Gene co-option, duplication and divergence of cement proteins underpin the evolution of bioadhesives across barnacle life histories.

Acquisition of new genes often results in the emergence of novel functions and is a key step in lineage-specific adaptation. As a group of sessile crustaceans, barnacles establish permanent attachment through initial cement secretion at the larval phase followed by continuous cement secretion in juveniles and adults. However, the origins and evolution of barnacle larval and adult cement proteins remain poorly understood. By performing microdissection of larval cement glands, transcriptome and shotgun proteomics and immunohistochemistry validation, we identified 30 larval and 27 adult cement proteins of the epibiotic turtle barnacle Chelonibia testudinaria, of which the majority are stage- and barnacle-specific. While only two proteins, SIPC and CP100K, were expressed in both larvae and adults, detection of protease inhibitors and the cross-linking enzyme lysyl oxidase paralogs in larvae and adult cement. Other barnacle-specific cement proteins such as CP100k and CP52k likely share a common origin dating back at least to the divergence of Rhizocephala and Thoracica. Different CP52k paralogues could be detected in larval and adult cement, suggesting stage-specific cement proteins may arise from duplication followed by changes in expression timing of the duplicates. Interestingly, the biochemical properties of larval- and adult-specific CP52k paralogues exhibited remarkable differences. We conclude that barnacle larval and adult cement systems evolved independently, and both emerged from co-option of existing genes and de novo formation, duplication and functional divergence of lineage-specific cement protein genes. Our findings provide important insights into the evolutionary mechanisms of bioadhesives in sessile marine invertebrates.

Single-specimen systematics resolves the phylogeny and diversity conundrum of enigmatic crustacean y-larvae.

Resolving the evolutionary history of organisms is a major goal in biology. Yet for some taxa the diversity, phylogeny, and even adult stages remain unknown. The enigmatic crustacean "y-larvae" (Facetotecta) are one particularly striking example. Here, we use extensive video-imaging and single-specimen molecular sequencing of >200 y-larval specimens to comprehensively explore for the first time their evolutionary history and diversity. This integrative approach revealed five major clades of Facetotecta, four of which encompass a considerable larval diversity. Whereas morphological analyses recognized 35 y-naupliar "morphospecies", molecular species-delimitation analyses suggested the existence of between 88 and 127 species. The phenotypic and genetic diversity between the morphospecies suggests that a more elaborate classification than the current one-genus approach is needed. Morphology and molecular data were highly congruent at shallower phylogenetic levels, but no morphological synapomorphies could be unambiguously identified for major clades, which mostly comprise both planktotrophic and lecithotrophic y-nauplii. We argue that lecithotrophy arose several times independently whereas planktotrophic y-nauplii, which are structurally more similar across clades, most likely display the ancestral feeding mode of Facetotecta. We document a remarkably complex and highly diverse phylogenetic backbone for a taxon of larval marine crustaceans, the full life cycle of which remains a mystery.

Phylogeny and shell form evolution of the hydrothermal vent asymmetrical barnacles (Cirripedia, Thoracicalcarea, Neoverrucidae).

Imbricaverruca and Neoverruca are two genera of hydrothermal vent asymmetrical barnacles in Neoverrucidae, but found in vents of the Southwest Pacific and Northwest Pacific Oceans, respectively. Imbricaverruca has a flattened operculum and the shell base with multiple whorls of imbricating plates, while Neoverruca has an inclined operculum and the shell base with fewer developed imbricating plates. It has been hypothesized that Imbricaverruca has apomorphic shell characters in Neoverrucidae. Although the monophyletic relationship of the vent barnacle members in the superfamily Neolepadoidea were confirmed based on molecular phylogeny, the relationships between Neobrachylepadidae and Neoverrucidae, and between Neoverruca and Imbricaverruca have not been determined because there are no molecular data on Imbricaverruca. In this study, we sequenced three nuclear (18S rDNA, 28S rDNA, histone 3) and one mitochondrial (CO1) genes of I. yamaguchii from the Southwest Pacific. Our phylogenetic results showed that Neobrahchylepadidae is the sister taxon to Neoverrucidae (Imbricaverruca + Neoverruca), and Imbricaverruca and Neoverruca are monophyletic sister taxa each other, which not supporting the previous hypothesis that Neoverruca is sister to the clade containing Neobrahchylepadidae and Neolepadidae. These implied that the differences in shell forms between Neoverruca and Imbricaverruca are a result of independent divergent evolution in different deep-sea basins.

