Phylogenomic analyses reveal a single deep-water colonisation in Patellogastropoda.

Patellogastropoda, the true limpets, is a major group of gastropods widely distributed in marine habitats from the intertidal to deep sea. Though important for understanding their evolutionary radiation, the phylogenetic relationships among the patellogastropod families have always been challenging to reconstruct, with contradictory results likely due to insufficient sampling. Here, we obtained mitogenomic and phylogenomic data (transcriptomic or genomic) from six species representing the three predominantly deep-water patellogastropod families: Lepetidae, Neolepetopsidae, and Pectinodontidae. By using various phylogenetic methods, we show that mitogenome phylogeny recovers monophyly of eight families in most of the trees, though the relationships among families remain contentious. Meanwhile, a more robust family-level topology consistent with morphology was achieved by phylogenomics. This also reveals that these mainly deep-water families are monophyletic, suggesting a single colonisation of the deep water around the Jurassic. We also found a lack of significant correlation between genome size and habitat depth, despite some deep-water species exhibiting larger genome sizes. Our phylogenomic tree provides a stable phylogenetic backbone for Patellogastropoda that includes seven of the nine recognized families and paves the way for future evolutionary analyses in this major group of molluscs.

High Abundances of Microplastic Pollution in Deep-Sea Sediments: Evidence from Antarctica and the Southern Ocean.

Plastic pollution in Antarctica and the Southern Ocean has been recorded in scientific literature since the 1980s; however, the presence of microplastic particles (<5 mm) is less understood. Here, we aimed to determine whether microplastic accumulation would vary among Antarctic and Southern Ocean regions through studying 30 deep-sea sediment cores. Additionally, we aimed to highlight whether microplastic accumulation was related to sample depth or the sediment characteristics within each core. Sediment cores were digested and separated using a high-density sodium polytungstate solution (SPT) and microplastic particles were identified using micro-Fourier-transform infrared spectroscopy (µFTIR). Microplastic pollution was found in 93% of the sediment cores (28/30). The mean (±SE) microplastics per gram of sediment was 1.30 ± 0.51, 1.09 ± 0.22, and 1.04 ± 0.39 MP/g, for the Antarctic Peninsula, South Sandwich Islands, and South Georgia, respectively. Microplastic fragment accumulation correlated significantly with the percentage of clay within cores, suggesting that microplastics have similar dispersion behavior to low density sediments. Although no difference in microplastic abundance was found among regions, the values were much higher in comparison to less remote ecosystems, suggesting that the Antarctic and Southern Ocean deep-sea accumulates higher numbers of microplastic pollution than previously expected.

Cryptic niche switching in a chemosymbiotic gastropod.

Life stages of some animals, including amphibians and insects, are so different that they have historically been seen as different species. 'Metamorphosis' broadly encompasses major changes in organism bodies and, importantly, concomitant shifts in trophic strategies. Many marine animals have a biphasic lifestyle, with small pelagic larvae undergoing one or more metamorphic transformations before settling into a permanent, adult morphology on the benthos. Post-settlement, the hydrothermal vent gastropod Gigantopelta chessoia experiences a further, cryptic metamorphosis at body sizes around 5-7 mm. The terminal adult stage is entirely dependent on chemoautotrophic symbionts; smaller individuals do not house symbionts and presumably depend on grazing. Using high-resolution X-ray microtomography to reconstruct the internal organs in a growth series, we show that this sudden transition in small but sexually mature individuals dramatically reconfigures the organs, but is in no way apparent from external morphology. We introduce the term 'cryptometamorphosis' to identify this novel phenomenon of a major body change and trophic shift, not related to sexual maturity, transforming only the internal anatomy. Understanding energy flow in ecosystems depends on the feeding ecology of species; the present study highlights the possibility for adult animals to make profound shifts in biology that influence energy dynamics.

By more ways than one: Rapid convergence at hydrothermal vents shown by 3D anatomical reconstruction of Gigantopelta (Mollusca: Neomphalina).

