Revision of the coral genus Acropora (Scleractinia: Astrocoeniina: Acroporidae) in Indonesia

The coral genus Acropora is reviewed from Indonesia for the first time, following detailed collections made at 131 sites and additional material collected from approximately 40 sites throughout the archipelago during the period 1993–6. Eighty-three species are recorded, four of these ( Acropora halmaherae, A. awi, A. plumosa and A. simplex ) new to science, six first described in 1994 and six in 1997. Records are compared with specimen-based records from localities worldwide. The species of Acrokora occurring in Indonesian waters include five recorded only from the Indian Ocean and Indonesia, seven recorded only from the Pacific Ocean, South China Sea and Indonesia, and a further 10 species apparently endemic to Indonesia, as well as widespread Indo-Pacific species. Two species ( A. jacquelineae Wallace, 1994 and A. batunai Wallace, 1996) are recorded only from north central Indonesia and Papua New Guinea, and two species ( A. russelli Wallace, 1994 and A. turaki Wallace, 1994) only from north central Indonesia and north western Australia. The findings contribute to a new view of the corals of the Indo-Pacific 'centre of diversity' as a composite fauna with origins in a number of events in space and time.     

A revision of the hard coral genus Acropora Oken, 1815 (Scleractinia: Astrocoeniina: Acroporidae) in south-east Africa

All species of the scleractinian coral genus Acropora presently known to occur in south-east Africa (denoting the African coast south of the Tropic of Capricorn, and including the atoll Bassas da India in the Mozambique channel) are reviewed. Twenty-three species are discussed, most of which are of wide Indo-Pacific distribution. Field and laboratory characteristics of all species are described in detail. Species richness in southern Mozambique is much higher (23 species) than in South Africa (14 species), probably due to the higher ecological differentiation of Mozambiquan reefs. The Acropora fauna of the atoll Bassas da India is similar to that of Mozambique but with two additional species A. paniculata and A. cf. striata. Compared to other coral reef areas in similar latitudes, the south-east African Acropora fauna is of average diversity. An identification key to all species is provided.     

Necrotic patches affect Acropora palmata (Scleractinia: Acroporidae) in the Mexican Caribbean.

An outbreak of necrotic patches was observed affecting Acropora palmata in the Mexican Caribbean in the summer of 1999. This study documents the tissue loss produced by these patches. Following a marked initial increase in the number of patches, there was a decrease in the appearance of new patches but the size of the patches increased throughout the study. In some cases patches expanded but in most cases they enlarged due to fusion of 2 or more patches. Patches recovered but not sufficiently to overcome damage in most colonies surveyed. Percentage tissue loss does not appear to be directly related to temperature but may be related to a combination of factors associated with prolonged summer doldrum-like conditions. The necrotic patch syndrome can have a substantial impact in tissue loss in affected A. palmata colonies.     

Recognition of separate genera within Acropora based on new morphological, reproductive and genetic evidence from Acropora togianensis , and elevation of the subgenus Isopora Studer, 1878 to genus (Scleractinia: Astrocoeniidae; Acroporidae)

Many attempts have been made to recognise divisions within Acropora, the most diverse reef building coral genus on modern reefs, but only subgenera Acropora and Isopora are currently recognised. In this paper, morphological and genetic analyses, and study of reproductive mode and anatomy, demonstrate that an endemic Indonesian species A. (Acropora) togianensis, Wallace, 1997, belongs to Isopora. Despite the presence of a clear central axial corallite (indicating sub-genus Acropora), this species has supplementary axial corallites, broods planula larvae rather than broadcast-spawning for external fertilisation and develops stalked ova: all characters in common with the type species of subgenus Isopora A. (Isopora) palifera and the other species for which such data are available, A. (I.) cuneata and A. (I.) brueggemanni. Phylogenies are based on the protein-coding genes, mitochondrial cytochrome b (cytb) and nuclear histone 2a and 2b (h2ab) also group A. togianensis with these Isoporans. High bootstrapping and Bayesian support in the major lineages of the family Acroporidae demonstrate significant differences between Isopora (including A. togianensis) and Acropora. As the type species of both subgenera, A. (Acropora) muricata (Linneaus 1758) and A. (Isopora) palifera (Lamarck, 1816) are used in these analyses, elevation of Isopora Studer, 1878 to genus is formally proposed.     

