Systematics and character evolution of capitate hydrozoans.

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

A hybrid-capture approach to reconstruct the phylogeny of Scleractinia (Cnidaria: Hexacorallia).

A well-supported evolutionary tree representing most major lineages of scleractinian corals is in sight with the development and application of phylogenomic approaches. Specifically, hybrid-capture techniques are shedding light on the evolution and systematics of corals. Here, we reconstructed a broad phylogeny of Scleractinia to test previous phylogenetic hypotheses inferred from a few molecular markers, in particular, the relationships among major scleractinian families and genera, and to identify clades that require further research. We analysed 449 nuclear loci from 422 corals, comprising 266 species spanning 26 families, combining data across whole genomes, transcriptomes, hybrid capture and low-coverage sequencing to reconstruct the largest phylogenomic tree of scleractinians to date. Due to the large number of loci and data completeness (less than 38% missing data), node supports were high across shallow and deep nodes with incongruences observed in only a few shallow nodes. The "Robust" and "Complex" clades were recovered unequivocally, and our analyses confirmed that Micrabaciidae Vaughan, 1905 is sister to the "Robust" clade, transforming our understanding of the "Basal" clade. Several families remain polyphyletic in our phylogeny, including Deltocyathiidae Kitahara, Cairns, Stolarski & Miller, 2012, Caryophylliidae Dana, 1846, and Coscinaraeidae Benzoni, Arrigoni, Stefani & Stolarski, 2012, and we hereby formally proposed the family name Pachyseridae Benzoni & Hoeksema to accommodate Pachyseris Milne Edwards & Haime, 1849, which is phylogenetically distinct from Agariciidae Gray, 1847. Results also revealed species misidentifications and inconsistencies within morphologically complex clades, such as Acropora Oken, 1815 and Platygyra Ehrenberg, 1834, underscoring the need for reference skeletal material and topotypes, as well as the importance of detailed taxonomic work. The approach and findings here provide much promise for further stabilising the topology of the scleractinian tree of life and advancing our understanding of coral evolution.

Phylogeography of recent Plesiastrea (Scleractinia: Plesiastreidae) based on an integrated taxonomic approach.

Scleractinian corals are a diverse group of ecologically important yet highly threatened marine invertebrates, which can be challenging to identify to the species level. An influx of molecular studies has transformed scleractinian systematics, highlighting that cryptic species may be more common than previously understood. In this study, we test the hypothesis that Plesiastrea versipora (Lamarck, 1816), a species currently considered to occur throughout the Indo-Pacific in tropical, sub-tropical and temperate waters, is a single species. Molecular and morphological analyses were conducted on 80 samples collected from 31 sites spanning the majority of the species putative range and twelve mitogenomes were assembled to identify informative regions for phylogenetic reconstruction. Congruent genetic data across three gene regions supports the existence of two monophyletic clades aligning with distinct tropical and temperate provenances. Multivariate macromorphological analyses based on 13 corallite characters provided additional support for the phylogeographic split, with the number of septa and corallite density varying across this biogeographic divide. Furthermore, micromorphological and microstructural analyses identified that the temperate representatives typically develop sub-cerioid corallites with sparse or absent coenosteal features and smooth septal faces. In contrast, tropical representatives typically develop plocoid corallites separated by a porous dissepimental coenosteum and have granulated septal faces. These data suggest that at least two species exist within the genus PlesiastreaMilne Edwards & Haime, 1848. Based on examination of type material, we retain the name Plesiastrea versipora (Lamarck, 1816) for the temperate representatives of the genus and resurrect the name Plesiastrea peroniMilne Edwards & Haime, 1857 for the tropical members. This study highlights how broadly distributed hard coral taxa still need careful re-examination through an integrated systematics approach to better understand their phylogeographic patterns. Furthermore, it demonstrates the utility of integrating micro-, macro-morphological and genetic datasets, and the importance of type specimens when dealing with taxonomic revisions of scleractinian taxa.

Diversity, host specificity and biogeography in the Cladocorynidae (Hydrozoa, Capitata), with description of a new genus.

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.

Global raster dataset on historical coastline positions and shelf sea extents since the Last Glacial Maximum.

