BACKGROUND: Coral-associated microbiomes vary greatly between colonies and localities with functional consequences on the host. However, the full extent of variability across the ranges of most coral species remains unknown, especially for corals living in deep waters which span greater ranges. Here, we characterized the microbiomes of four octocoral species from mesophotic and bathyal deep-sea habitats in the northern Gulf of Mexico, Muricea pendula, Swiftia exserta, Callogorgia delta, and Paramuricea biscaya, using 16S rRNA gene metabarcoding. We sampled extensively across their ranges to test for microbiome differentiation between and within species, examining the influence of environmental factors that vary with depth (53-2224 m) and geographic location (over 680 m) as well as the host coral\'s genotype using RAD-sequencing.
RESULTS: Coral microbiomes were often dominated by amplicon sequence variants whose abundances varied across their hosts\' ranges, including symbiotic taxa: corallicolids, Endozoicomonas, members of the Mollicutes, and the BD1-7 clade. Coral species, depth, and geographic location significantly affected diversity, microbial community composition, and the relative abundance of individual microbes. Depth was the strongest environmental factor determining microbiome structure within species, which influenced the abundance of most dominant symbiotic taxa. Differences in host genotype, bottom temperature, and surface primary productivity could explain a significant part of the microbiome variation associated with depth and geographic location.
CONCLUSIONS: Altogether, this work demonstrates that the microbiomes of corals in deep waters vary substantially across their ranges in accordance with depth and other environmental conditions. It reveals that the influence of depth on the ecology of mesophotic and deep-sea corals extends to its effects on their microbiomes which may have functional consequences. This work also identifies the distributions of microbes including potential parasites which can be used to inform restoration plans in response to the Deepwater Horizon oil spill.

A new family of antipatharian corals, Ameripathidae (Cnidaria: Anthozoa: Antipatharia), is established for Ameripathespseudomyriophylla Opresko & Horowitz, gen. et sp. nov. The new family resembles Myriopathidae and Stylopathidae in terms of the morphology of the polyps and tentacles and the pinnulate branching of the corallum. Phylogenetic analysis using a genomic data set of 741 conserved element loci indicates that the new family is sister to a clade containing the Myriopathidae, Stylopathidae, Antipathidae, and Aphanipathidae.

The distribution of symbiotic scleractinian corals is driven, in part, by light availability, as host energy demands are partially met through translocation of photosynthate. Physiological plasticity in response to environmental conditions, such as light, enables the expansion of resilient phenotypes in the face of changing environmental conditions. Here we compared the physiology, morphology, and taxonomy of the host and endosymbionts of individual Madracis pharensis corals exposed to dramatically different light conditions based on colony orientation on the surface of a shipwreck at 30 m depth in the Bay of Haifa, Israel. We found significant differences in symbiont species consortia, photophysiology, and stable isotopes, suggesting that these corals can adjust multiple aspects of host and symbiont physiology in response to light availability. These results highlight the potential of corals to switch to a predominantly heterotrophic diet when light availability and/or symbiont densities are too low to sustain sufficient photosynthesis, which may provide resilience for corals in the face of climate change.

Over the last 2 decades, routine collections in the Hawaiian Archipelago have expanded to mesophotic reefs, leading to the discovery of a new red algal genus and species, here described as Anunuuluaehu liula gen. et sp. nov. This study provides a detailed genus and species description and characterizes chloroplast and mitochondrial organellar genomes. The new genus, Anunuuluaehu, shares many characteristics with the family Phyllophoraceae and shows close similarities to Archestennogramma and Stenogramma, including habit morphology, nemathecia forming proliferations at the outer cortex with terminal chains of tetrasporangia, and carposporophytes with multi-layered pericarps. The single species in this genus exhibits distinctive features within the Phyllophoraceae: the presence of single-layer construction of large medullary cells and the development of long, tubular gonimoblastic filaments. Multi-gene phylogenetic analyses confirmed it as a unique, monophyletic lineage within the family. Cis-splicing genes, interrupted by intron-encoded proteins within group II introns, are present in both the chloroplast and mitochondrial genomes of A. liula. Notably, a specific region of the coxI group II intron exhibits similarity to fungal introns. Anunuuluaehu liula is presumed to be endemic to the Hawaiian Archipelago and thus far is known to live solely at mesophotic depths from Hōlanikū to Kaho\'olawe ranging from 54 to 201 m, which is the deepest collection record of any representative in the family. Overall, this study enhances our understanding of the genomic and taxonomic complexities of red algae in mesophotic habitats, emphasizing the significance of continued research in this area to uncover further insights into evolutionary processes and biogeographic patterns.

