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The scleractinian coral Cyphastrea is a common and widespread genus throughout the coral reefs of the Indo-Pacific. Little is known about the phylogenetic relationships within this taxon and species identification is based mainly on traditional skeletal characters, such as the number of septa, septa cycles, growth form and corallite dimensions. Here we present the first focussed reconstruction of phylogenetic relationships among Cyphastrea species, analysing 57 colonies from the Red Sea, where five morphospecies live in sympatry. Analyses based on three loci (nuclear histone H3, 28S rDNA and a mitochondrial intergenic region) reveal the existence of three well-supported molecular lineages. None of the five previously defined morphospecies are monophyletic and they cluster into two clades, suggesting the need of a systematic revision in Cyphastrea. The third lineage is described as C. magna Benzoni & Arrigoni, sp. nov., a new reef coral species collected from the northern and central Red Sea. Cyphastrea magna Benzoni & Arrigoni, sp. nov. is characterised by the largest corallite diameter among known Cyphastrea species, a wide trabecular columella >1/4 of calice width, and 12 equal primary septa. This study suggests that morphology-based taxonomy in Cyphastrea may not identify monophyletic units and strengthens the application of genetics in coral systematics.

Screening of the Arabidopsis thaliana genome revealed three potential homologues of mammalian and yeast mitochondrial DICs (dicarboxylate carriers) designated as DIC1, DIC2 and DIC3, each belonging to the mitochondrial carrier protein family. DIC1 and DIC2 are broadly expressed at comparable levels in all the tissues investigated. DIC1-DIC3 have been reported previously as uncoupling proteins, but direct transport assays with recombinant and reconstituted DIC proteins clearly demonstrate that their substrate specificity is unique to plants, showing the combined characteristics of the DIC and oxaloacetate carrier in yeast. Indeed, the Arabidopsis DICs transported a wide range of dicarboxylic acids including malate, oxaloacetate and succinate as well as phosphate, sulfate and thiosulfate at high rates, whereas 2-oxoglutarate was revealed to be a very poor substrate. The role of these plant mitochondrial DICs is discussed with respect to other known mitochondrial carrier family members including uncoupling proteins. It is proposed that plant DICs constitute the membrane component of several metabolic processes including the malate-oxaloacetate shuttle, the most import

To shed light on the metabolic role of two mitochondrial transporters for basic amino acids in Arabidopsis, we compared their functional properties in liposomes and expression during germination. Recombinant and purified BAC2, as previously reported for BAC1, transported various basic l-amino acids upon reconstitution in phospholipid vesicles. Both displayed highest affinity for arginine with similar K(m) and V(max). However, BAC2 transported citrulline for which BAC1 had little or no affinity. Furthermore, BAC2 was less stereospecific than BAC1, transporting d-arginine and d-lysine at significant rates, and displayed a striking alkaline pH optimum (pH 8.0) whereas BAC1 activity was unaltered from pH 7.0 to 9.0. By semi-quantitative RT-PCR BAC1 transcript levels were found to be higher than those of BAC2 in germinated seeds. However, BAC2 expression transiently increased 2 days after germination. Disruption of the Arabidopsis arginase structural genes (ARGAH1 or ARGAH2) accentuated the increases of transcript levels of BAC1 at germination and of BAC2 2 days after germination and from 6 days on. Early expression of BAC1 and BAC2 is consistent with the delivery of arginine, released from seed reserves, to mitochondrial arginase and the export of ornithine. Increase of BAC2 transcript levels later in seedling development is consistent with roles in NO, polyamine or proline metabolism-processes involving arginine, citrulline and/or ornithine.

The dicarboxylate carrier (DIC) is a nuclear-encoded protein located in the mitochondrial inner membrane. It catalyses the transport of dicarboxylates such as malate and succinate across the mitochondrial membrane in exchange for phosphate, sulphate and thiosulphate. We have determined the sequences of the human cDNA and gene for the DIC. The gene sequence was established from overlapping genomic clones generated by PCRs by use of primers and probes based upon the human cDNA sequence. It is spread over 8.6 kb of human DNA and is divided into 11 exons. Five short interspersed repetitive Alu sequences are found in intron I. The protein encoded by the gene is 287 amino acids long. In common with the rat protein, it does not have a processed presequence to help to target it into mitochondria. It has been demonstrated by Northern- and Western-blot analyses that the DIC is present in high amounts in liver and kidney, and at lower levels in all the other tissues analysed. The positions of introns contribute towards an understanding of the processes involved in the evolution of human genes for carrier proteins.

