Search

Crustaceans are one of the most widespread and speciose groups of marine organisms, fulfilling multiple ecological roles in numerous ecosystems. On coral reefs, many crustacean species form associations with scleractinian corals. Although the Red Sea is considered a biodiversity hotspot, few studies examined the diversity of coral-associated crustacean communities to date. In this study, 460 decapod crustaceans were recovered from 67 coral colonies of the three branching genera Acropora, Pocillopora and Stylophora in the central Saudi Arabian Red Sea. Crabs and shrimps were morphologically identified to the lowest taxonomic level possible, and portions of the mitochondrial COI and 16S rRNA genes were amplified with the objective of assessing their diversity and phylogenetic relationships. Finally, patterns of co-occurrence were evaluated to investigate the presence of species-specific symbiotic epifauna on different host corals. Overall, we recovered four families, five genera, and nine species of Red Sea crabs, nested into 11 molecular clades, and two families, eight genera and 11 species of shrimps, grouped within 12 lineages. Crabs of the species Trapezia tigrina were found to be exclusively associated with Pocillopora corals, while Tetralia crabs and the shrimps Jocaste japonica and Harpilius lutescens only occurred on Acropora colonies, providing evidence that potential loss of host corals due to local and global impacts could lead to consequent shifts in the symbiotic communities on reefs and to the loss of certain associated taxa. This study represents an advancement towards the understanding and molecular characterization of coral-associated benthic communities in the Red Sea and lays the ground for further research assessing the patterns of biodiversity, evolution, and ecological preferences of these organisms in the area.

The family Euphylliidae consists of reef-building zooxanthellate scleractinian corals distributed across the Indo-Pacific. Seven extant genera comprising a total of 22 valid species are currently recognised. Recent studies have re-organised the taxonomy of the family at the genus level based on molecular and morphological data, including a comprehensive revision of Euphyllia and the resurrection of Fimbriaphyllia. Here, three mitochondrial loci (coi, 12S rRNA, and 16S rRNA) were sequenced and morphological examinations were conducted at three scales (macro/micromorphology and microstructure of the skeleton, and polyp morphology) to study the phylogeny and taxonomy of Euphylliidae. We analysed a total of 11 valid species collected from seven Indo-Pacific localities. The monotypic genus Coeloseris, currently in Agariciidae, was also investigated since previous molecular data suggested a close relationship with the Euphylliidae. Molecular and morphological phylogenetic trees were broadly concordant in the definition of genus-level clades. All analysed genera, i.e., Ctenella, Euphyllia, Fimbriaphyllia, Galaxea, and Gyrosmilia, were reciprocally monophyletic based on molecular results. Coeloseris was nested within the family and, therefore, is formally moved into Euphylliidae. Updated morphological diagnoses are provided for each investigated genus. This study further demonstrated that a phylogenetic classification of scleractinian corals can be achieved by applying a combined morpho-molecular approach. Finally, we encourage phylogenetic and taxonomic studies of the euphylliid taxa not yet analysed molecularly, such as the monotypic genera Montigyra and Simplastrea.

The hydrozoan family Cladocorynidae inhabits tropical to temperate waters and comprises the two genera Pteroclava and Cladocoryne. Pteroclava lives in association with some octocorals and hydrozoans, whereas Cladocoryne is more generalist in terms of substrate choice. This work provides a thorough morpho-molecular reassessment of the Cladocorynidae by presenting the first well-supported phylogeny of the family based on the analyses of three mitochondrial and four nuclear markers. Notably, the two nominal genera were confirmed to be monophyletic and both morphological and genetic data led to the formal description of a new genus exclusively associated with octocorals, Pseudozanclea gen. nov. Maggioni & Montano. Accordingly, the diagnosis of the family was updated. The ancestral state reconstruction of selected characters revealed that the symbiosis with octocorals likely appeared in the most recent common ancestor of Pteroclava and Pseudozanclea. Additionally, the presence of euryteles aggregation in the polyp stage and the exumbrellar nematocyst pouches with euryteles represent synapomorphies of all cladocorynid taxa and probably emerged in their most recent common ancestor. The analysis of several Pteroclava krempfi colonies from Indo-Pacific and Caribbean localities associated with several host octocorals revealed a high intra-specific genetic variability. Single- and multi-locus species delimitations resulted in three to five species hypotheses, but the statistical analysis of morphometric data showed only limited distinction among the clades of P. krempfi. However, P. krempfi clades showed differences in both host specificity, mostly at the octocoral family level, and geographic distribution, with one clade found exclusively in the Caribbean Sea and the others found in the Indo-Pacific.

