A methodological exploration to study 2D arm kinematics in Ophiuroidea (Echinodermata).

Brittle stars, unlike most other echinoderms, do not use their small tube feet for locomotion but instead use their flexible arms to produce a rowing or reverse rowing movement. They are among the fastest-moving echinoderms with the ability of complex locomotory behaviors. Considering the high species diversity and variability in morphotypes, a proper understanding of intra- and interspecies variation in arm flexibility and movement is lacking. This study focuses on the exploration of the methods to investigate the variability in brittle star locomotion and individual arm use. We performed a two-dimensional (2D) image processing on horizontal movement only. The result indicated that sinuosity, disc displacement and arm angle are important parameters to interpret ophiuroid locomotion. A dedicated Python script to calculate the studied movement parameters and visualize the results applicable to all 5-armed brittle stars was developed. These results can serve as the basis for further research in robotics inspired by brittle star locomotion.

Order Euryalida (Echinodermata, Ophiuroidea), new species and new records from the South China Sea and the Northwest Pacific seamounts.

Ophiuroids were collected by the manned submersible 'Shenhaiyongshi' from the deep-sea seamounts in the South China Sea and Northwest Pacific regions at 602-1920 m depth, during 2018 to 2020. A total of nine species was identified, including two new species and seven new records from the South China Sea and one new record from the Northwest Pacific region. Two new species are described as Asteroschemashenhaiyongshii sp. nov. and Asteroschemadomogranulatum sp. nov. The seven new records included five species from the genus Asteroschema, and one species each from the genera Asterostegus and Astrodendrum. Comprehensive descriptions of morphological features are provided, including characteristics of the arm skeleton, as well as a phylogenetic analysis based on 16S and COI sequences. Intraspecific genetic distance ranges of Euryalida species from the present study were 0.34% to 1.38%, which was relatively low compared to other orders in the class Ophiuroidea. The present study suggests a high probability that species of the order Euryalida are more widely spread around the Indo-Pacific region than previously expected.

The evolutionary relationship between arm vertebrae shape and ecological lifestyle in brittle stars (Echinodermata: Ophiuroidea).

Ophiuroidea are one of the most diverse classes among extant echinoderms, characterized by their flexible arms composed of a series of ossicles called vertebrae, articulating with each other proximally and distally. Their arms show a wide range of motion, important for feeding and locomotion, associated with their epizoic and non-epizoic lifestyles. It remains to be explored to what degree the phenotypic variation in these ossicles also reflects adaptations to these lifestyles, rather than only their phylogenetic affinity. In this study, we analyzed the 3D shape variation of six arm vertebrae from the middle and distal parts of an arm in 12 species, belonging to the intertidal, subtidal and bathyal zones and showing epizoic and non-epizoic behaviors. A PERMANOVA indicated a significant difference in ossicle morphology between species and between lifestyles. A principal component analysis showed that the morphology of epizoic ophiuroids is distinct from non-epizoic ones; which may reflect variation in arm function related to these different lifestyles. The Phylogenetic MANOVA and phylogenetic signal analysis showed that shape variation in the vertebral articulation seems to reflect ecological and functional adaptations, whereas phylogeny controls more the lateral morphology of the vertebrae. This suggests a convergent evolution through ecological adaptation to some degree, indicating that some of these characters may have limited taxonomic value.

Review of Ophioplinthaca Verrill, 1899 (Echinodermata, Ophiuroidea, Ophiacanthidae), description of new species in Ophioplinthaca and Ophiophthalmus, and new records from the Northwest Pacific and the South China Sea.

The ophiuroid genus Ophioplinthaca is well characterized by the deep incisions in the disc. Prior to this study, it contained 32 accepted species, but species limits and geographic distributions were not well understood. The manned submersible vehicle 'Shenhaiyongshi' was used to collect ophiuroid specimens from the deep-sea seamounts and cold seeps in the South China Sea and Northwest Pacific at 602-3600 m depth, during 2018 to 2020. The genus Ophioplinthaca was reviewed using both morphological data and a phylogenetic analysis, based on COI sequences. The taxonomic status of the genus Ophiophthalmus Matsumoto, 1917, a junior homonym of Ophiophthalmus Fitzinger, 1843 (a reptile) was clarified by proving prevailing usage of the ophiuroid name. A total of eight species were identified, including two new species, described asOphioplinthacabrachispina sp. nov. and Ophiophthalmusserratus sp. nov., and two new records. The new species are characterized by unique features of the arm skeletons. Tabular keys to all Ophioplinthaca and Ophiophthalmus species are provided. Interspecific and intraspecific genetic distance of Ophioplinthaca species ranged from 2.32% to 19.72%, and from 0.26% to 0.90%, respectively. The data suggest that species of the genus Ophioplinthaca are more widely spread around the Northwest Pacific region deep-sea seamounts than previously known.