The authors thank all the people involved in collecting/providing material or organising sampling campaigns: Peter Schuchert (MHNG, Switzerland), Tullia Isotta Terraneo (KAUST, Saudi Arabia), Malek Amr Gusti (KAUST, Saudi Arabia), Timothy Ravasi (OIST, Japan), the captain and crew of the MV Dream-Master (Saudi Arabia), the KAUST Coastal and Marine Resources Core Lab, Inga Dehnert (UNIMIB, Italy), Nicholas WL Yap (NUS, Singapore), Sudhanshi S Jain (NUS, Singapore), Stephen Keable (Australian Museum), Penny Berents (Australian Museum), Anne Hoggett (Australian Museum), Lyle Vail (Australian Museum). Additionally, we wish to thank Leen P. van Ofwegen (Naturalis, The Netherlands) for his valuable help in identifying the octocoral Paralemnalia sp., Peter Schuchert for his comments on an earlier version of the manuscript, and two anonymous referees for their thorough revision of this work. Permissions relevant to undertake the research have been obtained from the applicable governmental agencies. Fieldwork at St. Eustatius was funded through a Martin Fellowship from Naturalis Biodiversity Center to SM, while logistic support was supplied by St. Eustatius Marine Parks (STENAPA), the Caribbean Netherlands Science Institute (CNSI) and Scubaqua Dive Centre. Samples from Eilat (Israel) were collected during the HyDRa Project funded by the EU FP7 Research Infrastructure Initiative ‘ASSEMBLE’ (Grant #227799) to DP. Financial support to DP for collecting samples at Lizard Island (Australia) was provided by the 2018 John and Laurine Proud Fellowship and the Australian Museum’s Lizard Island Research Station. Fieldwork in Mozambique was conducted during the Green Bubbles financed by EU’s H2020 research and innovation programme to DP, under the Marie Sklodowska-Curie grant agreement no 643712 (Permit n° 09/2018 ANAC). Fieldwork in Singapore was partially funded by the National Research Foundation, Prime Minister’s Office, Singapore under its Marine Science R&D Programme (MSRDP-P03) to DH.

Scleractinian corals provide habitats for a broad variety of cryptofauna, which in turn may contribute to the overall functioning of coral symbiomes. Among these invertebrates, hydrozoans belonging to the genus Zanclea represent an increasingly known and ecologically important group of coral symbionts. In this study, we analysed 321 Zanclea colonies associated with 31 coral genera collected from 11 localities across the Indo-Pacific and Caribbean regions, and used a multi-disciplinary approach to shed light on the evolution and biogeography of the group. Overall, we found high genetic diversity of hydrozoans that spans nine clades corresponding to cryptic or pseudo-cryptic species. All but two clades are associated with one or two coral genera belonging to the Complex clade, whereas the remaining ones are generalists associated with both Complex and Robust corals. Despite the observed specificity patterns, no congruence between Zanclea and coral phylogenies was observed, suggesting a lack of coevolutionary events. Most Zanclea clades have a wide distribution across the Indo-Pacific, including a generalist group extending also into the Caribbean, while two host-specific clades are possibly found exclusively in the Red Sea, confirming the importance of this peripheral region as an endemicity hotspot. Ancestral state reconstruction suggests that the most recent common ancestor of all extant coral-associated Zanclea was a specialist species with a perisarc, occurring in what is now known as the Indo-Pacific. Ultimately, a mixture of geography- and host-related diversification processes is likely responsible for the observed enigmatic phylogenetic structure of coral-associated Zanclea.

Modern systematics integrating molecular and morphological data has greatly improved our understanding of coral evolutionary relationships during the last two decades and led to a deeply revised taxonomy of the order Scleractinia. The family Merulinidae (Cnidaria: Scleractinia) was recently subjected to a series of revisions following this integrated approach but the phylogenetic affinities of several genera ascribed to it remain unknown. Here, we partially fill this gap through the study of 89 specimens belonging to all 10 valid species from four genera (Caulastraea, Erythrastrea, Oulophyllia and Dipsastraea) collected from 14 localities across the Indo‐Pacific realm. Four molecular loci (histone H3, COI, ITS and IGR) were sequenced, and a total of 44 skeletal morphological characters (macromorphology, micromorphology and microstructure) were analysed. Molecular phylogenetic analyses revealed that the phaceloid Caulastraea species are split into two distinct lineages. A species previously ascribed to the genus Dipsastraea, Dipsastraea maxima, is also recovered in one on these lineages. Furthermore, Erythrastrea is nested within Oulophyllia. The molecular reconstructions of evolutionary relationships are further corroborated by multiscale morphological evidence. To resolve the taxonomy of these genera, Astraeosmilia is resurrected to accommodate Astraeosmilia connata, Astraeosmilia curvata, Astraeosmilia tumida and Astraeosmilia maxima, with Caulastraea retaining Caulastraea furcata and Caulastraea echinulata. Based on the examination of type material, Erythrastrea flabellata is considered an objective synonym of Lobophyllia wellsi, which is transferred to Oulophyllia following the obtained morpho‐molecular results. This work further confirms that an integrated morpho‐molecular approach based on a rigorous phylogenetic framework is fundamental for an objective classification that reflects the evolutionary history of scleractinian corals.