New species and new records of recently described species of the coral genus Acropora (Scleractinia: Astrocoeniina: Acroporidae) from Indonesia

Six new species of the coral genus Acropora arc described from Indonesia. These include a species which is remarkable for tubercular cocnostcal structures similar to those of the confamilial genus Montipora. The new species include three regional endemics (A. togianensis and A. batunai from central east Sulawesi and A. derawanensis from east Kalimantan), one species with broad distribution across the southern island chains (A. sukarnoi) and two species which occur throughout most of the Indonesian archipelago (A. Indonesia and A. hoeksemai). A further two species described from Western Australia and Papua New Guinea in 1994 (A. turaki and A. jacquelineae respectively) are recorded from Indonesia for the first time, as common members of an unusual assemblage type in the Togian Islands. The range of another species described from Lombok in 1994 (A. suharsonoi) is extended into Bali. With A. desalwii, A. lokani and A. indiana , this brings to 12 the number of Acropora species newly recorded as being endemic to the Indonesian archipelago or to Indonesia and one adjoining region (either the Indian Ocean or the western Pacific).     

Description of four new species in the hard coral genus Acropora Oken, 1815 (Scleractinia: Astrocoeniina: Acroporidae) from south-east Africa

Four new species in the hermatypic scleractinian coral genus Acropora art described from southeast Africa. Two species, A. natalensis sp. nov. and A. mossambica sp. nov. , occur in both South Africa and Mozambique. Two species, A. sordiensis sp. nov. , and A. branchi sp. nov. , are only known from northern Natal in South Africa. A. branchi sp. nov. and A. mossambica sp. nov. are intertidal species and were collected from permanent tidal pools. A. sordiensis sp. nov. and A. natalensis sp. nov. are strictly subtidal species.     

Taxonomic classification of the reef coral family Mussidae (Cnidaria: Anthozoa: Scleractinia)