Five hundred million years to mobility: directed locomotion and its ecological function in a turtle barnacle.

Movement is a fundamental characteristic of life, yet some invertebrate taxa, such as barnacles, permanently affix to a substratum as adults. Adult barnacles became 'sessile' over 500 Ma; however, we confirm that the epizoic sea turtle barnacle, Chelonibia testudinaria, has evolved the capacity for self-directed locomotion as adults. We also assess how these movements are affected by water currents and the distance between conspecifics. Finally, we microscopically examine the barnacle cement. Chelonibia testudinaria moved distances up to 78.6 mm yr-1 on loggerhead and green sea turtle hosts. Movements on live hosts and on acrylic panels occasionally involved abrupt course alterations of up to 90°. Our findings showed that barnacles tended to move directly against water flow and independent of nearby conspecifics. This suggests that these movements are not passively driven by external forces and instead are behaviourally directed. In addition, it indicates that these movements function primarily to facilitate feeding, not reproduction. While the mechanism enabling movement remained elusive, we observed that trails of cement bore signs of multi-layered, episodic secretion. We speculate that proximal causes of movement involve one or a combination of rapid shell growth, cement secretion coordinated with basal membrane lifting, and directed contraction of basal perimeter muscles.

Swimming kinematics and hydrodynamics of barnacle larvae throughout development.

Changes in size strongly influence organisms' ecological performances. For aquatic organisms, they can transition from viscosity- to inertia-dominated fluid regimes as they grow. Such transitions are often associated with changes in morphology, swimming speed and kinematics. Barnacles do not fit into this norm as they have two morphologically distinct planktonic larval phases that swim differently but are of comparable sizes and operate in the same fluid regime (Reynolds number 100-101). We quantified the hydrodynamics of the rocky intertidal stalked barnacle Capitulum mitella from the nauplius II to cyprid stage and examined how kinematics and size increases affect its swimming performance. Cyprids beat their appendages in a metachronal wave to swim faster, more smoothly, and with less backwards slip per beat cycle than did all naupliar stages. Micro-particle image velocimetry showed that cyprids generated trailing viscous vortex rings that pushed water backwards for propulsion, contrary to the nauplii's forward suction current for particle capture. Our observations highlight that zooplankton swimming performance can shift via morphological and kinematic modifications without a significant size increase. The divergence in ecological functions through ontogeny in barnacles and the removal of feeding requirement likely contributed to the evolution of the specialized, taxonomically unique cyprid phase.

Phylogenetic relationships of Darwin's "Mr. Arthrobalanus": The burrowing barnacles (Cirripedia: Acrothoracica).

The barnacles of the superorder Acrothoracica are small, burrowing, epibiotic, and dioecious (large female with dwarf male) crustaceans largely found in the carbonate sediments and skeletons of marine invertebrates. The acrothoracicans represent the Cirripedia with the most plesiomorphic characters and have prominently featured in phylogenetic speculations concerning these crustaceans. Traditionally, Acrothoracica was divided into two main orders, Pygophora and Apygophora. The Apygophora had uniramus cirri and no anus. The Pygophora had biramus terminal cirri and an anus and was further divided into two families, Lithoglyptidae and Cryptophialidae. Kolbasov (2009) revised the superorder Acrothoracica on the basis of morphological examinations of females, dwarf males, and cyprids and rearranged the acrothoracican species into two new orders, Lithoglyptida and Cryptophialida. The present study is the first attempt to reconstruct the phylogenetic relationships of acrothoracican barnacles by sequencing two mitochondrial (cytochrome C oxidase I and 16S ribosomal DNA) and two nuclear (18S ribosomal DNA and histone H3) markers of 8 of the 11 genera comprising 23 acrothoracican species. All monophylies of the eight acrothoracican genera sampled in this study were strongly supported. The deep interfamilial relationship constructed is consistent with the recent morphological phylogenetic relationship proposed by Kolbasov, Newman, and Høeg (Kolbasov, 2009) that Cryptophialidae (order Cryptophialida) is the sister group to all other acrothoracicans (order Lithoglyptida). According to an ancestral character state reconstruction analysis, the posterior lobes of females; armament of opercular bars, attachment stalk, lateral projections of the body, and aperture slits in dwarf males; and habitat use appear to have phylogenetic importance.