Zanclea divergens is a tropical hydrozoan living in symbiotic association with bryozoans and currently reported from Papua New Guinea, Indonesia, and Maldives. Here, we used an integrative approach to assess the morpho-molecular diversity of the species across the Indo-Pacific. Phylogenetic and species delimitation analyses based on seven mitochondrial and nuclear loci revealed four well-supported molecular lineages corresponding to cryptic species, and representing a Pacific clade, an Indian clade, and two Red Sea clades. Since the general polyp morphology was almost identical in all samples, the nematocyst capsules were measured and analysed to search for possible fine-scale differences, and their statistical treatment revealed a significant difference in terms of length and width among the clades investigated. All Zanclea divergens specimens were specifically associated with cheilostome bryozoans belonging to the genus Celleporaria. The Pacific and Indian clades were associated with Celleporaria sp. and C. vermiformis, respectively, whereas both Red Sea lineages were associated with C. pigmentaria. Nevertheless, the sequencing of host bryozoans revealed that one of the Red Sea hydrozoan clades is associated with two morphologically undistinguishable, but genetically divergent, bryozoan species. Overall, our results show that Z. divergens is a species complex composed of morphologically cryptic lineages showing partially disjunct distributions and host specificity. The presence of two sympatric lineages living on the same host species reveal complex dynamics of diversification, and future research aimed at understanding their diversification process will likely improve our knowledge on the mechanisms of speciation among currently sympatric cryptic species.

Accurate delimitation of species and their relationships is a fundamental issue in evolutionary biology and taxonomy and provides essential implications for conservation management. Scleractinian corals are difficult to identify because of their ecophenotypic and geographic variation and their morphological plasticity. Furthermore, phylogenies based on traditional loci are often unresolved at the species level because of uninformative loci. Here, we attempted to resolve these issues and proposed a consistent species definition method for corals by applying the genome-wide technique Restriction-site Associated DNA sequencing (RADseq) to investigate phylogenetic relationships and species delimitation within the genus Leptastrea. We collected 77 colonies from nine localities of the Indo-Pacific and subjected them to genomic analyses. Based on de novo clustering, we obtained 44,162 SNPs (3701 loci) from the holobiont dataset and 62,728 SNPs (9573 loci) from the reads that map to coral transcriptome to reconstruct a robust phylogenetic hypothesis of the genus. Moreover, nearly complete mitochondrial genomes and ribosomal DNA arrays were retrieved by reference mapping. We combined concatenation-based phylogenetic analyses with coalescent-based species tree and species delimitation methods. Phylogenies suggest the presence of six distinct species, three corresponding to known taxa, namely Leptastrea bottae, Leptastrea inaequalis, Leptastrea transversa, one characterized by a remarkable skeletal variability encompassing the typical morphologies of Leptastrea purpurea and Leptastrea pruinosa, and two distinct and currently undescribed species. Therefore, based on the combination of genomic, morphological, morphometric, and distributional data, we herein described Leptastrea gibbosa sp. n. from the Pacific Ocean and Leptastrea magaloni sp. n. from the southwestern Indian Ocean and formally considered L. pruinosa as a junior synonym of L. purpurea. Notably, mitogenomes and rDNA yielded a concordant yet less resolved phylogeny reconstruction compared to the ones based on SNPs. This aspect demonstrates the strength and utility of RADseq technology for disentangling species boundaries in closely related species and in a challenging group such as scleractinian corals.

Extant biodiversity can easily be underestimated owing to the presence of cryptic taxa, even among commonly observed species. Scleractinian corals are challenging to identify because of their ecophenotypic variation and morphological plasticity. In addition, molecular analyses have revealed the occurrence of cryptic speciation. Here, we describe a new cryptic lobophylliid genus and species Paraechinophyllia variabilis gen. nov., sp. nov., which is morphologically similar to Echinophyllia aspera and E. orpheensis. The new taxon occurs in Mayotte Island, Madagascar, the Gulf of Aden and the Red Sea. Six molecular markers (COI, 12S, ATP6-NAD4, NAD3-NAD5, histone H3 and ITS) and 46 morphological characters at three different levels (macromorphology, micromorphology and microstructure) were examined. The resulting molecular phylogenetic reconstruction showed that Paraechinophyllia gen. nov. represents a distinct group within the Lobophylliidae that diverged from the lineage leading to Echinophyllia and Oxypora in the Early Miocene, approximately 21.5 Ma. The morphological phylogenetic reconstruction clustered Paraechinophyllia gen. nov., Echinophyllia and Oxypora together in a single clade. A sole morphological character, calice relief, discriminated Paraechinophyllia gen. nov. from the latter two genera, suggesting that limited morphological variation has occurred over a long period. These results highlight the importance of cryptic taxa in reef corals, with implications for population genetics, ecological studies and conservation.