Motivation: Historical changes in sea level caused shifting coastlines that affected the distribution and evolution of marine and terrestrial biota. At the onset of the Last Glacial Maximum (LGM) 26 ka, sea levels were >130 m lower than at present, resulting in seaward-shifted coastlines and shallow shelf seas, with emerging land bridges leading to the isolation of marine biota and the connection of land-bridge islands to the continents. At the end of the last ice age, sea levels started to rise at unprecedented rates, leading to coastal retreat, drowning of land bridges and contraction of island areas. Although a growing number of studies take historical coastline dynamics into consideration, they are mostly based on past global sea-level stands and present-day water depths and neglect the influence of global geophysical changes on historical coastline positions. Here, we present a novel geophysically corrected global historical coastline position raster for the period from 26 ka to the present. This coastline raster allows, for the first time, calculation of global and regional coastline retreat rates and land loss rates. Additionally, we produced, per time step, 53 shelf sea rasters to present shelf sea positions and to calculate the shelf sea expansion rates. These metrics are essential to assess the role of isolation and connectivity in shaping marine and insular biodiversity patterns and evolutionary signatures within species and species assemblages. Main types of variables contained: The coastline age raster contains cells with ages in thousands of years before present (bp), representing the time since the coastline was positioned in the raster cells, for the period between 26 ka and the present. A total of 53 shelf sea rasters (sea levels <140 m) are presented, showing the extent of land (1), shelf sea (0) and deep sea (NULL) per time step of 0.5 kyr from 26 ka to the present. Spatial location and grain: The coastline age raster and shelf sea rasters have a global representation. The spatial resolution is scaled to 120 arcsec (0.333° × 0.333°), implying cells of c. 3,704 m around the equator, 3,207 m around the tropics (±30°) and 1,853 m in the temperate zone (±60°). Time period and temporal resolution: The coastline age raster shows the age of coastline positions since the onset of the LGM 26 ka, with time steps of 0.5 kyr. The 53 shelf sea rasters show, for each time step of 0.5 kyr, the position of the shelf seas (seas shallower than 140 m) and the extent of land. Level of measurement: Both the coastline age raster and the 53 shelf sea rasters are provided as TIFF files with spatial reference system WGS84 (SRID 4326). The values of the coastline age raster per grid cell correspond to the most recent coastline position (in steps of 0.5 kyr). Values range from 0 (0 ka, i.e., present day) to 260 (26 ka) in bins of 5 (0.5 kyr). A value of "no data" is ascribed to pixels that have remained below sea level since 26 ka. Software format: All data processing was done using the R programming language.

Coral diversity matches marine park zonation but not economic value of coral reef sites at St. Eustatius, eastern Caribbean.

Stony corals play a key role in the marine biodiversity of many tropical coastal areas as suppliers of substrate, food and shelter for other reef organisms. Therefore, it is remarkable that coral diversity usually does not play a role in the planning of protected areas in coral reef areas. In the present study we examine how stony coral diversity patterns relate to marine park zonation and the economic value of reefs around St. Eustatius, a small island in the eastern Caribbean, with fisheries and tourism as important sources of income. The marine park contains two no-take reserves. A biodiversity survey was performed at 39 sites, 24 inside the reserves and 15 outside; 22 had a maximum depth >18 m and 17 were shallower. Data on economic value per site were obtained from the literature. Corals were photographed for the verification of identifications made in the field. Coral species richness (n = 49) was highest in the no-take reserves and species composition was mainly affected by maximum depth. No distinct relation is observed between coral diversity and fishery value or total economic value. Based on the outcome of this study we suggest that in future designs of marine park zonation in reef areas, coral diversity should be taken into consideration. This is best served by including reef areas with a continuous depth gradient from shallow flats to deep slopes.

3D Chitin Scaffolds of Marine Demosponge Origin for Biomimetic Mollusk Hemolymph-Associated Biomineralization Ex-Vivo.

