More evidence for pervasive paraphyly in scleractinian corals: Systematic study of Southeast Asian Faviidae (Cnidaria; Scleractinia) based on molecular and morphological data

Coral taxonomy and systematics continue to be plagued by a host of problems. Due to high phenotypic variability within species, morphological approaches have often failed to recognize natural taxa, and molecular techniques have yet to be applied to many groups. Here, we summarize the levels of paraphyly found for scleractinian corals and test, based on new data, whether paraphyly is also a significant problem in Faviidae, the second-most speciose hermatypic scleractinian family. Using both DNA sequence and morphological data we find that, regardless of analysis technique (maximum parsimony, maximum likelihood and Bayesian likelihood), many conventional taxonomic groups are not monophyletic. Based on two mitochondrial markers (COI and a noncoding region) that we amplified for 81 samples representing 41 faviid species and 13 genera, five genera that are represented by more than one species are paraphyletic, as is the family Faviidae. The morphological characters currently used to identify these corals similarly fail to recover many genera. Furthermore, trees based on both data types are incongruent, and total evidence analysis does little to salvage conventional taxonomic groupings. Morphological convergence, phenotypic variability in response to the environment, and recent speciation are likely causes for these conflicts, which suggest that the present classification of corals is in need of a major overhaul. We propose more detailed studies of problematic faviid taxa using standardized morphological, mitochondrial, and nuclear genetic markers to facilitate combining of data.     

Exploring the role of Micronesian islands in the maintenance of coral genetic diversity in the Pacific Ocean

Understanding how genetic diversity is maintained across patchy marine environments remains a fundamental problem in marine biology. The Coral Triangle, located in the Indo‐West Pacific, is the centre of marine biodiversity and has been proposed as an important source of genetic diversity for remote Pacific reefs. Several studies highlight Micronesia, a scattering of hundreds of small islands situated within the North Equatorial Counter Current, as a potentially important migration corridor. To test this hypothesis, we characterized the population genetic structure of two ecologically important congeneric species of reef‐building corals across greater Micronesia, from Palau to the Marshall Islands. Genetic divergences between islands followed an isolation‐by‐distance pattern, with Acropora hyacinthus exhibiting greater genetic divergences than A. digitifera, suggesting different migration capabilities or different effective population sizes for these closely related species. We inferred dispersal distance using a biophysical larval transport model, which explained an additional 15–21% of the observed genetic variation compared to between‐island geographical distance alone. For both species, genetic divergence accumulates and genetic diversity diminishes with distance from the Coral Triangle, supporting the hypothesis that Micronesian islands act as important stepping stones connecting the central Pacific with the species‐rich Coral Triangle. However, for A. hyacinthus, the species with lower genetic connectivity, immigration from the subequatorial Pacific begins to play a larger role in shaping diversity than input from the Coral Triangle. This work highlights the enormous dispersal potential of broadcast‐spawning corals and identifies the biological and physical drivers that influence coral genetic diversity on a regional scale.     

Taxonomic status of Euphylliidae corals in Peninsular Malaysia based on morphological structures and phylogenetic analyses using mitochondrial COI gene

The morphological and molecular studies provide greater taxonomic resolution for the scleractinian coral identification. The Euphylliidae corals are among the scleractinian family for which their corallite and polyp morphologies have been examined for species identification. However, knowledge on the molecular study for coral identification in Malaysia is very limited. Therefore, this study aimed to examine the morphological structures and phylogenetic analyses for six Euphylliidae coral species using the mitochondrial gene of cytochrome oxidase subunit I (COI). The results showed that the Euphylliidae corals are present under both “complex” and “robust” coral clades as supported by many researchers. The result also revealed that the species phylogeny of Euphylliidae corals is in concordances with its morphological structures of corallites. It can be concluded the combination between morphological structures and phylogenetic analyses provide more accurate identification than relying on morphological study alone. Hence, it provides a future direction for the scleractinian research progress in species identification.     