AimThe aim of the study was to assess the diversity of algal symbionts of the family Symbiodiniaceae associated with the coral genus Porites in the Red Sea, and to test for host‐specificity and environmental variables driving biogeographical patterns of algal symbiont distribution.LocationSaudi Arabian Red Sea.TaxonEndosymbiotic dinoflagellates of the family Symbiodiniaceae in association with the reef‐building coral genus Porites.MethodsEighty Porites coral specimens were collected along the Saudi Arabian Red Sea coast. Species boundaries were assessed morphologically and genetically (putative Control Region – mtCR; ITS region – ITS). Community composition of symbiotic dinoflagellates of the family Symbiodiniaceae was also assessed. Using the ITS2 marker with the SymPortal framework, Symbiodiniaceae data at the genus, majority ITS2 sequence and ITS2 type profile were used to assess symbiont diversity and distribution patterns. These were analysed in relation to coral host diversity, geographic location and environmental variables.ResultsAmong the 80 Porites samples, 10 morphologies were identified. These corals were clustered into five lineages (clades I–V) by each of the markers independently. Clades I, II and III each comprised of a single Porites morphology, while clades IV and V contained up to five distinct morphologies. The diversity of Symbiodiniaceae associated with Porites was high and latitudinal differentiation was observed. In particular, a shift from a Cladocopium‐dominated to a Durusdinium‐dominated community was found along the north–south gradient. Symbiont diversity showed the patterns of geographic‐specific association at Symbiodiniaceae genus, majority ITS2 sequence and ITS2 type profile level. Specific associations with host genotypes (but not morphological species) were also recovered when considering Symbiodiniaceae majority ITS2 sequence and ITS2 type profiles.Main conclusionsThis study provides the first large‐scale molecular characterization of Symbiodiniaceae communities associated with Porites corals from the Saudi Arabian Red Sea. The use of intragenomic diversity data enabled the resolution of host‐symbiont specificity and biogeographical patterns of distribution, previously unachievable with the ITS2 marker alone. Finally, correlation among symbiont diversity and Red Sea environmental gradients was documented. published

The biodiversity of Red Sea corals captured the attention of some of the earliest European natural historians. Many of the first descriptions of tropical reef corals were based on Red Sea material. Modern approaches to resolve the notorious challenges of coral taxonomy have only recently been applied to Red Sea taxa. This chapter reviews current knowledge of the distributions of coral species in the Arabian region, including assessments of endemism. We also review new species described (or resurrected) since the last major assessment (in 2002). Where sufficient data is available, we highlight within-region distribution patterns. The Red Sea has the highest levels of endemism among all regions of the Indian Ocean. Analysis of the similarity of species composition among the Arabian subregions shows that the Red Sea and Socotra Island are the most speciose, but also have distinct community compositions. The regional diversity of Red Sea corals is likely influenced by the unique environmental gradients of the Arabian region. Despite evolving in testing conditions, Red Sea corals have been impacted by global climate change. Recent thermal bleaching events in the Red Sea highlight the pressures and challenges to future recovery.

The colonial stony coral genus Madracis is cosmopolitan, lives in shallow and deep water habitats, and includes zooxanthellate, azooxanthellate and facultative symbiotic species. One of its species, Madracis pharensis, has been recorded from the Mediterranean and East Atlantic, where it forms small knobby and facultative zooxanthellate colonies (also named M. pharensis f. pharensis), and from the tropical Caribbean, where it also occurs in a massive and zooxanthellate form (named M. pharensis f. luciphila by some). These two forms have been previously found to host different Symbiodinium species. In this study, species boundaries and phylogenetic relationships between these two Madracis pharensis forms (from the Mediterranean Sea and the Caribbean), M. senaria, and the Indo-west Pacific M. kirbyi were analyzed through an integrated systematics approach, including corallite dimensions, micromorphology and two molecular markers (ITS and ATP8). Significant genetic and morphological differences were found between all the examined Madracis species, and between M. pharensis from the Mediterranean Sea and M. pharensis f. luciphila from the Caribbean in particular. Based on these results, the latter does not represent a zooxanthellate ecomorph of the former but a different species. Its identity remains to be ascertained and its relationship with the Caribbean M. decactis, with which it bears morphologic resemblance, must be investigated in further studies. Overall, the presence of cryptic Madracis species in the Easter and Central Atlantic Ocean remains to be evaluated.

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.

The scleractinian coral Cyphastrea is a common and widespread genus throughout the coral reefs of the Indo-Pacific. Little is known about the phylogenetic relationships within this taxon and species identification is based mainly on traditional skeletal characters, such as the number of septa, septa cycles, growth form and corallite dimensions. Here we present the first focussed reconstruction of phylogenetic relationships among Cyphastrea species, analysing 57 colonies from the Red Sea, where five morphospecies live in sympatry. Analyses based on three loci (nuclear histone H3, 28S rDNA and a mitochondrial intergenic region) reveal the existence of three well-supported molecular lineages. None of the five previously defined morphospecies are monophyletic and they cluster into two clades, suggesting the need of a systematic revision in Cyphastrea. The third lineage is described as C. magna Benzoni & Arrigoni, sp. nov., a new reef coral species collected from the northern and central Red Sea. Cyphastrea magna Benzoni & Arrigoni, sp. nov. is characterised by the largest corallite diameter among known Cyphastrea species, a wide trabecular columella >1/4 of calice width, and 12 equal primary septa. This study suggests that morphology-based taxonomy in Cyphastrea may not identify monophyletic units and strengthens the application of genetics in coral systematics.