Megafauna of the German exploration licence area for seafloor massive sulphides along the Central and South East Indian Ridge (Indian Ocean).

BACKGROUND: The growing interest in mineral resources of the deep sea, such as seafloor massive sulphide deposits, has led to an increasing number of exploration licences issued by the International Seabed Authority. In the Indian Ocean, four licence areas exist, resulting in an increasing number of new hydrothermal vent fields and the discovery of new species. Most studies focus on active venting areas including their ecology, but the non-vent megafauna of the Central Indian Ridge and South East Indian Ridge remains poorly known.In the framework of the Indian Ocean Exploration project in the German license area for seafloor massive sulphides, baseline imagery and sampling surveys were conducted yearly during research expeditions from 2013 to 2018, using video sledges and Remotely Operated Vehicles. NEW INFORMATION: This is the first report of an imagery collection of megafauna from the southern Central Indian- and South East Indian Ridge, reporting the taxonomic richness and their distribution. A total of 218 taxa were recorded and identified, based on imagery, with additional morphological and molecular confirmed identifications of 20 taxa from 89 sampled specimens. The compiled fauna catalogue is a synthesis of megafauna occurrences aiming at a consistent morphological identification of taxa and showing their regional distribution. The imagery data were collected during multiple research cruises in different exploration clusters of the German licence area, located 500 km north of the Rodriguez Triple Junction along the Central Indian Ridge and 500 km southeast of it along the Southeast Indian Ridge.

Deep-sea Ophiuroidea (Echinodermata) from the Danish Galathea II Expedition, 1950-52, with taxonomic revisions.

The brittle star samples collected by the Danish cruise 'Galathea II' (1950-52) had not been studied completely. We examined the remaining deep-sea samples (400 m) and present the species inventory, discussing taxonomic issues in relation to recent phylogenetic data. About 235 samples were examined, over 9,300 individuals, from 67 species and 74 sampling localities, at depths of 425-5340 m. The species complex Amphiophiura bullata (Thomson, 1877) is morphologically not well separated, but molecular data suggest at least two clades. We propose to apply A. bullata for Atlantic and Australian populations and A. convexa (Lyman, 1878) for the North Pacific clade. We consider A. bullata pacifica Litvinova, 1971 conspecific with A. convexa. Ophiuroglypha irrorata (Lyman, 1878) and its subspecies are a polyphyletic group with unclear morphological boundaries. We propose to transfer Ophiura ossiculata (Koehler, 1908), Ophiura plana (Lütken Mortensen, 1899) and Ophiura scomba Paterson, 1985 to Ophiuroglypha. Silax Fell, 1962, until now synonymised with Amphioplus Verrill, 1899, is proposed as a valid genus with the species S. verrilli (Lyman, 1879), S. consors (Koehler, 1908), S. daleus (Lyman, 1879), S. patulus (Lyman, 1879) and S. magnificus (Koehler, 1907). Triplodia Turner Hallen, 2011 (a replacement name for Triodia A. M. Clark, 1970, due to homonymy) is synonymised with Silax, and possible specimens of its type species Triodia abdita A. M. Clark, 1970 are analysed. The species limits of Ophiacantha cosmica Lyman, 1879 and Ophiacantha pacifica Lütken Mortensen, 1899 could not be confirmed morphologically, but published molecular data suggest two clades. We propose to apply O. pacifica to the Northern/Central Pacific population and O. cosmica to the Southern Pacific/Antarctic population.

Species delimitation in the presence of strong incomplete lineage sorting and hybridization: Lessons from Ophioderma (Ophiuroidea: Echinodermata).

