Incorporation of deep-sea and small-sized species provides new insights into gastropods phylogeny.

The use of phylogeny with uneven or limited taxon sampling may bias our interpretation of organismal evolution, for instance, the origin(s) of the deep-sea animals. The Mollusca is the second most speciose phylum, in which the Gastropoda forms the largest group. However, the currently proposed hypotheses of gastropod phylogeny are mainly based on part of their taxonomic diversity, notably on the large-sized and shallow-water species. In this study, we aimed at correcting this bias by reconstructing the phylogeny with new mitogenomes of deep-sea gastropods including Anatoma sp., Bathysciadiidae sp., Bayerotrochus teramachii, Calliotropis micraulax, Coccocrater sp., Cocculina subcompressa, Lepetodrilus guaymasensis, Peltospira smaragdina, Perotrochus caledonicus, Pseudococculinidae sp., and Shinkailepas briandi. This dataset provided the first reports of the mitogenomes for the Cocculiniformia, three vetigastropod superfamilies: Pleurotomarioidea, Lepetelloidea, and Scissurelloidea, and the neritimorph family Phenacolepadidae. The addition of deep-sea representatives also allowed us to evaluate the evolution of habitat use in gastropods. Our results showed a strongly supported sister-group relationship between the deep-sea lineages Cocculiniformia and Neomphalina. Within the Vetigastropoda, the Pleurotomarioidea was revealed as the sister-group of the remaining vetigastropods. Although this clade was presently restricted to the deep sea, fossil records showed that it has only recently invaded this habitat, thus suggesting that shallow waters was the ancestral habitat for the Vetigastropoda. The deep-sea Lepetelloidea and Lepetodriloidea formed a well-supported clade, with the Scissurelloidea sister to it, suggesting an early transition from shallow water to deep sea in this lineage. In addition, the switch between different chemosynthetic habitats was also observed in deep-sea gastropod lineages, notably in Neomphalina and Lepetelloidea. In both cases, the biogenic substrates appeared as the putative ancestral habitat, confirming the previously proposed hypothesis of a wooden-step to deep-sea vents scenario of evolution of habitat use for these taxa.

Invasion history and demographic processes associated with rapid morphological changes in the Red-whiskered bulbul established on tropical islands.

The Red-whiskered bulbul is a very successful invasive bird species. Morphological differences have been reported among individuals inhabiting the humid and dry coasts of Reunion Island, in a 30-year-old population. This suggests a capacity for rapid local adaptation which could explain the general invasive success of this species. However, the origin and invasion history of this population is unknown. It is therefore not possible to establish with certainty the cause of these morphological differences. Here, we investigated the invasion history of populations of Red-whiskered bulbul established on Reunion Island, Mauritius and Oahu (three geographically similar tropical islands) to assess the link between invasion history and morphological changes in these populations. We first assessed the source(s) of the invasive populations. We then compared the morphology of the individuals between the invasive and native populations and between the dry and humid coasts of invaded islands. Finally, we inferred the invasion history of the invasive populations to investigate the role of neutral processes (e.g. founder effect and drift) on morphology. We found that the invasive populations have a similar origin and that the morphology of the individuals in these populations has diverged in a similar way from the native range, suggesting a convergent adaptation to tropical islands. Like on Reunion, we found differences in morphology between the dry and humid coasts on Mauritius. These morphological differences can be explained by invasion history on Reunion but not on Mauritius. Both neutral evolution and adaptation thus shape the morphology of invasive Red-whiskered bulbuls.

Species are hypotheses: avoid connectivity assessments based on pillars of sand.

Connectivity among populations determines the dynamics and evolution of populations, and its assessment is essential in ecology in general and in conservation biology in particular. The robust basis of any ecological study is the accurate delimitation of evolutionary units, such as populations, metapopulations and species. Yet a disconnect still persists between the work of taxonomists describing species as working hypotheses and the use of species delimitation by molecular ecologists interested in describing patterns of gene flow. This problem is particularly acute in the marine environment where the inventory of biodiversity is relatively delayed, while for the past two decades, molecular studies have shown a high prevalence of cryptic species. In this study, we illustrate, based on marine case studies, how the failure to recognize boundaries of evolutionary-relevant unit leads to heavily biased estimates of connectivity. We review the conceptual framework within which species delimitation can be formalized as falsifiable hypotheses and show how connectivity studies can feed integrative taxonomic work and vice versa. Finally, we suggest strategies for spatial, temporal and phylogenetic sampling to reduce the probability of inadequately delimiting evolutionary units when engaging in connectivity studies.