To date, four nominal species and several other unidentified species of Zanclea hydrozoans are known to live symbiotically with scleractinians, and recent surveys reported this association also in the Red Sea. Previous molecular studies showed that each coral genus involved in this association hosts only one species or molecular clade of Zanclea, with the only exception being the genus Acropora, which hosts at least two Zanclea species. Moreover, some of the detected genetic lineages were morphologically undistinguishable in the polyp stage, suggesting the presence of cryptic species. In this study, we investigated the morphology and genetic diversity of Acropora-associated Zanclea specimens collected in previous studies in Egypt and Israel, as well as new samples collected in Saudi Arabia. Based on the current data, all the analysed samples were morphologically identical to Zanclea gallii, a species associated with Acropora corals from the Maldives. However, molecular analyses separated the samples collected in the Red Sea from all other coral-associated hydroids. Therefore, phylogenetic reconstructions, haplotype networks, genetic distance analyses and distribution data allowed us to identify a previously unknown cryptic species of Acropora-associated hydroid, here named Zanclea gallii IIa, following a recently proposed molecular nomenclature.

In this study, we sequenced the complete mitochondrial genome of Porites harrisoni using ezRAD and Illumina technology. Genome length consisted of 18,630 bp, with a base composition of 25.92% A, 13.28% T, 23.06% G, and 37.73% C. Consistent with other hard corals, P. harrisoni mitogenome was arranged in 13 protein-coding genes, 2 rRNA, and 2 tRNA genes. nad5 and cox1 contained embedded Group I Introns of 11,133 bp and 965 bp, respectively.

Corals in the genus Porites are among the major framework builders of reef structures worldwide, yet the genus has been challenging to study due to a lack of informative molecular markers. Here, we used ezRAD sequencing to reconstruct the complete mitochondrial genome of Porites fontanesii (GenBank accession number MG754069), a widespread coral species endemic to the Red Sea and Gulf of Aden. The gene arrangement of P. fontanesii did not differ from other Scleractinia and consisted of 18,658 bp, organized in 13 protein-coding genes, 2 rRNA genes, and 2 tRNA genes. This mitochondrial genome contributes essential data to work towards a better understanding of evolutionary relationships within Porites.

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.

The complete nucleotide sequence of the mitochondrial genome of the scleractinian coral Acanthastrea maxima has been obtained, representing the first sequenced mitogenome of a member of the Lobophylliidae. The mitochondrial genome is 18,278 bp in length, the longest sequence among the robust corals sequenced mitogenome to date. The overall GC composition (33.7%) and the gene arrangement are similar to those of the other scleractinian corals, including 13 protein-coding genes, 2 rRNA genes (rnl and rns) and 2 tRNA genes (tRNA-Met and tRNA-Trp). All genes except tRNA-Trp, atp8, cox1, tRNA-Met and rnl are engulfed by a large group I intron in the nad5 gene. A second group I intron of 1077 bp in length is inserted in the cox1 gene and it encodes a putative homing endonuclease. There are four regions of gene overlaps totalling 22 bp and nine intergenic spacer regions for a total of 2220 bp, of which the cox3-cox2 region may correspond to the putative control region.

The scleractinian family Lobophylliidae is undergoing a major taxonomic revision thanks to the combination of molecular and morphological data. In this study, we investigate the evolutionary relationships and the macro- and micromorphology of six nominal coral species belonging to two of the nine molecular clades of the Lobophylliidae, clades A and B, and of Symphyllia wilsoni, a lobophylliid species analyzed from a molecular point of view for the first time. Sequence data from mitochondrial DNA (COI and the intergenic spacer between COI and l-rRNA), and nuclear DNA (histone H3 and ITS region) are used to generate robust molecular phylogenies and a median-joining haplotype network. Molecular results are strongly in agreement with detailed observations of gross- and fine-scale morphology of skeletons, leading to the formal revision of the genera Micromussa and Homophyllia and the description of two newly discovered zooxanthellate shallow-water species, Micromussa pacifica sp. nov. Benzoni & Arrigoni and Micromussa indiana sp. nov. Benzoni & Arrigoni, and a new genus, Australophyllia gen. nov. Benzoni & Arrigoni. In particular, Acanthastrea lordhowensis and Montastraea multipunctata are moved into Micromussa, A. hillae is synonymized with A. bowerbanki and is transferred to Homophyllia, and a revised diagnosis for both genera is provided. Micromussa pacifica sp. nov. is described from the Gambier Islands with its distribution spanning New Caledonia and eastern Australia. Despite a superficial resemblance with Homophyllia australis, it has distinctive macroand micromorphological septal features. Micromussa indiana sp. nov., previously identified as M. amakusensis, is here described from the Gulf of Aden and the southern Red Sea as a distinct species that is genetically separated from M. amakusensis and is morphologically distinct from the latter due to its smaller corallite size and lower number of septa. Finally, molecular trees show that S. wilsoni is closely related, but molecularly separated from clades A and B, and, also based on a unique combination of corallite and sub-corallite characters, the species is moved into Australophyllia gen. nov. These findings confirm the need for using both genetic and morphological datasets for the ongoing taxonomic revision of scleractinian corals.