Indo-Pacific echinoids in the tropical eastern Pacific

The existing literature reports that only one species of Indo-Pacific echinoid (Echinometra oblonga), occurs in the eastern Pacific. In this study we confirm the presence of this species at Islas Revillagigedo and also report the presence of two species ofEchinothrix (a genus hitherto unknown outside the Indo-Pacific) at Isla del Coco and at Clipperton Island. We also present evidence from isozymes and from mitochondrial DNA sequences indicating that at least one individual ofDiadema at Clipperton may belong to a maternal lineage characteristic of the west Pacific speciesD. savignyi. These data are consistent with the hypothesis that the observed populations of Indo-Pacific echinoid species are recent arrivals to the eastern Pacific, as opposed to the view that they are relicts of Tethyan pan-tropical distributions.Echinothrix diadema, in particular, may have arrived at Isla del Coco during the 1982-1983 El Nifio. In addition to Indo-Pacific species, Clipperton, Isla del Coco and the Revillagigedos contain a complement of eastern Pacific echinoids. The echinoid faunas of these islands should, therefore, be regarded as mixtures of two biogeographic provinces. Though none of the Indo-Pacific species are known to have reached the coast of the American mainland, their presence at the offshore islands of the eastern Pacific suggests that, for some echinoids, the East Pacific Barrier is not as formidable an obstacle to migration as was previously thought.     

Coral growth with thermal stress and ocean acidification: lessons from the eastern tropical Pacific

The rapid growth of scleractinian corals is responsible for the persistence of coral reefs through time. Coral growth rates have declined over the past 30 years in the western Pacific, Indian, and North Atlantic Oceans. The spatial scale of this decline has led researchers to suggest that a global phenomenon like ocean acidification may be responsible. A multi-species inventory of coral growth from Pacific Panamá confirms that declines have occurred in some, but not all species. Linear extension declined significantly in the most important reef builder of the eastern tropical Pacific, Pocillopora damicornis, by nearly one-third from 1974 to 2006. The rate of decline in skeletal extension for P. damicornis from Pacific Panamá (0.9% year⁻¹) was nearly identical to massive Porites in the Indo-Pacific over the past 20–30 years (0.89–1.23% year⁻¹). The branching pocilloporid corals have shown an increased tolerance to recurrent thermal stress events in Panamá, but appear to be susceptible to acidification. In contrast, the massive pavonid corals have shown less tolerance to thermal stress, but may be less sensitive to acidification. These differing sensitivities will be a fundamental determinant of eastern tropical Pacific coral reef community structure with accelerating climate change that has implications for the future of reef communities worldwide.     

A unique reproductive strategy in the mushroom coral Fungia fungites

Coral Reefs - The vast majority of scleractinian corals are either simultaneous hermaphrodites or gonochoric. Exceptions to these are rare. Nevertheless, species belonging to the family Fungiidae...     

Environmental drivers of distribution and reef development of the Mediterranean coral Cladocora caespitosa

Coral Reefs - Cladocora caespitosa is the only Mediterranean scleractinian similar to tropical reef-building corals. While this species is part of the recent fossil history of the Mediterranean...     

168 Cell Wall Proteomics of the Green Alga Haematococcus Pluvialis (Chlorophyceae)

Coral reefs of US-held islands in the central Pacific Ocean are among the most pristine in the world and represent over 93% of the reef systems under United States jurisdiction. The remote location of many islands has limited past algal research, resulting in incomplete understanding of species diversity, quantity, and ecology. Starting in 2000, the Coral Reef Ecosystem Investigation (CREI) began rapid ecological assessments on many Pacific island reefs to monitor ecological changes in reef biota over time. During the past year, algal efforts have concentrated on the French Frigate Shoals (Northwestern Hawaiian Islands) where we have increased the number of algal species reported by 1000%. Additionally, species new to science, including Acrosymphyton brainardii and Scinaia huismanii, have been described. Quantitative field sampling using a photoquadrat method is revealing species of the green algae Halimeda and Microdicyton to be ecological dominants in many areas during late summer/early autumn. Preliminary analyses with Primer software show species composition and abundance of all benthic organisms to differ significantly between most field sites sampled. Additional benthic habitat mapping of Pacific island reefs by CREI researchers is breaking the long-held paradigm that macroalgal cover is minimal in healthy tropical reef systems. Videotape analyses of benthic communities often find over 50% algal cover from 1 to 20 meter depths in many locations. Common ratios of macroalgae, turf algae, and crustose coralline algae to corals, other benthic organisms and substrate types on US Pacific reefs are being calculated for the first time.     

