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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.

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.

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.

For millennia, coastal and marine ecosystems have adapted and flourished in the Red Sea’s unique environment. Surrounded by deserts on all sides, the Red Sea is subjected to high dust inputs and receives very little freshwater input, and so harbors a high salinity. Coral reefs, seagrass meadows, and mangroves flourish in this environment and provide socio-economic and environmental benefits to the bordering coastlines and countries. Interestingly, while coral reef ecosystems are currently experiencing rapid decline on a global scale, those in the Red Sea appear to be in relatively better shape. That said, they are certainly not immune to the stressors that cause degradation, such as increasing ocean temperature, acidification and pollution. In many regions, ecosystems are already severely deteriorating and are further threatened by increasing population pressure and large coastal development projects. Degradation of these marine habitats will lead to environmental costs, as well as significant economic losses. Therefore, it will result in a missed opportunity for the bordering countries to develop a sustainable blue economy and integrate innovative nature-based solutions. Recognizing that securing the Red Sea ecosystems’ future must occur in synergy with continued social and economic growth, we developed an action plan for the conservation, restoration, and growth of marine environments of the Red Sea. We then investigated the level of resources for financial and economic investment that may incentivize these activities. This study presents a set of commercially viable financial investment strategies, ecological innovations, and sustainable development opportunities, which can, if implemented strategically, help ensure long-term economic benefits while promoting environmental conservation. We make a case for investing in blue natural capital and propose a strategic development model that relies on maintaining the health of natural ecosystems to s

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.

Corals and the reef ecosystems that they support are in global decline due to increasing anthropogenic pressures such as climate change1. However, effective reef conservation strategies are hampered by a limited mechanistic understanding of coral biology and the functional roles of the diverse microbial communities that underpin coral health2,3. Here, we present an integrated genomic characterization of the coral species Porites lutea and its microbial partners. High-quality genomes were recovered from P. lutea, as well as a metagenome-assembled Cladocopium C15 (the dinoflagellate symbiont) and 52 bacterial and archaeal populations. Comparative genomic analysis revealed that many of the bacterial and archaeal genomes encode motifs that may be involved in maintaining association with the coral host and in supplying fixed carbon, B-vitamins and amino acids to their eukaryotic partners. Furthermore, mechanisms for ammonia, urea, nitrate, dimethylsulfoniopropionate and taurine transformation were identified that interlink members of the holobiont and may be important for nutrient acquisition and retention in oligotrophic waters. Our findings demonstrate the critical and diverse roles that microorganisms play within the coral holobiont and underscore the need to consider all of the components of the holobiont if we are to effectively inform reef conservation strategies.

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