Accurate species delimitation is essential to properly assess biodiversity, but also for management and conservation purposes. Yet, it is not always trivial to accurately define species boundaries in closely related species due to incomplete lineage sorting. Additional difficulties may be caused by hybridization, now evidenced as a frequent phenomenon. The brittle star cryptic species complex Ophioderma longicauda encompasses six mitochondrial lineages, including broadcast spawners and internal brooders, yet the actual species boundaries are unknown. Here, we combined three methods to delimit species in the Ophioderma longicauda complex and to infer its divergence history: (i) unsupervised species discovery based on multilocus genotypes; (ii) divergence time estimation using the multi-species coalescent; (iii) divergence scenario testing (including gene flow) using Approximate Bayesian Computation (ABC) methods. 30 sequence markers (transcriptome-based, mitochondrial or non-coding) for 89 O. longicauda and outgroup individuals were used. First, multivariate analyses revealed six genetic clusters, which globally corresponded to the mitochondrial lineages, yet with many exceptions, suggesting ancient hybridization events and challenging traditional mitochondrial barcoding approaches. Second, multi-species coalescent-based analyses confirmed the occurrence of six species and provided divergence time estimates, but the sole use of this method failed to accurately delimit species, highlighting the power of multilocus genotype clustering to delimit recently diverged species. Finally, Approximate Bayesian Computation showed that the most likely scenario involves hybridization between brooders and broadcasters. Our study shows that despite strong incomplete lineage sorting and past hybridization, accurate species delimitation in Ophioderma was possible using a combination of complementary methods. We propose that these methods, especially multilocus genotype clustering, may be useful to resolve other complex speciation histories.

Annotated species list of Ophiuroidea (Echinodermata) from the Persian Gulf and Gulf of Oman, with new records.

Published records on the Ophiuroidea fauna of the Persian Gulf and Gulf of Oman are scattered in difficult to access journals and books. This study presents a compilation of all published records, complemented with data from new samples. Distribution, habitat and depth in the study area, as well as known Indian Ocean distributions, are included. The taxonomic status of all species was evaluated, critical comments were added as applicable, and several previous records were reassigned to other species. Ophiocoma erinaceus was removed from the fauna of the Persian Gulf and Gulf of Oman, because the only published record was instead Ophiocoma schoenleinii. Previous studies assumed that 46 species of brittle star were known from the study area, but only 38 species were confirmed by this re-assessment, including two new records (Ophiothrix (Ophiothrix) foveolata and Ophiocomella sexradia). Diagnostic features are supplied for difficult to distinguish species. Five species (Macrophiothrix elongata, Amphiura fasciata, Amphiura (Ophiopeltis) hexactis, Amphioplus echinulatus, and Amphioplus seminudus) are so far worldwide known only from the Persian Gulf area. A hexamerous, fissiparous species of Ophiothela that does not concur with any known species was found and may represent a new species. All newly collected species are illustrated with photographs.

A New Fissiparous Brittle Star, Ophiacantha scissionis sp. nov. (Echinodermata, Ophiuroidea, Ophiacanthida), from Jeju Island, Korea.

A new species of Ophiuroidea, Ophiacantha scissionis sp. nov., is described. It was collected from Munseom Island, Jeju-do, South Korea at a depth of 28-36 m, by SCUBA diving. Ophiacantha scissionis has unique and distinct morphological characteristics: small disk, densely covered by small trifid spines, radial shields completely concealed by disk spines, six arms with four arm spines proximally, fissiparous. In addition, this species is clearly distinguished from other species of Ophiacantha by mitochondrial COI. This species is usually hexamerous and reproduces asexually by fissiparity. It is the first report of fissiparous reproduction in the genus Ophiacantha.

Unravelling the origin of the basket stars and their allies (Echinodermata, Ophiuroidea, Euryalida).