Capitate hydrozoans are a morphologically and ecologically diverse hydrozoan suborder, currently including about 200 species. Being grouped in two clades, Corynida and Zancleida, these hydrozoans still show a number of taxonomic uncertainties at the species, genus and family levels. Many Capitata species established symbiotic relationships with other benthic organisms, including bryozoans, other cnidarians, molluscs and poriferans, as well as with planktonic dinoflagellates for mixotrophic relationships and with bacteria for thiotrophic ectosymbioses. Our study aimed at providing an updated and comprehensive phylogeny reconstruction of the suborder, at modelling the evolution of selected morphological and ecological characters, and at testing evolutionary relationships between the symbiotic lifestyle and the other characters, by integrating taxonomic, ecological and evolutionary data. The phylogenetic hypotheses here presented shed light on the evolutionary relationships within Capitata, with most families and genera being recovered as monophyletic. The genus Zanclea and family Zancleidae, however, were divided into four divergent clades, requiring the establishment of the new genus Apatizanclea and the new combinations for species in Zanclea and Halocoryne genera. The ancestral state reconstructions revealed that symbiosis arose multiple times in the evolutionary history of the Capitata, and that homoplasy is a common phenomenon in the group. Correlations were found between the evolution of symbiosis and morphological characters, such as the perisarc. Overall, our results highlighted that the use of genetic data and a complete knowledge of the life cycles are strongly needed to disentangle taxonomic and systematic issues in capitate hydrozoans. Finally, the colonization of tropical habitat appears to have influenced the evolution of a symbiotic lifestyle, playing important roles in the evolution of the group.

Reef fishes are a treasured part of marine biodiversity, and also provide needed protein for many millions of people. Although most reef fishes might survive projected increases in ocean temperatures, corals are less tolerant. A few fish species strictly depend on corals for food and shelter, suggesting that coral extinctions could lead to some secondary fish extinctions. However, secondary extinctions could extend far beyond those few coral-dependent species. Furthermore, it is yet unknown how such fish declines might vary around the world. Current coral mass mortalities led us to ask how fish communities would respond to coral loss within and across oceans. We mapped 6964 coral-reef-fish species and 119 coral genera, and then regressed reef-fish species richness against coral generic richness at the 1° scale (after controlling for biogeographic factors that drive species diversification). Consistent with small-scale studies, statistical extrapolations suggested that local fish richness across the globe would be around half its current value in a hypothetical world without coral, leading to more areas with low or intermediate fish species richness and fewer fish diversity hotspots.

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.

An integrated approach using morphological and genetic data is needed to disentangle taxonomic uncertainties affecting the hydrozoan families Sphaerocorynidae and Zancleopsidae. Here we used this approach to accurately characterise species in these families, identify the previously unknown polypstages of the genera Euphysilla and Zancleopsis, which were originally described exclusively based on the medusa stages, describe a new sphaerocorynid genus and species, and assess the phylogenetic position of the two families within the Capitata. The monotypic genus Astrocoryne was found to be a synonym of Zancleopsis. Astrocoryne cabela was therefore transferred to the genus Zancleopsis as Zancleopsis cabela comb. nov. The new polyp-based genus and species Kudacoryne diaphana gen. nov. sp. nov. was erected within the Sphaerocorynidae. Both taxa are primarily based on genetic data, but the introduction of this new genus was made necessary by the fact that it clustered with the genera Heterocoryne and Euphysilla, despite showing Sphaerocoryne-like polyps. Interestingly, the species analysed in this work showed contrasting biogeographical patterns. Based on our data and literature records, some species appear to have a wide circumtropical range, whereas others are limited to few localities. Overall, these results lay the ground for future investigations aimed at resolving the taxonomy and systematics of these two enigmatic families.

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.

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 rDN

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.

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.

