The skeletal structure of Desmophyllum cristagalli: the use of deep-water corals in sclerochronology

Skeletal banding has been found in the deep-water scleractinian coral Desmophyllum cristagalli , an important animal in studies of climate change. This banding pattern sheds light on skeletogenesis and suggests methods by which the record of climate change contained within the coral skeletons may be interpreted. A central wall built of trabeculae forms the interior of the septa and rings the theca. Lamellae form a sheath over the trabecular frame, showing continuity from thecal edge to septum. Skeletal bands are added by the tissue layer, which overlaps and seals the internal coral and upper portion of the outer theca. Truncated inner bands on the outer theca indicate a pattern of skeletal deposition and dissolution dependent on the presence or absence of the live tissue layer. A long-term record will be difficult to collect from D. cristagalli since lamellae are less than 10 μm thick and band position is unpredictable. Density banding in shallow-water coral skeletons has long been recognized as a valuable paleo-oceanographic tool, and deep-water corals are now being used to reconstruct deep-ocean environments. Pattern of skeletal growth must be carefully considered if deep-water corals are to be used as proxy climate recorders.     

Late Paleozoic reef mounds of the Carnic Alps (Austria/Italy)

The Late Paleozoic (early Kasimovian-late Artinskian) sedimentary sequence of the Carnic Alps (Austria/Italy) is composed of cyclic, shallow-marine, mixed siliciclastic-carbonate sedimentary rocks. It contains different types of skeletal mounds in different stratigraphic levels. The oldest mounds occur at the base of the Auernig Group, within a transgressive sequence of the basal Meledis Formation. These mounds are small and built by auloporid corals. Algal mounds are developed in the Auernig Formation of the Auernig Group, forming biostromes, and Lower Pseudoschwagerina Limestone of the Rattendorf Group forming biostromes and bioherms. The dominant mound-forming organism of these mounds is the dasycladacean alga Anthracoporella spectabilis. In mounds of the Auernig Formation subordinately the ancestral corallinacean alga Archaeolithophyllum missouriense is present, whereas in mounds of the Lower Pseudoschwagerina Limestone a few calcisponges and phylloid algae occur locally at the base and on top of some Anthracoporella mounds. Mounds of the Auernig Formation formed during relative sea level highstands whereas mounds of the Lower Pseudoschwagerina Limestone formed during transgression. The depositional environment was in the shallow marine, low-turbulence photic zone, just below the active wave base and lacking siliciclastic influx. The algal mounds of the Carnic Alps differ significantly from all other algal mounds in composition, structure, zonation and diagenesis; the formation of the mounds cannot be explained by the model proposed by Wilson (1975). The largest mounds occur in the Trogkofel Limestone, they are composed of Tubiphytes/Archaeolithoporella boundstone, which shows some similarities to the “Tubiphytes thickets” of stage 2 of the massive Capitan reef complex of the Guadalupe Mountains of New Mexico/West Texas.     

Calcium localisation by X-ray microanalysis and fluorescence microscopy in larvae of zooxanthellate and azooxanthellate corals

X-ray microanalysis and fluorescence microscopy (Calcium Orangetrade mark) was used to determine the distribution of intracellular calcium (I(Ca)), in the form of total and ionic calcium respectively, in planulae and settled larvae of a zooxanthellate coral. The distribution of total calcium only was determined in larvae of an azooxanthellate coral. In azooxanthellate planulae and settled larvae, total I(Ca) concentration in the oral ectoderm was high and similar to that in seawater (SW). Calcium concentration did not vary (P > 0.05) between planulae and settled larvae. However, settled larvae accumulated large amounts of calcium in gastrodermal lipid-containing cells. In contrast, zooxanthellate planulae possessed significantly (P < 0.01) lower concentrations of total I(Ca) within ectodermal cells in comparison to settled larvae. In addition, in settled zooxanthellate larvae total calcium concentration in the mesogloea and coelenteron was significantly (P < 0.05) higher than in the oral ectodermal and gastrodermal cells, respectively. Total I(Ca) concentrations in the oral ectoderm of settled larvae were also significantly (P < 0.01) lower than that of the calicoblastic ectoderm. In zooxanthellate settled larvae, ionic I(Ca) levels in the aboral epithelium surrounding rapidly growing septa were high. These levels increased significantly (P < 0.05) within the tissue surrounding growing septa after incubation in high-calcium SW.     