Molecular analyses are transforming our understanding of the evolution of scleractinian corals and conflict with traditional classification, which is based on skeletal morphology. A new classification system, which integrates molecular and morphological data, is essential for documenting patterns of biodiversity and establishing priorities for marine conservation, as well as providing the morphological characters needed for linking present‐day corals with fossil species. The present monograph is the first in a series whose goal is to develop such an integrated system. It addresses the taxonomic relationships of 55 Recent zooxanthellate genera (one new) in seven families (one new), which were previously assigned to the suborder Faviina (eight genera are transferred to incertae sedis). The present monograph has two objectives. First, we introduce the higher‐level classification system for the 46 genera whose relationships are clear. Second, we formally revise the taxonomy of those corals belonging to the newly discovered family‐level clade (restricted today to the western Atlantic and Caribbean regions); this revised family Mussidae consists of ten genera (one of which is new) and 26 species that were previously assigned to the ‘traditional’ families Faviidae and Mussidae. To guide in discovering morphologic characters diagnostic of higher‐level taxa, we mapped a total of 38 morphologic characters [19 macromorphology, eight micromorphology, 11 microstructure] onto a molecular tree consisting of 67 species [22 Indo‐Pacific and seven Atlantic species in the traditional family Faviidae; 13 Indo‐Pacific and ten Atlantic species in the traditional family Mussidae; 13 species in the traditional families Merulinidae (5), Pectiniidae (7), and Trachyphylliidae (1); two Atlantic species of traditional Montastraea], and trace character histories using parsimony. To evaluate the overall effectiveness of morphological data in phylogeny reconstruction, we performed morphology‐based phylogenetic analyses using 27 (80 states) of the 38 characters, and compared morphological trees with molecular trees. The results of the ancestral state reconstructions revealed extensive homoplasy in almost all morphological characters. Family‐ and subfamily‐level molecular clades [previously identified as XVII?XXI] are best distinguished on the basis of the shapes of septal teeth and corresponding microstructure. The newly revised family Mussidae (XXI) has septal teeth with regular pointed tips (a symplesiomorphy) and a stout blocky appearance. It has two subfamilies, Mussinae and Faviinae. The subfamily Mussinae is distinguished by spine‐shaped teeth and widely spaced costoseptal clusters of calcification centres. The subfamily Faviinae is distinguished by blocky, pointed tricorne or paddle‐shaped teeth with elliptical bases, transverse structures such as carinae that cross the septal plane, and well‐developed aligned granules. Defining diagnostic characters for the broader data set is more challenging. In analyses of taxonomic subsets of the data set that were defined by clade, morphological phylogenetic analyses clearly distinguished the families Mussidae (XXI) and Lobophylliidae (XIX), as well as the two subfamilies of Mussidae (Mussinae, Faviinae), with one exception (Homophyllia australis). However, analyses of the entire 67‐species data set distinguished the family Lobophylliidae (XIX), but not the Merulinidae (XVII) and not the newly defined Mussidae (XXI), although the subfamily Mussinae was recovered as monophyletic. Some lower‐level relationships within the Merulinidae (XVII) agree with molecular results, but this particular family is especially problematic and requires additional molecular and morphological study. Future work including fossils will not only allow estimation of divergence times but also facilitate examination of the relationship between these divergences and changes in the environment and biogeography. Published 2012. This article is a U.S. Government work and is in the public domain in the USA. Zoological Journal of the Linnean Society, 2012, 166 , 465–529.     

Taxonomic classification of the reef coral family Lobophylliidae (Cnidaria: Anthozoa: Scleractinia)

Lobophylliidae is a family‐level clade of corals within the ‘robust’ lineage of Scleractinia. It comprises species traditionally classified as Indo‐Pacific ‘mussids’, ‘faviids’, and ‘pectiniids’. Following detailed revisions of the closely related families Merulinidae, Mussidae, Montastraeidae, and Diploastraeidae, this monograph focuses on the taxonomy of Lobophylliidae. Specifically, we studied 44 of a total of 54 living lobophylliid species from all 11 genera based on an integrative analysis of colony, corallite, and subcorallite morphology with molecular sequence data. By examining coral skeletal features at three distinct levels – macromorphology, micromorphology, and microstructure – we built a morphological matrix comprising 46 characters. Data were analysed via maximum parsimony and transformed onto a robust molecular phylogeny inferred using two nuclear (histone H3 and internal transcribed spacers) and one mitochondrial (cytochrome c oxidase subunit I) DNA loci. The results suggest that micromorphological characters exhibit the lowest level of homoplasy within Lobophylliidae. Molecular and morphological trees show that Symphyllia, Parascolymia, and Australomussa should be considered junior synonyms of Lobophyllia, whereas Lobophyllia pachysepta needs to be transferred to Acanthastrea. Our analyses also lend strong support to recent revisions of Acanthastrea, which has been reorganized into five separate genera (Lobophyllia, Acanthastrea, Homophyllia, Sclerophyllia, and Micromussa), and to the establishment of Australophyllia. Cynarina and the monotypic Moseleya remain unchanged, and there are insufficient data to redefine Oxypora, Echinophyllia, and Echinomorpha. Finally, all lobophylliid genera are diagnosed under the phylogenetic classification system proposed here, which will facilitate the placement of extinct taxa on the scleractinian tree of life.     