How do coral barnacles start their life in their hosts?

Coral-associated invertebrates are the most significant contributors to the diversity of reef ecosystems, but no studies have examined how larvae manage to settle and grow in their coral hosts. Video recordings were used to document this process in the coral barnacle Darwiniella angularis associated with the coral Cyphastrea chalcidicum Settlement and metamorphosis in feeding juveniles lasted 8-11 days and comprised six phases. The settling cyprid starts by poking its antennules into the tissue of the prospective host (I: probing stage). The coral releases digestive filaments for defence, but tolerating such attack the cyprid penetrates further (II: battling stage). Ecdysis is completed 2 days after settlement (III: carapace detachment). The barnacle becomes embedded deep in the coral tissue while completing metamorphosis between 4 and 6 days (IV: embedding stage), but reappears as a feeding juvenile 8-11 days after settlement (V: emerging stage; VI: feeding stage). Cyprids preferably settle in areas between the coral polyps, where they have a much higher survival rate than on the polyp surfaces.

The origins and evolution of dwarf males and habitat use in thoracican barnacles.

Barnacles are exceptional in having various sexual systems (androdioecy, hermaphroditism, dioecy) and with a high morphological diversity of males, though these are always minute (dwarf) compared to their female or hermaphrodite partners. For the first time, we use a multiple DNA marker-based phylogeny to elucidate the ancestral states and evolution of (1) dwarf males, (2) their morphology when present, (3) their attachment site on the partner, and (4) habitat use in thoracican barnacles. Our taxon sampling was especially rich in rare deep-sea Scalpelliformes and comprised species with diverse sexual systems and dwarf male morphologies. Within the thoracican barnacles dwarf male evolution is subject to extensive convergence, but always correlated to similar ecological conditions. Males evolved convergently at least four times from purely hermaphroditic ancestors, in each case correlated with the invasion into habitats with low mating group sizes. The independent evolution of dwarf males in these lineages dovetails with the males having different morphologies and occurring in several different locations on their sexual partner.

Gorgonolaureus bicornutus sp. nov (Crustacea: Thecostraca: Ascothoracida) from off South-East Taiwan with notes on morphology and distribution.

A new ascothoracidan species, Gorgonolaureus bicornutus sp. nov., has been discovered off south-eastern Taiwan at a depth of 227 m. Five females were found in permanent cysts on the branches of a plexaurid octocoralian alcyonacean (former "gorgonian"), Echinogorgia sp. These specimens are assigned to the genus Gorgonolaureus on account of their having an enlarged and inflated carapace with a long, slit-like aperture, long dorsal thoracic horns, no filamentary appendages associated with the first pair of thoracopods, and rudimentary telsonic spines. Gorgonolaureus bicornutus differs from its congeners in having two long, naked dorsal horns on thoracomeres 2 and 3, the number of seminal receptacles in the thoracopods, and the higher number of stout setae on the fifth antennular segment. The characters that unite the genus Gorgonolaureus are redefined as follows: i) the absence of filamentary appendages associated with the first pair of thoracopods; ii) the possession of 1-3 dorsal horns distributed singly among thoracomeres 1-3; iii) the absence of prominent proximal teeth medially on the mandibles; iv) the possession of short or rudimentary telsonic spines; and v) host preference, with most species infecting octocorals of the suborder Halaxonia (mostly of the family Plexauridae) and never the calcaxonian families Chrysogorgiidae and Isididae. The gorgonian-infecting genera Gorgonolaureus and Isidascus are Tethyan relics that have survived only in the Western Pacific and the Eastern Atlantic, respectively, while their relatives in the genera Cardomanica and Thalassomembracis have disjunct Western Atlantic/Western Pacific distributions, thus exemplifying a major pattern of Tethyan reliction.

Worldwide genetic differentiation in the common fouling barnacle, Amphibalanus amphitrite.

Amphibalanus amphitrite is a common fouling barnacle distributed globally in tropical and subtropical waters. In the present study, the genetic (mitochondrial cytochrome oxidase subunit I) and morphological differentiation in A. amphitrite from 25 localities around the world were investigated. The results revealed three clades within A. amphitrite with a genetic divergence of ~ 4% among clades, whereas there were no diagnostic morphological differences among clades. Clade 1 is widely distributed in both temperate and tropical waters, whereas Clade 3 is currently restricted to the tropical region. The deep divergence among clades suggests historical isolation within A. amphitrite; thus, the present geographical overlaps are possibly a result of the combined effects of rising sea level and human-mediated dispersals. This study highlights the genetic differentiation that exists in a common, widely distributed fouling organism with great dispersal potential; future antifouling research should take into account the choice of lineages.

