated phylogeny of Scleractinia is provided, and mutation rate heterogeneity is demonstrated across main lineages. The use of this data set as a valuable reference for investigating aspects of ecology, biology, molecular taxonomy and evolution of scleractinian corals is discussed.     

Molecular and morphological supertree of stony corals (Anthozoa: Scleractinia) using matrix representation parsimony

The supertree algorithm matrix representation with parsimony was used to combine existing hypotheses of coral relationships and provide the most comprehensive species-level estimate of scleractinian phylogeny, comprised of 353 species (27% of extant species), 141 genera (63%) and 23 families (92%) from all seven suborders. The resulting supertree offers a guide for future studies in coral systematics by highlighting regions of concordance and conflict in existing source phylogenies. It should also prove useful in formal comparative studies of character evolution. Phylogenetic effort within Scleractinia has been taxonomically uneven, with a third of studies focussing on the Acroporidae or its most diverse genera. Sampling has also been geographically non-uniform, as tropical, reef-forming taxa have been considered twice as often as non-reef species. The supertree indicated that source trees concur on numerous aspects of coral relationships, such as the division between robust versus complex corals and the distant relationship between families in Archaeocoeniina. The supertree also supported the existence of a large, taxonomically diverse and monophyletic group of corals with many Atlantic representatives having exsert corallites. Another large, unanticipated clade consisted entirely of solitary deep-water species from three families. Important areas of ambiguity include the relationship of Astrocoeniidae to Pocilloporidae and the relative positions of several, mostly deep-water genera of Caryophylliidae. Conservative grafting of species at the base of congeneric groups with uncontroversial monophyletic status resulted in a more comprehensive, though less resolved tree of 1016 taxa.     

Effect of light and zooplankton on skeletal δ13C values in the eastern Pacific corals Pavona clavus and Pavona gigantea

 Skeletal δ¹³C levels in symbiotic reef corals are believed to be predominantly influenced by metabolic fractionation. Therefore, environmental variables influencing coral metabolism should also affect skeletal δ¹³C levels. To test this hypothesis, we measured the effects of light (which drives photosynthesis) and relative zooplankton levels (heterotrophy) on skeletal δ¹³C values in the corals Pavona clavus and P. gigantea at two depths (1 m and 7 m). For both species, decreases in light or increases in zooplankton resulted in significant decreases in skeletal δ¹³C levels. A significant decrease in δ¹³C values with depth was observed in Pavona gigantea only. Thus, light and zooplankton directly affect coral skeletal δ¹³C values, supporting the hypothesis that metabolic fractionation significantly contributes to skeletal δ¹³C levels. Simultaneous decreases in both light and zooplankton resulted in decreases in skeletal δ¹³C values, reflecting decreases in light. In Pavona clavus, intra-annual variation in skeletal δ¹³C values over one year correlated with seasonal changes in irradiance. Further study is needed to resolve how skeletal δ¹³C values vary at intermediate levels of irradiance and zooplankton, and in other coral species. Accepted: 14 July 1998     

Symbiont diversity in scleractinian corals from tropical reefs and subtropical non-reef communities in Taiwan

We examined zooxanthellae diversity in scleractinian corals from southern Taiwan and the Penghu Archipelago, a tropical coral reef and a subtropical non-reefal community, respectively. Zooxanthellae diversity was investigated in 52 species of scleractinian corals from 26 genera and 13 families, using restriction fragment length polymorphism (RFLP), and phylogenetic analyses of the nuclear small-subunit ribosomal DNA (nssrDNA) and large-subunit ribosomal DNA (nlsrDNA). RFLP and phylogenetic analyses of nuclear-encoded ribosomal RNA genes showed that Symbiodinium clade C was the dominant zooxanthellae in scleractinian corals in the seas around Taiwan; Symbiodinium clade D was also found in some species. Both Symbiodinium clade C and D were found in colonies of seven species of scleractinian corals. Symbiodinium clade D was associated with corals that inhabit either shallow water or the reef edge in deep water, supporting the hypothesis that Symbiodinium clade D is a relatively stress-tolerant zooxanthellae found in marginal habitats.Communicated by Biological Editor H.R. Lasker     

Recruitment patterns of scleractinian corals in an isolated sub-tropical reef system

The density of recruits of scleractinian corals on settlement plates at Lord Howe Island, a small isolated sub-tropical island 630 km off the Australian coastline, was within the range of values reported for comparable studies on the Great Barrier Reef. However, there was a difference in the relative abundance of taxonomic groups, with recruitment at Lord Howe Island during the summer of 1990/91 dominated by corals from the Family Pocilloporidae, Family Poritidae, and sub-genus Acropora (Isopora) (in order of abundance). By contrast, on the Great Barrier Reef, recruits are generally predominantly species from the Family Acroporidae (other than the Acropora (Isopora) group). Both the recruits and the established coral communities at Lord Howe Island are dominanted by corals which release brooded planulae, as opposed to the pattern of mass-spawning with external fertilisation more typical of Great Barrier Reef corals. I hypothesise that the release of brooded planulae would be advantageous in an isolated reef community because (a) brooded larvae can travel large distances and survive the journey to the isolated reef and/or (b) brooded larvae have a shorter period before they are competent to settle and are therefore more likely to be retained on the parental reef once a population has been established.     

