The Mitogenomic Landscape of Hexacorallia Corals: Insight into Their Slow Evolution.

The utility of the mitochondrial genomes (mitogenomes) in analyzing the evolutionary history of animals has been proven. Five deep-sea corals (Bathypathes sp.1, Bathypathes sp.2, Schizopathidae 1, Trissopathes sp., and Leiopathes sp.) were collected in the South China Sea (SCS). Initially, the structures and collinearity of the five deep-sea coral mitogenomes were analyzed. The gene arrangements in the five deep-sea coral mitogenomes were similar to those in the order Antipatharia, which evidenced their conservation throughout evolutionary history. Additionally, to elucidate the slow evolutionary rates in Hexacorallia mitogenomes, we conducted comprehensive analyses, including examining phylogenetic relationships, performing average nucleotide identity (ANI) analysis, and assessing GC-skew dissimilarity combining five deep-sea coral mitogenomes and 522 reference Hexacorallia mitogenomes. Phylogenetic analysis using 13 conserved proteins revealed that species clustered together at the order level, and they exhibited interspersed distributions at the family level. The ANI results revealed that species had significant similarities (identity > 85%) within the same order, while species from different orders showed notable differences (identity < 80%). The investigation of the Hexacorallia mitogenomes also highlighted that the GC-skew dissimilarity was highly significant at the order level, but not as pronounced at the family level. These results might be attributed to the slow evolution rate of Hexacorallia mitogenomes and provide evidence of mitogenomic diversity. Furthermore, divergence time analysis revealed older divergence times assessed via mitogenomes compared with nuclear data, shedding light on significant evolutionary events shaping distinct orders within Hexacorallia corals. Those findings provide new insights into understanding the slow evolutionary rates of deep-sea corals in all lineages of Hexacorallia using their mitogenomes.

Diversity patterns of the South African azooxanthellate scleractinians (Cnidaria: Anthozoa), with considerations of environmental correlates.

Azooxanthellate scleractinian corals, a group of species that lack a symbiotic relationship with dinoflagellates, are influenced by environmental variables at various scales. As the global commitment to sustainably manage ocean ecosystems and resources rises, there is a growing need to describe biodiversity trends in previously unsampled areas. Benthic invertebrate research in South Africa is a developing field, and many taxa in deep water environments remain inadequately characterized. Recently, the South African azooxanthellate scleractinian fauna was taxonomically reviewed, but their distributional correlations with physical parameters have not been studied. Here we aim to understand the biodiversity gradients of the South African azooxanthellate coral fauna by analysing the environmental correlates of museum samples. The associated coordinate data were georeferenced and depth obtained from a national bathymetric dataset, prior to undertaking a multivariate analysis. This analysis encompassed several steps, including the grouping of the longitude and depth data (environmental data), identifying families characteristic of the group variability, and examining the correlation of the associated data with the biological data. Additionally, the analysis involved quantifying diversity patterns along the environmental gradients. Overall, our results confirmed two longitudinal groups (eastern margin [Group A] vs southern and western margin [Group B]) and 11 depth categories represented within two bathymetric zones (shallow [50-200 m] and deep [300-1000 m]). Caryophylliids, flabellids, and dendrophylliids contributed the most towards distinguishing longitudinal and depth gradients. Both abiotic variable (longitudinal and depth) partially explained coral distribution patterns, with depth being highly correlated to the species variation observed. Data limitations within our data set resulted to unexplained variance, however, despite these limitations, the study demonstrates that historical museum samples provide a valuable data source that can fill research sampling gaps and help improve the understanding of biodiversity patterns of the coral fauna in under sampled marine ecosystems.

Cryptic diversity of shallow and mesophotic Stephanocoenia intersepta corals across Florida Keys National Marine Sanctuary.