Disappearance of <Emphasis Type="Italic">Acropora</Emphasis> from the Marquesas (French Polynesia) during the last deglacial period

The major reef-building coral genus Acropora has never been recorded, living or fossil, from the Marquesas Islands in the central Pacific Ocean, which are characterized by limited modern reef formations. During the “Musorstom 9” cruise in 1997, investigations of marine platforms representing drowned reef systems revealed for the first time the presence of two Acropora species as fossils at seven Marquesas islands. The predominant species was Acropora valida, which was widespread in the archipelago and dated between 7.4 and 48.6 ka, providing evidence of an earlier Pacific distribution pattern broader than previously observed. It is proposed that disappearance of Acropora after 7.4 ka was linked to climatic events probably ENSO events controlling the distribution of corals and coral reefs in the eastern Pacific without excluding alternatively the effects of an increase in sea-level rise.     

Endemism in birds of tropical Pacific islands

Summary I analysed avifaunal data from 30 archipelagos and isolated islands in the tropical Pacific Ocean to examine the effects of geography on endemism. I divided the total bird species list (pelagic and migrant species excluded) for each island group into continental (also found outside of the study area), Pacific (found only within the study area but within more than one archipelago), and endemic (found only within a single archipelago) species and estimated ten variables related to the geography of each archipelago. I used multiple linear regression analysis to relate numbers and proportions of species in each category to the geographical variables. Total land area of an archipelago was the most improtant variable in explaining variation in the number of species in each category, with elevation and isolation also being important. The relationships between the proportions of species in each category and the geographical variables underscore the importance of isolation and the number of large islands in promoting endemism, presumably by allowing both inter- and intra-archipelagal speciation to proceed.     

Uniform phenotype conceals double colonization by reed-warblers of a remote Pacific archipelago

Aim Remote oceanic islands often provide good illustrations of adaptive radiations, but phylogenetic studies have also demonstrated unexpected multiple colonization events for a given archipelago. In this study we investigate the relationships among endemic populations of the Marquesas reed‐warbler, Acrocephalus mendanae Tristram, 1883, which have colonized nearly all islands of this remote Polynesian archipelago, and which exhibit a very uniform plumage pattern. We study the phylogeny and morphology of all subspecies in the Marquesas, providing an examination of the position of the Marquesas lineages in relation to reed‐warblers distributed across multiple Polynesian archipelagos. Location This study focused on all the main islands of the Marquesas archipelago, along with samples from other Polynesian archipelagos (Society, Tuamotu, Austral, Cook, Kiribati) and Australia. Methods We used mitochondrial DNA markers (cytochrome b and ND2 genes) to develop a phylogeny of the main eastern Polynesian taxa. All subspecies for the Marquesas were investigated, including multiple individuals per island. Phylogenetic analyses using maximum‐likelihood and Bayesian approaches were employed to infer relationships among A. mendanae populations and between the main Polynesian archipelagos. Morphometric analyses based on 110 specimens from museum collections were performed on external characters to investigate the differences between islands, and these results were compared to the phylogeny. Results Our data indicate that the Marquesas reed‐warbler is in fact a polyphyletic taxon including two independent lineages: the northern Marquesas reed‐warbler, closely related to the Tuamotu reed‐warbler, and the southern Marquesas reed‐warbler, sister taxon to that endemic to the Kiribati. Analyses of morphological characters show that the size and shape features of the Marquesas reed‐warblers exhibit high plasticity linked to adaptation to ecological factors, particularly habitat richness (the diversity of vegetation structure that provides suitable resources and habitat for reed‐warblers, simplified here as the number of indigenous plant species). Main conclusions Our results suggest that reed‐warblers have successfully colonized the Marquesas archipelago, one of the most remote groups of islands in the Pacific Ocean, at least twice. Both events occurred more or less simultaneously at ca. 0.6 Ma, and are more recent than the islands' formation. We outline the taxonomic consequences of our phylogeny and discuss the supertramp strategy of reed‐warblers in the Pacific.     