Scleractinian corals are affected by environment-induced phenotypic plasticity and intraspecific morphological variation caused by genotype. In an effort to identify new strategies for resolving this taxonomic issue, we applied a molecular approach for species evaluation to two closely related genera, Echinophyllia and Oxypora, for which few molecular data are available. A robust multi-locus phylogeny using DNA sequence data across four loci of both mitochondrial (COI, ATP6-NAD4) and nuclear (histone H3, ITS region) origin from 109 coral colonies was coupled with three independent putative species delimitation methods based on barcoding threshold (ABGD) and coalescence theory (PTP, GMYC). Observed overall congruence across multiple genetic analyses distinguished two traditional species (E. echinoporoides and O. convoluta), a species complex composed of E. aspera, E. orpheensis, E. tarae, and O. glabra, whereas O. lacera and E. echinata were indistinguishable with the sequenced loci. The combination of molecular species delimitation approaches and skeletal character observations allowed the description of two new reef coral species, E. bulbosa sp. n. from the Red Sea and E. gallii sp. n. from the Maldives and Mayotte. This work demonstrated the efficiency of multi-locus phylogenetic analyses and recently developed molecular species delimitation approaches as valuable tools to disentangle taxonomic issues caused by morphological ambiguities and to re-assess the diversity of scleractinian corals.

Traditional classification of scleractinian corals (Cnidaria, Anthozoa, Scleractinia) have been conducted on the basis of skeleton macromorphology. However, the commonly used skeletal characters are plagued by phenotypic plasticity, intraspecific variation, and morphological convergence that fail to recognize most natural evolutionary lineages. Recent molecular studies have revolutionized the conventional taxonomic schemes, suggesting remarkably different phylogenetic relationships when compared with those based on macromorphology. In the last decade, the integration of this increasing amount of genetic data and new micromorphological and microstructural traits has led to a better understanding of evolutionary relationships between hard corals and opened the way to the new era of coral taxonomy. The Indo-Pacific family Lobophylliidae potentially represents an interesting case study of “reciprocal illumination” between genetics and morphology. This taxon has been defined on a combination of phylogenetic analyses and micromorphological observations, although it still remains poorly understood.This dissertation aims to fill this gap in knowledge through the investigation of evolutionary relationships of the Lobophylliidae and starting the impending process of taxonomic revision of this family as a result of an integrated molecular and micromorphological approach. A molecular phylogeny of one of the three major groups recovered in the order Scleractinia, the Robusta, to which the Lobophylliidae belong is presented. The representatives of this family were recovered in a cohesive lineage for the first time and the phylogenetic relationships with the other closely related families were discussed. The analysis was then expanded analyzing more species and samples and focused exclusively on the Lobophylliidae in order to produce the most comprehensive molecular phylogeny reconstruction of this group to date. The monophyly of the family was strongly supported and nine main monophyletic genus-level lineages were recovered within the Lobophylliidae. All analyzed polytypic genera, were not monophyletic and resulted in need of a formal taxonomic revision. Subsequently, the complete mitochondrial genome of Acanthastrea maxima was sequenced. This species based on a combination of data including its restricted geographic distribution, evolutionary distinctiveness, and small population size was revealed be a case of priority for future conservation strategies. Being 18,278 bp in length, it is the longest sequence among the robust corals sequenced mitogenome to date, while the GC content and the gene arrangement are similar to those of the other scleractinian corals. Finally, integrating multi-locus molecular phylogenies and detailed gross- and fine-scale morphologic observations three cases analyzed in the family provided examples of how reverse taxonomy can be useful in understanding the evolutionary history of the Lobophylliidae: I) a taxonomic revision for the monotypic genus Australomussa, revealed to be a junior synonym of Parascolymia, was proposed; II) the long-ignored monospecific genus Sclerophyllia was resurrected and the unforeseen sister relationships between Sclerophyllia margariticola and A. maxima led to the placement of the latter species in Sclerophyllia; III) the closely related genera Homophyllia and Micromussa were revised with the description of two new species of Micromussa and Symphyllia wilsoni, an Australian endemism, was placed in a new genus based on an unique combination of molecular and micromorphological data. Overall, the results stemming from the results obtained in the framework of my thesis significantly improve our understanding of evolution of the family Lobophylliidae and provide a solid case for the importance of an integrated morpho-molecular approach in resolving taxonomy of this group of marine animals.