BACKGROUND: Extreme environments prompt the evolution of characteristic adaptations. Yet questions remain about whether radiations in extreme environments originate from a single lineage that masters a key adaptive pathway, or if the same features can arise in parallel through convergence. Species endemic to deep-sea hydrothermal vents must accommodate high temperature and low pH. The most successful vent species share a constrained pathway to successful energy exploitation: hosting symbionts. The vent-endemic gastropod genus Gigantopelta, from the Southern and Indian Oceans, shares unusual features with a co-occurring peltospirid, the 'scaly-foot gastropod' Chrysomallon squamiferum. Both are unusually large for the clade and share other adaptive features such as a prominent enlarged trophosome-like oesophageal gland, not found in any other vent molluscs. RESULTS: Transmission electron microscopy confirmed endosymbiont bacteria in the oesophageal gland of Gigantopelta, as also seen in Chrysomallon. They are the only known members of their phylum in vent ecosystems hosting internal endosymbionts; other vent molluscs host endosymbionts in or on their gills, or in the mantle cavity. A five-gene phylogenetic reconstruction demonstrated that Gigantopelta and Chrysomallon are not phylogenetically sister-taxa, despite their superficial similarity. Both genera have specialist adaptations to accommodate internalised endosymbionts, but with anatomical differences that indicate separate evolutionary origins. Hosting endosymbionts in an internal organ within the host means that all resources required by the bacteria must be supplied by the animal, rather than directly by the vent fluid. Unlike Chrysomallon, which has an enlarged oesophageal gland throughout post-settlement life, the oesophageal gland in Gigantopelta is proportionally much smaller in juveniles and the animals likely undergo a trophic shift during ontogeny. The circulatory system is hypertrophied in both but the overall size is smaller in Gigantopelta. In contrast with Chrysomallon, Gigantopelta possesses true ganglia and is gonochoristic. CONCLUSIONS: Key anatomical differences between Gigantopelta and Chrysomallon demonstrate these two genera acquired a similar way of life through independent and convergent adaptive pathways. What appear to be the holobiont's adaptations to an extreme environment, are driven by optimising bacteria's access to vent nutrients. By comparing Gigantopelta and Chrysomallon, we show that metazoans are capable of rapidly and repeatedly evolving equivalent anatomical adaptations and close-knit relationships with chemoautotrophic bacteria, achieving the same end-product through parallel evolutionary trajectories.

Commonness and rarity in the marine biosphere.

Explaining patterns of commonness and rarity is fundamental for understanding and managing biodiversity. Consequently, a key test of biodiversity theory has been how well ecological models reproduce empirical distributions of species abundances. However, ecological models with very different assumptions can predict similar species abundance distributions, whereas models with similar assumptions may generate very different predictions. This complicates inferring processes driving community structure from model fits to data. Here, we use an approximation that captures common features of "neutral" biodiversity models--which assume ecological equivalence of species--to test whether neutrality is consistent with patterns of commonness and rarity in the marine biosphere. We do this by analyzing 1,185 species abundance distributions from 14 marine ecosystems ranging from intertidal habitats to abyssal depths, and from the tropics to polar regions. Neutrality performs substantially worse than a classical nonneutral alternative: empirical data consistently show greater heterogeneity of species abundances than expected under neutrality. Poor performance of neutral theory is driven by its consistent inability to capture the dominance of the communities' most-abundant species. Previous tests showing poor performance of a neutral model for a particular system often have been followed by controversy about whether an alternative formulation of neutral theory could explain the data after all. However, our approach focuses on common features of neutral models, revealing discrepancies with a broad range of empirical abundance distributions. These findings highlight the need for biodiversity theory in which ecological differences among species, such as niche differences and demographic trade-offs, play a central role.

The heart of a dragon: 3D anatomical reconstruction of the 'scaly-foot gastropod' (Mollusca: Gastropoda: Neomphalina) reveals its extraordinary circulatory system.