The genetic identity of dinoflagellate symbionts in Caribbean octocorals

Many cnidarians (e.g., corals, octocorals, sea anemones) maintain a symbiosis with dinoflagellates (zooxanthellae). Zooxanthellae are grouped into clades, with studies focusing on scleractinian corals. We characterized zooxanthellae in 35 species of Caribbean octocorals. Most Caribbean octocoral species (88.6%) hosted clade B zooxanthellae, 8.6% hosted clade C, and one species (2.9%) hosted clades B and C. Erythropodium caribaeorum harbored clade C and a unique RFLP pattern, which, when sequenced, fell within clade C. Five octocoral species displayed no zooxanthella cladal variation with depth. Nine of the ten octocoral species sampled throughout the Caribbean exhibited no regional zooxanthella cladal differences. The exception, Briareum asbestinum, had some colonies from the Dry Tortugas exhibiting the E. caribaeorum RFLP pattern while elsewhere hosting clade B. In the Caribbean, octocorals show more symbiont specificity at the cladal level than scleractinian corals. Both octocorals and scleractinian corals, however, exhibited taxonomic affinity between zooxanthella clade and host suborder.Communicated by R.C. Carpenter     

Cheilostomatous Bryozoa from Vanuatu

The present account of some Bryozoa from Vanuatu is the first study of reef-flat species from the eastern Coral Sea. Based on samples collected from several shallow water localities on the island of Efate, a total of 92 species is described, including a new family, three new genera and 20 new species. Some of the newly recognized species are the result of taxonomic revision stimulated by these new samples, while others are entirely new to science. Of the new taxa described here, 16 are presently known only from Vanuatu. The total number of species recorded does not accurately represent the true taxonomic diversity of the Cheilostomatida of Vanuatu, and further sampling, at other localities and in other habitats, is needed before it can be reasonably estimated. The fauna is predominantly Indo-West Pacific in character with a high proportion of species apparently known only in the Southwest Pacific.     

Mesophotic coral ecosystems in the Hawaiian Archipelago

Efforts to map coral reef ecosystems in the Hawaiian Archipelago using optical imagery have revealed the presence of numerous scleractinian, zoothanthellate coral reefs at depths of 30–130+ m, most of which were previously undiscovered. Such coral reefs and their associated communities have been recently defined as mesophotic coral ecosystems (MCEs). Several types of MCEs are found in Hawai‘i, each of which dominates a different depth range and is characterized by a unique pattern of coral community structure and colony morphology. Although MCEs are documented near both ends of the archipelago and on many of the islands in between, the maximum depth and prevalence of MCEs in Hawai‘i were found to decline with increasing latitude. The Main Hawaiian Islands (MHI) had significantly deeper and greater percentages of scleractinian coral, and peaks in cover of both scleractinian corals and macroalgae occurred within depth bins 20 m deeper than in the Northwestern Hawaiian Islands (NWHI). Across the archipelago, as depth increased the combined percentage of living cover of mega benthic taxa declined sharply with increasing depth below 70 m, despite the widespread availability of hard substrate.     