Next generation sequencing for characterizing biodiversity: promises and challenges.

DNA barcoding approaches are used to describe biodiversity by analysing specimens or environmental samples in taxonomic, phylogenetic and ecological studies. While sharing data among these disciplines would be highly valuable, this remains difficult because of contradictory requirements. The properties making a DNA barcode efficient for specimen identification or species delimitation are hardly reconcilable with those required for a powerful analysis of degraded DNA from environmental samples. The use of next generation sequencing methods open up the way towards the development of new markers (e.g., multilocus barcodes) that would overcome such limitations. However, several challenges should be taken up for coordinating actions at the interface between taxonomy, ecology, molecular biology and bioinformatics in order to develop methods and protocols compatible with both taxonomic and ecological studies.

The coral sea: physical environment, ecosystem status and biodiversity assets.

The Coral Sea, located at the southwestern rim of the Pacific Ocean, is the only tropical marginal sea where human impacts remain relatively minor. Patterns and processes identified within the region have global relevance as a baseline for understanding impacts in more disturbed tropical locations. Despite 70 years of documented research, the Coral Sea has been relatively neglected, with a slower rate of increase in publications over the past 20 years than total marine research globally. We review current knowledge of the Coral Sea to provide an overview of regional geology, oceanography, ecology and fisheries. Interactions between physical features and biological assemblages influence ecological processes and the direction and strength of connectivity among Coral Sea ecosystems. To inform management effectively, we will need to fill some major knowledge gaps, including geographic gaps in sampling and a lack of integration of research themes, which hinder the understanding of most ecosystem processes.

New insights into diversity and evolution of deep-sea Mytilidae (Mollusca: Bivalvia).

Bathymodiolinae mussels have been used as a biological model to better understand the evolutionary origin of faunas associated with deep-sea hydrothermal vents and cold seeps. Most studies to date, however, have sampled with a strong bias towards vent and seep species, mainly because of a lack of knowledge of closely related species from organic falls. Here we reassess the species diversity of deep-sea mussels using two genes and a large taxon sample from the South-Western Pacific. This new taxonomic framework serves as a basis for a phylogenetic investigation of their evolutionary history. We first highlight an unexpected allopatric pattern and suggest that mussels usually reported from organic falls are in fact poorly specialized with regard to their environment. This challenges the adaptive scenarios proposed to explain the diversification of the group. Second, we confirm that deep-sea mussels arose from organic falls and then colonized hydrothermal vents and cold seeps in multiple events. Overall, this study constitutes a new basis for further phylogenetic investigations and a global systematic revision of deep-sea mussels.

Several deep-sea mussels and their associated symbionts are able to live both on wood and on whale falls.

Bathymodiolin mussels occur at hydrothermal vents and cold seeps, where they thrive thanks to symbiotic associations with chemotrophic bacteria. Closely related genera Idas and Adipicola are associated with organic falls, ecosystems that have been suggested as potential evolutionary 'stepping stones' in the colonization of deeper and more sulphide-rich environments. Such a scenario should result from specializations to given environments from species with larger ecological niches. This study provides molecular-based evidence for the existence of two mussel species found both on sunken wood and bones. Each species specifically harbours one bacterial phylotype corresponding to thioautotrophic bacteria related to other bathymodiolin symbionts. Phylogenetic patterns between hosts and symbionts are partially congruent. However, active endocytosis and occurrences of minor symbiont lineages within species which are not their usual host suggest an environmental or horizontal rather than strictly vertical transmission of symbionts. Although the bacteria are close relatives, their localization is intracellular in one mussel species and extracellular in the other, suggesting that habitat choice is independent of the symbiont localization. The variation of bacterial densities in host tissues is related to the substrate on which specimens were sampled and could explain the abilities of host species to adapt to various substrates.

Molecular and ultrastructural characterization of two ascomycetes found on sunken wood off Vanuatu islands in the deep Pacific ocean.

A new genus of a deep-sea ascomycete with one new species, Alisea longicolla, is described based on analyses of 18S and 28S rDNA sequences and morphological characters. A. longicolla was found together with Oceanitis scuticella, on small twigs and sugar cane debris trawled from the bottom of the Pacific Ocean off Vanuatu Islands. Molecular and morphological characters indicate that both fungi are members of Halosphaeriaceae. Within this family, O. scuticella is phylogenetically related to Ascosalsum and shares similar ascospore morphology and appendage ontogeny. The genus Ascosalsum is considered congeneric with Oceanitis and Ascosalsum cincinnatulum, Ascosalsum unicaudatum and Ascosalsum viscidulum are transferred to Oceanitis, an earlier generic name.