INTRODUCTION: The 'scaly-foot gastropod' (Chrysomallon squamiferum Chen et al., 2015) from deep-sea hydrothermal vent ecosystems of the Indian Ocean is an active mobile gastropod occurring in locally high densities, and it is distinctive for the dermal scales covering the exterior surface of its foot. These iron-sulfide coated sclerites, and its nutritional dependence on endosymbiotic bacteria, are both noted as adaptations to the extreme environment in the flow of hydrogen sulfide. We present evidence for other adaptations of the 'scaly-foot gastropod' to life in an extreme environment, investigated through dissection and 3D tomographic reconstruction of the internal anatomy. RESULTS: Our anatomical investigations of juvenile and adult specimens reveal a large unganglionated nervous system, a simple and reduced digestive system, and that the animal is a simultaneous hermaphrodite. We show that Chrysomallon squamiferum relies on endosymbiotic bacteria throughout post-larval life. Of particular interest is the circulatory system: Chrysomallon has a very large ctenidium supported by extensive blood sinuses filled with haemocoel. The ctenidium provides oxygen for the host but the circulatory system is enlarged beyond the scope of other similar vent gastropods. At the posterior of the ctenidium is a remarkably large and well-developed heart. Based on the volume of the auricle and ventricle, the heart complex represents approximately 4 % of the body volume. This proportionally giant heart primarily sucks blood through the ctenidium and supplies the highly vascularised oesophageal gland. Thus we infer the elaborate cardiovascular system most likely evolved to oxygenate the endosymbionts in an oxygen poor environment and/or to supply hydrogen sulfide to the endosymbionts. CONCLUSIONS: This study exemplifies how understanding the autecology of an organism can be enhanced by detailed investigation of internal anatomy. This gastropod is a large and active species that is abundant in its hydrothermal vent field ecosystem. Yet all of its remarkable features-protective dermal sclerites, circulatory system, high fecundity-can be viewed as adaptations beneficial to its endosymbiont microbes. We interpret these results to show that, as a result of specialisation to resolve energetic needs in an extreme chemosynthetic environment, this dramatic dragon-like species has become a carrying vessel for its bacteria.

The discovery of new deep-sea hydrothermal vent communities in the southern ocean and implications for biogeography.

Since the first discovery of deep-sea hydrothermal vents along the Galápagos Rift in 1977, numerous vent sites and endemic faunal assemblages have been found along mid-ocean ridges and back-arc basins at low to mid latitudes. These discoveries have suggested the existence of separate biogeographic provinces in the Atlantic and the North West Pacific, the existence of a province including the South West Pacific and Indian Ocean, and a separation of the North East Pacific, North East Pacific Rise, and South East Pacific Rise. The Southern Ocean is known to be a region of high deep-sea species diversity and centre of origin for the global deep-sea fauna. It has also been proposed as a gateway connecting hydrothermal vents in different oceans but is little explored because of extreme conditions. Since 2009 we have explored two segments of the East Scotia Ridge (ESR) in the Southern Ocean using a remotely operated vehicle. In each segment we located deep-sea hydrothermal vents hosting high-temperature black smokers up to 382.8°C and diffuse venting. The chemosynthetic ecosystems hosted by these vents are dominated by a new yeti crab (Kiwa n. sp.), stalked barnacles, limpets, peltospiroid gastropods, anemones, and a predatory sea star. Taxa abundant in vent ecosystems in other oceans, including polychaete worms (Siboglinidae), bathymodiolid mussels, and alvinocaridid shrimps, are absent from the ESR vents. These groups, except the Siboglinidae, possess planktotrophic larvae, rare in Antarctic marine invertebrates, suggesting that the environmental conditions of the Southern Ocean may act as a dispersal filter for vent taxa. Evidence from the distinctive fauna, the unique community structure, and multivariate analyses suggest that the Antarctic vent ecosystems represent a new vent biogeographic province. However, multivariate analyses of species present at the ESR and at other deep-sea hydrothermal vents globally indicate that vent biogeography is more complex than previously recognised.