Regional biogeography of shallow reef fish and macro-invertebrate communities in the Galapagos archipelago

Aim To delineate biogeographical patterns in Galapagos shallow‐water reef fauna at regional scales. Location Galapagos Islands. Methods Fishes and macro‐invertebrates were quantitatively censused using underwater visual techniques along more than 500 transects at defined depth strata across the Galapagos archipelago. Data were analysed using multivariate techniques to define regional patterns and identify species typical of different regions. Results Subtidal communities of fishes and macro‐invertebrates on shallow reefs differed consistently in species composition across the Galapagos archipelago, with three major biogeographical groupings: (1) the ‘far‐northern area’ containing the islands of Darwin and Wolf, (2) the ‘central/south‐eastern area’, including the east coast of Isabela, and (3) the ‘western area’, encompassing Fernandina and western Isabela. In addition, the northern islands of Pinta, Marchena and Genovesa form a separate region in the central/south‐eastern area, and Bahia Elizabeth and Canal Bolivar separate from other parts of the western area. The far‐northern bioregion is characterized by high fish species richness overall, including a high proportion of species of Indo‐Pacific origin. However, very few endemic fishes or species with distributions extending south from Ecuador (‘Peruvian’ species) are present, and the bioregion also possesses relatively low species richness of mobile macro‐invertebrate taxa. By contrast, the ‘western’ bioregion possesses disproportionately high numbers of endemic fish taxa, high numbers of cool‐temperate Peruvian fish species, and high invertebrate species richness, but very few species of Indo‐Pacific origin. The Bahia Elizabeth/Canal Bolivar bioregion possesses more endemic species and fewer species with Peruvian affinities than coasts within the western bioregion. The northern bioregion of Pinta, Marchena and Genovesa represents an overlap zone with affinities to both the far‐northern and south‐eastern islands. The south‐eastern bioregion includes species from a variety of different sources, particularly ‘Panamic’ species with distributions extending north to Central America. Main conclusions On the basis of congruent divisions for reef fish and macro‐invertebrate communities, the Galapagos archipelago can be separated into three major biogeographical areas, two of which can be further subdivided into two regions. Each of these five bioregions possesses communities characterized by a distinctive mix of species derived from Indo‐Pacific, Panamic, Peruvian and endemic source areas. The conservation significance of different regions is not reflected in counts of total species richness. The regions with the lowest overall fish species richness possess a temperate rather than tropical climate and highest levels of endemism.     

Rethinking the phylogeny of scleractinian corals: a review of morphological and molecular data

Scleractinian corals, which include the architects of coral reefs, are found throughout the world's oceans and have left a rich fossil record over their 240 million year history. Their classification has been marked by confusion but recently developed molecular and morphological tools are now leading to a better understanding of the evolutionary history of this important group. Although morphological characters have been the basis of traditional classification in the group, they are relatively few in number. In addition, our current understanding of skeletal growth and homology is limited, and homoplasy is rampant, limiting the usefulness of morphological phylogenetics. Molecular phylogenetic hypotheses for the order, which have been primarily focused on reef-building corals, differ significantly from traditional classification. They suggest that the group is represented by two major lineages and do not support the monophyly of traditional suborders and most traditional families. It appears that once a substantial number of azooxanthellate taxa are included in molecular phylogenetic analyses, basal relationships within the group will be clearly defined. Understanding of relationships at lower taxonomic levels will be best clarified by combined analyses of morphological and molecular characters. Molecular phylogenies are being used to inform our understanding of the evolution of morphological characters in the Scleractinia. Better understanding of the evolution of these characters will help to integrate the systematics of fossil and extant taxa. We demonstrate how the combined use of morphological and molecular tools holds great promise for ending confusion in scleractinian systematics.     

Free-living scleractinian corals on reefs of the Seychelles Islands

Thirty-nine species of unattached scleractinian corals that belong to 22 genera of 9 families were found on the Seychelles reefs. Variations of the colony form of corals living on soft sediments under continuous wave action are described. Irrespective of their initial growth form and taxonomic position, corals assume a form close to spherical. Because of the worldwide deterioration of coral reefs, the adaptation to changing ecological conditions by reef-building corals needs to be studied.     