Genotypic diversity and gene flow in brooding and spawning corals along the Great Barrier Reef, Australia

Marine organisms exhibit great variation in reproductive modes, larval types, and other life-history traits that may have major evolutionary consequences. We measured local and regional patterns of genetic variation in corals along Australia's Great Barrier Reef to determine the relative contributions of sexual and asexual reproduction to recruitment and to infer levels of gene flow both locally (among adjacent sites, < 5 km apart) and regionally (among reefs separated by 500-1,200 km). We selected five common brooding species (Acropora cuneata, A. palifera, Pocillopora damicornis, Seriatopora hystrix, and Stylophora pistillata) and four broadcast spawners (Acropora hyacinthus, A. cytherea, A. millepora, and A. valida), which encompassed a wide range of larval types and potential dispersal capabilities. We found substantial genotypic diversity at local scales in six of the nine species (four brooders, two spawners). For these six, each local population displayed approximately the levels of multilocus genotypic diversity (Go) expected for outcrossed sexual reproduction (mean values of Go:Ge ranged from 0.85 to 1.02), although consistent single-locus heterozygous deficits indicate that inbreeding occurs at the scale of whole reefs. The remaining three species, the brooder S. hystrix and the spawners A. valida and A. millepora displayed significantly less multilocus genotypic diversity (Go) than was expected for outcrossed sexual reproduction (Ge) within each of several sites. Acropora valida and A. millepora showed evidence of extensive localized asexual replication: (1) a small number of multilocus (clonal) genotypes were numerically dominant within some sites (Go:Ge values were as low as 0.17 and 0.20): (2) single-locus genotype frequencies were characterized by both excesses and deficits of heterozygotes (cf. Hardy-Weinberg expectations), and (3) significant linkage disequilibria occurred. For the brooding S. hystrix Go:Ge values were also low within each of four sites (x = 0.48). However, this result most likely reflects the highly restricted dispersal of gametes or larvae, because levels of genetic variation among sites within reefs were extremely high (FSR = 0.28). For all species, we detected considerable genetic subdivision among sites within each reef (high FSR-values), and we infer that larval dispersal is surprisingly limited (i.e., Nem among sites ranging from 0.6 to 3.3 migrants per generation), even in species that have relatively long planktonic durations. Nevertheless, our estimates of allelic variation among reefs (FRT) also imply that for all four broadcast spawning species and three of the brooders, larval dispersal is sufficient to maintain moderate to high levels of gene flow along the entire Great Barrier Reef (i.e., Nem among reefs ranged from 5 to 31). In contrast, widespread populations of S. hystrix and S. pistilata (the two remaining brooders) are relatively weakly connected (Nem among reefs was 1.4 and 2.5, respectively). We conclude that most recruitment by corals is very local, particularly in brooders, but that enough propagules are widely dispersed to ensure that both broadcast spawning and brooding species form vast effectively panmictic populations on the Great Barrier Reef.     

"Species" radiations of symbiotic dinoflagellates in the Atlantic and Indo-Pacific since the Miocene-Pliocene transition

Endosymbiotic dinoflagellates, or "zooxanthellae," are required for the survival of a diverse community of invertebrates that construct and dominate shallow, tropical coral reef ecosystems. Molecular systematics applied to this once understudied symbiont partner, Symbiodinium spp., divide the group into divergent lineages or subgeneric "clades." Within each clade, numerous closely related "types," or species, exhibit distinctive host taxon, geographic, and/or environmental distributions. This diversity is greatest in clade C, which dominates the Indo-Pacific host fauna and shares dominance in the Atlantic-Caribbean with clade B. Two "living" ancestors in this group, C1 and C3, are common to both the Indo-Pacific and Atlantic-Caribbean. With these exceptions, each ocean possesses a diverse clade C assemblage that appears to have independently evolved (adaptively radiated) through host specialization and allopatric differentiation. This phylogeographic evidence suggests that a worldwide selective sweep of C1/C3, or their progenitor, must have occurred before both oceans separated. The probable timing of this event corresponds with the major climactic changes and low CO(2) levels of the late Miocene and/or early Pliocene. Subsequent bursts of diversification have proceeded in each ocean since this transition. An ecoevolutionary expansion to numerous and taxonomically diverse hosts by a select host-generalist symbiont followed by the onset of rapid diversification suggests a radical process through which coral-algal symbioses respond and persist through the vicissitudes of planetary climate change.     