The Early Miocene Cape Melville Formation fossil assemblage and the evolution of modern Antarctic marine communities.

The fossil community from the Early Miocene Cape Melville Formation (King George Island, Antarctica) does not show the archaic retrograde nature of modern Antarctic marine communities, despite evidence, such as the presence of dropstones, diamictites and striated rocks, that it was deposited in a glacial environment. Unlike modern Antarctic settings, and the upper units of the Eocene La Meseta Formation on Seymour Island, Antarctica, which are 10 million years older, the Cape Melville Formation community is not dominated by sessile suspension feeding ophiuroids, crinoids or brachiopods. Instead, it is dominated by infaunal bivalves, with a significant component of decapods, similar to present day South American settings. It is possible that the archaic retrograde structure of the modern community did not fully evolve until relatively recently, maybe due to factors such as further cooling and isolation of the continent leading to glaciations, which resulted in a loss of shallow shelf habitats.

First insights into the biodiversity and biogeography of the Southern Ocean deep sea.

Shallow marine benthic communities around Antarctica show high levels of endemism, gigantism, slow growth, longevity and late maturity, as well as adaptive radiations that have generated considerable biodiversity in some taxa. The deeper parts of the Southern Ocean exhibit some unique environmental features, including a very deep continental shelf and a weakly stratified water column, and are the source for much of the deep water in the world ocean. These features suggest that deep-sea faunas around the Antarctic may be related both to adjacent shelf communities and to those in other oceans. Unlike shallow-water Antarctic benthic communities, however, little is known about life in this vast deep-sea region. Here, we report new data from recent sampling expeditions in the deep Weddell Sea and adjacent areas (748-6,348 m water depth) that reveal high levels of new biodiversity; for example, 674 isopods species, of which 585 were new to science. Bathymetric and biogeographic trends varied between taxa. In groups such as the isopods and polychaetes, slope assemblages included species that have invaded from the shelf. In other taxa, the shelf and slope assemblages were more distinct. Abyssal faunas tended to have stronger links to other oceans, particularly the Atlantic, but mainly in taxa with good dispersal capabilities, such as the Foraminifera. The isopods, ostracods and nematodes, which are poor dispersers, include many species currently known only from the Southern Ocean. Our findings challenge suggestions that deep-sea diversity is depressed in the Southern Ocean and provide a basis for exploring the evolutionary significance of the varied biogeographic patterns observed in this remote environment.

Vulcanolepas scotiaensis sp. nov., a new deep-sea scalpelliform barnacle (Eolepadidae: Neolepadinae) from hydrothermal vents in the Scotia Sea, Antarctica.

A new deep-sea stalked barnacle, Vulcanolepas scotiaensis sp. nov. is described from hydrothermal vents at depths of 2400-2600 metres along segments of the East Scotia Ridge and from 1400 metres in the Kemp Caldera. Both locations are areas of volcanic activity that lie on the Antarctic-South American Ocean Ridge complex near the South Sandwich Islands. This discovery confirms a wide distribution in southern seas for Vulcanolepas, complementing the previous records from deep-sea vents in the Lau Basin and Kermadec Ridge in the southwest Pacific, and the Pacific Antarctic Ridge in the southeast Pacific. V. scotiaensis sp. nov., the third described species of Vulcanolepas shows an extraordinary range in morphology, requiring a reassessment of the original diagnosis for Vulcanolepas. Although the morphological envelope of V. scotiaensis sp. nov. includes representatives with a peduncle to capitulum ratio similar to that observed in most neolepadines, the peduncle generally shows greater proportional length than in species in any neolepadine genus except Leucolepas; it is distinguished from other species of Vulcanolepas by a broader capitulum, much smaller imbricating scales on the peduncle and more ornamented capitulum plates. The morphological diversity of V. scotiaensis sp. nov. is interpreted as having arisen due to abrupt changes in water temperature.LSID: urn:lsid:zoobank.org:act:AA2AFDA5-0B08-466A-A584-D3FDBDE9DA61.