Deep-sea benthic communities in the largest oceanic desert are structured by the presence of polymetallic crust.

Based on the specimens collected during three deep-sea cruises, and deposited at the Muséum National d'Histoire Naturelle (MNHN) in Paris, we analysed the diversity of benthic communities within the EEZ of French Polynesia. The literature and the MNHN database allowed us to inventory 471 species of invertebrates, among which 169 were newly described. We mainly found data for Mollusca, Crustacea, Brachiopoda and Crinoidea. We also found samples from other taxa, which still remain unidentified within the collections of the MNHN. Although this inventory is incomplete, we demonstrate that the deep waters of French Polynesia host unique benthic communities and endemic species. Using diversity and multivariate analyses, we show that the deep-sea benthic communities are structured by depth, habitats, geography and also by the presence of polymetallic crust. Furthermore, by focusing on the molluscs of the central area of French Polynesia, we show that the spectrum of shell size differs among deep-sea habitats. Specifically, shells tend to be smaller on encrusted seamounts than on island slopes. Together with the size range of organisms, low abundance, rarity and endemism designate these habitats as sensitive. These results should thus be taken into account in the evaluation of the expected impact of mining activities on biological communities.

Local variation within marinas: Effects of pollutants and implications for invasive species.

Urban structures like marinas are dominant features of our coasts, often hotspots for invasive species. The processes that govern the distribution of invasive species within and between marinas are not well understood. We therefore investigated the impacts of local-scale variability within and between marinas, analysing fouling communities at two zones (inner and outer) within three close marinas in accordance with pollutants recorded in the water and sediment. Communities varied between zones, however no significant differences in abundances of invasive species was recorded. The inner zones contained higher levels of copper and other pollutants and were correlated with lower biodiversity and abundances of many species in comparison to the outer zones. Only the native Ascidiella aspersa was found in greater abundances in the inner zones. This local-scale variability and how it impacts biodiversity is important for consideration for coastal managers in mitigating the build-up of pollutants and spread of invasive species.

Rhodopsin gene evolution in early teleost fishes.

Rhodopsin mediates an essential step in image capture and is tightly associated with visual adaptations of aquatic organisms, especially species that live in dim light environments (e.g., the deep sea). The rh1 gene encoding rhodopsin was formerly considered a single-copy gene in genomes of vertebrates, but increasing exceptional cases have been found in teleost fish species. The main objective of this study was to determine to what extent the visual adaptation of teleosts might have been shaped by the duplication and loss of rh1 genes. For that purpose, homologous rh1/rh1-like sequences in genomes of ray-finned fishes from a wide taxonomic range were explored using a PCR-based method, data mining of public genetic/genomic databases, and subsequent phylogenomic analyses of the retrieved sequences. We show that a second copy of the fish-specific intron-less rh1 is present in the genomes of most anguillids (Elopomorpha), Hiodon alosoides (Osteoglossomorpha), and several clupeocephalan lineages. The phylogenetic analysis and comparisons of alternative scenarios for putative events of gene duplication and loss suggested that fish rh1 was likely duplicated twice during the early evolutionary history of teleosts, with one event coinciding with the hypothesized fish-specific genome duplication and the other in the common ancestor of the Clupeocephala. After these gene duplication events, duplicated genes were maintained in several teleost lineages, whereas some were secondarily lost in specific lineages. Alternative evolutionary schemes of rh1 and comparison with previous studies of gene evolution are also reviewed.

Molecular phylogeny in mytilids supports the wooden steps to deep-sea vents hypothesis.

Molecular data were used to study the diversity of mytilids associated with sunken-woods sampled in the Solomon Islands and discuss the 'wooden steps to deep-sea vent' hypothesis proposed by Distel et al. First, COI data used in a barcoding approach confirm the presence of four distinct species. Analyses of the 18S rDNA and COI dataset then confirmed that these sunken-wood mytilids belonged to a monophyletic group including all species from deep-sea reducing environments. Finally, we analyzed the relationships within this monophyletic group that include the Bathymodiolinae using a COI dataset and a combined analysis of mitochondrial COI and ND4 genes and nuclear rDNA 18S and 28S. Our study supported the 'wooden steps to deep-sea vent' hypothesis: one of the sunken-wood species had a basal position within the Bathymodiolionae, and all described vent and seep mussels included in our analyses were derived taxa within Bathymodiolinae.

Assembly of the mitochondrial genome of the hydrothermal vent crab Segonzacia mesatlantica and detection of potential nuclear pseudogenes.