Euryalids, which include the spectacular basket stars, form a morphologically aberrant group of brittle stars. Surprisingly, the most recent molecular work found them to be sister to ophiurid brittle stars, thus challenging the traditional dichotomy between euryalids and non-euryalids, and leaving an enormous ghost lineage of more than 100 million years between the oldest unambiguous euryalid fossils and their predicted divergence from ophiurids during the Triassic. Here we examine both previously known and newly collected fossils to explore the evolutionary history of euryalids. A morphology-based phylogenetic estimate confirms the Triassic Aspiduriella as a basal member of the euryalid clade that superficially resembles members of the living ophiurid sister clades. Furthermore, we use lateral arm plates and vertebrae to identify two new Jurassic ophiuroids, Melusinaster alissawhitegluzae and Melusinaster arcusinimicus, as early euryalids that are morphologically intermediate between Aspiduriella and extant euryalids. Our phylogenetic analysis is the first to combine data from completely preserved skeletons and from microfossils in order to bridge morphological and stratigraphical gaps between the sampled taxa. It fills a major gap in the fossil record of euryalids and sets a robust phylogenetic framework to understand the morphological transition from ophiurid-like ancestors to the typical modern euryalids better.

New records of the brittle stars Ophiothela venusta and Ophiactis modesta (Echinodermata: Ophiuroidea) from the northern Persian Gulf, with morphological details.

During a faunistic survey on two shipwrecks in the northern Persian Gulf, several species of ophiuroid were collected in 2015 and 2016. Ophiactis modesta is reported for the first time in the Persian Gulf and Ophiothela venusta is re-described, because it has likely been misidentified in the past, possibly confused with Ophiopsammium semperi. Both species were epizoic: Ophiothela venusta was found on octocorals, Ophiactis modesta on sponges. Ophiothela venusta is characterized by often bright and striking colour patterns with banded arms and large irregular patches on the disc, varying between combinations of yellow, red, blue, orange, black and grey. Its dorsal disc and arms are covered by rugose granules that vary in size and density. In comparison, O. semperi has a denser cover of smaller granules and a subdued uniform colouration. Ophiactis modesta is a hexamerous, fissiparous species that may be mistaken for Ophiactis savignyi. It can be distinguished by its large round to oval oral papilla and its smaller radial shields.

Restructuring higher taxonomy using broad-scale phylogenomics: The living Ophiuroidea.

The power and throughput of next-generation sequencing is instigating a major transformation in our understanding of evolution and classification of life on our planet. The new trees of life are robust and comprehensive. Here we provide a landmark phylogeny of the living ophiuroids and use it as the basis for a major revision of the higher classification of this class of marine invertebrates. We used an exon-capture system to generate a 1484 exon (273kbp) data-matrix from DNA extracted from ethanol-preserved museum samples. We successfully obtained an average of 90% of our target sequence from 576 species spread across the known taxonomic diversity. The topology of the major lineages was robust to taxon sampling, exon-sampling, models and methods. However, estimates of node age were much less precise, varying by about a quarter of mean age. We used a combination of phylogenetic distinctiveness and temporal-banding to guide our revision of the family-level classification. Empirically, we determined that limiting family crown age to 110±10Ma (mid Cretaceous) selected phylogenetically distinct nodes while minimising disruption to the existing taxonomy. The resulting scheme of 32 families and six orders considerably expands the number of higher taxa. The families are generally longitudinally widespread across the world's oceans, although 17 are largely confined to temperate and equatorial latitudes and six to relatively shallow water (less than 1000m depth).

Paedomorphosis as an Evolutionary Driving Force: Insights from Deep-Sea Brittle Stars.

Heterochronic development has been proposed to have played an important role in the evolution of echinoderms. In the class Ophiuroidea, paedomorphosis (retention of juvenile characters into adulthood) has been documented in the families Ophiuridae and Ophiolepididae but not been investigated on a broader taxonomic scale. Historical errors, confusing juvenile stages with paedomorphic species, show the difficulties in correctly identifying the effects of heterochrony on development and evolution. This study presents a detailed analysis of 40 species with morphologies showing various degrees of juvenile appearance in late ontogeny. They are compared to a range of early ontogenetic stages from paedomorphic and non-paedomorphic species. Both quantitative and qualitative measurements are taken and analysed. The results suggest that strongly paedomorphic species are usually larger than other species at comparable developmental stage. The findings support recent notions of polyphyletic origin of the families Ophiuridae and Ophiolepididae. The importance of paedomorphosis and its correct recognition for the practice of taxonomy and phylogeny are emphasized.

A New Morphological Phylogeny of the Ophiuroidea (Echinodermata) Accords with Molecular Evidence and Renders Microfossils Accessible for Cladistics.