Structure-based tissue engineering requires large-scale 3D cell/tissue manufacture technologies, to produce biologically active scaffolds. Special attention is currently paid to naturally pre-designed scaffolds found in skeletons of marine sponges, which represent a renewable resource of biomaterials. Here, an innovative approach to the production of mineralized scaffolds of natural origin is proposed. For the first time, a method to obtain calcium carbonate deposition ex vivo, using living mollusks hemolymph and a marine-sponge-derived template, is specifically described. For this purpose, the marine sponge Aplysin aarcheri and the terrestrial snail Cornu aspersum were selected as appropriate 3D chitinous scaffold and as hemolymph donor, respectively. The formation of calcium-based phase on the surface of chitinous matrix after its immersion into hemolymph was confirmed by Alizarin Red staining. A direct role of mollusks hemocytes is proposed in the creation of fine-tuned microenvironment necessary for calcification ex vivo. The X-ray diffraction pattern of the sample showed a high CaCO3 amorphous content. Raman spectroscopy evidenced also a crystalline component, with spectra corresponding to biogenic calcite. This study resulted in the development of a new biomimetic product based on ex vivo synthetized ACC and calcite tightly bound to the surface of 3D sponge chitin structure.

Lack of host specificity of copepod crustaceans associated with mushroom corals in the Red Sea.

The radiation of symbiotic copepods (Crustacea: Copepoda) living in association with stony corals (Cnidaria: Scleractinia) is considered host-specific and linked to the phylogenetic diversification of their hosts. However, symbiotic copepods are poorly investigated, occurrence records are mostly anecdotal, and no explicit analysis exists regarding their relationship with the hosts. Here, we analysed the occurrence of symbiotic copepods on different co-occurring and phylogenetically closely related scleractinian corals. We used an innovative approach of DNA extraction from single microscopic specimens that preserves the shape of the organisms for integrative morphological studies. The rationale of the study involved: (i) sampling of mushroom corals (Fungiidae) belonging to 13 species and eight genera on different reefs along the Saudi coastline in the Red Sea, (ii) extraction of all the associated copepods, (iii) morphological screening and identification of copepod species, (iv) use of DNA taxonomy on mitochondrial and nuclear markers to determine species boundaries for morphologically unknown copepod species, (v) reconstruction of phylogenies to understand their evolutionary relationships, and (vi) analysis of the ecological drivers of the occurrence, diversity and host specificity of the copepods. The seven species of coral-associated copepods, all new to science, did not show any statistically significant evidence of host-specificity or other pattern of ecological association. We thus suggest that, contrary to most assumptions and previous anecdotal evidence on this coral-copepod host-symbiont system, the association between copepods and their host corals is rather labile, not strict, and not phylogenetically constrained, changing our perception on evolutionary patterns and processes in symbiotic copepods.

Host specificity of coral-associated fauna and its relevance for coral reef biodiversity.

Coral-associated fauna predominantly consists of invertebrates and constitutes an important component of coral reef biodiversity. The symbionts depend on their hosts for food, shelter and substrate. They may act as parasites by feeding on their hosts, by overgowing their polyps, or by excavating their skeletons. Because some of these species partly reside inside their hosts, they may be cryptic and can easily be overlooked in biodiversity surveys. Since no quantitative overview is available about these inter-specific relationships, this present study adresses variation in host ranges and specificity across four large coral-associated taxa and between the Atlantic and Indo-Pacific oceans. These taxa are: coral barnacles (Pyrgomatidae, n = 95), coral gall crabs (Cryptochiridae, n = 54), tubeworms (Serpulidae, n = 31), and date mussels (Lithophaginae, n = 23). A total of 335 host coral species was recorded. An index of host specificity (STD) was calculated per symbiont species, based on distinctness in taxonomic host range levels (species, genus, family, etc.). Mean indices were statistically compared among the four associated taxa and the two oceanic coral reef regions. Barnacles were the most host-specific, tubeworms the least. Indo-Pacific associates were approximately 10 times richer in species and two times more host-specific than their Atlantic counterparts. Coral families varied in the number of associates, with some hosting none. This variation could be linked to host traits (coral growth form, maximum host size) and is most probably also a result of the evolutionary history of the interspecific relationships.

In the aftermath of Hurricane Irma: Colonization of a 4-year-old shipwreck by native and non-native corals, including a new cryptogenic species for the Caribbean.

Little is known about early coral settlement on shipwrecks with regard to their species and size compositions. Hurricanes in the Caribbean have a long history of sinking ships but a link with new coral settlement is understudied. In 2017, Hurricane Irma caused the sinking of over 300 vessels in the coastal waters of Saint Martin, eastern Caribbean. In 2021, coral settlement was studied on one of them, which included two native, one non-native, and two cryptogenic species. The corals were smaller than 8 cm in diameter. The invasive Tubastraea coccinea was the most abundant scleractinian and was predominantly represented by juveniles. A cryptogenic species, Stragulum bicolor, new for the Caribbean, was the most common octocoral. Because they can be harmful to the environment, shipwrecks should be monitored frequently for the occurrence of non-native species, especially when they are only a few years old.