First study on gene expression of cement proteins and potential adhesion-related genes of a membranous-based barnacle as revealed from Next-Generation Sequencing technology.

This is the first study applying Next-Generation Sequencing (NGS) technology to survey the kinds, expression location, and pattern of adhesion-related genes in a membranous-based barnacle. A total of 77,528,326 and 59,244,468 raw sequence reads of total RNA were generated from the prosoma and the basis of Tetraclita japonica formosana, respectively. In addition, 55,441 and 67,774 genes were further assembled and analyzed. The combined sequence data from both body parts generates a total of 79,833 genes of which 47.7% were shared. Homologues of barnacle cement proteins - CP-19K, -52K, and -100K - were found and all were dominantly expressed at the basis where the cement gland complex is located. This is the main area where transcripts of cement proteins and other potential adhesion-related genes were detected. The absence of another common barnacle cement protein, CP-20K, in the adult transcriptome suggested a possible life-stage restricted gene function and/or a different mechanism in adhesion between membranous-based and calcareous-based barnacles.

First stage zoeal morphology of four ghost crabs Ocypode ceratophthalmus (Pallas, 1772), O. cordimanus Latreille, 1818, O. sinensis Dai, Song & Yang, 1985 and O. stimpsoni Ortmann, 1897 (Crustacea, Decapoda, Ocypodidae).

Light and scanning electron microscopy (SEM) were used to examine the first zoeal stage of four ghost crabs, Ocypode ceratophthalmus (Pallas, 1772), O. cordimanus Latreille, 1818, O. sinensis Dai, Song & Yang, 1985 and O. stimpsoni Ortmann, 1897. Finding diagnostic characters to distinguish between the four species proved difficult because their setal appendage patterns were identical. However the rectangular and pockmarked patterns on the ventral carapace are rather pronounced in O. ceratophthalmus and O. stimpsoni but weak in O. cordimanus and O. sinensis. The spinulation on the furca of the telson is less in O. cordimanus than in the other three species.

Comparative genomics reveals the dynamic evolutionary history of cement protein genes of barnacles from intertidal to deep-sea hydrothermal vents.

Thoracican barnacles are a diverse group of marine organisms for which the availability of genome assemblies is currently limited. In this study, we sequenced the genomes of two neolepadoid species (Ashinkailepas kermadecensis, Imbricaverruca yamaguchii) from hydrothermal vents, in addition to two intertidal species. Genome sizes ranged from 481 to 1054 Mb, with repetitive sequence contents of 21.2% to 50.7%. Concordance rates of orthologs and heterozygosity rates were between 82.4% and 91.7% and between 1.0% and 2.1%, respectively, indicating high genetic diversity and heterozygosity. Based on phylogenomic analyses, we revised the nomenclature of cement genes encoding cement proteins that are not homologous to any known proteins. The major cement gene, CP100A, was found in all thoracican species, including vent-associated neolepadoids, and was hypothesised to be essential for thoracican settlement. Duplicated genes, CP100B and CP100C, were found only in balanids, suggesting potential functional redundancy or acquisition of new functions associated with the calcareous base. An ancestor of CP52 genes was duplicated dynamically among lepadids, pollicipedids with multiple copies on a single scaffold, and balanids with multiple sequential repeats of the conserved regions, but no CP52 genes were found in neolepadoids, providing insights into cement gene evolution among thoracican lineages. This study enhances our understanding of the adhesion mechanisms of thoracicans in underwater environments. The newly sequenced genomes provide opportunities for studying their evolution and ecology, shedding light on their adaptation to diverse marine environments, and contributing to our knowledge of barnacle biology with valuable genomic resources for further studies in this field.

Novel molecular resources for single-specimen barcoding of enigmatic crustacean y-larvae.