Aim: The 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. Location: Saudi Arabian Red Sea. Taxon: Endosymbiotic dinoflagellates of the family Symbiodiniaceae in association with the reef-building coral genus Porites. Methods: Eighty 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. Results: Among 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 conclusions: This study provides the first large-scale molecula

Two new reef coral species, Porites farasani sp. nov. and Porites hadramauti sp. nov. (Scleractinia, Poritidae), are described from the Red Sea and the Gulf of Aden. Porites farasani sp. nov. only occurs in the Farasan Islands in the southern Red Sea, while P. hadramauti sp. nov. has been collected in the Yemen Hadramaut region in the Gulf of Aden. Both species presented striking in situ differences with respect to other Porites species, and were characterized by small encrusting colonies and unusual polyp colouration. In order to test the genetic distinctiveness of P. farasani sp. nov. and P. hadramauti sp. nov. between each other and with respect to other representatives in the genus Porites, we investigated their evolutionary relationships with eight other morphological species of Porites occurring in the Red Sea and in the Gulf of Aden. Two DNA loci, the mitochondrial putative control region and the nuclear ribosomal ITS region, were sequenced, and three species delimitation approaches based on barcoding threshold (Automated Barcoding Gap Discovery) and coalescence theory (Poisson-Tree process, Generalized Mixed Yule Coalescent) were applied. Phylogenetic and species delimitation analyses were overall concordant, resolving P. farasani sp. nov. and P. hadramauti sp. nov. as two divergent but closely related lineages. Of the other morphologically defined Porites species, three were genetically differentiated (P. rus, P. columnaris and P. fontanesii), but five were genetically indistinguishable. The discovery of two regional endemics confirms the importance of the Red Sea and the Gulf of Aden as regions of high biodiversity and suggests the need for an integration of genome-wide molecular data with the re-evaluation of skeletal structures in the systematics of Porites.

Interspecific associations are common in coral reefs, but those involving hydrozoans and octocorals have not been widely investigated. The hydroid Pteroclava krempfi (Hydrozoa, Cladocorynidae) lives in association with different soft coral taxa (Alcyonacea), showing a widespread distribution. However, very little information is available on the ecology of these relationships. Here, we tested for differences in the taxon-specific prevalence and habitat preference of the symbiosis and determined ecological traits of the P. krempfi-host associations in central Red Sea reefs. P. krempfi was found associated with the alcyonacean genera Lobophytum, Rhytisma, Sarcophyton and Sinularia, updating its host range and geographic distribution. The symbiosis prevalence was high in the area and especially at inshore sites compared to midshore and offshore sites. Rhytisma was the most common host, while the association with Lobophytum showed the lowest taxon-specific prevalence. P. krempfi did not show a clear preference for a specific alcyonacean size, and an increase in host size automatically led to an increase in the surface occupied by hydrozoans, although they rarely colonized more than 50% of the upper surface of the host. The spatial distribution of the hydroids on the host surface appeared related to the host genus and size as well as to the coverage of the hydroids. Despite the nature of this symbiosis requiring further investigation, P. krempfi did not seem to play a role in affecting the bleaching susceptibilities of the host colonies. The study shows that the Red Sea coral reef symbioses are more widespread than previously known and therefore deserve more attention.

Coral diseases are emerging as a major threat to coral reefs worldwide, and although many of them have been described, knowledge on their epizootiology is still limited. This is the case of the Halofolliculina ciliate infections, recognized as the skeletal eroding band (SEB) and Caribbean ciliate infection (CCI), two diseases caused by ciliates belonging to the genus Halofolliculina (Class Heterotrichea). Despite their similar macroscopic appearance, the two diseases are considered different and their pathogens have been hypothesized to belong to different Halofolliculina species. In this work, we analysed the morphology and genetic diversity of Halofolliculina ciliates collected in the Caribbean Sea, Red Sea and Indo-Pacific Ocean. Our analyses showed a strong macroscopic similarity of the lesions and similar settlement patterns of the halofolliculinids from the collection localities. In particular, the unique erosion patterns typical of the SEB were observed also in the Caribbean corals. Fine-scale morphological and morphometric examinations revealed a common phenotype in all analysed ciliates, unequivocally identified as Halofolliculina corallasia. Phylogenetic analyses based on nuclear and mitochondrial (COI) molecular markers consistently found all samples as monophyletic. However, although the nuclear marker displayed an extremely low intra-specific diversity, consistent with the morphological recognition of a single species, the analyses based on COI showed a certain level of divergence between samples from different localities. Genetic distances between localities fall within the intra-specific range found in other heterotrich ciliates, but they may also suggest the presence of a H. corallasia species complex. In conclusion, the presented morpho-molecular characterization of Halofolliculina reveals strong similarities between the pathogens causing SEB and CCI and call for further detailed studies about the distinction of these two coral diseases.

