Fossil evidence for the ancient link between clonal fragmentation, six-fold symmetry and an epizoic lifestyle in asterozoan echinoderms.

Asexual reproduction by means of splitting, also called fissiparity, is a common feature in some asterozoan groups, especially in ophiactid brittle stars. Most fissiparous brittle stars show six instead of the usual five rays, live as epibionts on host organisms, and use clonal fragmentation to rapidly colonize secluded habitats and effectively expand the margins of their distribution area. While the biology and ecology of clonal fragmentation are comparatively well understood, virtually nothing is known about the evolution and geological history of that phenomenon. Here, we describe an exceptional fossil of an articulated six-armed brittle star from the Late Jurassic of Germany, showing one body half in the process of regeneration, and assign it to the new species Ophiactis hex sp. nov. Phylogenetic inference shows that the fossil represents the oldest member of the extant family Ophiactidae. Because the Ophiactis hex specimen shows an original six-fold symmetry combined with a morphology typically found in epizoic ophiuroids, in line with recent fissiparous ophiactid relatives, we assume that the regenerating body half is an indication for fissiparity. Ophiactis hex thus shows that fissiparity was established as a means of asexual reproduction in asterozoan echinoderms by the Late Jurassic.

Miniaturization during a Silurian environmental crisis generated the modern brittle star body plan.

Pivotal anatomical innovations often seem to appear by chance when viewed through the lens of the fossil record. As a consequence, specific driving forces behind the origination of major organismal clades generally remain speculative. Here, we present a rare exception to this axiom by constraining the appearance of a diverse animal group (the living Ophiuroidea) to a single speciation event rather than hypothetical ancestors. Fossils belonging to a new pair of temporally consecutive species of brittle stars (Ophiopetagno paicei gen. et sp. nov. and Muldaster haakei gen. et sp. nov.) from the Silurian (444-419 Mya) of Sweden reveal a process of miniaturization that temporally coincides with a global extinction and environmental perturbation known as the Mulde Event. The reduction in size from O. paicei to M. haakei forced a structural simplification of the ophiuroid skeleton through ontogenetic retention of juvenile traits, thereby generating the modern brittle star bauplan.

New fossils of Jurassic ophiurid brittle stars (Ophiuroidea; Ophiurida) provide evidence for early clade evolution in the deep sea.

Understanding of the evolutionary history of the ophiuroids, or brittle stars, is hampered by a patchy knowledge of the fossil record. Especially, the stem members of the living clades are poorly known, resulting in blurry concepts of the early clade evolution and imprecise estimates of divergence ages. Here, we describe new ophiuroid fossil from the Lower Jurassic of France, Luxembourg and Austria and introduce the new taxa Ophiogojira labadiei gen. et sp. nov. from lower Pliensbachian shallow sublittoral deposits, Ophiogojira andreui gen. et sp. nov. from lower Toarcian shallow sublittoral deposits and Ophioduplantiera noctiluca gen. et sp. nov. from late Sinemurian to lower Pliensbachian bathyal deposits. A Bayesian morphological phylogenetic analysis shows that Ophiogojira holds a basal position within the order Ophiurida, whereas Ophioduplantiera has a more crownward position within the ophiurid family Ophiuridae. The position of Ophioduplantiera in the evolutionary tree suggests that family-level divergences within the Ophiurida must have occurred before the late Sinemurian, and that ancient slope environments played an important role in fostering early clade evolution.

A new bathyal ophiacanthid brittle star (Ophiuroidea: Ophiacanthidae) with Caribbean affinities from the Plio-Pleistocene of the Mediterranean.

Identifiable remains of large deep-sea invertebrates are exceedingly rare in the fossil record. Thus, every new discovery adds to a better understanding of ancient deep-sea environments based on direct fossil evidence. Here we describe a collection of dissociated skeletal parts of ophiuroids (brittle stars) from the latest Pliocene to earliest Pleistocene of Sicily, Italy, preserved as microfossils in sediments deposited at shallow bathyal depths. The material belongs to a previously unknown species of ophiacanthid brittle star, Ophiacantha oceani sp. nov. On the basis of morphological comparison of skeletal microstructures, in particular spine articulations and vertebral articular structures of the lateral arm plates, we conclude that the new species shares closest ties with Ophiacantha stellata, a recent species living in the present-day Caribbean at bathyal depths. Since colonization of the deep Mediterranean following the Messinian crisis at the end of the Miocene was only possibly via the Gibraltar Sill, the presence of tropical western Atlantic clades in the Plio-Pleistocene of the Mediterranean suggests a major deep-sea faunal turnover yet to be explored.

Brittle-star mass occurrence on a Late Cretaceous methane seep from South Dakota, USA.

Articulated brittle stars are rare fossils because the skeleton rapidly disintegrates after death and only fossilises intact under special conditions. Here, we describe an extraordinary mass occurrence of the ophiacanthid ophiuroid Brezinacantha tolis gen. et sp. nov., preserved as articulated skeletons from an upper Campanian (Late Cretaceous) methane seep of South Dakota. It is uniquely the first fossil case of a seep-associated ophiuroid. The articulated skeletons overlie centimeter-thick accumulations of dissociated skeletal parts, suggesting lifetime densities of approximately 1000 individuals per m2, persisting at that particular location for several generations. The ophiuroid skeletons on top of the occurrence were preserved intact most probably because of increased methane seepage, killing the individuals and inducing rapid cementation, rather than due to storm-induced burial or slumping. The mass occurrence described herein is an unambiguous case of an autochthonous, dense ophiuroid community that persisted at a particular spot for some time. Thus, it represents a true fossil equivalent of a recent ophiuroid dense bed, unlike other cases that were used in the past to substantiate the claim of a mid-Mesozoic predation-induced decline of ophiuroid dense beds.

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.

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.