Population genetic analyses can provide useful data on species' regional connectivity and diversity which can inform conservation and restoration efforts. In this study, we quantified the genetic connectivity and diversity of Stephanocoenia intersepta corals from shallow (<30 m) to mesophotic (30-45 m) depths across Florida Keys National Marine Sanctuary. We generated single nucleotide polymorphism (SNP) markers to identify genetic structuring of shallow and mesophotic S. intersepta corals. We uncovered four distinct, cryptic genetic lineages with varying levels of depth-specificity. Shallow-specific lineages exhibited lower heterozygosity and higher inbreeding relative to depth-generalist lineages found across both shallow and mesophotic reefs. Estimation of recent genetic migration rates demonstrated that mesophotic sites are more prolific sources than shallow sites, particularly in the Lower Keys and Upper Keys. Additionally, we compared endosymbiotic Symbiodiniaceae among sampled S. intersepta using the ITS2 region and SYMPORTAL analysis framework, identifying symbionts from the genera Symbiodinium, Breviolum, and Cladocopium. Symbiodiniaceae varied significantly across depth and location and exhibited significant, but weak correlation with host lineage and genotype. Together, these data demonstrate that despite population genetic structuring across depth, some mesophotic populations may provide refuge for shallow populations moving forward and remain important contributors to the overall genetic diversity of this species throughout the region. This study highlights the importance of including mesophotic as well as shallow corals in population genetic assessments and informs future science-based management, conservation, and restoration efforts within Florida Keys National Marine Sanctuary.

Mucus carbohydrate composition correlates with scleractinian coral phylogeny.

The mucus surface layer serves vital functions for scleractinian corals and consists mainly of carbohydrates. Its carbohydrate composition has been suggested to be influenced by environmental conditions (e.g., temperature, nutrients) and microbial pressures (e.g., microbial degradation, microbial coral symbionts), yet to what extend the coral mucus composition is determined by phylogeny remains to be tested. To investigate the variation of mucus carbohydrate compositions among coral species, we analyzed the composition of mucosal carbohydrate building blocks (i.e., monosaccharides) for five species of scleractinian corals, supplemented with previously reported data, to discern overall patterns using cluster analysis. Monosaccharide composition from a total of 23 species (belonging to 14 genera and 11 families) revealed significant differences between two phylogenetic clades that diverged early in the evolutionary history of scleractinian corals (i.e., complex and robust; p = 0.001, R2 = 0.20), mainly driven by the absence of arabinose in the robust clade. Despite considerable differences in environmental conditions and sample analysis protocols applied, coral phylogeny significantly correlated with monosaccharide composition (Mantel test: p < 0.001, R2 = 0.70). These results suggest that coral mucus carbohydrates display phylogenetic dependence and support their essential role in the functioning of corals.

300 million years apart: the extreme case of macromorphological skeletal convergence between deltocyathids and a turbinoliid coral (Anthozoa, Scleractinia).

The integration of morphological and molecular lines of evidence has enabled the family Deltocyathidae to be erected to accommodate Deltocyathus species that were previously ascribed to the family Caryophylliidae. However, although displaying the same morphological characteristics as other species of Deltocyathus , molecular data suggested that D. magnificus was phylogenetically distant from Deltocyathidae, falling within the family Turbinoliidae instead. To elucidate the enigmatic evolutionary history of this species and skeletal microstructural features, the phylogenetic relationships of Deltocyathidae and Turbinoliidae were investigated using nuclear ultraconserved and exon loci and complete mitochondrial genomes. Both nuclear and mitochondrial phylogenomic reconstructions confirmed the position of D. magnificus within turbinolids. Furthermore, a novel mitochondrial gene order was uncovered for Deltocyathidae species. This gene order was not present in Turbinoliidae or in D. magnificus that both have the scleractinian canonical gene order, further indicating the taxonomic utility of mitochondrial gene order. D. magnificus is therefore formally moved to the family Turbinoliidae and accommodated in a new genus (Dennantotrochus Kitahara, Vaga & Stolarski, gen. nov.). Surprisingly, turbinolids and deltocyathids do not differ in microstructural organisation of the skeleton that consists of densely packed, individualised rapid accretion deposits and thickening deposits composed of fibres perpendicular to the skeleton surface. Therefore, although both families are clearly evolutionarily divergent, macromorphological features indicate a case of skeletal convergence while these may still share conservative biomineralisation mechanisms. ZooBank: urn:lsid:zoobank.org:pub:5F1C0E25-3CC6-4D1F-B1F0-CD9D0014678E.

Molecular diversity of black corals from the Saudi Arabian Red Sea: a first assessment.