Coral bleaching represents the most serious threat to contemporary coral reefs. In response, focus is being laid on understanding the cellular processes involved in the response of corals to the environmental stresses and the molecular mechanisms that determine the bleaching patterns. In the present study, a component of the cellular stress response such as the expression of the heat shock proteins (Hsps) was analyzed following the coral bleaching event which occurred in the central Red Sea (Saudi Arabia) in 2015. During this event, corals of different species, growth forms and sites showed variable bleaching susceptibility. In particular, we investigated the expression of Hsp70, Hsp60 and Hsp32 in both healthy and bleached colonies belonging to four different coral species (Goniopora lobata, Porites lobata, Seriatopora hystrix and Stylophora pistillata), in order to explore the intra- and inter-specific modulation of these biomarkers as well as the existence of spatial patterns of Hsp expression. In healthy colonies, the level of all the biomarkers was significantly different among the different species, although within each species it remained similar regardless of the distance from the shore. All the coral species showed a significant modulation of the Hsp expression in response to bleaching, whose typology and amplitude were species-specific. In all the species, Hsp70 and Hsp60 showed a coordinated dual expression, which, in response to bleaching resulted in an up-regulation in G. lobata and P. lobata and in a down-regulation in S. hystrix and S. pistillata. Hsp32 was up-regulated in all four species following bleaching, indicative of elevated oxidative stress. Overall, the protein expression profiles of each species contribute to assess the role of Hsps in regulating the susceptibility to thermal stresses of the various coral taxa of the Red Sea.

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

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

The colonial stony coral genus Madracis is cosmopolitan, lives in shallow and deep water habitats, and includes zooxan-thellate, 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 inte-grated 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 resem-blance, must be investigated in further studies. Overall, the presence of cryptic Madracis species in the Easter and Central Atlantic Ocean remains to be evaluated

Fire corals of the hydrocoral genus Millepora provide an important ecological role as framework builders of coral reefs in the Indo-Pacific and the Atlantic. Recent works have demonstrated the incongruence between molecular data and the traditional taxonomy of Millepora spp. based on overall skeleton growth form and pores. In an attempt to establish a reliable and standardized approach for defining species boundaries in Millepora, we focused on those from the Red Sea. In this region, three species are currently recognized: the fan-shaped branching M. dichotoma, the blade-like M. platyphylla, and the massive/encrusting M. exaesa. A total of 412 colonies were collected from six localities. Two mitochondrial marker genes (COI and 16S rDNA) were sequenced to obtain phylogeny reconstructions and haplotype networks. Eight morphological traits of pores and the nematocysts of both polyp and eumedusoid stages were measured to determine whether significant morphological differences occur among the three species. Both markers clearly resolved M. dichotoma, M. platyphylla, and M. exaesa as distinct, monophyletic lineages in the Red Sea. Nevertheless, they also revealed deep genetic breaks with Southwestern Indian Ocean populations of the three species. In the Red Sea, the three species were further distinguished based on their pore and nematocyst features. A discriminant analysis revealed dactylopore density, number of dactylopores per gastropore, dactylopore distance, and gastropore diameter as the most informative discriminative characters. The heteronemes, the large and small stenoteles of polyps, and the distribution of mastigophores of eumedusoids also showed significant interspecific differences. An integrated morpho-molecular approach proved to be decisive in defining species boundaries of Millepora supported by a combination of pore and nematocyst characters, which may be phylogenetically informative.

