A pioneer morphological and genetic study of the intertidal fauna of the Gerlache Strait (Antarctic Peninsula).

The underexplored intertidal ecosystems of Antarctica are facing rapid changes in important environmental factors. Associated with temperature increase, reduction in coastal ice will soon expose new ice-free areas that will be colonized by local or distant biota. To enable detection of future changes in faunal composition, a biodiversity baseline is urgently required. Here, we evaluated intertidal faunal diversity at 13 locations around the Gerlache Strait (western Antarctic Peninsula), using a combination of a quadrat approach, morphological identification and genetic characterization. Our data highlight a community structure comprising four generally distributed and highly abundant species (the flatworm Obrimoposthia wandeli, the bivalve Kidderia subquadrata, and the gastropods Laevilitorina umbilicata and Laevilitorina caliginosa) as well as 79 rarer and less widely encountered species. The most abundant species thrive in the intertidal zone due to their ability to either survive overwinter in situ or to rapidly colonize this zone when conditions allow. In addition, we confirmed the presence of multiple trophic levels at nearly all locations, suggesting that complex inter-specific interactions occur within these communities. Diversity indices contrasted between sampling locations (from 3 to 32 species) and multivariate approaches identified three main groups. This confirms the importance of environmental heterogeneity in shaping diversity patterns within the investigated area. Finally, we provide the first genetic and photographic baseline of the Antarctic intertidal fauna (106 sequences, 137 macrophotographs), as well as preliminary insights on the biogeography of several species. Taken together, these results provide a timely catalyst to assess the diversity and to inform studies of the potential resilience of these intertidal communities.

Trophic markers and biometric measurements in Southern Ocean sea stars (1985-2017).

Sea stars (Echinodermata: Asteroidea) are a key component of Southern Ocean benthos, with 16% of the known sea star species living there. In temperate marine environments, sea stars commonly play an important role in food webs, acting as keystone species. However, trophic ecology and functional role of Southern Ocean sea stars are still poorly known, notably due to the scarcity of large-scale studies. Here, we report 24,332 trophic marker (stable isotopes and elemental contents of C, N, and S of tegument and/or tube feet) and biometric (arm length, disk radius, arm to disk ratio) measurements in 2,456 specimens of sea stars. Samples were collected between 12 January 1985 and 8 October 2017 in numerous locations along the Antarctic littoral and subantarctic islands. The spatial scope of the data set covers a significant portion of the Southern Ocean (47.717° S to 86.273° S; 127.767° W to 162.201° E; depth, 6-5,338 m). The data set contains 133 distinct taxa, including 72 currently accepted species spanning 51 genera, 20 families, and multiple feeding guilds/functional groups (suspension feeders, sediment feeders, omnivores, predators of mobile or sessile prey). For 505 specimens, mitochondrial CO1 genes were sequenced to confirm and/or refine taxonomic identifications, and those sequences are already publicly available through the Barcode of Life Data System. This number will grow in the future, as molecular analyses are still in progress. Overall, thanks to its large taxonomic, spatial, and temporal extent, as well as its integrative nature (combining genetic, morphological, and ecological data), this data set can be of wide interest to Southern Ocean ecologists, invertebrate zoologists, benthic ecologists, and environmental managers dealing with associated areas. Please cite this data paper in research products derived from the data set, which is freely available without copyright restrictions.

Complete distribution of the genus Laevilitorina (Littorinimorpha, Littorinidae) in the Southern Hemisphere: remarks and natural history.

Littorinid snails are present in most coastal areas globally, playing a significant role in the ecology of intertidal communities. Laevilitorina is a marine gastropod genus distributed exclusively in the Southern Hemisphere, with 21 species reported from South America, the sub-Antarctic islands, Antarctica, New Zealand, Australia and Tasmania. Here, an updated database of 21 species generated from a combination of sources is presented: 1) new field sampling data; 2) published records; 3) the Global Biodiversity Information Facility (GBIF) and The Atlas of Living Australia (ALA), to provide a comprehensive description of the known geographic distribution of the genus and detailed occurrences for each of the 21 species. The database includes 813 records (occurrences), 53 from field sampling, 174 from the literature, 128 from GBIF, and 458 from ALA. West Antarctica had the highest species richness (8 species), followed by sub-Antarctic islands of New Zealand (4 species) and the south-east shelf of Australia (4 species). The provinces of Magellan, New Zealand South Island, and sub-Antarctic Islands of the Indian Ocean include two species each. This study specifically highlights reports of L.pygmaea and L.venusta, species that have been almost unrecorded since their description. Recent advances in molecular studies of L.caliginosa showed that this species does not correspond to a widely distributed taxon, but to multiple divergent lineages distributed throughout the Southern Ocean. Ongoing molecular and taxonomic studies are necessary for a better understanding of the diversity and biogeography of this genus.