Despite discovery more than 100years ago and documented global occurrence from shallow waters to the deep sea, the life cycle of the enigmatic crustacean y-larvae isincompletely understood and adult forms remain unknown. To date, only 2 of the 17 formally described species, all based on larval stages, have been investigated using an integrative taxonomic approach. This approach provided descriptions of the morphology of the naupliar and cyprid stages, and made use of exuvial voucher material and DNA barcodes. To improve our knowledge about the evolutionary history and ecological importance of y-larvae, we developed a novel protocol that maximises the amount of morpho-ecological and molecular data that can be harvested from single larval specimens. This includes single-specimen DNA barcoding and daily imaging of y-nauplii reared in culture dishes, mounting of the last naupliar exuviae on a slide as a reference voucher, live imaging of the y-cyprid instar that follows, and fixation, DNA extraction, amplification and sequencing of the y-cyprid specimen. Through development and testing of a suite of new primers for both nuclear and mitochondrial protein-coding and ribosomal genes, we showcase how new sequence data can be used to estimate the phylogeny of Facetotecta. We expect that our novel procedure will help to unravel the complex systematics of y-larvae and show how these fascinating larval forms have evolved. Moreover, we posit that our protocols should work on larval specimens from a diverse array of moulting marine invertebrate taxa.

Potential PCR amplification bias in identifying complex ecological patterns: Higher species compositional homogeneity revealed in smaller-size coral reef zooplankton by metatranscriptomics.

PCR-based high-throughput sequencing has permitted comprehensive resolution analyses of zooplankton diversity dynamics. However, significant methodological issues still surround analyses of complex bulk community samples, not least as in prevailing PCR-based approaches. Marine drifting animals-zooplankton-play essential ecological roles in the pelagic ecosystem, transferring energy and elements to higher trophic levels, such as fishes, cetaceans and others. In the present study, we collected 48 size-fractionated zooplankton samples in the vicinity of a coral reef island with environmental gradients. To investigate the spatiotemporal dynamics of zooplankton diversity patterns and the effect of PCR amplification biases across these complex communities, we first took metatranscriptomics approach. Comprehensive computational analyses revealed a clear pattern of higher/lower homogeneity in smaller/larger zooplankton compositions across samples respectively. Our study thus suggests changes in the role of dispersal across the sizes. Next, we applied in silico PCR to the metatranscriptomics datasets, in order to estimate the extent of PCR amplification bias. Irrespective of stringency criteria, we observed clear separations of size fraction sample clusters in both metatranscriptomics and in silico datasets. In contrast, the pattern-smaller-fractioned communities had higher compositional homogeneity than larger ones-was observed in the metatranscriptomics data but not in the in silico datasets. To investigate this discrepancy further, we analysed the mismatches of widely used mitochondrial CO1 primers and identified priming site mismatches likely driving PCR-based biases. Our results suggest the use of metatranscriptomics or, although less ideal, redesigning the CO1 primers is necessary to circumvent these issues.

Phylogenetic position and evolutionary history of the turtle and whale barnacles (Cirripedia: Balanomorpha: Coronuloidea).

Barnacles of the superfamily Coronuloidea are obligate epibionts of various marine mammals, marine reptiles and large crustaceans. We used five molecular markers: 12S rDNA, 16S rDNA, 18S rDNA, 28S rDNA and Histone 3 to infer phylogenetic relationships among sixteen coronuloids, representing most of the recent genera of barnacles of this superfamily. Our analyses confirm the monophyly of Coronuloidea and that this superfamily and Tetraclitoidea are sister groups. The six-plated Austrobalanus clusters with these two superfamilies. Based on BEAST and ML trees, Austrobalanus is basal and sister to the Coronuloidea, but the NJ tree places Austrobalanus within the Tetraclitoidae, and in the MP tree it is sister to both Coronuloidea and Tetraclitoidae. Hence the position of Austrobalanus remains unresolved. Within the Coronuloidea we identified four clades. Chelonibia occupies a basal position within the Coronuloidea which is in agreement with previous studies. The grouping of the other clades does not conform to previous studies. Divergence time analyses show that some of the time estimates are congruent with the fossil record while some others are older, suggesting the possibility of gaps in the fossil record.

Morphometric and molecular identification of individual barnacle cyprids from wild plankton: an approach to detecting fouling and invasive barnacle species.