Mollusca collected by Agassiz trawl from the 2016 SO-AntEco (JR15005) expedition to the South Orkney Islands, Antarctica - data.

Background: This dataset contributes to the knowledge of macro- and megafaunal Mollusca associated with a range of benthic habitat types in the South Orkney Islands, Antarctica, an exceptionally diverse region of the Southern Ocean. The information presented is derived from Agassiz trawl samples collected on the archipelago's shelf plateau and slope, within and outside of the South Orkney Islands Southern Shelf Marine Protected Area (SOISS MPA). Sampling was conducted in the framework of the British Antarctic Survey/SCAR "South Orkneys - State of the Antarctic Ecosystem" (SO-AntEco) project aboard RRS James Clark Ross during expedition JR15005 in Austral summer 2016. This dataset is published by the British Antarctic Survey under the licence CC-BY 4.0. We would appreciate it if you could follow the guidelines from the SCAR Data Policy (SCAR 2023) when using the data. If you have any questions regarding this dataset, do not hesitate to contact us via the contact information provided in the metadata or via data-biodiversity-aq@naturalsciences.be. Issues with the dataset can be reported at https://github.com/biodiversity-aq/data-publication/. This dataset is part of the Biodiversity, Evolution and Adaptation Project of the Environmental Change and Evolution Program of the British Antarctic Survey. The cruise report of the expedition is available at https://www.bodc.ac.uk/resources/inventories/cruise_inventory/reports/jr15005.pdf. New information: We report occurrences of Mollusca from individual samples taken with a 2 m-wide Agassiz trawl (AGT) in the framework of the February - March 2016 research expedition JR15005 of RRS James Clark Ross to the SOISS MPA and adjacent shelf and slope areas. Of 78 successful AGT deployments, 44 trawls at depths ranging from 235-2194 m yielded living Mollusca, totalling 2276 individuals, 67 morphospecies and 163 distributional records. One hundred and fifteen empty shells were also collected and recorded in the dataset. Three morphospecies (one Bivalvia and two Gastropoda) were sampled exclusively as empty shells, yielding a total of 70 morphospecies and 2391 specimens represented in the dataset. All specimens were preserved in 96% undenatured ethanol and are stored as vouchers in the collections of the British Antarctic Survey (BAS), Cambridge, United Kingdom. The publication of this dataset aims at increasing the knowledge on the biodiversity, abundance and geographical and bathymetric distribution of larger-sized epi- and shallow infaunal Mollusca of the South Orkney Islands.

Genomic evidence for West Antarctic Ice Sheet collapse during the Last Interglacial.

The marine-based West Antarctic Ice Sheet (WAIS) is considered vulnerable to irreversible collapse under future climate trajectories, and its tipping point may lie within the mitigated warming scenarios of 1.5° to 2°C of the United Nations Paris Agreement. Knowledge of ice loss during similarly warm past climates could resolve this uncertainty, including the Last Interglacial when global sea levels were 5 to 10 meters higher than today and global average temperatures were 0.5° to 1.5°C warmer than preindustrial levels. Using a panel of genome-wide, single-nucleotide polymorphisms of a circum-Antarctic octopus, we show persistent, historic signals of gene flow only possible with complete WAIS collapse. Our results provide the first empirical evidence that the tipping point of WAIS loss could be reached even under stringent climate mitigation scenarios.

Habitat variability and faunal zonation at the Ægir Ridge, a canyon-like structure in the deep Norwegian Sea.