The Belgica 121 expedition to the Western Antarctic Peninsula: a detailed biodiversity census.

BACKGROUND: This dataset relates to the biodiversity census carried out during the Belgica 121 (B121) expedition to the Western Antarctic Peninsula from February to March 2019. One of the aims of the campaign was to explore the surroundings of the Gerlache Strait and to carry out a detailed biodiversity census focusing on inter- and subtidal shallow-water areas using both classic descriptive marine ecology methods, as well as state-of-the art techniques (habitat mapping, genetics, trophic ecology). The biodiversity census was carried out onboard a nimble research vessel, RV Australis. This dataset will offer access to the raw data on biodiversity occurrences, obtained using a range of methods described in this data paper. NEW INFORMATION: New raw biodiversity data for a poorly-sampled region (Western Antarctic Peninsula) with a special focus on shallow ecosystems.

Facilitating population genomics of non-model organisms through optimized experimental design for reduced representation sequencing.

BACKGROUND: Genome-wide data are invaluable to characterize differentiation and adaptation of natural populations. Reduced representation sequencing (RRS) subsamples a genome repeatedly across many individuals. However, RRS requires careful optimization and fine-tuning to deliver high marker density while being cost-efficient. The number of genomic fragments created through restriction enzyme digestion and the sequencing library setup must match to achieve sufficient sequencing coverage per locus. Here, we present a workflow based on published information and computational and experimental procedures to investigate and streamline the applicability of RRS. RESULTS: In an iterative process genome size estimates, restriction enzymes and size selection windows were tested and scaled in six classes of Antarctic animals (Ostracoda, Malacostraca, Bivalvia, Asteroidea, Actinopterygii, Aves). Achieving high marker density would be expensive in amphipods, the malacostracan target taxon, due to the large genome size. We propose alternative approaches such as mitogenome or target capture sequencing for this group. Pilot libraries were sequenced for all other target taxa. Ostracods, bivalves, sea stars, and fish showed overall good coverage and marker numbers for downstream population genomic analyses. In contrast, the bird test library produced low coverage and few polymorphic loci, likely due to degraded DNA. CONCLUSIONS: Prior testing and optimization are important to identify which groups are amenable for RRS and where alternative methods may currently offer better cost-benefit ratios. The steps outlined here are easy to follow for other non-model taxa with little genomic resources, thus stimulating efficient resource use for the many pressing research questions in molecular ecology.

Is reproductive strategy a key factor in understanding the evolutionary history of Southern Ocean Asteroidea (Echinodermata)?

Life traits such as reproductive strategy can be determining factors of species evolutionary history and explain the resulting diversity patterns. This can be investigated using phylogeographic analyses of genetic units. In this work, the genetic structure of five asteroid genera with contrasting reproductive strategies (brooding: Diplasterias, Notasterias and Lysasterias versus broadcasting: Psilaster and Bathybiaster) was investigated in the Southern Ocean. Over 1,400 mtDNA cytochrome C oxidase subunit I (COI) sequences were analysed using five species delineation methods (ABGD, ASAP, mPTP, sGMYC and mGMYC), two phylogenetic reconstructions (ML and BA), and molecular clock calibrations, in order to examine the weight of reproductive strategy in the observed differences among phylogeographic patterns. We hypothesised that brooding species would show higher levels of genetic diversity and species richness along with a clearer geographic structuring than broadcasting species. In contrast, genetic diversity and species richness were not found to be significantly different between brooders and broadcasters, but broadcasters are less spatially structured than brooders supporting our initial hypothesis and suggesting more complex evolutionary histories associated to this reproductive strategy. Broadcasters' phylogeography can be explained by different scenarios including deep-sea colonisation routes, bipolarity or cosmopolitanism, and sub-Antarctic emergence for the genus Bathybiaster; Antarctic- New Zealand faunal exchanges across the Polar Front for the genus Psilaster. Brooders' phylogeography could support the previously formulated hypothesis of a past trans-Antarctic seaway established between the Ross and the Weddell seas during the Plio-Pleistocene. Our results also show, for the first time, that the Weddell Sea is populated by a mixed asteroid fauna originating from both the East and West Antarctic.