Sexual reproduction and early development of the solitary coral Fungia scutaria (Anthozoa: Scleractinia)

Fungia scutaria spawned vigorously with a lunar beriodicity during the summer months of 1981 and 1982. Spawning activity declined in the fall of both years and was absent in winter and spring (1983). There was only one short spawning event per lunar cycle. Each event occurred in the evening between 1700 and 1900 hours 1 to 4 days following the full moon. Fungia scutaria exhibits gonochorism. Females ejected eggs through their mouths into the seawater above. Many of these negatively buoyant eggs settled onto the oral discs and were moved off the edge by ciliary-mucoid activity. Spermatoza from males were similarly expelled in a jet of gastrovascular fluid. Spawned eggs were small and lacked endosymbiotic zooxanthellae. Rapid development led to ciliated solid planulae by the next morning. Within 24 h a mouth had begun to develop. Planulae may have been able to feed within 39 h. Infection with zooxanthellae occurred 4–5 days following spawning. Planulae may have become competent for settlement by 7 days, but attempts to docment settlement produced ambiguous results.     

Assessment of current rates of Diadema antillarum larval settlement

The generally slow and incomplete recovery of the long-spined sea urchin, Diadema antillarum, from the 1983–84 Caribbean-wide die-off, particularly in the Florida Keys, USA, raises the question of factors limiting population recovery. This study sought to quantify larval settlement rates as an indicator of larval supply at two sites in the Florida Keys, utilizing methods comparable to an historic study. Settlement at two sites in southwest Puerto Rico was also examined as a comparison of present-day settlement rates at a site where D. antillarum recovery has been moderate. Monthly settlement rates were low (max < 2 m⁻²) and did not differ between the two sites examined in the Florida Keys. Settlement was significantly higher at only one of the Puerto Rico sites (max 16 m⁻²), but still an order of magnitude lower than that reported for historic populations in Cura?ao (1982–83). Results are consistent with the hypothesis of low larval supply limiting D. antillarum recovery in the Florida Keys.     

Hiding in plain sight: invasive coral Tubastraea tagusensis (Scleractinia:Hexacorallia) in the Gulf of Mexico

Coral Reefs - Our research presents the first record of Tubastraea tagusensis (Wells, Notes on Indo-Pacific scleractinian corals. Part 9. New corals from the Galápagos Islands, 1982) in the...     

Metabolic tolerance of the cold-water coral Lophelia pertusa (Scleractinia) to temperature and dissolved oxygen change

Lophelia pertusa is the world's most common and widespread framework-forming cold-water coral. It forms deep-water coral reefs and carbonate mounds supporting diverse animal communities on the continental shelf and on seamounts. These recently discovered ecosystems have been damaged by deep-sea fishing and are threatened by predicted shallowing of the aragonite saturation horizon. Despite this, very little is known about the ecophysiology of L. pertusa and its likely response to environmental changes. Here we describe the first study of the respiratory physiology of L. pertusa and the effects of altered temperature and oxygen level. This study shows that L. pertusa can maintain respiratory independence over a range of PO₂ illustrated by a high regulation value (R = 78%). The critical PO₂ value of 9-10 kPa is very similar to the lower values of oxygen concentration recorded in the field. This suggests that oxygen level may be a limiting factor in the distribution of this cold-water coral. L. pertusa survived periods of anoxia (1 h), hypoxia (up to 96 h), but high Q₁₀ values revealed sensitivity to short-term temperature changes (6.5-11 °C). For the first time vital data have been gathered on the physiology of this species that is essential to understand distribution and underpin future climate change studies.     

