Tyler
Paul A.
Tyler
Paul A.
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ArticleDeep, diverse and definitely different : unique attributes of the world's largest ecosystem(Copernicus Publications on behalf of the European Geosciences Union, 2010-09-22) Ramirez-Llodra, Eva ; Brandt, A. ; Danovaro, Roberto ; De Mol, B. ; Escobar Briones, Elva ; German, Christopher R. ; Levin, Lisa A. ; Arbizu, P. Martinez ; Menot, Lenaick ; Buhl-Mortensen, P. ; Narayanaswamy, Bhavani E. ; Smith, Craig R. ; Tittensor, D. P. ; Tyler, Paul A. ; Vanreusel, A. ; Vecchione, M.The deep sea, the largest biome on Earth, has a series of characteristics that make this environment both distinct from other marine and land ecosystems and unique for the entire planet. This review describes these patterns and processes, from geological settings to biological processes, biodiversity and biogeographical patterns. It concludes with a brief discussion of current threats from anthropogenic activities to deep-sea habitats and their fauna. Investigations of deep-sea habitats and their fauna began in the late 19th century. In the intervening years, technological developments and stimulating discoveries have promoted deep-sea research and changed our way of understanding life on the planet. Nevertheless, the deep sea is still mostly unknown and current discovery rates of both habitats and species remain high. The geological, physical and geochemical settings of the deep-sea floor and the water column form a series of different habitats with unique characteristics that support specific faunal communities. Since 1840, 28 new habitats/ecosystems have been discovered from the shelf break to the deep trenches and discoveries of new habitats are still happening in the early 21st century. However, for most of these habitats the global area covered is unknown or has been only very roughly estimated; an even smaller – indeed, minimal – proportion has actually been sampled and investigated. We currently perceive most of the deep-sea ecosystems as heterotrophic, depending ultimately on the flux on organic matter produced in the overlying surface ocean through photosynthesis. The resulting strong food limitation thus shapes deep-sea biota and communities, with exceptions only in reducing ecosystems such as inter alia hydrothermal vents or cold seeps. Here, chemoautolithotrophic bacteria play the role of primary producers fuelled by chemical energy sources rather than sunlight. Other ecosystems, such as seamounts, canyons or cold-water corals have an increased productivity through specific physical processes, such as topographic modification of currents and enhanced transport of particles and detrital matter. Because of its unique abiotic attributes, the deep sea hosts a specialized fauna. Although there are no phyla unique to deep waters, at lower taxonomic levels the composition of the fauna is distinct from that found in the upper ocean. Amongst other characteristic patterns, deep-sea species may exhibit either gigantism or dwarfism, related to the decrease in food availability with depth. Food limitation on the seafloor and water column is also reflected in the trophic structure of heterotrophic deep-sea communities, which are adapted to low energy availability. In most of these heterotrophic habitats, the dominant megafauna is composed of detritivores, while filter feeders are abundant in habitats with hard substrata (e.g. mid-ocean ridges, seamounts, canyon walls and coral reefs). Chemoautotrophy through symbiotic relationships is dominant in reducing habitats. Deep-sea biodiversity is among of the highest on the planet, mainly composed of macro and meiofauna, with high evenness. This is true for most of the continental margins and abyssal plains with hot spots of diversity such as seamounts or cold-water corals. However, in some ecosystems with particularly "extreme" physicochemical processes (e.g. hydrothermal vents), biodiversity is low but abundance and biomass are high and the communities are dominated by a few species. Two large-scale diversity patterns have been discussed for deep-sea benthic communities. First, a unimodal relationship between diversity and depth is observed, with a peak at intermediate depths (2000–3000 m), although this is not universal and particular abiotic processes can modify the trend. Secondly, a poleward trend of decreasing diversity has been discussed, but this remains controversial and studies with larger and more robust data sets are needed. Because of the paucity in our knowledge of habitat coverage and species composition, biogeographic studies are mostly based on regional data or on specific taxonomic groups. Recently, global biogeographic provinces for the pelagic and benthic deep ocean have been described, using environmental and, where data were available, taxonomic information. This classification described 30 pelagic provinces and 38 benthic provinces divided into 4 depth ranges, as well as 10 hydrothermal vent provinces. One of the major issues faced by deep-sea biodiversity and biogeographical studies is related to the high number of species new to science that are collected regularly, together with the slow description rates for these new species. Taxonomic coordination at the global scale is particularly difficult, but is essential if we are to analyse large diversity and biogeographic trends. Because of their remoteness, anthropogenic impacts on deep-sea ecosystems have not been addressed very thoroughly until recently. The depletion of biological and mineral resources on land and in shallow waters, coupled with technological developments, are promoting the increased interest in services provided by deep-water resources. Although often largely unknown, evidence for the effects of human activities in deep-water ecosystems – such as deep-sea mining, hydrocarbon exploration and exploitation, fishing, dumping and littering – is already accumulating. Because of our limited knowledge of deep-sea biodiversity and ecosystem functioning and because of the specific life-history adaptations of many deep-sea species (e.g. slow growth and delayed maturity), it is essential that the scientific community works closely with industry, conservation organisations and policy makers to develop robust and efficient conservation and management options.