Dwarf brooder versus giant broadcaster: combining genetic and reproductive data to unravel cryptic diversity in an Antarctic brittle star.

Poecilogony, or multiple developmental modes in a single species, is exceedingly rare. Several species described as poecilogenous were later demonstrated to be multiple (cryptic) species with a different developmental mode. The Southern Ocean is known to harbor a high proportion of brooders (Thorson's Rule) but with an increasing number of counter examples over recent years. Here we evaluated poecilogony vs. crypticism in the brittle star Astrotoma agassizii across the Southern Ocean. This species was initially described from South America as a brooder before some pelagic stages were identified in Antarctica. Reproductive and mitochondrial data were combined to unravel geographic and genetic variation of developmental modes. Our results indicate that A. agassizii is composed of seven well-supported and deeply divergent clades (I: Antarctica and South Georgia; II: South Georgia and Sub-Antarctic locations including Kerguelen, Patagonian shelf, and New Zealand; III-VI-VII: Patagonian shelf, IV-V: South Georgia). Two of these clades demonstrated strong size dimorphism when in sympatry and can be linked to differing developmental modes (Clade V: dwarf brooder vs. Clade I: giant broadcaster). Based on their restricted geographic distributions and on previous studies, it is likely that Clades III-VI-VII are brooders. Clade II is composed of different morphological species, A. agassizii and A. drachi, the latter originally used as the outgroup. By integrating morphology, reproductive, and molecular data we conclude that the variation identified in A. agassizii is best described as crypticism rather than poecilogony.

Highly contrasted population genetic structures in a host-parasite pair in the Caribbean Sea.

Evolution and population genetic structure of marine species across the Caribbean Sea are shaped by two complex factors: the geological history and the present pattern of marine currents. Characterizing and comparing the genetic structures of codistributed species, such as host-parasite associations, allow discriminating the relative importance of environmental factors and life history traits that influenced gene flow and demographic events. Using microsatellite and Cytochrome Oxidase I markers, we investigated if a host-parasite pair (the heart urchin Meoma ventricosa and its parasitic pea crab Dissodactylus primitivus) exhibits comparable population genetic structures in the Caribbean Sea and how the observed patterns match connectivity regions from predictive models and other taxa. Highly contrasting patterns were found: the host showed genetic homogeneity across the whole studied area, whereas the parasite displayed significant differentiation at regional and local scales. The genetic diversity of the parasitic crabs (both in microsatellites and COI) was distributed in two main groups, Panama-Jamaica-St Croix on the one hand, and the South-Eastern Caribbean on the other. At a smaller geographical scale, Panamanian and Jamaican parasite populations were genetically more similar, while more genetic differentiation was found within the Lesser Antilles. Both species showed a signature of population expansion during the Quaternary. Some results match predictive models or data from previous studies (e.g., the Western-Eastern dichotomy in the parasite) while others do not (e.g., genetic differentiation within the Lesser Antilles). The sharp dissimilarity of genetic structure of these codistributed species outlines the importance of population expansion events and/or contrasted patterns of gene flow. This might be linked to differences in several life history traits such as fecundity (higher for the host), swimming capacity of larval stages (higher for the parasite), and habitat availability (higher for the host).

Southern Ocean Asteroidea: a proposed update for the Register of Antarctic Marine Species.

BACKGROUND: The Register of Antarctic Marine Species (RAMS, De Broyer et al. 2015) is the regional component of the World Register of Marine Species (WoRMS Editorial Board 2015) in the Southern Ocean. It has been operating for the last ten years, with a special effort devoted towards its completion after the International Polar Year (IPY) in 2007-2008, in the framework of the Census of Antarctic Marine Life (CAML, 2005 - 2010). Its objective is to offer free and open access to a complete register of all known species living in the Southern Ocean, building a workbench of the present taxonomic knowledge for that region. The Antarctic zone defined by this dynamic and community-based tool has been investigated with a particular interest. The Sub-Antarctic zone was a secondary objective during the establishment of the RAMS and is still lacking the impulse of the scientific community for some taxa. NEW INFORMATION: In the present study, more than 13,000 occurrences records of Asteroidea (Echinodermata) have been compiled within the RAMS area of interest and checked against the RAMS species list of sea stars, using WoRMS Taxon Match tool. Few mismatches (basionym mistakes : i.e. original name misspelled or incorrect) were found within the existing list and 97 unregistered species are actually occurring within the RAMS boundaries. After this update, the number of Asteroidea species was increased by around 50%, now reaching 295 accepted species.