We assembled the mitogenome of the Bythograeid crab Segonzacia mesatlantica, using long-range amplification of the mitochondrial genome. The mitogenome is 15,521 base pair long (33.8% A, 21.7% C, 10.5% G, 34% T) with 13 protein-coding genes, 2 ribosomal RNA genes, 22 transfer RNAs, and a 624 bp AT-rich region. The gene arrangement is similar to other Brachyuran species. A whole genome shotgun sequencing approach revealed the presence of mitochondrial pseudogenes in the nuclear genome. This fifth mitogenome for a species of Bythograeidae should help resolve the puzzling question of the evolutionary origin of a family limited to deep-sea hydrothermal vents.

WITHDRAWN: An overview of the present status of hospital waste management in Kerman, Iran.

This article has been withdrawn at the request of the author(s) and/or editor. The Publisher apologizes for any inconvenience this may cause. The full Elsevier Policy on Article Withdrawal can be found at http://www.elsevier.com/locate/withdrawalpolicy.

The contrasted evolutionary fates of deep-sea chemosynthetic mussels (Bivalvia, Bathymodiolinae).

Bathymodiolinae are giant mussels that were discovered at hydrothermal vents and harboring chemosynthetic symbionts. Due to their close phylogenetic relationship with seep species and tiny mussels from organic substrates, it was hypothesized that they gradually evolved from shallow to deeper environments, and specialized in decaying organic remains, then in seeps, and finally colonized deep-sea vents. Here, we present a multigene phylogeny that reveals that most of the genera are polyphyletic and/or paraphyletic. The robustness of the phylogeny allows us to revise the genus-level classification. Organic remains are robustly supported as the ancestral habitat for Bathymodiolinae. However, rather than a single step toward colonization of vents and seeps, recurrent habitat shifts from organic substrates to vents and seeps occurred during evolution, and never the reverse. This new phylogenetic framework challenges the gradualist scenarios "from shallow to deep." Mussels from organic remains tolerate a large range of ecological conditions and display a spectacular species diversity contrary to vent mussels, although such habitats are yet underexplored compared to vents and seeps. Overall, our data suggest that for deep-sea mussels, the high specialization to vent habitats provides ecological success in this harsh habitat but also brings the lineage to a kind of evolutionary dead end.

Integrative biology of Idas iwaotakii (Habe, 1958), a 'model species' associated with sunken organic substrates.

The giant bathymodioline mussels from vents have been studied as models to understand the adaptation of organisms to deep-sea chemosynthetic environments. These mussels are closely related to minute mussels associated to organic remains decaying on the deep-sea floor. Whereas biological data accumulate for the giant mussels, the small mussels remain poorly studied. Despite this lack of data for species living on organic remains it has been hypothesized that during evolution, contrary to their relatives from vents or seeps, they did not acquire highly specialized biological features. We aim at testing this hypothesis by providing new biological data for species associated with organic falls. Within Bathymodiolinae a close phylogenetic relationship was revealed between the Bathymodiolus sensu stricto lineage (i.e. "thermophilus" lineage) which includes exclusively vent and seep species, and a diversified lineage of small mussels, attributed to the genus Idas, that includes mostly species from organic falls. We selected Idas iwaotakii (Habe, 1958) from this latter lineage to analyse population structure and to document biological features. Mitochondrial and nuclear markers reveal a north-south genetic structure at an oceanic scale in the Western Pacific but no structure was revealed at a regional scale or as correlated with the kind of substrate or depth. The morphology of larval shells suggests substantial dispersal abilities. Nutritional features were assessed by examining bacterial diversity coupled by a microscopic analysis of the digestive tract. Molecular data demonstrated the presence of sulphur-oxidizing bacteria resembling those identified in other Bathymodiolinae. In contrast with most Bathymodiolus s.s. species the digestive tract of I. iwaotakii is not reduced. Combining data from literature with the present data shows that most of the important biological features are shared between Bathymodiolus s.s. species and its sister-lineage. However Bathymodiolus s.s. species are ecologically more restricted and also display a lower species richness than Idas species.

Is the species flock concept operational? The Antarctic shelf case.