Hydroids in the genus Zanclea are a recently discovered component of the fauna associated with reef-building corals. The phylogenetic relationships among these species are not well known. The present work is based on field surveys in the Republic of Maldives, and for the first time, morphological and molecular analyses are integrated to distinguish a new hydroid species and provide new information on the ecology of this symbiosis. This new hydroid, Zanclea gallii sp. n., was associated with the scleractinian Acropora muricata; it was living sympatrically with its congener Zanclea sango, which was observed for the first time at this locality on the new scleractinian host Pavona varians. The relationships between these two hydroids and other available scleractinian-associated Zanclea were investigated using two molecular markers, nuclear 28S rDNA and mitochondrial 16S rRNA. Zanclea gallii sp. n. and Z. sango were recovered as distinct lineages within a monophyletic group of scleractinian-associated Zanclea based on both molecular and morphological data. All Zanclea species that were observed living in association with scleractinians belong to the ‘polymorpha group’ and share the morphological characteristic ‘polymorphic colony’. The genus Leptoseris is the 16th host coral identified for Zanclea. Compared with the frequency of the Z. gallii sp. n. association with A. muricata and Z. sango with the scleractinian P. varians, the latter is twice as common; however, the former exhibited higher Zanclea polyps concentrations over the colony surface. Overall, the Zanclea survey indicates that these diminutive hydroids are more commonly associated with coral than previously known.

Recent molecular phylogenetic studies have shown that most traditional families of zooxanthellate shallow-water scleractinians are polyphyletic, whereas most families mainly composed of deep-sea and azooxanthellate species are monophyletic. In this context, the family Dendrophylliidae (Cnidaria, Scleractinia) has unique features. It shows a remarkable variation of morphological and ecological traits by including species that are either colonial or solitary, zooxanthellate or azooxanthellate, and inhabiting shallow or deep water. Despite this morphological heterogeneity, recent molecular works have confirmed that this family is monophyletic. Nevertheless, what so far is known about the evolutionary relationships within this family, is predominantly based on skeleton macromorphology, while most of its species have remained unstudied from a molecular point of view. Therefore, we analysed 11 dendrophylliid genera, four of which were investigated for the first time, and 30 species at molecular, micromorphological and microstructural levels. We present a robust molecular phylogeny reconstruction based on two mitochondrial markers (COI and the intergenic spacer between COI and 16S) and one nuclear (rDNA), which is used as basis to compare micromorphogical and microstructural character states within the family. The monophyly of the Dendrophylliidae is well supported by molecular data and also by the presence of rapid accretion deposits, which are ca. 5 μm in diameter and arranged in irregular clusters, and fibres that thicken the skeleton organized in small patches of a few micrometres in diameter. However, all genera represented by at least two species are not monophyletic, Tubastraea excluded. They were defined by traditional macromorphological characters that appear affected by convergence, homoplasy and intraspecific variation. Micromorphogical and microstructural analyses do not support the distinction of clades, with the exception of the organization of thickening deposits for the Tubastraea clade.

Recent advances in scleractinian systematics and taxonomy have been achieved through the integration of molecular and morphological data, as well as rigorous analysis using phylogenetic methods. In this study, we continue in our pursuit of a phylogenetic classification by examining the evolutionary relationships between the closely related reef coral genera Merulina, Goniastrea, Paraclavarina and Scapophyllia (Merulinidae). In particular, we address the extreme polyphyly of Favites and Goniastrea that was discovered a decade ago. We sampled 145 specimens belonging to 16 species from a wide geographic range in the Indo-Pacific, focusing especially on type localities, including the Red Sea, western Indian Ocean and central Pacific. Tree reconstructions based on both nuclear and mitochondrial markers reveal a novel lineage composed of three species previously placed in Favites and Goniastrea. Morphological analyses indicate that this clade, Paragoniastrea Huang, Benzoni & Budd, gen. n., has a unique combination of corallite and subcorallite features observable with scanning electron microscopy and thin sections. Molecular and morphological evidence furthermore indicates that the monotypic genus Paraclavarina is nested within Merulina, and the former is therefore synonymised.