Non-native coral species dominate the fouling community on a semi-submersible platform in the southern Caribbean.

A coral community was examined on a semi-submersible platform that was moored at the leeward side of Curaçao, in the southern Caribbean, from August 2016 until August 2017. This community included several non-native or cryptogenic species. Among them were two scleractinian corals (Tubastraea coccinea and T. tagusensis) and two octocorals (Chromonephthea sp. and an unidentified Nephtheidae sp.). This is the first reported presence of T. tagusensis in the southern Caribbean, and the genus Chromonephthea in the Caribbean region. An ascidian, Perophora cf. regina, is also reported from the southern Caribbean for the first time, as well as a coral-associated vermetid gastropod, Petaloconchus sp., first recorded in the Caribbean in 2014. Lack of biofouling management could potentially harm indigenous marine fauna through the introduction of non-native species. Therefore monitoring communities associated with semi-submersible platforms is essential to track the presence and dispersal of non-native, potentially invasive species.

Abundance of coral-associated fauna in relation to depth and eutrophication along the leeward side of Curaçao, southern Caribbean.

Coral-associated invertebrates contribute much to the biodiversity of Caribbean coral reefs. Although the nature of their symbiotic relation is usually not fully understood, they can cause damage to their hosts, especially when they occur in high densities. The abundance of seven groups of coral-associated invertebrates was investigated on reefs along the leeward side of Curaçao, southern Caribbean. In particular, coral barnacles (Pyrgomatidae), boring mussels (Mytilidae: Leiosolenus spp.), gall crabs (Cryptochiridae), and Christmas tree worms (Serpulidae: Spirobranchus spp.) were recorded together with their host corals by means of a photo survey at four depths (5, 10, 15, 20 m) and across seven sites with high and five sites with low eutrophication values (based on δ15N isotope data). Feather duster worms (Sabellidae: Anamobaea), coral blennies (Chaenopsidae: Acanthemblemaria), and worm snails (Vermetidae: Petaloconchus) were insufficiently abundant for thorough quantitative analyses. The results show a decrease in the number of barnacles and Christmas tree worms per host over depth, which could be related to the availability of their host corals. Sites with high δ15N values show a higher abundance of barnacles and Christmas tree worms per host than sites with low values. This indicates that eutrophication could be favourable for these filter feeding organisms but when their densities become too high, they tend to overgrow their hosts and may become a threat to them.

A centuries-old manmade reef in the Caribbean does not substitute natural reefs in terms of species assemblages and interspecific competition.

With increasing maritime activities in the proximity of coral reefs, a growing number of manmade structures are becoming available for coral colonisation. Yet, little is known about the sessile community composition of such artificial reefs in comparison with that of natural coral reefs. Here, we compared the diversity of corals and their competitors for substrate space between a centuries-old manmade structure and the nearest natural reef at St. Eustatius, eastern Caribbean. The artificial reef had a significantly lower species richness and fewer competitive interactions than the natural reef. The artificial reef was dominated by a cover of crustose coralline algae and zoantharians, instead of turf algae and fire corals on the natural reef. Significant differences in species composition were also found between exposed and sheltered sites on both reefs. Our study indicates that even a centuries-old manmade reef cannot serve as a surrogate for natural reefs.

Downslope migration of free-living corals (Scleractinia: Fungiidae) in typhoon-exposed reef habitats at Okinawa, Japan.

Offshore Onna Village, Okinawa Island, Japan, there is a large and densely covered coral assemblage of free-living mushroom corals (Scleractinia: Fungiidae) on a reef slope at depths from 20 m to 32 m, covering an area of approximately 350 × 40 m2. From previous research, it is known that migration distances of mushroom corals may depend on coral shapes, coral sizes, substrate, and bottom inclination. However, until now there have been no published examples of regular Fungiidae movement and behavior from typhoon-exposed coastlines, such as those in the western Pacific Ocean. Our surveys across three years offshore Onna Village show that mushroom corals always move in down-slope direction from shallow to deeper reef zones. The results indicated that mushroom corals migrated faster in autumn than in other seasons, and that oval-elongate fungiids, and particularly those with a smooth underside, migrated more quickly than species with other shapes. Surprisingly, we observed a negative relationship between the presence of typhoons and migration rates. We also observed active migration by fungiid individuals to escape situations in which they were threatened to become overgrown by Acropora corals, or when they needed to escape from burial underneath coral debris.