Black corals occur as part of benthic assemblages from shallow to deep waters in all oceans. Despite the importance in many benthic ecosystems, where these act as biodiversity aggregators, antipatharians remain poorly studied, with 75% of the known species occurring below recreational SCUBA diving depth limits. Currently, information regarding the diversity and evolutionary history is limited, with most studies focusing on Hawaii and the South Pacific Ocean. Other regions of the world have received less attention, such as the Red Sea, where only two black coral families and four genera have been recorded. We provide the first analysis of the molecular diversity of black corals in the eastern Gulf of Aqaba and the northern and central Saudi Arabian Red Sea, based on a dataset of 161 antipatharian colonies collected down to 627 m deep. Based on specimen morphology, we ascribed our material to 11 genera belonging to 4 of the 7 known Antipatharia families, i.e. Antipathidae, Aphanipathidae, Myriopathidae and Schizopathidae. The genus level phylogeny of three intergenic mitochondrial regions, the trnW-IGR-nad2 (IgrW ), nad5-IGR-nad1 (IgrN ) and cox3-IGR-cox1 was reconstructed including previously published material. Overall, we recovered six molecular clades that included exclusively Red Sea sequences, with the highest diversity occurring at mesophotic depths. This study highlights that diversity of black corals in the Red Sea is much higher than previously known, with seven new generic records, suggesting that this basin may be a hotspot for antipatharian diversity as is known for other taxa. Our results recovered unresolved relationships within the order at the familial and generic levels. This emphasises the urgent need for an integration of genomic-wide data with a re-examination of informative morphological features necessary to revise the systematics of the order at all taxonomic levels.

Unravelling the relationships among Madrepora Linnaeus, 1758, Oculina Lamark, 1816 and Cladocora Ehrenberg, 1834 (Cnidaria: Anthozoa: Scleractinia).

Despite the widespread use of integrative taxonomic approaches, many scleractinian coral genera and species remain grouped in polyphyletic families, classified as incertae sedis or simply understudied. Oculinidae Gray, 1847 represents a family for which many taxonomic questions remain unresolved, particularly those related to some of the current genera, such as Oculina Lamark, 1816 or recently removed genera, including Cladocora Ehrenberg, 1834 and Madrepora Linnaeus, 1758. Cladocora is currently assigned to the family Cladocoridae Milne Edwards & Haime, 1857 and a new family, Bathyporidae Kitahara, Capel, Zilberberg & Cairns, 2024, was recently raised to accommodate Madrepora . However, the name Bathyporidae is not valid because this was not formed on the basis of a type genus name. To resolve taxonomic questions related to these three genera, the evolutionary relationships are explored through phylogenetic analyses of 18 molecular markers. The results of these analyses support a close relationship between the species Oculina patagonica and Cladocora caespitosa , indicating that these may belong to the same family (and possibly genus), and highlighting the need for detailed revisions of Oculina and Cladocora . By contrast, a distant relationship is found between these two species and Madrepora oculata , with the overall evidence supporting the placement of Madrepora in the resurrected family Madreporidae Ehrenberg, 1834. This study advances our knowledge of coral systematics and highlights the need for a comprehensive review of the genera Oculina , Cladocora and Madrepora .

Biodiversity patterns of the coral reef cryptobiota around the Arabian Peninsula.

The Arabian Peninsula accounts for approximately 6% of the world's coral reefs. Some thrive in extreme environments of temperature and salinity. Using 51 Autonomous Reef Monitoring Structure (ARMS), a standardized non-destructive monitoring device, we investigated the spatial patterns of coral reef cryptobenthic diversity in four ecoregions around the Arabian Peninsula and analyzed how geographical and/or environmental drivers shape those patterns. The mitochondrial cytochrome c oxidase subunit I (COI) gene was used to identify Amplicon Sequence Variants and assign taxonomy of the cryptobenthic organisms collected from the sessile and mobile fractions of each ARMS. Cryptobenthic communities sampled from the two ecoregions in the Red Sea showed to be more diverse than those inhabiting the Arabian (Persian) Gulf and the Gulf of Oman. Geographic distance revealed a stronger relationship with beta diversity in the Mantel partial correlation than environmental distance. However, the two mobile fractions (106-500 µm and 500-2000 µm) also had a significant correlation between environmental distance and beta diversity. In our study, dispersal limitations explained the beta diversity patterns in the selected reefs, supporting the neutral theory of ecology. Still, increasing differences in environmental variables (environmental filtering) also had an effect on the distribution patterns of assemblages inhabiting reefs within short geographic distances. The influence of geographical distance in the cryptofauna assemblages makes these relevant, yet usually ignored, communities in reef functioning vulnerable to large scale coastal development and should be considered in ecosystem management of such projects.

