Ben Thuy

Global Diversity of Brittle Stars (Echinodermata: Ophiuroidea)

This review presents a comprehensive overview of the current status regarding the global diversity of the echinoderm class Ophiuroidea, focussing on taxonomy and distribution patterns, with brief introduction to their anatomy, biology, phylogeny, and palaeontological history. A glossary of terms is provided. Species names and taxonomic decisions have been extracted from the literature and compiled in The World Ophiuroidea Database, part of the World Register of Marine Species (WoRMS). Ophiuroidea, with 2064 known species, are the largest class of Echinodermata. A table presents 16 families with numbers of genera and species. The largest are Amphiuridae (467), Ophiuridae (344 species) and Ophiacanthidae (319 species). A biogeographic analysis for all world oceans and all accepted species was performed, based on published distribution records. Approximately similar numbers of species were recorded from the shelf (n = 1313) and bathyal depth strata (1297). The Indo-Pacific region had the highest species richness overall (825 species) and at all depths. Adjacent regions were also relatively species rich, including the North Pacific (398), South Pacific (355) and Indian (316) due to the presence of many Indo-Pacific species that partially extended into these regions. A secondary region of enhanced species richness was found in the West Atlantic (335). Regions of relatively low species richness include the Arctic (73 species), East Atlantic (118), South America (124) and Antarctic (126).

Phylogenomic Resolution of the Class Ophiuroidea Unlocks a Global Microfossil Record

Our understanding of the origin, evolution, and biogeography of seafloor fauna is limited because we have insufficient spatial and temporal data to resolve underlying processes. The abundance and wide distribution of modern and disarticulated fossil Ophiuroidea, including brittle stars and basket stars, make them an ideal model system for global marine biogeography if we have the phylogenetic framework necessary to link extant and fossil morphology in an evolutionary context. Here we construct a phylogeny from a highly complete 425-gene, 61-taxa transcriptome-based data set covering 15 of the 18 ophiuroid families and representatives of all extant echinoderm classes. We calibrate our phylogeny with a series of novel fossil discoveries from the early Mesozoic. We confirm the traditional paleontological view that ophiuroids are sister to the asteroids and date the crown group Ophiuroidea to the mid-Permian (270 ± 30 mega-annum). We refute all historical classification schemes of the Ophiuroidea based on gross structural characters but find strong congruence with schemes based on lateral arm plate microstructure and the temporal appearance of various plate morphologies in the fossil record. The verification that these microfossils contain phylogenetically informative characters unlocks their potential to advance our understanding of marine biogeographical processes.

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.

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.

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 worlds oceans, although 17 are largely confined to temperate and equatorial latitudes and six to relatively shallow water (less than 1000m depth).

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.

The pitfalls of extrapolating modern depth ranges to fossil assemblages: new insights from Middle Jurassic brittle stars (Echinodermata: Ophiuroidea) from Switzerland

Depth reconstruction based on the extrapolation to fossil assemblages of present-day depth ranges of closely related groups is one of the most widely used approaches in palaeobathymetry. Here, we assess the ophiuroid fauna of the Bajocian to Bathonian (Middle Jurassic) Hauptrogenstein Formation and coeval formations in Switzerland with respect to the depth ranges of extant members of the groups identified. In addition to previously known taxa, we describe three new species, one assignable to the extant genus Ophiotholia within the family Ophiomycetidae (resurrected herein), and two belonging to new genera within the family Ophiacanthidae. The Hauptrogenstein ophiuroid fauna is shown to display a striking similarity to modern bathyal brittle star assemblages. In combination with taphonomic evidence of the autochthonous nature of the ophiuroid occurrences, the direct extrapolation of present-day depth ranges, as performed in various previous studies, would imply the Hauptrogenstein Formation to have been deposited in a bathyal setting. This, however, is in stark contrast with the generally accepted, sedimentology-based concept of this unit as a very shallow, high-energy carbonate platform deposit. Evidently, direct extrapolation of modern depth distribution patterns fails to provide a reliable palaeobathymetrical assessment here. In this respect, the case of the Hauptrogenstein ophiuroid fauna serves as a remarkable example to stress the pitfalls of assemblage-based palaeodepth estimates: (1) depth distribution patterns might not be controlled by water depth, or not even by a factor directly related to depth, (2) habitat preferences of a group might have changed through time without being reflected by morphological modifications and (3) shifts in depth ranges might occur due to the rise or extinction of groups interacting with the organism in question. Thus, extrapolation of present-day depth ranges to ancient communities can only produce reliable palaeodepth estimates if there is a mechanistic explanation why organisms are confined to a particular depth.

New ophiacanthid brittle stars (Echinodermata: Ophiuroidea) from the Upper Triassic of Japan: first insights into the origin and evolution of an extant deep-sea group

Well preserved, articulated brittle star skeletons from the early Carnian (early Late Triassic) Halobia Shales of Yamaguchi, Japan, are described as a new genus and species of the family Ophiacanthidae: Leadagmara gracilispina. The new form constitutes the oldest unequivocal representative of this extant ophiuroid family, significantly extending its stratigraphical range. The first cladistic analysis of the Ophiacanthidae including all extant genera suggests L. gracilispina holds a basal, but not the basalmost, position within the family, forming a sister group with the extant Ophiomedea and Ophiopristis to the other ophiacanthids except for Ophiologimus, which itself is sister to all other ophiacanthids. These results imply that ophiacanthids originated in the pre-Carnian. A critical re-evaluation of the ophiacanthid fossil record reveals that the vast majority of records consist of dissociated lateral arm plates, which provide no insights into divergence times of the lineages. Among the few records of articulated specimens, only Inexpectacantha acrobatica from the Pliensbachian of France proved sufficiently well-known to yield phylogenetically relevant information. Its position within the ophiacanthid clade indicates that more than half of the extant ophiacanthid lineages, in particular the former ophiacanthinids and ophiotominids, must have diverged by the Pliensbachian. http://zoobank.org/urn:lsid:zoobank.org:pub:F64E44C3-7275-4EC8-B048-4C7B2C6D57EF

A starfish bed in the Middle Miocene Grand Bay Formation of Carriacou, The Grenadines (West Indies)

A starfish bed in the Middle Miocene Grand Bay Formation of Carriacou, The Grenadines (West Indies)

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.