The Register of Antarctic Marine Species (RAMS): a ten-year appraisal.

The Register of Antarctic Marine Species (RAMS) is a marine species database that manages an authoritative taxonomic list of species occurring in the Southern Ocean. RAMS links with several other initiatives managing biogeographic or genomics information. The current paper aims to briefly present RAMS and provides an updated snapshot of its contents, in the form of a DarwinCore checklist (available through http://ipt.biodiversity.aq/resource.do?r=rams) and illustrative barplots. Moreover, this article presents a ten year appraisal (since the creation of RAMS). This appraisal first focuses on RAMS bibliometrics. We observed that RAMS was cited (Google Scholar) in 50 distinct publications among which 32 were peer-reviewed in 18 different journals. Three journals (Antarctic Science, Polar Biology, ZooKeys) represent almost 40% of these peer-review publications. The second appraisal focuses on the evolution of new RAMS records. We observed an important decrease in data additions since 2011. As a case study, we focused on an original dataset for a specific group (Asteroidea, Echinodermata). It appears that around one hundred species of asteroids are lacking in RAMS despite the relatively high availability of these data. This suggests that the users' community (or collaborative projects such as AquaRES) could be helpful in order to maintain the RAMS database over the long term.

Parameter Estimations of Dynamic Energy Budget (DEB) Model over the Life History of a Key Antarctic Species: The Antarctic Sea Star Odontaster validus Koehler, 1906.

Marine organisms in Antarctica are adapted to an extreme ecosystem including extremely stable temperatures and strong seasonality due to changes in day length. It is now largely accepted that Southern Ocean organisms are particularly vulnerable to global warming with some regions already being challenged by a rapid increase of temperature. Climate change affects both the physical and biotic components of marine ecosystems and will have an impact on the distribution and population dynamics of Antarctic marine organisms. To predict and assess the effect of climate change on marine ecosystems a more comprehensive knowledge of the life history and physiology of key species is urgently needed. In this study we estimate the Dynamic Energy Budget (DEB) model parameters for key benthic Antarctic species the sea star Odontaster validus using available information from literature and experiments. The DEB theory is unique in capturing the metabolic processes of an organism through its entire life cycle as a function of temperature and food availability. The DEB model allows for the inclusion of the different life history stages, and thus, becomes a tool that can be used to model lifetime feeding, growth, reproduction, and their responses to changes in biotic and abiotic conditions. The DEB model presented here includes the estimation of reproduction handling rules for the development of simultaneous oocyte cohorts within the gonad. Additionally it links the DEB model reserves to the pyloric caeca an organ whose function has long been ascribed to energy storage. Model parameters described a slowed down metabolism of long living animals that mature slowly. O. validus has a large reserve that-matching low maintenance costs- allow withstanding long periods of starvation. Gonad development is continuous and individual cohorts developed within the gonads grow in biomass following a power function of the age of the cohort. The DEB model developed here for O. validus allowed us to increase our knowledge on the ecophysiology of this species, providing new insights on the role of food availability and temperature on its life cycle and reproduction strategy.

Genetic evidence confirms polygamous mating system in a crustacean parasite with multiple hosts.

Mating systems are diverse in animals, notably in crustaceans, but can be inferred from a limited set of parameters. Baeza and Thiel (2007) proposed a model predicting mating systems of symbiotic crustaceans with three host characteristics and the risk of predation. These authors proposed five mating systems, ranging from monogamy to polygynandry (where multiple mating occurs for both genders). Using microsatellite loci, we tested the putatively mating system of the ectoparasite crab Dissodactylus primitivus. We determined the mating frequencies of males and females, parentage assignment (COLONY & GERUD software) as well as the contents of female spermathecae. Our results are globally consistent with the model of Baeza and Thiel and showed, together with previous aquarium experiments, that this ectoparasite evolved a polygamous mating system where males and females move between hosts for mate search. Parentage analyses revealed that polyandry is frequent and concerns more than 60% of clutches, with clutches being fertilized by up to 6 different fathers. Polygyny is supported by the detection of eight males having sired two different broods. We also detected a significant paternity skew in 92% of the multipaternal broods. Moreover, this skew is probably higher than the estimation from the brood because additional alleles were detected in most of spermathecae. This high skew could be explained by several factors as sperm competition or cryptic female choice. Our genetic data, combined with previous anatomic analyses, provide consistent arguments to suggest sperm precedence in D. primitivus.