Elevated temperature reduces survivorship and settlement of the larvae of the Caribbean scleractinian coral, Favia?fragum (Esper)

The effect of elevated seawater temperatures, such as those plaguing tropical seas during the summers of anomalously warm years, on early life stages of reef corals remains poorly studied. To redress this situation, survivorship of larvae of the brooding coral, Favia fragum, was studied in the laboratory, using both short term (48 h) and long term (156–191 h) exposures to 28, 29, and 31°C. Ability to settle when presented with induction substrates and survival after settlement, at the same exposure temperature and after reciprocal transfers to the other experimental temperatures, were also measured. No significant effect of temperature on survivorship was detected after 48 h of exposure, but larvae incubated for 156 h at the highest temperature (31°C) exhibited a 13% reduced survivorship compared to larvae at 28°C. Induction of settlement further increased mortality at the highest temperature (31°C); survivorship after settlement at 31°C was 27% lower than when larvae were simply maintained at the elevated temperature. These results indicate that elevated temperatures are more detrimental to coral larvae undergoing the developmentally complex settlement process than to the swimming planula stage. This may bode poorly for Caribbean corals with late summer reproductive seasons. Communicated by Ecology Editor Prof. Peter Mumby     

Diel vertical migration thresholds of Karenia brevis (Dinophyceae)

Light and nutrient availability change throughout dinoflagellate diel vertical migration (DVM) and/or with sub-population location in the water column along the west Florida shelf. Typically, the vertical depth of the shelf is greater than the distance a sub-population can vertically migrate during a diel cycle, limiting the ability of a sub-population to photosynthetically fix carbon toward the surface and access nutrients sub-surface. This project investigated changes of Karenia brevis (C.C. Davis) G. Hansen et Moestrup intracellular carbon, nitrogen, internal nitrate (iNO₃), free amino acid (FAA), and total lipid concentrations in high-light, nitrate-replete (960 μmol quanta m⁻² s⁻¹, 80 μM NO₃), and high-light, nitrate-reduced (960 μmol quanta m⁻² s⁻¹, <5 μM NO₃) mesocosms. The nitrate-reduced mesocosm had a slowed cell division rate when compared to the nitrate-replete mesocosm. Minimum intracellular carbon, nitrogen, iNO₃, FAA, and total lipid concentrations during the largest surface sub-population aggregations led to the conclusion that daughter cells resulting from cell division received unequal shares of the parental resources and that this inequality influenced migration behavior. Nutrient reduced daughter cells were more strongly influenced by light and phototaxis for carbon production than their replete same cell division sister cells during vertical migration thus rapidly increasing the fulfillment of constituents through photosynthesis. Vertical migration was consistent with an optimization scheme based on threshold limits through utilization or formation of photosynthate. We propose a simplified conceptual model describing how K. brevis is transported along the benthos of the west Florida shelf from off-shore to on-shore. Dynamic carbon thresholds are also suggested for future DVM modeling efforts on K. brevis populations transported between nitrogen replete and nitrogen reduced environmental conditions.     

Taxonomy and species boundaries in the coral genus Favia Milne Edwards and Haime, 1857 (Cnidaria: Scleractinia) from Thailand revealed by morphological and genetic data

While Faviidae is a widely and uniformly distributed coral family throughout the Indo-Pacific, the extensive phenotypic plasticity of colony surface and corallite features often confounds the use of macromorphological characters in species identification, and contributes to conflict between traditional classification and molecular analyses of the group. Recent advances in morphological and molecular techniques now provide a suite of methods to re-address coral taxonomy in complex groups, such as that represented by the Faviidae. This study combines morphologic measurements including “3D coordinates landmarks” data with phylogenetic assessments of nuclear (ITS) and mitochondrial (COI-trnM) DNA to assess species boundaries in nine species of Faviidae with para-septothecal walls from Thailand. Strong concordance was found between morphological features and a priori groupings based on both morphospecies and genetically defined groups (ITS and COI-trnM). Favia truncatus was the most well-defined species based on morphological analyses, and it was also shown to be monophyletic using phylogenetic analyses. Besides F. truncatus, the only other species that was found to be monophyletic in analyses of both genes was F. cf. helianthoides, but its skeletal morphology overlapped with the F. favus species complex (comprised of F. favus, F. speciosa, F. matthaii and F. rotumana). Although not genetically monophyletic, the F. favus species complex and F. pallida were fairly well delineated morphologically. Morphospecies within the F. favus species complex are therefore possibly a result of genetic drift and/or stable polymorphisms driven by divergent selection. These results represent a first step toward a taxonomic revision of the Indo-Pacific Favia, which will integrate morphological methods with the study of type material, genetic information, reproductive data, and tests of phenotypic plasticity—given that multiple lines of evidence are needed to resolve ambiguous species and assign species names.     