A biogeographic analysis and review of the far eastern Pacific coral reef region

New information on the presence and relative abundances of 41 reef-building (zooxanthellate) coral species at 11 eastern Pacific and 3 central Pacific localities is examined in a biogeographic analysis and review of the eastern Pacific coral reef region. The composition and origin of the coral fauna and other reef-associated taxa are assessed in the context of dispersal and vicariance hypotheses. A minimum variance cluster analysis using coral species presence–absence classification data at the 14 localities revealed three eastern Pacific reef-coral provinces: (1) equatorial– mainland Ecuador to Costa Rica, including the Galápagos and Cocos Islands; (2) northern– mainland México and the Revillagigedo Islands; (3) island group– eastern Pacific Malpelo Island and Clipperton Atoll, and central Pacific Hawaiian, Johnston and Fanning Islands. Coral species richness is relatively high in the equatorial (17–26 species per locality) and northern (18–24 species) provinces, and low at two small offshore island localities (7–10 species). A high proportion (36.6%, 15 species) of eastern Pacific coral species occurs at only one or two localities; of these, three disappeared following the 1982–83 ENSO event, three occur as death assemblages at several localities, and five are endangered with known populations of ten or fewer colonies. Principal component analysis using ordinal relative density data for the 41 species at the 14 localities indicated three main species groupings, i.e., those with high, mid, and narrow spatial distributions. These groupings correlated with species population-dynamic characteristics. These results were compared with data for riverine discharges, ocean circulation patterns, shoreline habitat characteristics, and regional sea surface temperature data to help clarify the analyses as these measures of environmental variability affect coral community composition. Local richness was highest at localities with the highest environmental variability. Recent information regarding the strong affinity between eastern and central Pacific coral faunas, abundance of teleplanic larvae in oceanic currents, high genetic similarity of numerous reef-associated species, and appearances of numerous Indo-west Pacific species in the east Pacific following ENSO activity, suggest the bridging of the east Pacific filter bridge (formerly east Pacific barrier). Accepted: 20 September 1999     

Framework of barrier reefs threatened by ocean acidification

To date, studies of ocean acidification (OA) on coral reefs have focused on organisms rather than communities, and the few community effects that have been addressed have focused on shallow back reef habitats. The effects of OA on outer barrier reefs, which are the most striking of coral reef habitats and are functionally and physically different from back reefs, are unknown. Using 5‐m long outdoor flumes to create treatment conditions, we constructed coral reef communities comprised of calcified algae, corals, and reef pavement that were assembled to match the community structure at 17 m depth on the outer barrier reef of Moorea, French Polynesia. Communities were maintained under ambient and 1200 μatm pCO₂ for 7 weeks, and net calcification rates were measured at different flow speeds. Community net calcification was significantly affected by OA, especially at night when net calcification was depressed ~78% compared to ambient pCO₂. Flow speed (2–14 cm s^?1) enhanced net calcification only at night under elevated pCO₂. Reef pavement also was affected by OA, with dissolution ~86% higher under elevated pCO₂ compared to ambient pCO₂. These results suggest that net accretion of outer barrier reef communities will decline under OA conditions predicted within the next 100 years, largely because of increased dissolution of reef pavement. Such extensive dissolution poses a threat to the carbonate foundation of barrier reef communities.     

Dinosaur Highways

A specimen of Calceola sandalina (Linne 1771) from the Givetian of the Czech Republic shows severe injury on its right side when observed from its cardinal side. Approximately one-half of the counter side is missing between the counter septum and the alar corallite angle. The injury is healed within the calice, as visible also on deformed septa close to injury, while the outer flat ventral side shows no signs of healing. The operculum is not preserved but the damage clearly must have affected it. It is difficult to envision how such damage might have occurred by abiotic means in a rather low-energy environment without the influx of grains larger than silt. We consider the injury a result of attack by a predator. When speculating on the animal capable of producing such injury, one must consider preferably a vertebrate with strong jaws: fish-like animals capable of durophagy (placoderms and chondrichthyans) appeared in the Devonian.     

The multidimensionality of the niche reveals functional diversity changes in benthic marine biotas across geological time

Despite growing attention on the influence of functional diversity changes on ecosystem functioning, a palaeoecological perspective on the long-term dynamic of functional diversity, including mass extinction crises, is still lacking. Here, using a novel multidimensional functional framework and comprehensive null-models, we compare the functional structure of Cambrian, Silurian and modern benthic marine biotas. We demonstrate that, after controlling for increases in taxonomic diversity, functional richness increased incrementally between each time interval with benthic taxa filling progressively more functional space, combined with a significant functional dissimilarity between periods. The modern benthic biota functionally overlaps with fossil biotas but some modern taxa, especially large predators, have new trait combinations that may allow more functions to be performed. From a methodological perspective, these results illustrate the benefits of using multidimensional instead of lower dimensional functional frameworks when studying changes in functional diversity over space and time.