The Ægir Ridge System (ARS) is an ancient extinct spreading axis in the Nordic seas extending from the upper slope east of Iceland (∼550 m depth), as part of its Exclusive Economic Zone (EEZ), to a depth of ∼3,800 m in the Norwegian basin. Geomorphologically a rift valley, the ARS has a canyon-like structure that may promote increased diversity and faunal density. The main objective of this study was to characterize benthic habitats and related macro- and megabenthic communities along the ARS, and the influence of water mass variables and depth on them. During the IceAGE3 expedition (Icelandic marine Animals: Genetics and Ecology) on RV Sonne in June 2020, benthic communities of the ARS were surveyed by means of a remotely-operated vehicle (ROV) and epibenthic sledge (EBS). For this purpose, two working areas were selected, including abyssal stations in the northeast and bathyal stations in the southwest of the ARS. Video and still images of the seabed were usedtoqualitatively describebenthic habitats based on the presence of habitat-forming taxa and the physical environment. Patterns of diversity and community composition of the soft-sediment macrofauna, retrieved from the EBS, were analyzed in a semiquantitative manner. These biological data were complemented by producing high-resolution bathymetric maps using the vessel's multi-beam echosounder system. As suspected, we were able to identify differences in species composition and number of macro- and megafaunal communities associated with a depth gradient. A biological canyon effect became evident in dense aggregates of megafaunal filter feeders and elevated macrofaunal densities. Analysis of videos and still images from the ROV transects also led to the discovery of a number ofVulnerable Marine Ecosystems (VMEs) dominated by sponges and soft corals characteristic of the Arctic region. Directions for future research encompass a more detailed, quantitative study of the megafauna and more coherent sampling over the entire depth range in order to fully capture the diversity of the habitats and biota of the region. The presence of sensitive biogenic habitats, alongside seemingly high biodiversity and naturalness are supportive of ongoing considerations of designating part of the ARS as an "Ecologically and Biologically Significant Area" (EBSA).

Strong population genetic structure in a broadcast-spawning Antarctic marine invertebrate.

Although studies of population genetic structure are commonplace, a strong bias exists toward species from low latitudes and with relatively poor dispersal capabilities. Consequently, we used 280 amplified fragment length polymorphism bands to explore patterns of genetic differentiation among 8 populations of a high latitude broadcast-spawning marine mollusc, the Antarctic limpet Nacella concinna. Over 300 individuals were sampled along a latitudinal gradient spanning the Antarctic Peninsula from Adelaide Island to King George Island (67°-62°S), then to Signy Island (60°S) and South Georgia (54°S). Populations from the Antarctic Peninsula exhibited little genetic structure but were themselves strongly differentiated from both Signy and South Georgia. This finding was analytically highly robust and implies the presence of significant oceanographic barriers to gene flow in a species long regarded as a classic example of a widely dispersing broadcast spawner.

Abundance data of benthic peracarid crustaceans from the South Atlantic and Southern Ocean.

Peracarid data were collected in the Southern Ocean and South Atlantic Ocean. Sampling was performed during nine different expeditions on board of RRS James Clark Ross and RV Polarstern, using epibenthic sledges (EBS) at depth ranging between 160-6348 m at 109 locations. The correlation between environmental variables and peracarid abundance was investigated. Abundance data comprise a total of 128570 peracarids (52366 were amphipods, 28516 were cumaceans, 36142 isopods, 5676 mysidaceans and 5870 were tanaidaceans). The presented data are useful to investigate the composition and abundance patterns of peracarid orders at a wide depth range and spatial scale in the Southern Ocean. They can also be reused to compare their abundance with that of other taxa in broader ecological surveys.

Latitudinal Biogeographic Structuring in the Globally Distributed Moss Ceratodon purpureus.