The present study used DNA barcodes to identify individual cyprids to species. This enables accurate quantification of larvae of potential fouling species in the plankton. In addition, it explains the settlement patterns of barnacles and serves as an early warning system of unwanted immigrant species. Sequences from a total of 540 individual cypris larvae from Taiwanese waters formed 36 monophyletic clades (species) in a phylogenetic tree. Of these clades, 26 were identified to species, but 10 unknown monophyletic clades represented non-native species. Cyprids of the invasive barnacle, Megabalanus cocopoma, were identified. Multivariate analysis of antennular morphometric characters revealed three significant clusters in a nMDS plot, viz. a bell-shaped attachment organ (most species), a shoe-shaped attachment organ (some species), and a spear-shaped attachment organ (coral barnacles only). These differences in attachment organ structure indicate that antennular structures interact directly with the diverse substrata involved in cirripede settlement.

Acrothoracican barnacles (Lithoglyptida) in Taiwan, including the taxonomic status of Balanodytes taiwanus Utinomi, 1950 and cryptic diversity of Auritoglyptes bicornis (Aurivillius, 1892).

We list five acrothoracican barnacles of the order Lithoglyptida currently found in Taiwanese waters, including two new records, Trypetesa habei Utinomi, 1962 and Berndtiapurpurea Utinomi, 1950 and a new undescribed species of the genus Lithoglyptes. We also investigate the morphology and molecular genetics of Balanodytes taiwanus Utinomi, from its type locality, Taiwan. The original description of B. taiwanus in Utinomi (1950a) reported the absence of caudal appendages. Re-examination of the somatic body of one of the syntype specimens (the other syntype only having an opercular bar remaining) of B. taiwanus deposited in the Seto Marine Laboratory in Japan, revealed the presence of caudal appendages. The morphology of our specimens collected in various locations in Taiwan fits the description in Utinomi (1950a) and all have caudal appendages. The diagnosis of Balanodytes Utinomi, 1950 is revised herein. The genus Armatoglyptes Kolbasov & Newman, 2005 is a nomen nudum and ajunior synonym of Balanodytes. Our results also show that the widespread species Lithoglyptes habei Tomlinson, 1963 is a junior synonym of B. taiwanensis. The molecular sequence divergencse of the 16S RNA and COI genes were studied for samples of another monotypic genus Auritoglyptes (A. bicornis) from different regions of Taiwan. It was shown that there are at least three phylogenetic clades in Taiwan, suggesting that Auritoglyptes represents a cryptic species complex.

Host specificity and adaptive evolution in settlement behaviour of coral-associated barnacle larvae (Cirripedia: Pyrgomatidae).

Coral-associated organisms often exhibit a continuum of host specificities. We do not know whether the variation in host specificity is related to the settlement organs or preferential settlement behaviours of the larvae. We examined the morphology of attachment discs, the settlement and metamorphosis of coral barnacles-Pyrgoma cancellatum (lives in a single coral species), Nobia grandis (two families of corals), and Armatobalanus allium (six families of corals). Our results revealed that the attachment organ of all three species are spear-shaped with sparse villi, indicating that the morphology of the attachment organs does not vary among species with different host specificities. Larvae of P. cancellatum and N. grandis only settle on their specific hosts, suggesting that chemical cues are involved in the settlement. Cyprids of N. grandis display close searching behaviour before settlement. Cyprids of P. cancellatum settle immediately on their specific host corals, without any exploratory behaviour. The host specificity and exploratory behaviours of coral barnacle cyprids are results of adaptive evolution. We argue that there is a trade-off between exploration and energy conservation for metamorphosis processes. Coral barnacle metamorphosis is longer when compared to free-living species, likely because it involves the development of a tube-shaped base on the coral surface.

A draft genome assembly of reef-building octocoral Heliopora coerulea.

Coral reefs are under existential threat from climate change and anthropogenic impacts. Genomic studies have enhanced our knowledge of resilience and responses of some coral species to environmental stress, but reference genomes are lacking for many coral species. The blue coral Heliopora is the only reef-building octocoral genus and exhibits optimal growth at a temperature close to the bleaching threshold of scleractinian corals. Local and high-latitude expansions of Heliopora coerulea were reported in the last decade, but little is known about the molecular mechanisms underlying its thermal resistance. We generated a draft genome of H. coerulea with an assembled size of 429.9 Mb, scaffold N50 of 1.42 Mb and BUSCO completeness of 94.9%. The genome contains 239.1 Mb repetitive sequences, 27,108 protein coding genes, 6,225 lncRNAs, and 79 miRNAs. This reference genome provides a valuable resource for in-depth studies on the adaptive mechanisms of corals under climate change and the evolution of skeleton in cnidarian.