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ArticleDeep-water chemosynthetic ecosystem research during the Census of Marine Life decade and beyond : a proposed deep-ocean road map(Public Library of Science, 2011-08-04) German, Christopher R. ; Ramirez-Llodra, Eva ; Baker, Maria C. ; Tyler, Paul A. ; ChEss Scientific Steering CommitteeThe ChEss project of the Census of Marine Life (2002–2010) helped foster internationally-coordinated studies worldwide focusing on exploration for, and characterization of new deep-sea chemosynthetic ecosystem sites. This work has advanced our understanding of the nature and factors controlling the biogeography and biodiversity of these ecosystems in four geographic locations: the Atlantic Equatorial Belt (AEB), the New Zealand region, the Arctic and Antarctic and the SE Pacific off Chile. In the AEB, major discoveries include hydrothermal seeps on the Costa Rica margin, deepest vents found on the Mid-Cayman Rise and the hottest vents found on the Southern Mid-Atlantic Ridge. It was also shown that the major fracture zones on the MAR do not create barriers for the dispersal but may act as trans-Atlantic conduits for larvae. In New Zealand, investigations of a newly found large cold-seep area suggest that this region may be a new biogeographic province. In the Arctic, the newly discovered sites on the Mohns Ridge (71°N) showed extensive mats of sulfur-oxidisng bacteria, but only one gastropod potentially bears chemosynthetic symbionts, while cold seeps on the Haakon Mossby Mud Volcano (72°N) are dominated by siboglinid worms. In the Antarctic region, the first hydrothermal vents south of the Polar Front were located and biological results indicate that they may represent a new biogeographic province. The recent exploration of the South Pacific region has provided evidence for a sediment hosted hydrothermal source near a methane-rich cold-seep area. Based on our 8 years of investigations of deep-water chemosynthetic ecosystems worldwide, we suggest highest priorities for future research: (i) continued exploration of the deep-ocean ridge-crest; (ii) increased focus on anthropogenic impacts; (iii) concerted effort to coordinate a major investigation of the deep South Pacific Ocean – the largest contiguous habitat for life within Earth's biosphere, but also the world's least investigated deep-ocean basin.
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ArticleThe discovery of new deep-sea hydrothermal vent communities in the Southern Ocean and implications for biogeography(Public Library of Science, 2012-01-03) Rogers, Alex D. ; Tyler, Paul A. ; Connelly, Douglas P. ; Copley, Jonathan T. ; James, Rachael H. ; Larter, Robert D. ; Linse, Katrin ; Mills, Rachel A. ; Naveira Garabato, Alberto C. ; Pancost, Richard D. ; Pearce, David A. ; Polunin, Nicholas V. C. ; German, Christopher R. ; Shank, Timothy M. ; Boersch-Supan, Philipp H. ; Alker, Belinda J. ; Aquilina, Alfred ; Bennett, Sarah A. ; Clarke, Andrew ; Dinley, Robert J. J. ; Graham, Alastair G. C. ; Green, Darryl R. H. ; Hawkes, Jeffrey A. ; Hepburn, Laura ; Hilario, Ana ; Huvenne, Veerle A. I. ; Marsh, Leigh ; Ramirez-Llodra, Eva ; Reid, William D. K. ; Roterman, Christopher N. ; Sweeting, Christopher J. ; Thatje, Sven ; Zwirglmaier, KatrinSince 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.