Biogeographic patterns of globally widespread species are expected to reflect regional structure, as well as connectivity caused by occasional long-distance dispersal. We assessed the level and drivers of population structure, connectivity, and timescales of population isolation in one of the most widespread and ruderal plants in the world - the common moss Ceratodon purpureus. We applied phylogenetic, population genetic, and molecular dating analyses to a global (n = 147) sampling data set, using three chloroplast loci and one nuclear locus. The plastid data revealed several distinct and geographically structured lineages, with connectivity patterns associated with worldwide, latitudinal "bands." These imply that connectivity is strongly influenced by global atmospheric circulation patterns, with dispersal and establishment beyond these latitudinal bands less common. Biogeographic patterns were less clear within the nuclear marker, with gene duplication likely hindering the detection of these. Divergence time analyses indicated that the current matrilineal population structure in C. purpureus has developed over the past six million years, with lineages diverging during the late Miocene, Pliocene, and Quaternary. Several colonization events in the Antarctic were apparent, as well as one old and distinct Antarctic clade, possibly isolated on the continent since the Pliocene. As C. purpureus is considered a model organism, the matrilineal biogeographic structure identified here provides a useful framework for future genetic and developmental studies on bryophytes. Our general findings may also be relevant to understanding global environmental influences on the biogeography of other organisms with microscopic propagules (e.g., spores) dispersed by wind.

Fauna of the Kemp Caldera and its upper bathyal hydrothermal vents (South Sandwich Arc, Antarctica).

Faunal assemblages at hydrothermal vents associated with island-arc volcanism are less well known than those at vents on mid-ocean ridges and back-arc spreading centres. This study characterizes chemosynthetic biotopes at active hydrothermal vents discovered at the Kemp Caldera in the South Sandwich Arc. The caldera hosts sulfur and anhydrite vent chimneys in 1375-1487 m depth, which emit sulfide-rich fluids with temperatures up to 212°C, and the microbial community of water samples in the buoyant plume rising from the vents was dominated by sulfur-oxidizing Gammaproteobacteria. A total of 12 macro- and megafaunal taxa depending on hydrothermal activity were collected in these biotopes, of which seven species were known from the East Scotia Ridge (ESR) vents and three species from vents outside the Southern Ocean. Faunal assemblages were dominated by large vesicomyid clams, actinostolid anemones, Sericosura sea spiders and lepetodrilid and cocculinid limpets, but several taxa abundant at nearby ESR hydrothermal vents were rare such as the stalked barnacle Neolepas scotiaensis. Multivariate analysis of fauna at Kemp Caldera and vents in neighbouring areas indicated that the Kemp Caldera is most similar to vent fields in the previously established Southern Ocean vent biogeographic province, showing that the species composition at island-arc hydrothermal vents can be distinct from nearby seafloor-spreading systems. δ 13C and δ 15N isotope values of megafaunal species analysed from the Kemp Caldera were similar to those of the same or related species at other vent fields, but none of the fauna sampled at Kemp Caldera had δ 13C values, indicating nutritional dependence on Epsilonproteobacteria, unlike fauna at other island-arc hydrothermal vents.

Phylum Tardigrada: an "individual" approach.

Phylum Tardigrada consists of ∼1000 tiny, hardy metazoan species distributed throughout terrestrial, limno-terrestrial and oceanic habitats. Their phylogenetic status has been debated, with current evidence placing them in the Ecdysozoa. Although there have been efforts to explore tardigrade phylogeny using both morphological and molecular data, limitations such as their few morphological characters and low genomic DNA concentrations have resulted in restricted taxonomic coverage. Using a protocol that allows us to identify and extract DNA from individuals, we have sequenced 18S rDNA from 343 tardigrades from across the globe. Using maximum parsimony and Bayesian analyses we have found support for dividing Order Parachela into three super-families and further evidence that indicates the traditional taxonomic perspective of families in the class Eutardigrada are nonmonophyletic and require re-working. It appears that conserved morphology within Tardigrada has resulted in conservative taxonomy as we have found cases of several discrete lineages grouped into single genera. Although this work substantially adds to the understanding of the evolution and taxonomy of the phylum, we highlight that inferences gained from this work are likely to be refined with the inclusion of further taxa-specifically representatives of the nine families yet to be sampled. © The Willi Hennig Society 2008.