Ophiuroid systematics is currently in a state of upheaval, with recent molecular estimates fundamentally clashing with traditional, morphology-based classifications. Here, we attempt a long overdue recast of a morphological phylogeny estimate of the Ophiuroidea taking into account latest insights on microstructural features of the arm skeleton. Our final estimate is based on a total of 45 ingroup taxa, including 41 recent species covering the full range of extant ophiuroid higher taxon diversity and 4 fossil species known from exceptionally preserved material, and the Lower Carboniferous Aganaster gregarius as the outgroup. A total of 130 characters were scored directly on specimens. The tree resulting from the Bayesian inference analysis of the full data matrix is reasonably well resolved and well supported, and refutes all previous classifications, with most traditional families discredited as poly- or paraphyletic. In contrast, our tree agrees remarkably well with the latest molecular estimate, thus paving the way towards an integrated new classification of the Ophiuroidea. Among the characters which were qualitatively found to accord best with our tree topology, we selected a list of potential synapomorphies for future formal clade definitions. Furthermore, an analysis with 13 of the ingroup taxa reduced to the lateral arm plate characters produced a tree which was essentially similar to the full dataset tree. This suggests that dissociated lateral arm plates can be analysed in combination with fully known taxa and thus effectively unlocks the extensive record of fossil lateral arm plates for phylogenetic estimates. Finally, the age and position within our tree implies that the ophiuroid crown-group had started to diversify by the Early Triassic.

Sometimes two arms are enough--an unusual life-stage in brittle stars (Echinodermata: Ophiuroidea).

Off West Africa (Angola-Morocco), benthos samples were collected in the years 2005-2012. These contained 124 specimens of brittle stars with two long arms and three extremely short or absent arms and an elongated, narrow disc. These unusual brittle stars, as well as 33 specimens with five fully developed arms, were identified as Amphiura ungulata. The specimens with unequal arms were juvenile stages, whereas adults had five equal arms. The large number of specimens with unequal arms suggests that this condition is not the result of damage and regeneration, but a normal growth pattern in this species. This study documents the morphology by SEM, amends the species description, and discusses possible explanations for the evolution of this condition. Although brittle star species with unequal arm growth have been reported, this is an extreme case that was unknown before this study.

Redescription of Hemieuryale pustulata von Martens, 1867 (Echinodermata, Ophiuroidea) based on Brazilian specimens, with notes on systematics and habitat association.

We redescribe Hemieuryale pustulata on the basis of 325 specimens obtained from the continental shelf off northeastern and southeastern Brazil. This is the first record of the species for Brazil. We illustrate for the first time details of the dental plate, oral plate, and vertebrae. Few morphological variations were observed in our extensive material. All specimens were associated with the gorgonian Nicella guadalupensis, suggesting an obligatory association in the Brazilian littoral. H. pustulata did not reveal a particular pattern of distribution on its host gorgonian, but larger gorgonians tend to support a larger number of ophiuroids. In addition to expanding the knowledge on morphology and ecology, we extend the known bathymetric distribution of the species, which is now known from 18 to 330 m.

Genetic data, reproduction season and reproductive strategy data support the existence of biological species in Ophioderma longicauda.

Cryptic species are numerous in the marine environment. The brittle star Ophioderma longicauda is composed of six mitochondrial lineages, encompassing brooders, which form a monophyletic group, and broadcasters, from which the brooders are derived. To clarify the species limits within O. longicauda, we compared the reproductive status of the sympatric lineages L1 and L3 (defined after sequencing a portion of the mitochondrial gene COI) during the month of May in Greece. In addition, we genotyped a nuclear marker, intron i51. Each L3 female was brooding, whereas all L1 specimens displayed full gonads, suggesting temporal pre-zygotic isolation between brooders and broadcasters. Statistical differences were found among lineages in morphology and bathymetric distribution. Finally, the intron i51 was polymorphic in L1 (60 individuals), but monomorphic in L3 (109 individuals), confirming the absence of gene flow between brooders and broadcasters. In conclusion, the broadcasting lineage L1 and the brooding lineage L3 are different biological species.

First glimpse into Lower Jurassic deep-sea biodiversity: in situ diversification and resilience against extinction.