Taxonomic classification of the reef coral families Merulinidae,Montastraeidae, and Diploastraeidae (Cnidaria: Anthozoa: Scleractinia)

Modern coral taxonomy has begun to resolve many long‐standing problems in traditional systematics stemming from its reliance on skeletal macromorphology. By integrating examinations of colony, corallite, and subcorallite morphology with the molecular sequence data that have proliferated in the last decade, many taxa spread across the scleractinian tree of life have been incorporated into a rigorous classification underpinned by greater phylogenetic understanding. This monograph focuses on one of the most challenging clades recovered to date – its disarray epitomized by the informal name ‘Bigmessidae’. This group of predominantly Indo‐Pacific species previously comprised families Merulinidae, Faviidae, Pectiniidae, and Trachyphylliidae, but in a recent study these have been incorporated within Merulinidae. We studied 84 living merulinid species by examining morphological traits at three different scales of coral skeletal structure ? macromorphology, micromorphology, and microstructure ? to construct a morphological matrix comprising 44 characters. Data were analysed via maximum parsimony and also transformed onto a robust molecular phylogeny under the parsimony and maximum likelihood criteria. Comparisons amongst morphological character types suggest that although many characters at every scale are homoplastic, some to a greater extent than others, several can aid in distinguishing genus‐level clades. Our resulting trees and character analyses form the basis of a revised classification that spans a total of 139 species contained within 24 genera. The tree topologies necessitate the synonymization of Barabattoia as Dipsastraea, and Phymastrea as Favites. Furthermore, Astrea and Coelastrea are resurrected, and one new genus, P aramontastraea Huang & Budd gen. nov. , is described. All the genera in Merulinidae, along with the monotypic Montastraeidae and Diploastraeidae, are diagnosed based on the characters examined. The integrative classification system proposed here will form the framework for more accurate biodiversity estimates and guide the taxonomic placement of extinct species. © 2014 The Linnean Society of London     

The effects of a coral disease on the reproductive output of Montastraea faveolata (Scleractinia: Faviidae)

The direct impacts of coral diseases on coral populations have been assessed by quantifying coral tissue loss and colony mortality, but the determination of the indirect effects of diseases, such as disruptions in life history functions (e.g. reproduction, growth and maintenance), are more difficult to ascertain and have been scant. This study involved a comparison of various measures of reproductive output from histological slides of healthy tissue samples of Montastraea faveolata and tissue samples from colonies with white plague (WP) infections in Dominica (West Indies). Although the variability in the reproductive data was high, WP had significant negative impacts on the percentage of reproductive polyps per cm2, the percentage of reproductive mesenteries within a polyp, oocyte quantity per polyp, mean oocyte volume (mm3), and fecundity (oocyte volume per cm2 of tissue). However, these effects were only observed in the tissue directly impacted by the WP disease "band" and were not observed in tissue samples taken 20 cm away from the lesion. Therefore, the effects of a coral disease (WP) on reproductive output are localized and not expressed colony-wide.     

Diel thermoregulation of the crawfish Procambarus Clarkii (crustacea, cambaridae)

1. 1. In a diel cycle Procambarus clarkii has two preferred temperatures: 24.0 ± 0.15 SEM °C during the day and 26.7 ± 0.13 SEM °C at night.

2. 2. The preferred temperatures are independent from the weight of the organisms.

3. 3. In the photophase the animals are dispersed, in the scotophase they congregate.

4. 4. The crawfish seem to feed during the thermal interphases.

5. 5. Animals in a constantly dark condition maintain a diel preferendum of temperature.