The widespread decline of shallow-water coral reefs has fueled interest in assessing whether mesophotic reefs can act as refugia replenishing deteriorated shallower reefs through larval exchange. Here we explore the morphological and molecular basis facilitating survival of planulae and adults of the coral Porites astreoides (Lamarck, 1816; Hexacorallia: Poritidae) along the vertical depth gradient in Bermuda. We found differences in micro-skeletal features such as bigger calyxes and coarser surface of the skeletal spines in shallow corals. Yet, tomographic reconstructions reveal an analogous mineral distribution between shallow and mesophotic adults, pointing to similar skeleton growth dynamics. Our study reveals patterns of host genetic connectivity and minimal symbiont depth-zonation across a broader depth range than previously known for this species in Bermuda. Transcriptional variations across life stages showed different regulation of metabolism and stress response functions, unraveling molecular responses to environmental conditions at different depths. Overall, these findings increase our understanding of coral acclimatory capability across broad vertical gradients, ultimately allowing better evaluation of the refugia potential of mesophotic reefs.

Tropical deep reefs (~40-300 m) are diverse ecosystems that serve as habitats for diverse communities of reef-associated fishes. Deep-reef fish communities are taxonomically and ecologically distinct from those on shallow reefs, but like those on shallow reefs, they are home to a species-rich assemblage of small, cryptobenthic reef fishes, including many species from the family Gobiidae (gobies). Here we describe two new species of deep-reef gobies, Varicusprometheussp. nov. and V.roatanensissp. nov., that were collected using the submersible Idabel from rariphotic reefs off the island of Roatan (Honduras) in the Caribbean. The new species are the 11th and 12th species of the genus Varicus, and their placement in the genus is supported by morphological data and molecular phylogenetic analyses. Additionally, we also collected new specimens of the closely-related genus and species Pinnichthysaimoriensis during submersible collections off the islands of Bonaire and St. Eustatius (Netherland Antilles) and included them in this study to expand the current description of that species and document its range extension from Brazil into the Caribbean. Collectively, the two new species of Varicus and new records of P.aimoriensis add to our growing knowledge of cryptobenthic fish diversity on deep reefs of the Caribbean.

Mesophotic coral communities are increasingly gaining attention for the unique biological diversity they host, exemplified by the numerous mesophotic fish species that continue to be discovered. In contrast, many of the photosynthetic scleractinian corals observed at mesophotic depths are assumed to be depth-generalists, with very few species characterised as mesophotic-specialists. This presumed lack of a specialised community remains largely untested, as phylogenetic studies on corals have rarely included mesophotic samples and have long suffered from resolution issues associated with traditional sequence markers.
Here, we used reduced-representation genome sequencing to conduct a phylogenomic assessment of the two dominant mesophotic genera of plating corals in the Indo-Pacific and Western Atlantic, respectively, Leptoseris and Agaricia. While these genome-wide phylogenies broadly corroborated the morphological taxonomy, they also exposed deep divergences within the two genera and undescribed diversity across the current taxonomic species. Five of the eight focal species consisted of at least two sympatric and genetically distinct lineages, which were consistently detected across different methods.
The repeated observation of genetically divergent lineages associated with mesophotic depths highlights that there may be many more mesophotic-specialist coral species than currently acknowledged and that an urgent assessment of this largely unstudied biological diversity is warranted.

