Sabine Stöhr

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).

Evolution of mate-choice copying : a dynamic model

Mate-choice copying has recently been demonstrated in several species. Two, not mutually exclusive, explanations for copying have been proposed: it reduces sampling costs and/or error of mate choice. In guppies, Poecilia reticulata, and black grouse, Tetrao tetrix, young females seem most likely to copy. Therefore, copying may teach inexperienced females what attractive males look like. I developed a 2-year dynamic model, to investigate under which conditions a mate-copying strategy might first evolve. An original population of pure choosers was assumed, which was invaded by a mutant female, able to copy during her first mating season, thereby instantly improving her ability to assess male quality. Alternatively, she could either wait and learn by observing males, just as non-copiers may do, but incurring some time costs, or choose, relying on her own abilities. The degree to which copying occurred among these mutant, young, inexperienced females increased with an increasing proportion of old, experienced females in the population, and with decreasing time left until the end of the season. The model demonstrates that mate-choice copying may evolve, when young females are poor at discrimination and need to learn what high-quality males look like. Male quality proved to be unimportant for copying to evolve, as long as there are sufficient differences in quality for mate choice to be meaningful. As with previous models, time constraints are an important assumption for copying to be advantageous over non-copying. Copyright 1998 The Association for the Study of Animal Behaviour. Copyright 1998 The Association for the Study of Animal Behaviour.

The fauna of hydrothermal vents on the Mohn Ridge (North Atlantic)

Abstract The macrofauna of the newly discovered hydrothermal vent field on the Mohn Ridge at 71°N was investigated. Samples were collected during the cruise BIODEEP 2006 using the ROV ‘Bathysaurus’. A total of 180 species-level taxa were identified. The region contains very few vent-endemic species, but some species of Porifera, Crustacea and Mollusca may be vent-associated. Dense aggregations of motile non-vent species such as Heliometra glacialis and Gorgonocephalus eucnemis surrounded the vent area, but the area in general only held small numbers of sedentary animals. Calcareous sponges comprised an unusually high portion of the sponge species found and they constitute one of the first pioneers among the sessile invertebrates settling on these vents. Possible explanations for the structure of the fauna in the region are discussed.

Did vicariance and adaptation drive cryptic speciation and evolution of brooding in Ophioderma longicauda (Echinodermata: Ophiuroidea), a common Atlanto- Mediterranean ophiuroid?

Over the last decade, cryptic speciation has been discovered in an increasing number of taxa. Species complexes are useful models for the understanding of speciation processes. Motivated by the discovery of brooding specimens in the common Atlanto‐Mediterranean broadcast spawning brittle star, Ophioderma longicauda, a recent study revealed the occurrence of divergent mitochondrial lineages. We analysed 218 specimens from 23 locations spread over the geographic range of the species with partial Cytochrome c Oxidase subunit I (COI) sequences. A subset of this sample was also surveyed with the internal transcribed spacer of the ribosomal DNA cluster (nuclear ITS‐1). Our study revealed six highly divergent mitochondrial lineages, and the ITS‐1 data confirmed that they most likely represent a species complex. Geographic ranges, abundances and genetic structures are contrasted among the putative cryptic species. Lineages in which brooding specimens have been found form a monophyletic group and are restricted to the Eastern Mediterranean basin, an oligotrophic zone. A phylogeny‐trait association analysis revealed a phylogenetic signal for low ‘chlorophyll a’ values (our proxy for oligotrophy). An ecological shift related to the hyper oligotrophy of the Eastern Mediterranean region is therefore likely to have played a role in the evolution of brooding. This study revealed that a complex mixture of vicariance, population expansion, adaptive divergence and possibly high local diversification rates resulting from brooding has shaped the evolution of this species complex. The dating analysis showed that these events probably occurred in the Pleistocene epoch.

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.

A non-lekking population of black grouse Tetrao tetrix

In a two-year study of an area in central Sweden a population of Black Grouse Tetrao tetrix was found in which displaying males were not clustered on leks. The study area which covered approximately 44 km(2) contained 19 and 30-35 displaying males in 1995

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).

Megafaunal Community Structure of Andaman Seamounts Including the Back-Arc Basin – A Quantitative Exploration from the Indian Ocean

Species rich benthic communities have been reported from some seamounts, predominantly from the Atlantic and Pacific Oceans, but the fauna and habitats on Indian Ocean seamounts are still poorly known. This study focuses on two seamounts, a submarine volcano (cratered seamount – CSM) and a non-volcano (SM2) in the Andaman Back–arc Basin (ABB), and the basin itself. The main purpose was to explore and generate regional biodiversity data from summit and flank (upper slope) of the Andaman seamounts for comparison with other seamounts worldwide. We also investigated how substratum types affect the megafaunal community structure along the ABB. Underwater video recordings from TeleVision guided Gripper (TVG) lowerings were used to describe the benthic community structure along the ABB and both seamounts. We found 13 varieties of substratum in the study area. The CSM has hard substratum, such as boulders and cobbles, whereas the SM2 was dominated by cobbles and fine sediment. The highest abundance of megabenthic communities was recorded on the flank of the CSM. Species richness and diversity were higher at the flank of the CSM than other are of ABB. Non-metric multi-dimensional scaling (nMDS) analysis of substratum types showed 50% similarity between the flanks of both seamounts, because both sites have a component of cobbles mixed with fine sediments in their substratum. Further, nMDS of faunal abundance revealed two groups, each restricted to one of the seamounts, suggesting faunal distinctness between them. The sessile fauna corals and poriferans showed a significant positive relation with cobbles and fine sediments substratum, while the mobile categories echinoderms and arthropods showed a significant positive relation with fine sediments only.

Morphological diagnosis of the two genetic lineages of Acrocnida brachiata (Echinodermata: Ophiuroidea), with description of a new species

The burrowing brittle-star Acrocnida brachiata has so far been regarded as a single, easily identifiable species. Recent studies showed habitat-related differences in maximum size, life span, breeding time and recruitment between intertidal and subtidal populations, which at first were attributed to environmental effects on individuals within the same species. Molecular data, however, strongly suggested the existence of two distinct lineages and ultimately two cryptic species with clear bathymetric segregation. Morphological evidence had so far not been presented, because any differences were interpreted as intraspecific variation. We collected A. brachiata from intertidal and subtidal habitats at the coast of Brittany, France, and examined 15 specimens of each group externally by SEM. A key character of A. brachiata is that the scales at the edge of the disc and on the ventral side are conically enlarged. Intertidal individuals showed a sparser disc scalation, more spine-like than conical ventral disc scales and spatulate, distally widened arm spines. In addition, we dissected several specimens of different size and examined the internal skeleton by SEM. The oral plates showed a rib-like structure on their abradial face that differs between individuals from either habitat. Subtidal specimens have fewer ribs than intertidal ones. These consistent differences support the existence of two species within A. brachiata . To describe the second species, we needed to establish the identity of A. brachiata . We describe a neotype, because no type material has been preserved since it was first described; it corresponds mainly to subtidal samples. The new species is described as Acrocnida spatulispina sp. nov. The taxonomic status of Acrocnida has been debated over the years with reference to its close affinities with Amphiura chiajei . We compared the species of Acrocnida with A. chiajei and Amphiura filiformis and found that Acrocnida is indeed morphologically similar to A. chiajei , among other characters by a similar oral plate structure, whereas A. filiformis differs greatly from Acrocnida as well as A. chiajei . Most strikingly, it has a different type of oral plate. These findings indicate that fundamental taxonomic changes may need to be made in the family Amphiuridae in the future.