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ArticleHabitat associations in gastropod species at East Pacific Rise hydrothermal vents (9°50'N)(Marine Biological Laboratory, 2007-06) Mills, Susan W. ; Mullineaux, Lauren S. ; Tyler, Paul A.At deep-sea hydrothermal vents on the East Pacific Rise (9°50'N), distinct megafaunal assemblages are positioned along strong thermal and chemical gradients. We investigated the distribution of gastropod species to determine whether they associate with specific megafaunal zones and to determine the thermal boundaries of their habitats. Gastropods colonized a series of basalt blocks that were placed into three different zones characterized by vestimentiferan tubeworms, bivalves, and suspension-feeders, respectively. Additional gastropods were collected on selected blocks from higher temperature vestimentiferan habitat and from grab samples of alvinellid polychaetes. On the blocks, gastropod species clustered into a "Cool" group (Clypeosectus delectus, Eulepetopsis vitrea, Gorgoleptis spiralis, and Lepetodrilus ovalis) whose species tended to be most abundant in the suspension-feeder zone, and a "Warm" group (Lepetodrilus cristatus, L. elevatus, L. pustulosus, and Cyathermia naticoides) whose species all were significantly more abundant in the vestimentiferan zone than elsewhere. The temperature ranges of Cool species were generally lower than the ranges of Warm ones, although both groups were present at 3 to 6 °C; also present was Bathymargarites symplector, which clustered with neither group. Three additional species, Rhynchopelta concentrica, Neomphalus fretterae, and Nodopelta rigneae, co-occurred with Warm-group species on selected blocks from hotter habitats. Although a few species were found only in alvinellid collections, most species were not exclusive to a specific megafaunal zone. We propose that species in the Cool and Warm groups occupy specific microhabitats that are present in more than one zone.
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ArticleHydrothermal vent fields and chemosynthetic biota on the world's deepest seafloor spreading centre(Nature Publishing Group, 2012-01-10) Connelly, Douglas P. ; Copley, Jonathan T. ; Murton, Bramley J. ; Stansfield, Kate ; Tyler, Paul A. ; German, Christopher R. ; Van Dover, Cindy L. ; Amon, Diva ; Furlong, Maaten ; Grindlay, Nancy ; Hayman, Nicholas W. ; Huhnerbach, Veit ; Judge, Maria ; Le Bas, Tim ; McPhail, Stephen ; Meier, Alexandra ; Nakamura, Ko-ichi ; Nye, Verity ; Pebody, Miles ; Pedersen, Rolf B. ; Plouviez, Sophie ; Sands, Carla M. ; Searle, Roger C. ; Stevenson, Peter ; Taws, Sarah ; Wilcox, SallyThe Mid-Cayman spreading centre is an ultraslow-spreading ridge in the Caribbean Sea. Its extreme depth and geographic isolation from other mid-ocean ridges offer insights into the effects of pressure on hydrothermal venting, and the biogeography of vent fauna. Here we report the discovery of two hydrothermal vent fields on the Mid-Cayman spreading centre. The Von Damm Vent Field is located on the upper slopes of an oceanic core complex at a depth of 2,300 m. High-temperature venting in this off-axis setting suggests that the global incidence of vent fields may be underestimated. At a depth of 4,960 m on the Mid-Cayman spreading centre axis, the Beebe Vent Field emits copper-enriched fluids and a buoyant plume that rises 1,100 m, consistent with > 400 °C venting from the world’s deepest known hydrothermal system. At both sites, a new morphospecies of alvinocaridid shrimp dominates faunal assemblages, which exhibit similarities to those of Mid-Atlantic vents.
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ArticleReproduction of gastropods from vents on the East Pacific Rise and the Mid-Atlantic Ridge(National Shellfisheries Association, 2008-03) Tyler, Paul A. ; Pendlebury, Sophie ; Mills, Susan W. ; Mullineaux, Lauren S. ; Eckelbarger, Kevin J. ; Baker, Maria C. ; Young, Craig M.The gametogenic biology is described for seven species of gastropod from hydrothermal vents in the East Pacific and from the Mid-Atlantic Ridge. Species of the limpet genus Lepetodrilus (Family Lepetodrilidae) had a maximum unfertilized oocyte size of <90 μm and there was no evidence of reproductive periodicity or spatial variation in reproductive pattern. Individuals showed early maturity with females undergoing gametogenesis at less than one third maximum body size. There was a power relationship between shell length and fecundity, with a maximum of 1,800 oocytes being found in one individual, although individual fecundity was usually <1,000. Such an egg size might be indicative of planktotrophic larval development, but there was never any indication of shell growth in larvae from species in this genus. Cyathermia naticoides (Family Neomphalidea) had a maximum oocyte size of 120 μm and a fecundity of <400 oocytes per individual. Rhynchopelta concentrica (Family Peltospiridae) had a maximum oocyte size of 184 μm and a fecundity <600, whereas in Eulepetopsis vitrea (Family Neolepetopsidae) maximum oocyte size was 232 μm with a fecundity of <200 oocytes per individual. In none of these three species was there any indication of episodicity in oocyte production. From our observations we support the paradigm that there is no reproductive pattern typical of vent systems but is more related to species' phylogeny.