The Zancleidae is a hydrozoan family that currently comprises three genera and 42 nominal species. The validity of numerous taxa in this family still needs to be assessed with integrative analyses and complete life cycle descriptions. The vast majority of its species live symbiotically with other organisms, among which cheilostomate bryozoans are the most common hosts. These bryozoan-associated zancleids are host-specific and encompass all species of the genera Halocoryne and Zanclella, as well as several species in the genus Zanclea. Zancleids show variable morphologies, including highly reduced polyps and medusae. Their phylogenetic history is uncertain due to the often intergrading morphologies and the shortage of molecular data. In the present study, two species of Zanclea from the Indian Ocean and the Red Sea are analysed, using morphological and DNA-based approaches. Morphological analyses of the polyp and medusa stages show that, despite a general resemblance with each other and with Zanclella diabolica, the two species differ in some characters, and this is supported also by molecular investigations. The DNA analyses show that the two species are monophyletic and closely related, but divergent from other Zanclea lineages. This newly recovered clade may correspond with the genus Zanclella or with a cryptic genus. The lack of both morphological and molecular data for several zancleid species did not allow to address this issue. Additionally, the singleand multi-locus phylogeny reconstructions reveal that both the family Zancleidae and the genus Zanclea are polyphyletic taxa, since they are composed of at least three divergent lineages. Most zancleid species have polyps and medusae similar to other closely related taxa, and this conserved general morphology poses a challenge in the delimitation of species, genera and even families in this group. Consequently, further conjunct morphological and molecular efforts are strongly needed to clarify the diversity and e

In spite of growing evidence that climate change may dramatically affect networks of interacting species, whether-and to what extent-ecological interactions can mediate species’ responses to disturbances is an open question. Here we show how a largely overseen association such as that between hydrozoans and scleractinian corals could be possibly associated with a reduction in coral susceptibility to ever-increasing predator and disease outbreaks. We examined 2455 scleractinian colonies (from both Maldivian and the Saudi Arabian coral reefs) searching for non-randompatterns in the occurrence of hydrozoans on corals showing signs of different health conditions (i.e. bleaching, algal overgrowth, corallivory and different coral diseases). We show that, after accounting for geographical, ecological and co-evolutionary factors, signs of disease and corallivory are significantly lower in coral colonies hosting hydrozoans than in hydrozoan-free ones. This finding has important implications for our understanding of the ecology of coral reefs, and for their conservation in the current scenario of global change, because it suggests that symbiotic hydrozoans may play an active role in protecting their scleractinian hosts from stresses induced by warming water temperatures.

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.

The family Sphaerocorynidae includes two valid genera and five species, most of which have a confusing taxonomic history. Here, a new genus and species, Astrocoryne cabela, gen. et sp. nov., is described from the Maldives and the Red Sea, based on both morphological and molecular evidence. Astrocoryne cabela has an apomorphy represented by the type of tentacles, here named 'dicapitate', and consisting of capitate tentacles with a proximal capitulum-like cluster of nematocysts. Molecular analyses confirmed the monophyly of this species, as well as its belonging to the Sphaerocorynidae, together with Sphaerocoryne spp. and Heterocoryne caribbensis Wedler & Larson, 1986, for which we present molecular data for the first time. Moreover, the high divergence of A. cabela from other species of the family justifies the establishment of a new genus. Interestingly, specimens from the Maldives and the Red Sea showed marked morphological variation in the polyp stage, although only a slight genetic divergence was detected. This study highlights that a comprehensive morpho-molecular assessment of Sphaerocorynidae is strongly needed in order to clarify the taxonomic issues and the diversity of this taxon.

Scleractinian corals ascribed to the family Agariciidae represent an important component of Red Sea coral reef fauna, though little genetic data are currently available for this group, and existing information shows polyphyly in the examined mesophotic taxa from the Pacific Ocean. In this work, we provide a first genetic survey of Agariciidae from the Saudi Arabian Red Sea, based on a collection of shallow-water material (<30 m) from the Gulf of Aqaba to the Farasan Islands. Two molecular markers were sequenced to infer morphospecies monophyly and relationships, the intergenic region between COI and 16S rRNA from mitochondrial DNA and the ribosomal ITS1 region from nuclear DNA. A total of 20 morphospecies were identified based on classical macromorphological characters. Six, namely Gardineroseris planulata, Pavona maldivensis, Pavona clavus, Pavona decussata, Leptoseris fragilis, and Leptoseris yabei, were resolved with both DNA loci. The molecular boundaries among the remaining 14 species remain unclear. Our results further confirm that the morphology-based taxonomy of most agariciid species is in disagreement with genetics. In order to disentangle the systematics of these taxa, the inclusion of more sampling locations, additional variable loci, and a micromophological approach are likely needed. Our genetic data represent a first step towards the comparison of biodiversity and connectivity between the Red Sea and the rest of the Indo-Pacific.

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.