SEA PENS (CNIDARIA: OCTOCORALLIA: PENNATULOIDEA) FROM THE MAR DEL PLATA SUBMARINE CANYON AND OUTSKIRTS.

During this work, over 200 specimens of sea pens (Anthozoa: Octocorallia: Pennatuloidea), collected in the Mar del Plata submarine canyon and outskirts (38S, up to 3500 m depth) during 2012 and 2013 were analyzed. Of the total, about 160 were identified in seven species of Anthoptilum, Distichoptilum, Kophobelemnon, Pseudumbellula, Solumbellula and Umbellula. Three of those represent new records for the southwestern Atlantic Ocean off Argentina.

Mediterranean Sea shelters for the gold coral Savalia savaglia (Bertoloni, 1819): An assessment of potential distribution of a rare parasitic species.

Savalia savaglia is an ecosystem engineer listed as Near-Threatened by the IUCN, even though effective management and proper monitoring efforts to assess its distribution is still lacking. The record of large, long-established colonies can indicate the occurrence of areas with limited human local pressure. These areas may be considered as proxies for the creation of baselines of reference useful to design restoration strategies. The aim of this work was to update the distribution of S. savaglia Mediterranean populations to develop an Ecological Niche Model, highlighting potential areas for future monitoring programs. Occurrence data were collected and harmonized into a single dataset using the scientific literature and validated observations to feed a presence-only MaxEnt model, obtaining a basin-level potential distribution of the species. The results of our study can support decision-makers in marine spatial planning measures including the preservation of mesophotic environments and prioritizing areas for conservation.

Effects of low pH on the coral reef cryptic invertebrate communities near CO2 vents in Papua New Guinea.

Small cryptic invertebrates (the cryptofauna) are extremely abundant, ecologically important, and species rich on coral reefs. Ongoing ocean acidification is likely to have both direct effects on the biology of these organisms, as well as indirect effects through cascading impacts on their habitats and trophic relationships. Naturally acidified habitats have been important model systems for studying these complex interactions because entire communities that are adapted to these environmental conditions can be analyzed. However, few studies have examined the cryptofauna because they are difficult to census quantitatively in topographically complex habitats and are challenging to identify. We addressed these challenges by using Autonomous Reef Monitoring Structures (ARMS) for sampling reef-dwelling invertebrates >2 mm in size and by using DNA barcoding for taxonomic identifications. The study took place in Papua New Guinea at two reef localities, each with three sites at varying distances from carbon dioxide seeps, thereby sampling across a natural gradient in acidification. We observed sharp overall declines in both the abundance (34-56%) and diversity (42-45%) of organisms in ARMS under the lowest pH conditions sampled (7.64-7.75). However, the overall abundance of gastropods increased slightly in lower pH conditions, and crustacean and gastropod families exhibited varying patterns. There was also variability in response between the two localities, despite their close proximity, as one control pH site displayed unusually low diversity and abundances for all invertebrate groups. The data illustrate the complexity of responses of the reef fauna to pH conditions, and the role of additional factors that influence the diversity and abundance of cryptic reef invertebrates.

First insights into coral recruit and juvenile abundances at remote Aldabra Atoll, Seychelles.