Biogeography of mammals on tropical Pacific islands

Aim We examine the influence of geography on species richness and endemism of mammals on tropical Pacific archipelagos to determine the importance of intra‐ and inter‐archipelago speciation in promoting local and regional species richness. Location Thirty tropical Pacific archipelagos. Methods A distributional list of mammals on 30 archipelagos was compiled, and values for 10 geographical variables were estimated for each archipelago. Mammal species were placed in three different categories (continental, Pacific and endemic) based on their distribution. The total number of species and numbers of species within each category were related to the geographical variables using Poisson regression analysis. Results Species richness was related positively to variables describing land area, numbers of large islands and elevation; and negatively to variables describing isolation. Levels of endemism did not differ between volant and non‐volant species, but differed between mega‐ and microchiropterans. Main conclusions Variation in species richness of mammals in the tropical Pacific region can be accounted for by a combination of intra‐archipelago speciation within archipelagos composed of large islands, and inter‐archipelago speciation, particularly among more isolated archipelagos. Mammals were less widely distributed throughout the study area than previously found for butterflies, skinks or birds. However, the level of endemism was similar to that of skinks and birds on the same archipelagos, and was higher than that of butterflies.     

Variation in Symbiodinium ITS2 sequence assemblages among coral colonies

Endosymbiotic dinoflagellates in the genus Symbiodinium are fundamentally important to the biology of scleractinian corals, as well as to a variety of other marine organisms. The genus Symbiodinium is genetically and functionally diverse and the taxonomic nature of the union between Symbiodinium and corals is implicated as a key trait determining the environmental tolerance of the symbiosis. Surprisingly, the question of how Symbiodinium diversity partitions within a species across spatial scales of meters to kilometers has received little attention, but is important to understanding the intrinsic biological scope of a given coral population and adaptations to the local environment. Here we address this gap by describing the Symbiodinium ITS2 sequence assemblages recovered from colonies of the reef building coral Montipora capitata sampled across Kāne'ohe Bay, Hawai'i. A total of 52 corals were sampled in a nested design of Coral Colony(Site(Region)) reflecting spatial scales of meters to kilometers. A diversity of Symbiodinium ITS2 sequences was recovered with the majority of variance partitioning at the level of the Coral Colony. To confirm this result, the Symbiodinium ITS2 sequence diversity in six M. capitata colonies were analyzed in much greater depth with 35 to 55 clones per colony. The ITS2 sequences and quantitative composition recovered from these colonies varied significantly, indicating that each coral hosted a different assemblage of Symbiodinium. The diversity of Symbiodinium ITS2 sequence assemblages retrieved from individual colonies of M. capitata here highlights the problems inherent in interpreting multi-copy and intra-genomically variable molecular markers, and serves as a context for discussing the utility and biological relevance of assigning species names based on Symbiodinium ITS2 genotyping.     

167 Algal Monitoring Studies on Remote Tropical Pacific Reefs

Coral reefs of US‐held islands in the central Pacific Ocean are among the most pristine in the world and represent over 93% of the reef systems under United States jurisdiction. The remote location of many islands has limited past algal research, resulting in incomplete understanding of species diversity, quantity, and ecology. Starting in 2000, the Coral Reef Ecosystem Investigation (CREI) began rapid ecological assessments on many Pacific island reefs to monitor ecological changes in reef biota over time. During the past year, algal efforts have concentrated on the French Frigate Shoals (Northwestern Hawaiian Islands) where we have increased the number of algal species reported by 1000%. Additionally, species new to science, including Acrosymphyton brainardii and Scinaia huismanii, have been described. Quantitative field sampling using a photoquadrat method is revealing species of the green algae Halimeda and Microdicyton to be ecological dominants in many areas during late summer/early autumn. Preliminary analyses with Primer software show species composition and abundance of all benthic organisms to differ significantly between most field sites sampled. Additional benthic habitat mapping of Pacific island reefs by CREI researchers is breaking the long‐held paradigm that macroalgal cover is minimal in healthy tropical reef systems. Videotape analyses of benthic communities often find over 50% algal cover from 1 to 20 meter depths in many locations. Common ratios of macroalgae, turf algae, and crustose coralline algae to corals, other benthic organisms and substrate types on US Pacific reefs are being calculated for the first time.