Assessing meiofaunal variation among individuals utilising morphological and molecular approaches: an example using the Tardigrada.

BACKGROUND: Meiofauna - multicellular animals captured between sieve size 45 mum and 1000 mum - are a fundamental component of terrestrial, and marine benthic ecosystems, forming an integral element of food webs, and playing a critical roll in nutrient recycling. Most phyla have meiofaunal representatives and studies of these taxa impact on a wide variety of sub-disciplines as well as having social and economic implications. However, studies of variation in meiofauna are presented with several important challenges. Isolating individuals from a sample substrate is a time consuming process, and identification requires increasingly scarce taxonomic expertise. Finding suitable morphological characters in many of these organisms is often difficult even for experts. Molecular markers are extremely useful for identifying variation in morphologically conserved organisms. However, for many species markers need to be developed de novo, while DNA can often only be extracted from pooled samples in order to obtain sufficient quantity and quality. Importantly, multiple independent markers are required to reconcile gene evolution with species evolution. In this primarily methodological paper we provide a proof of principle of a novel and effective protocol for the isolation of meiofauna from an environmental sample. We also go on to illustrate examples of the implications arising from subsequent screening for genetic variation at the level of the individual using ribosomal, mitochondrial and single copy nuclear markers. RESULTS: To isolate individual tardigrades from their habitat substrate we used a non-toxic density gradient media that did not interfere with downstream biochemical processes. Using a simple DNA release technique and nested polymerase chain reaction with universal primers we were able amplify multi-copy and, to some extent, single copy genes from individual tardigrades. Maximum likelihood trees from ribosomal 18S, mitochondrial cytochrome oxidase subunit 1, and the single copy nuclear gene Wingless support a recent study indicating that the family Hypsibiidae is a non-monophyletic group. From these sequences we were able to detect variation between individuals at each locus that allowed us to identify the presence of cryptic taxa that would otherwise have been overlooked. CONCLUSION: Molecular results obtained from individuals, rather than pooled samples, are a prerequisite to enable levels of variation to be placed into context. In this study we have provided a proof of principle of this approach for meiofaunal tardigrades, an important group of soil biota previously not considered amenable to such studies, thereby paving the way for more comprehensive phylogenetic studies using multiple nuclear markers, and population genetic studies.

On the ecological relevance of landscape mapping and its application in the spatial planning of very large marine protected areas.

In recent years very large marine protected areas (VLMPAs) have become the dominant form of spatial protection in the marine environment. Whilst seen as a holistic and geopolitically achievable approach to conservation, there is currently a mismatch between the size of VLMPAs, and the data available to underpin their establishment and inform on their management. Habitat mapping has increasingly been adopted as a means of addressing paucity in biological data, through use of environmental proxies to estimate species and community distribution. Small-scale studies have demonstrated environmental-biological links in marine systems. Such links, however, are rarely demonstrated across larger spatial scales in the benthic environment. As such, the utility of habitat mapping as an effective approach to the ecosystem-based management of VLMPAs remains, thus far, largely undetermined. The aim of this study was to assess the ecological relevance of broadscale landscape mapping. Specifically we test the relationship between broad-scale marine landscapes and the structure of their benthic faunal communities. We focussed our work at the sub-Antarctic island of South Georgia, site of one of the largest MPAs in the world. We demonstrate a statistically significant relationship between environmentally derived landscape mapping clusters, and the composition of presence-only species data from the region. To demonstrate this relationship required specific re-sampling of historical species occurrence data to balance biological rarity, biological cosmopolitism, range-restricted sampling and fine-scale heterogeneity between sampling stations. The relationship reveals a distinct biological signature in the faunal composition of individual landscapes, attributing ecological relevance to South Georgia's environmentally derived marine landscape map. We argue therefore, that landscape mapping represents an effective framework for ensuring representative protection of habitats in management plans. Such scientific underpinning of marine spatial planning is critical in balancing the needs of multiple stakeholders whilst maximising conservation payoff.