There has been a significant body of literature on species flock definition but not so much about practical means to appraise them. We here apply the five criteria of Eastman and McCune for detecting species flocks in four taxonomic components of the benthic fauna of the Antarctic shelf: teleost fishes, crinoids (feather stars), echinoids (sea urchins) and crustacean arthropods. Practical limitations led us to prioritize the three historical criteria (endemicity, monophyly, species richness) over the two ecological ones (ecological diversity and habitat dominance). We propose a new protocol which includes an iterative fine-tuning of the monophyly and endemicity criteria in order to discover unsuspected flocks. As a result nine « full ¼ species flocks (fulfilling the five criteria) are briefly described. Eight other flocks fit the three historical criteria but need to be further investigated from the ecological point of view (here called "core flocks"). The approach also shows that some candidate taxonomic components are no species flocks at all. The present study contradicts the paradigm that marine species flocks are rare. The hypothesis according to which the Antarctic shelf acts as a species flocks generator is supported, and the approach indicates paths for further ecological studies and may serve as a starting point to investigate the processes leading to flock-like patterning of biodiversity.

Deep-sea origin and in-situ diversification of chrysogorgiid octocorals.

The diversity, ubiquity and prevalence in deep waters of the octocoral family Chrysogorgiidae Verrill, 1883 make it noteworthy as a model system to study radiation and diversification in the deep sea. Here we provide the first comprehensive phylogenetic analysis of the Chrysogorgiidae, and compare phylogeny and depth distribution. Phylogenetic relationships among 10 of 14 currently-described Chrysogorgiidae genera were inferred based on mitochondrial (mtMutS, cox1) and nuclear (18S) markers. Bathymetric distribution was estimated from multiple sources, including museum records, a literature review, and our own sampling records (985 stations, 2345 specimens). Genetic analyses suggest that the Chrysogorgiidae as currently described is a polyphyletic family. Shallow-water genera, and two of eight deep-water genera, appear more closely related to other octocoral families than to the remainder of the monophyletic, deep-water chrysogorgiid genera. Monophyletic chrysogorgiids are composed of strictly (Iridogorgia Verrill, 1883, Metallogorgia Versluys, 1902, Radicipes Stearns, 1883, Pseudochrysogorgia Pante & France, 2010) and predominantly (Chrysogorgia Duchassaing & Michelotti, 1864) deep-sea genera that diversified in situ. This group is sister to gold corals (Primnoidae Milne Edwards, 1857) and deep-sea bamboo corals (Keratoisidinae Gray, 1870), whose diversity also peaks in the deep sea. Nine species of Chrysogorgia that were described from depths shallower than 200 m, and mtMutS haplotypes sequenced from specimens sampled as shallow as 101 m, suggest a shallow-water emergence of some Chrysogorgia species.

Species from Darwin onward.

Controversy regarding the species problem has been going on for many decades and no consensus has ever been reached about what a "species" really is and how best to define the concept. De Queiroz (1998) introduced a distinction between two aspects of this problem: on the one hand, the definition proper, and on the other, the criteria allowing biologists to recognize species in practice. This distinction is a first step on the way toward a solution of the problem. In the present paper, we show that de Queiroz's distinction is made possible by the radical theoretical change introduced by Darwin. We emphasize that the species problem did not appear in the 20th century, but long before, and that Darwin addresses it indirectly in the Origin of Species. It might seem paradoxical to refer to Darwin's views about species, because they are usually considered as unclear. However, we propose that an analysis of these views in the context of Darwin's own theory of evolution might reveal how a definition of the concept of species is made possible by being anchored to the very theory of evolution. To this aim, we present a plausible reconstruction of Darwin's implicit conception of species and show how this conception fits with the debates on species that took place in the 18th and 19th centuries. We then turn to today's biology and show what changes Darwin's implicit conception of species has brought about relative to the species concept and species delimitation.

Symbioses between deep-sea mussels (Mytilidae: Bathymodiolinae) and chemosynthetic bacteria: diversity, function and evolution.

Mussels of the subfamily Bathymodiolinae thrive around chimneys emitting hot fluids at deep sea hydrothermal vents, as well as at cold seeps and on sunken organic debris (sunken wood, whale falls). Despite the absence of light-driven primary production in these deep-sea ecosystems, mussels succeed reaching high biomasses in these harsh conditions thanks to chemosynthetic, carbon-fixing bacterial symbionts located in their gill tissue. Since the discovery of mussel symbioses about three decades ago our knowledge has increased, yet new findings are published regularly regarding their diversity, role and evolution. This article attempts to summarize current knowledge about symbiosis in Bathymodiolinae, focusing on mussel species for which information is available regarding both hosts and symbionts. Moreover, new data obtained from small mussels inhabiting sunken woods around the Philippines are provided. Indeed, mussel species from organic falls remain poorly studied compared to their vent and seep relatives despite their importance for the understanding of the evolution of symbiosis in the subfamily Bathymodiolinae.