Coral recruitment and successive growth are essential for post-disturbance reef recovery. As coral recruit and juvenile abundances vary across locations and under different environmental regimes, their assessment at remote, undisturbed reefs improves our understanding of early life stage dynamics of corals. Here, we first explored changes in coral juvenile abundance across three locations (lagoon, seaward west and east) at remote Aldabra Atoll (Seychelles) between 2015 and 2019, which spanned the 2015/16 global coral bleaching event. Secondly, we measured variation in coral recruit abundance on settlement tiles from two sites (lagoon, seaward reef) during August 2018-August 2019. Juvenile abundance decreased from 14.1 ± 1.2 to 7.4 ± 0.5 colonies m-2 (mean ± SE) during 2015-2016 and increased to 22.4 ± 1.2 colonies m-2 during 2016-2019. Whilst juvenile abundance increased two- to three-fold at the lagoonal and seaward western sites during 2016-2018 (from 7.7-8.3 to 17.3-24.7 colonies m-2), increases at the seaward eastern sites occurred later (2018-2019; from 5.8-6.9 to 16.6-24.1 colonies m-2). The composition of coral recruits on settlement tiles was dominated by Pocilloporidae (64-92% of all recruits), and recruit abundance was 7- to 47-fold higher inside than outside the lagoon. Recruit abundance was highest in October-December 2018 (2164 ± 453 recruits m-2) and lowest in June-August 2019 (240 ± 98 recruits m-2). As Acroporid recruit abundance corresponded to this trend, the results suggest that broadcast spawning occurred during October-December, when water temperature increased from 26 to 29°C. This study provides the first published record on coral recruit abundance in the Seychelles Outer Islands, indicates a rapid (2-3 years) increase of juvenile corals following a bleaching event, and provides crucial baseline data for future research on reef resilience and connectivity within the region.

A New Species of Sea Whip Gorgonian-Associated Zoantharian (Cnidaria: Anthozoa: Hexacorallia: Parazoanthidae) from the Ryukyu Islands, Japan, with Subgeneric Subdivision of Genus Umimayanthus.

Symbioses between invertebrates are common in the ocean although usually the diversity and specificity of their interactions are not well understood. Parazoanthidae (Cnidaria: Anthozoa: Zoantharia) is one of the most diverse zoantharian families in terms of numbers of genera and species. Species in this family are commonly associated with various other invertebrates that they utilize as their substrate. Previous studies have re-organized the taxonomy of Parazoanthidae and revealed a strong specificity between many parazoanthid species and genera and their substrates. However, our understanding of the species diversity of Parazoanthidae is far from complete, as parazoanthids are often overlooked in sampling surveys. In this study, we establish three subgenera under the genus Umimayanthus Montenegro, Sinniger, and Reimer, 2015; the nominotypical Umimayanthus, Paraumimayanthus subgen nov., and Gorgoniazoanthus subgen. nov., based on the finding of a new species, Umimayanthus (Gorgoniazoanthus) kanabou sp. nov., associated with the sea-whip gorgonian Ellisella sp. from approximately 30 m depth in shallow mesophotic coral reef communities in Oura Bay on Okinawajima Island and in Oshima Strait near Amami-Oshima Island, in the Ryukyu Islands, southern Japan. We additionally report on gastropods and crustaceans observed in association with U. kanabou, and these species are thought to potentially prey upon the zoantharians or on gorgonian polyps. Umimayanthus kanabou is phylogenetically closely related to congeneric sponge-associated Umimayanthus spp., further supporting the recent hypothesis that substrate preferences may change during the evolutionary history of zoantharians.

Early development and coloniality in Oligophylloides from the Devonian of Morocco-Are Heterocorallia Palaeozoic octocorals?

Heterocorals represent an enigmatic group of Palaeozoic corals, known from relatively short time intervals in the Devonian and Carboniferous periods. The major differences between Heterocorallia and other Palaeozoic corals are the lack of an external theca (epitheca), lack of calices and the presence of dichotomously dividing septa-like structures. Heterocoral skeleton was presumably externally covered by the soft tissue and each branch of their skeleton has, until now, been regarded as a corallite-a skeleton of a single polyp. We investigated upper Famennian Oligophylloides from Morocco, focussing on branching processes, wall structure, previously poorly known initial growth stages and the growing tip, described here for the first time. We demonstrate that Oligophylloides shows a unique colony development not known in any group of anthozoans possessing a septate-like architecture and suggest that the previously postulated homology between true septa in hexa- and rugose corals on one hand, and Oligophylloides on the other, must be rejected. Based on the skeleton structure and branching patterns, we postulate, contrary to former ideas, that the stem and branches of heterocorals represent the skeleton of a multi-polyp colonial coral, similar to many extant octocorals. We found numerous potential homologies with octocoral skeletons (notably the Keratoisidinae within the Isididae) and, as a result, we propose the inclusion of the order Heterocorallia within the subclass Octocorallia. This suggestion requires, however, further research on the other taxa of heterocorals. We also propose some changes to the morphological terminology for the Heterocorallia.