Spatio-temporal variation in coral recruitment at different scales on Heron Reef, southern Great Barrier Reef

 Recruitment of scleractinian corals on settlement plates at Heron Island, Great Barrier Reef, was examined over four years (September 1991–September 1995) to quantify spatio-temporal patterns at different scales and to assess post settlement mortality. Recruitment was dominated by pocilloporid corals which accounted for 80.1% of the 8627 spat counted, whereas non-isoporan acroporids represented only 16.4%. Poritids, faviids and isoporan acroporids rarely recruited to the plates (3.5%), despite their obvious abundance as adults on the reef. Recruitment patterns on the plates indicate strong space-time interactions as evidenced by patchy recruitment of both pocilloporid and acroporid spat. Interactions were found between space (on the scale of 10² m, i.e. sites within zones, and 10¹ m, i.e. racks within sites) and time (on the scale of years) for pocilloporids and between space (on the scale of 10³ m, i.e. zones, and 10² m) and time (on the scale of years) for acroporids. Post-recruitment mortality of acroporid spat in the period 3–10 months after their major spawning was dependent on their initial recruitment density, but pocilloporid mortality was either independent of initial recruitment density or, more likely, obscured by additional recruitment of pocilloporids to plates between late February and September. High rates of recruitment and growth by other sessile organisms, particularly bryozoans and oysters, appear to result in increased post-recruitment mortality and limit recruitment of scleractinian corals on settlement plates. The work reinforces an emerging picture that coral recruitment patterns are determined by mechanisms that manifest over a large range of spatial scales. Accepted: 1 September 1997     

Molecular evolution of a portion of the mitochondrial 16S ribosomal gene region in scleractinian corals

Relationships among families and suborders of scleractinian corals are poorly understood because of difficulties 1) in making inferences about the evolution of the morphological characters used in coral taxonomy and 2) in interpreting their 240-million-year fossil record. Here we describe patterns of molecular evolution in a segment of the mitochondrial (mt) 16S ribosomal gene from taxa of 14 families of corals and the use of this gene segment in a phylogenetic analysis of relationships within the order. We show that sequences obtained from scleractinians are homologous to other metazoan 16S ribosomal sequences and fall into two distinct clades defined by size of the amplified gene product. Comparisons of sequences from the two clades demonstrate that both sets of sequences are evolving under similar evolutionary constraints: they do not differ in nucleotide composition, numbers of transition and transversion substitutions, spatial patterns of substitutions, or in rates of divergence. The characteristics and patterns observed in these sequences as well as the secondary structures, are similar to those observed in mt 16S ribosomal DNA sequences from other taxa. Phylogenetic analysis of these sequences shows that they are useful for evaluating relationships within the order. The hypothesis generated from this analysis differs from traditional hypotheses for evolutionary relationships among the Scleractinia and suggests that a reevaluation of evolutionary affinities in the order is needed. Received: 4 September 1996 / Accepted: 7 April 1997     

Patterns of coral recruitment at the Gneering Shoals,southeast Queensland,Australia

Recruitment patterns of scleractinian corals were investigated at the Gneering Shoals, a coral-dominated rocky-reef south of the Great Barrier Reef, in subtropical Queensland. The density of recruits (mean of 0.8 to 6.3 recruits per tile (15 cm × 15 cm) pair from 4 sites) was the lowest ever recorded from six regions in tropical or subtropical eastern Australia that have been studied using directly comparable methods. Recruitment in summer was dominated by recruits from the Family Acroporidae, while corals from the Family Pocilloporidae recruited throughout the year. Recruits of massive corals andTurbinaria sp., which dominate the established coral communities, were absent. Possible explanations for the low recruitment rate in the region, include the depth of most sites (> 10m), competition for space with fouling organisms, and isolation, that is the failure of the south flowing East Australian Current to supply tropical larvae regularly from the Great Barrier Reef, 250 km to the north. The low coral recruitment rate at Gneering Shoals indicates that this region is unlikely to act as a stepping-stone for dispersal of tropical corals to more southern regions, which are more directly influenced by the East Australian Current.     