The family Poritidae formerly included 6 genera: Alveopora, Goniopora, Machadoporites, Porites, Poritipora, and Stylaraea. Morphologically, the genera can be differentiated based on the number of tentacles, the number of septa and their arrangement, the length of the polyp column, and the diameter of the corallites. However, the phylogenetic relationships within and between the genera are unknown or contentious. On the one hand, Alveopora has been transferred to the Acroporidae recently because it was shown to be more closely related to this family than to the Poritidae by previous molecular studies. On the other hand, Goniopora is morphologically similar to 2 recently described genera, Machadoporites and Poritipora, particularly with regard to the number of septa (approximately 24), but they have not yet been investigated at the molecular level. In this study, we analyzed 93 samples from all 5 poritid genera and Alveopora using 2 genetic markers (the barcoding region of the mitochondrial COI and the ITS region of the nuclear rDNA) to investigate their phylogenetic relationships and to revise their taxonomy. The reconstructed molecular trees confirmed that Alveopora is genetically distant from all poritid genera but closely related to the family Acroporidae, whereas the other genera are genetically closely related. The molecular trees also revealed that Machadoporites and Poritipora were indistinguishable from Goniopora. However, Goniopora stutchburyi was genetically isolated from the other congeneric species and formed a sister group to Goniopora together with Porites and Stylaraea , thus suggesting that 24 septa could be an ancestral feature in the Poritidae. Based on these data, we move G. stutchburyi into a new genus, Bernardpora gen. nov., whereas Machadoporites and Poritipora are merged with Goniopora. © 2014 Kitano et al.

Novel micromorphological characters in combination with molecular studies have led to an extensive revision of the taxonomy and systematics of scleractinian corals. In the present work, we investigate the macro- and micromorphology and the phylogenetic position of the genera Australomussa and Parascolymia, two monotypic genera ascribed to the family Lobophylliidae. The molecular phylogeny of both genera was addressed using three markers, the partial mitochondrial COI gene and the nuclear histone H3 and the ribosomal ITS region. Based on molecular data, Australomussa and Parascolymia belong to the Lobophylliidae and they cluster together with the genera Lobophyllia and Symphyllia within the same clade. While A. rowleyensis and P. vitiensis are closely related based on the three gene regions examined, their macro and micromorphology suggest that these species are distinct, differing in several characters, such as continuity and thickness of the costosepta, the number of septa, septal tooth height, spacing, and shape, and the distribution and shape of granules. Thus, we revise the taxonomic status of the genus Australomussa as a junior synonym of Parascolymia.

The Indo-Pacific scleractinian coral family Lobophylliidae was recently described on the basis of molecular data and micromorphological and microstructural characters. We present the most comprehensive molecular phylogeny reconstruction of the family to date based on COI and rDNA including 9 genera and 32 species, 14 of which were investigated for the first time. The monophyly of the family is now strongly supported, with the inclusion of the genera Acanthastrea and Micromussa, whereas previously it was based on uncertain molecular relationships. Nevertheless, these and the other lobophylliid genera Echinophyllia, Micromussa, Oxypora, and Symphyllia, are not themselves monophyletic and need to be investigated from a morphological point of view. Acanthastrea faviaformis is nested within the family Merulinidae. This study highlights the need for further analyses at species level and of formal taxonomic actions.

Caulerpa racemosa, a common and opportunistic species widely distributed in tropical and warm-temperate regions, is known to form monospecific stands outside its native range (Verlaque et al. 2003). In October 2011 we observed an alteration in benthic community due to a widespread overgrowth of C. racemosa around the inhabited island of Magoodhoo (3°04’N; 72°57’ E, Republic of Maldives). The mats formed a continuous dense meadow (Fig 1a) that occupied an area of 95 x 120 m (~11,000 m2) previously dominated by the branching coral Acropora muricata. Partial and total mortality (Fig 1 b-c) were recorded on 45% and 30% of A. muricata colonies, respectively. The total area of influence of C. racemosa was however much larger (~25,000 m2 ) including smaller coral patches near to the meadow, where mortality in contact with the algae was also observed on colonies of Isopora palifera, Lobophyllia corymbosa, Pavona varians, Pocillopora damicornis, and Porites solida. Although species of the genus Caulerpa are not usually abundant on oligotrophic coral reefs, nutrient enrichment from natural and/or anthropogenic sources are known to promote green algal blooms (Lapointe and Bedford 2009). Considering the current state of regression of many reefs in the Maldives (Lasagna 2010) we report an unusual phenomenon which could possibly become more common.

The scleractinian species Psammocora explanulata and Coscinaraea wellsi were originally classified in the family Siderastreidae, but in a recent morpho-molecular study it appeared that they are more closely related to each other and to the Fungiidae than to any siderastreid taxon. A subsequent morpho-molecular study of the Fungiidae provided new insights regarding the phylogenetic relationships within that family. In the present study existing molecular data sets of both families were analyzed jointly with those of new specimens and sequences of P. explanulata and C. wellsi. The results indicate that both species actually belong to the Cycloseris clade within the family Fungiidae. A reappraisal of their morphologic characters based on museum specimens and recently collected material substantiate the molecular results. Consequently, they are renamed Cycloseris explanulata and C. wellsi. They are polystomatous and encrusting like C. mokai, another species recently added to the genus, whereas all Cycloseris species were initially thought to be monostomatous and free-living. In the light of the new findings, the taxonomy and distribution data of C. explanulata and C. wellsi have been updated and revised. Finally, the ecological implications of the evolutionary history of the three encrusting polystomatous Cycloseris species and their free-living monostomatous congeners are discussed.