Owing to the assumed lack of deep-sea macrofossils older than the Late Cretaceous, very little is known about the geological history of deep-sea communities, and most inference-based hypotheses argue for repeated recolonizations of the deep sea from shelf habitats following major palaeoceanographic perturbations. We present a fossil deep-sea assemblage of echinoderms, gastropods, brachiopods and ostracods, from the Early Jurassic of the Glasenbach Gorge, Austria, which includes the oldest known representatives of a number of extant deep-sea groups, and thus implies that in situ diversification, in contrast to immigration from shelf habitats, played a much greater role in shaping modern deep-sea biodiversity than previously thought. A comparison with coeval shelf assemblages reveals that, at least in some of the analysed groups, significantly more extant families/superfamilies have endured in the deep sea since the Early Jurassic than in the shelf seas, which suggests that deep-sea biota are more resilient against extinction than shallow-water ones. In addition, a number of extant deep-sea families/superfamilies found in the Glasenbach assemblage lack post-Jurassic shelf occurrences, implying that if there was a complete extinction of the deep-sea fauna followed by replacement from the shelf, it must have happened before the Late Jurassic.

Global coordination and standardisation in marine biodiversity through the World Register of Marine Species (WoRMS) and related databases.

The World Register of Marine Species is an over 90% complete open-access inventory of all marine species names. Here we illustrate the scale of the problems with species names, synonyms, and their classification, and describe how WoRMS publishes online quality assured information on marine species. Within WoRMS, over 100 global, 12 regional and 4 thematic species databases are integrated with a common taxonomy. Over 240 editors from 133 institutions and 31 countries manage the content. To avoid duplication of effort, content is exchanged with 10 external databases. At present WoRMS contains 460,000 taxonomic names (from Kingdom to subspecies), 368,000 species level combinations of which 215,000 are currently accepted marine species names, and 26,000 related but non-marine species. Associated information includes 150,000 literature sources, 20,000 images, and locations of 44,000 specimens. Usage has grown linearly since its launch in 2007, with about 600,000 unique visitors to the website in 2011, and at least 90 organisations from 12 countries using WoRMS for their data management. By providing easy access to expert-validated content, WoRMS improves quality control in the use of species names, with consequent benefits to taxonomy, ecology, conservation and marine biodiversity research and management. The service manages information on species names that would otherwise be overly costly for individuals, and thus minimises errors in the application of nomenclature standards. WoRMS' content is expanding to include host-parasite relationships, additional literature sources, locations of specimens, images, distribution range, ecological, and biological data. Species are being categorised as introduced (alien, invasive), of conservation importance, and on other attributes. These developments have a multiplier effect on its potential as a resource for biodiversity research and management. As a consequence of WoRMS, we are witnessing improved communication within the scientific community, and anticipate increased taxonomic efficiency and quality control in marine biodiversity research and management.

Ancient origin of the modern deep-sea fauna.

The origin and possible antiquity of the spectacularly diverse modern deep-sea fauna has been debated since the beginning of deep-sea research in the mid-nineteenth century. Recent hypotheses, based on biogeographic patterns and molecular clock estimates, support a latest Mesozoic or early Cenozoic date for the origin of key groups of the present deep-sea fauna (echinoids, octopods). This relatively young age is consistent with hypotheses that argue for extensive extinction during Jurassic and Cretaceous Oceanic Anoxic Events (OAEs) and the mid-Cenozoic cooling of deep-water masses, implying repeated re-colonization by immigration of taxa from shallow-water habitats. Here we report on a well-preserved echinoderm assemblage from deep-sea (1000-1500 m paleodepth) sediments of the NE-Atlantic of Early Cretaceous age (114 Ma). The assemblage is strikingly similar to that of extant bathyal echinoderm communities in composition, including families and genera found exclusively in modern deep-sea habitats. A number of taxa found in the assemblage have no fossil record at shelf depths postdating the assemblage, which precludes the possibility of deep-sea recolonization from shallow habitats following episodic extinction at least for those groups. Our discovery provides the first key fossil evidence that a significant part of the modern deep-sea fauna is considerably older than previously assumed. As a consequence, most major paleoceanographic events had far less impact on the diversity of deep-sea faunas than has been implied. It also suggests that deep-sea biota are more resilient to extinction events than shallow-water forms, and that the unusual deep-sea environment, indeed, provides evolutionary stability which is very rarely punctuated on macroevolutionary time scales.