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 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

Reticulate evolution, introgressive hybridisation, and phenotypic plasticity have been documented in scleractinian corals and have challenged our ability to interpret speciation processes. Stylophora is a key model system in coral biology and physiology, but genetic analyses have revealed that cryptic lineages concealed by morphological stasis exist in the Stylophora pistillata species complex. The Red Sea represents a hotspot for Stylophora biodiversity with six morphospecies described, two of which are regionally endemic. We investigated Stylophora species boundaries from the Red Sea and the associated Symbiodinium by sequencing seven DNA loci. Stylophora morphospecies from the Red Sea were not resolved based on mitochondrial phylogenies and showed nuclear allele sharing. Low genetic differentiation, weak isolation, and strong gene flow were found among morphospecies although no signals of genetic recombination were evident among them. Stylophora mamillata harboured Symbiodinium clade C whereas the other two Stylophora morphospecies hosted either Symbiodinium clade A or C. These evolutionary patterns suggest that either gene exchange occurs through reticulate evolution or that multiple ecomorphs of a phenotypically plastic species occur in the Red Sea. The recent origin of the lineage leading to the Red Sea Stylophora may indicate an ongoing speciation driven by environmental changes and incomplete lineage sorting.

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

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.

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.

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 (tRNAMet 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 2220bp, of which the cox3-cox2 region may correspond to the putative control region.

Scleractinian reef corals have recently been acknowledged as the most numerous host group found in association with hydroids belonging to the Zanclea genus. However, knowledge of the molecular phylogenetic relationships among Zanclea species associated with scleractinians is just beginning. This study, using the nuclear 28S rDNA region and the fastevolving mitochondrial 16S rRNA and COI genes, provides the most comprehensive phylogenetic reconstruction of the genus Zanclea with a particular focus on the genetic diversity among Zanclea specimens associated with 13 scleractinian genera. The monophyly of Zanclea associated with scleractinians was strongly supported in all nuclear and mitochondrial phylogenetic reconstructions. Furthermore, a combined mitochondrial 16S and COI phylogenetic tree revealed a multitude of hidden molecular lineages within this group (Clades I, II, III, V, VI, VII, and VIII), suggesting the existence of both host-generalist and genus-specific lineages of Zanclea associated with scleractinians. In addition to Z. gallii living in association with the genus Acropora, we discovered four well-supported lineages (Clades I, II, III, and VII), each one forming a strict association with a single scleractinian genus, including sequences of Zanclea associated with Montipora from two geographically separated areas (Maldives and Taiwan). Two host-generalist Zanclea lineages were also observed, and one of them was formed by Zanclea specimens symbiotic with seven scleractinian genera (Clade VIII). We also found that the COI gene allows the recognition of separated hidden lineages in agreement with the commonly recommended mitochondrial 16S as a DNA barcoding gene for Hydrozoa and shows reasonable potential for phylogenetic and evolutionary analyses in the genus Zanclea. Finally, as no DNA sequences are available for the majority of the nominal Zanclea species known, we note that they will be necessary to elucidate the diversity of the Zanclea-scleractinian

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 mon

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.

The monospecific scleractinian coral genus Sclerophyllia Klunzinger, 1879 was originally described from Al-Qusayr (Egypt) in the Red Sea based on a series of solitary specimens. Thenceforth, it has been considered a junior synonym of Symphyllia and Cynarina based on corallum macromorphology. In this study, several specimens of Sclerophyllia margariticola were collected on the coasts of Saudi Arabia in the northern and central Red Sea. Four molecular markers were sequenced, COI and the intergenic spacer between COI and l-rRNA from mitochondrial DNA and Histone H3 and ribosomal ITS2 from nuclear DNA. Phylogenetic trees and haplotype network analyses show that S. margariticola belongs to the family Lobophylliidae and that it is closely related to Acanthastrea maxima, an uncommon species from waters around the Arabian peninsula (the Gulf of Aden, Arabian Sea, Gulf of Oman and Persian Gulf). Sclerophyllia margariticola and A. maxima share several macro- and micromorphological characters, such as the presence of free septa, high elliptical septal teeth perpendicular to the septal margin, irregular lobate tips, very wide tooth spacing, a very strong granulation with granules scattered all along the septal sides, and a palisade interarea structure, and their micromorphology differs substantially from that of Acanthastrea echinata, the type species of Acanthastrea. Therefore, we formally resurrect Sclerophyllia, provide a revised diagnosis for the genus, and move A. maxima into Sclerophyllia.