Genetic basis of stony coral biomineralization: History, trends and future prospects.

Despite their simple body plan, stony corals (order Scleractinia, phylum Cnidaria) can produce massive and complex exoskeletal structures in shallow, tropical and subtropical regions of Earth's oceans. The species-specific macromorphologies of their aragonite skeletons suggest a highly coordinated biomineralization process that is rooted in their genomes, and which has persisted across major climatic shifts over the past 400 + million years. The mechanisms by which stony corals produce their skeletons has been the subject of interest for at least the last 160 years, and the pace of understanding the process has increased dramatically in the past decade since the sequencing of the first coral genome in 2011. In this review, we detail what is known to date about the genetic basis of the stony coral biomineralization process, with a focus on advances in the last several years as well as ways that physical and chemical tools can be combined with genetics, and then propose next steps forward for the coming decade.

Microbial dysbiosis reflects disease resistance in diverse coral species.

Disease outbreaks have caused significant declines of keystone coral species. While forecasting disease outbreaks based on environmental factors has progressed, we still lack a comparative understanding of susceptibility among coral species that would help predict disease impacts on coral communities. The present study compared the phenotypic and microbial responses of seven Caribbean coral species with diverse life-history strategies after exposure to white plague disease. Disease incidence and lesion progression rates were evaluated over a seven-day exposure. Coral microbiomes were sampled after lesion appearance or at the end of the experiment if no disease signs appeared. A spectrum of disease susceptibility was observed among the coral species that corresponded to microbial dysbiosis. This dysbiosis promotes greater disease susceptiblity in coral perhaps through different tolerant thresholds for change in the microbiome. The different disease susceptibility can affect coral's ecological function and ultimately shape reef ecosystems.

Revised nomenclature of the sea pen genus Balticina Gray, 1870 (= Halipteris Kölliker, 1870) (Anthozoa: Octocorallia).

The pennatulacean genus Balticina has had a long and confusing taxonomic history, with serious nomenclatural problems that remain unresolved. Owing to disagreements about authorships and dates of publication, the names Pavonaria, Norticina and Halipteris have all been used as valid in place of Balticina, or else been regarded as its junior synonyms, even simultaneously. In this paper, after an extensive literature review, we determine the authorships and dates for all the taxa involved in accordance with the provisions of the International Code of Zoological Nomenclature and establish Balticina Gray, 1870 (=Halipteris) and Balticinidae Balss, 1910 (=Halipteridae) as the valid genus and family names, respectively, for this group of sea pens. We also propose the replacement name Rakollikeria for the preoccupied genus name Pavonaria Kölliker, 1870 (Balticinidae) (nec Schweigger, 1819 - Funiculinidae).

Description of two new genera and two new species of antipatharian corals in the family Aphanipathidae (Cnidaria: Anthozoa: Antipatharia).

Two new genera and two new species of black corals are recognized in the family Aphanipathidae. The new genus Anozopathes, with the species A. hawaiiensis sp. nov. and A. palauensis, sp. nov. is characterized by a sparsely and irregularly branched corallum with relatively long branches which can be straight, curved or crooked. The genus Aphanostichopathes, with the type species Cirripathes paucispina Brook, is characterized by an unbranched corallum with a long, curved stem with loose distal coils. Mitochondrial DNA data (nad5-IGR-nad1 for Anozopathes and cox3-cox1 for Aphanostichopathes) indicate that both taxa are related to the genera Aphanipathes, Phanopathes and Acanthopathes in the family Aphanipathidae, and morphologically they both share the characteristic of having spines with distinct conical tubercles. The two new species of Anozopathes are separated primarily by differences in colony growth form and in the size and shape of the skeletal spines. Species of Aphanostichopathes are separated primarily by differences in the size and shape of the spines and by size and density of the tubercles on the surface of the spines.

A brief review of the stony coral (Cnidaria: Anthozoa: Scleractinia) papers published by Zootaxa in its first 20 years.

In the 20 year history of Zootaxa, thirty-two papers have been published having Scleractinia as its primary focus. Twenty-four of the 32 scleractinian papers deal with Recent taxa, most of which belonging to shallow-water, reef corals. The 8 publications dealing with fossil Scleractinia include 3 monographic works, three papers discussing nomenclatural issues of individual taxa, and 2 papers deal with various aspects of select genera.