First report of folliculinid ciliates affecting Caribbean scleractinian corals

This is the first report of a ciliate of the genus Halofolliculina infecting hard coral species of six families (Acroporidae, Agaricidae, Astrocoeniidae, Faviidae, Meandrinidae and Poritidae) and milleporids in the Caribbean. Surveys conducted during 2004–2005 in Venezuela, Panama and México confirmed that this ciliate affects up to 25 scleractinian species. The prevalence of this ciliate at the coral community level was variable across sites, being most commonly found at Los Roques, Venezuela, and at Bocas del Toro, Panama (prevalence 0.2–2.5%), but rarely observed in the Mexican Caribbean. Ciliates were more prevalent within populations of acroporids (Acropora palmata, Acropora cervicornis and Acropora prolifera) in Los Roques. Recent observations also corroborate the presence of these ciliates in Curacao and Puerto Rico. Our observations indicate that ciliates affecting corals have a wider distribution than previously thought, and are no longer exclusively found in the Indo-Pacific and Red Sea.     

Contribution to the scleractinian corals of Hanifa Formation,Upper Jurassic,Jabal Al-Abakkayn,central Saudi Arabia

Eighteen scleractinian coral species belonging to 13 genera, 8 families and 4 suborders have been identified from the lower and upper parts of the Upper Jurassic (Oxfordian) Hanifa Formation at Jabal Al-Abakkayn, central Saudi Arabia. Actinastrea bernensis, A. crassoramosa, Coenastraea hyatti, Stylina kachensis, Cryptocoenia slovenica, C. wegeneri, Isastrea hemisphaerica, I. bernensis, Montlivaltia cornutiformis, M. frustriformis, Collignonastraea jumarensis, Ovalastrea michelini and Vallimeandropsis davidsoni are believed to be recorded for the first time from the Jurassic rocks of central Arabia. Most corals have massive hemispherical and globular forms, and few corals have dendroid and conical growth forms. They occur as small, isolated patches, about 0.5 m thick and about 10–30 m wide, in argillaceous reefal limestones. The identified corals show Africa, north America, northern, southern and western Europe, and southern or eastern Asia corals. The low diversity and abundance as well as the small size of colonies are attributed to inimical palaeoecological factors throughout the reefoids formation such as muddy substratum, water turbidity, high rate of sedimentation.     

Effect of naturally changing zooplankton concentrations on feeding rates of two coral species in the Eastern Pacific

Zooplankton concentrations are known to vary by as much as an order of magnitude over a lunar cycle. Here, we conducted an experiment to determine the effect of ambient zooplankton concentrations over a lunar cycle on feeding rates of the corals Pavona gigantea (Verrill) (mounding coral, 3.0 mm diameter polyps) and Pocillopora damicornis (Linnaeus) (branching coral, 1.0 mm diameter polyps) in situ on a shallow reef at Isla Contadora, Gulf of Panamá (Pacific), Panamá. Coral fragments exposed to either enhanced or ambient zooplankton concentrations were allowed to feed for 1 h, collected, and their gut contents dissected. The number of zooplankton captured was counted, feeding rates calculated per cm², and the species composition of captured zooplankton assemblages determined. Although both species captured the same zooplankton assemblage, feeding rates were always significantly higher for P. gigantea than for P. damicornis. Under ambient flow and zooplankton concentrations, feeding rates were highly correlated with zooplankton concentration in the 200-400 μm size class. Under constantly enhanced zooplankton concentrations in the control fragments, feeding rates did not vary significantly over the lunar cycle. As such, coral feeding rates vary not as a result of lunar phase per se, but with changes in zooplankton abundance over the lunar cycle. Coral feeding rates are directly proportional to ambient zooplankton concentrations and may vary by as much as 50% over a lunar cycle, suggesting that corals must cope with major swings in sources of fixed carbon and nutrients over relatively short timescales.     

Strategies for Amplification by Polymerase Chain Reaction of the Complete Sequence of the Gene Encoding Nuclear Large Subunit Ribosomal RNA in Corals

The nearly complete nuclear large subunit ribosomal RNA (LSU rRNA) gene in corals was amplified by primers designed from polymerase chain reaction (PCR) strategies. The motif of the putative 3′-terminus of the LSU rRNA gene was sequenced and identified from intergenic spacer (IGS) clones obtained by PCR using universal primers designed for corals. The 3′-end primer was constructed in tandem with the universal 5′-end primer for the LSU rRNA gene. PCR fragments of 3500 bp were amplified for octocorals and non-Acropora scleractinian corals. More than 80% of the Acropora LSU rRNA gene (3000 bp) was successfully amplified by modification of the 5′-end of the IGS primer. Analysis of the 5′-end of LSU rDNA sequences, including the D1 and D2 divergent domains, indicates that the evolutionary rate of the LSU rDNA differs among these taxonomic groups of corals. The genus Acropora showed the highest divergence pattern in the LSU rRNA gene, and the presence of a long branch of the Acropora clade from the other scleractinian corals in the phylogenetic tree indicates that the evolutionary rate of Acropora LSU rDNA might have accelerated after divergence from the common ancestor of scleractinian corals. Received February 17, 2000; accepted June 12, 2000.