Tropical deep reefs (>30 m) are biologically and ecologically unique ecosystems with a higher geographic reach to shallow (<30 m) reefs. Yet they are poorly understood and rarely considered in conservation practices. Here, we characterise benthic and fish communities across a depth gradient (10-350 m) in remote coral atolls in Seychelles, Western Indian Ocean. Using taxonomic and trait-based approaches we present the taxonomic and functional composition of shallow and deep reef communities, with distinct communities and traits dominating different depths. Depth-related changes in community metrics (taxa richness, abundance and biomass) and functional diversity metrics (richness, dispersion, and evenness) indicate complex relationships across different biological components (fish, benthos) that differ between shallow and deep reefs. These in turn translate into different patterns of reef resilience against disturbance or species invasions with depth. Notably, deep reefs host on average fewer and less abundant taxa but with higher functional contribution and originality scores, some of which are of conservation concern. Overall, the results highlight the unique nature of deep reefs that requires their explicit consideration in conservation and management activities.

Local thermal environment has a strong influence on the physiology of marine ectotherms. This is particularly relevant for tropical organisms living close to their thermal optimum, well exemplified by the increasing frequency of bleaching occurrence in shallow-water corals. Mesophotic Coral Ecosystems (MCEs) were suggested as potential oases, especially when they are submitted to internal waves inducing short-term cooling events. Indeed, probability of bleaching occurrence in scleractinians was reported to decrease with depth in Mo\'orea as temperature variability increases. However, ecophysiological data are currently lacking to understand the cause of lower susceptibility/increased plasticity of deeper corals. A growing interest has been devoted the last decade to MCEs, but our understanding of the physiological performance of benthic organisms living in this environment remains relatively unexplored. To tackle that question, we first compared the metabolic responses (dark respiration, net photosynthesis and photosynthetic efficiency) of the depth-generalist scleractinian Pachyseris speciosa from two heterogeneous thermal environment (25 and 85 m depths) to acute heat stress to determine if the local thermal environment could predict coral response to warming. Then, we tested the thermal performance of two sympatric species (the scleractinian P. speciosa and the antipatharian Stichopathes sp.) to determine if there are inter-species differences in performances in species experiencing identical levels of temperature variability, at mesophotic depths (85 m). Results revealed broader thermal performances in the mesophotic P. speciosa compared to mid-depth ones, and constrained performances in the mesophotic antipatharian compared to the scleractinian species. We hypothesize that the high fluctuations in temperature due to internal waves in deeper areas contribute to the broader thermal performances of mesophotic P. speciosa. However, the constrained performances of the mesophotic antipatharian compared to P. speciosa suggests that other processes than the symbiosis with zooxanthellae also influence thermal performances of these mesophotic organisms. Our results supported that Stichopathes sp. lives close to its thermal optimum, suggesting a (relatively) cold thermal specialist strategy. In this context, composition of MCEs in the future is unlikely to shift to antipatharian-dominated landscape and will remain coral-dominated landscape.

Despite a global prevalence of photosynthetic organisms in the ocean\'s mesophotic zone (30-200+ m depth), the mechanisms that enable photosynthesis to proceed in this low light environment are poorly defined. Red coralline algae are the deepest known marine benthic macroalgae - here we investigated the light harvesting mechanism and mesophotic acclimatory response of the red coralline alga Lithothamnion glaciale.
Following initial absorption by phycourobilin and phycoerythrobilin in phycoerythrin, energy was transferred from the phycobilisome to photosystems I and II within 120 ps. This enabled delivery of 94% of excitations to reaction centres. Low light intensity, and to a lesser extent a mesophotic spectrum, caused significant acclimatory change in chromophores and biliproteins, including a 10% increase in phycoerythrin light harvesting capacity and a 20% reduction in chlorophyll-a concentration and photon requirements for photosystems I and II. The rate of energy transfer remained consistent across experimental treatments, indicating an acclimatory response that maintains energy transfer.
Our results demonstrate that responsive light harvesting by phycobilisomes and photosystem functional acclimation are key to red algal success in the mesophotic zone.