Contrasting coral community structures between natural and artificial substrates at Koh Tao, Gulf of Thailand.

Concrete cubic frames and decommissioned steel naval vessels have been deployed in Thailand liberally to act as artificial substrates for coral restoration and marine recreation. We assessed recruitment at such substrate types at Koh Tao, Gulf of Thailand, and compared the community structure of scleractinian corals between artificial substrates and nearby natural reefs. Our results from a sample of 2677 recruits from nine sites highlighted significant differences in community structure between both reef types. Investigations of variables including time since deployment, distance from the natural reef, and seafloor depth revealed only the latter as a possible influencing factor. The diversity of recruits could not be explained by dynamics in coral spawning, and were found to represent groups with lower structural complexity. Our results suggest that coral community structure on artificial and natural reefs differs and supports distinct ecological and functional roles.

Selective consumption of sacoglossan sea slugs (Mollusca: Gastropoda) by scleractinian corals (Cnidaria: Anthozoa).

Recent studies revealed that reef corals can eat large-sized pelagic and benthic animals in addition to small planktonic prey. As follow-up, we document natural ingestion of sea slugs by corals and investigate the role of sacoglossan sea slugs as possible prey items of scleractinian corals. Feeding trials were carried out using six sacoglossan species as prey, two each from the genera Costasiella, Elysia and Plakobranchus, and four free-living solitary corals (Danafungia scruposa, Fungia fungites, Pleuractis paumotensis and Heteropsammia cochlea) as predators. Trials were carried out under both in-situ and ex-situ conditions with the aim to observe ingestion and assess signs of prey consumption based on tissue loss of prey individuals over time. Significant differences were observed in both ingestion time and consumption state of prey between prey species, with three of them being ingested more rapidly and preferentially consumed over the others. Additionally, prey size was found to be a significant factor with larger prey (>12 mm) being ingested more slowly and rarely than smaller ones (<6 mm and 6-12 mm). Comparisons of consumption capability among predators showed no significant difference with all coral species showing similar preferences for prey species. While no specific mechanism of prey capture is proposed, we also document instances of kleptoparisitism and resuspension of prey items by wrasses. This study highlights the important distinction between opportunistic prey capture and true predation events.

Morphological and phylogenetic diversity of Waminoa and similar flatworms (Acoelomorpha) in the western Pacific Ocean.

The genus Waminoa currently contains two described species, which each contains two types of endosymbiotic algae. Waminoa individuals are basically brown in body color, derived from these algal symbionts, and their body shape has been described as "discoid to obcordate". They have been found as associates of various anthozoans (Cnidaria) in the Indo-Pacific Ocean and the Red Sea. In order to reveal the diversity of the genus Waminoa and their hosts, we conducted phylogenetic and morphological analyses on acoelomate flatworms specimens collected from Japan, Palau and Indonesia. At least 18 Waminoa morphotypes were found on at least 20 anthozoan host species, and two specimens were found on species of two sea stars. Overall, there were two main body shapes of specimens; obcordate, as seen in W. litus and W. brickneri, and the other molar-like with an elongated body. These two body shapes each represented a separate clade in 18S rDNA and mitochondrial cytochrome c oxidase subunit 1 (COI) phylogenetic trees, with W. brickneri included in the obcordate subclade. Automatic Barcode Gap Discovery (ABGD) analyses on COI sequences of our specimens revealed the presence of at least five operational taxonomic units (OTUs). These five OTUs consisted of one large group of all obcordate animals, three OTUs consisting of one specimen each within the molar-like clade, and one large group of the remaining molar-like specimens. Both clades contain numerous morphotypes and were associated with a variety of hosts. Finally, based on genetic distances, the molar-like specimens are considered as an unnamed genus group separate from Waminoa, which needs to be clarified in future studies.