Ramirez-Llodra Eva

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A blueprint for an inclusive, global deep-sea ocean decade field program

(Frontiers Media, 2020-11-25) Howell, Kerry L. ; Hilario, Ana ; Allcock, A. Louise ; Bailey, David ; Baker, Maria C. ; Clark, Malcolm R. ; Colaço, Ana ; Copley, Jonathan T. ; Cordes, Erik E. ; Danovaro, Roberto ; Dissanayake, Awantha ; Escobar Briones, Elva ; Esquete, Patricia ; Gallagher, Austin J. ; Gates, Andrew R. ; Gaudron, Sylvie M. ; German, Christopher R. ; Gjerde, Kristina M. ; Higgs, Nicholas D. ; Le Bris, Nadine ; Levin, Lisa A ; Manea, Elisabetta ; McClain, Craig ; Menot, Lenaick ; Mestre, Mireia ; Metaxas, Anna ; Milligan, Rosanna J. ; Muthumbi, Agnes W. N. ; Narayanaswamy, Bhavani E. ; Ramalho, Sofia P. ; Ramirez-Llodra, Eva ; Robson, Laura M. ; Rogers, Alex D. ; Sellanes, Javier ; Sigwart, Julia D. ; Sink, Kerry ; Snelgrove, Paul V. R. ; Stefanoudis, Paris V. ; Sumida, Paulo Y. ; Taylor, Michelle L. ; Thurber, Andrew R. ; Vieira, Rui P. ; Watanabe, Hiromi K. ; Woodall, Lucy C. ; Xavier, Joana R.

The ocean plays a crucial role in the functioning of the Earth System and in the provision of vital goods and services. The United Nations (UN) declared 2021–2030 as the UN Decade of Ocean Science for Sustainable Development. The Roadmap for the Ocean Decade aims to achieve six critical societal outcomes (SOs) by 2030, through the pursuit of four objectives (Os). It specifically recognizes the scarcity of biological data for deep-sea biomes, and challenges the global scientific community to conduct research to advance understanding of deep-sea ecosystems to inform sustainable management. In this paper, we map four key scientific questions identified by the academic community to the Ocean Decade SOs: (i) What is the diversity of life in the deep ocean? (ii) How are populations and habitats connected? (iii) What is the role of living organisms in ecosystem function and service provision? and (iv) How do species, communities, and ecosystems respond to disturbance? We then consider the design of a global-scale program to address these questions by reviewing key drivers of ecological pattern and process. We recommend using the following criteria to stratify a global survey design: biogeographic region, depth, horizontal distance, substrate type, high and low climate hazard, fished/unfished, near/far from sources of pollution, licensed/protected from industry activities. We consider both spatial and temporal surveys, and emphasize new biological data collection that prioritizes southern and polar latitudes, deeper (> 2000 m) depths, and midwater environments. We provide guidance on observational, experimental, and monitoring needs for different benthic and pelagic ecosystems. We then review recent efforts to standardize biological data and specimen collection and archiving, making “sampling design to knowledge application” recommendations in the context of a new global program. We also review and comment on needs, and recommend actions, to develop capacity in deep-sea research; and the role of inclusivity - from accessing indigenous and local knowledge to the sharing of technologies - as part of such a global program. We discuss the concept of a new global deep-sea biological research program ‘Challenger 150,’ highlighting what it could deliver for the Ocean Decade and UN Sustainable Development Goal 14.
Deep, 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.
Deep-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 Committee

The 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.
The 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, Katrin

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.
Biodiversity and biogeography of hydrothermal vent species : thirty years of discovery and investigations

(Oceanography Society, 2007-03) Ramirez-Llodra, Eva ; Shank, Timothy M. ; German, Christopher R.

The discovery of hydrothermal vents and the unique, often endemic fauna that inhabit them represents one of the most extraordinary scientific discoveries of the latter twentieth century. Not surprisingly, after just 30 years of study of these remarkable—and extremely remote—systems, advances in understanding the animals and microbial communities living around hydrothermal vents seem to occur with every fresh expedition to the seafloor. On average, two new species are described each month—a rate of discovery that has been sustained over the past 25–30 years. Furthermore, the physical, geological, and geochemical features of each part of the ridge system and its associated hydrothermal-vent structures appear to dictate which novel biological species can live where. Only 10 percent of the ridge system has been explored for hydrothermal activity to date (Baker and German, 2004), yet we find different diversity patterns in that small fraction. While it is well known that species composition varies along discrete segments of the global ridge system, this “biogeographic puzzle” has more pieces missing than pieces in place.
Marine genetic resources in areas beyond national jurisdiction: promoting marine scientific research and enabling equitable benefit sharing

(Frontiers Media, 2021-03-31) Rogers, Alex D. ; Baco, Amy R. ; Escobar Briones, Elva ; Currie, Duncan ; Gjerde, Kristina M. ; Gobin, Judith ; Jaspars, Marcel ; Levin, Lisa A. ; Linse, Katrin ; Rabone, Muriel ; Ramirez-Llodra, Eva ; Sellanes, Javier ; Shank, Timothy M. ; Sink, Kerry ; Snelgrove, Paul V. R. ; Taylor, Michelle L. ; Wagner, Daniel ; Harden-Davies, Harriet

Growing human activity in areas beyond national jurisdiction (ABNJ) is driving increasing impacts on the biodiversity of this vast area of the ocean. As a result, the United Nations General Assembly committed to convening a series of intergovernmental conferences (IGCs) to develop an international legally-binding instrument (ILBI) for the conservation and sustainable use of marine biological diversity of ABNJ [the biodiversity beyond national jurisdiction (BBNJ) agreement] under the United Nations Convention on the Law of the Sea. The BBNJ agreement includes consideration of marine genetic resources (MGR) in ABNJ, including how to share benefits and promote marine scientific research whilst building capacity of developing states in science and technology. Three IGCs have been completed to date with the fourth delayed by the Covid pandemic. This delay has allowed a series of informal dialogues to take place between state parties, which have highlighted a number of areas related to MGR and benefit sharing that require technical guidance from ocean experts. These include: guiding principles on the access and use of MGR from ABNJ; the sharing of knowledge arising from research on MGR in ABNJ; and capacity building and technology transfer for developing states. In this paper, we explain what MGR are, the methods required to collect, study and archive them, including data arising from scientific investigation. We also explore the practical requirements of access by developing countries to scientific cruises, including the sharing of data, as well as participation in research and development on shore whilst promoting rather than hindering marine scientific research. We outline existing infrastructure and shared resources that facilitate access, research, development, and benefit sharing of MGR from ABNJ; and discuss existing gaps. We examine international capacity development and technology transfer schemes that might facilitate or complement non-monetary benefit sharing activities. We end the paper by highlighting what the ILBI can achieve in terms of access, utilization, and benefit sharing of MGR and how we might future-proof the BBNJ Agreement with respect to developments in science and technology.
sFDvent: a global trait database for deep-sea hydrothermal-vent fauna

(Wiley, 2019-07-30) Chapman, Abbie S. A. ; Beaulieu, Stace E. ; Colaço, Ana ; Gebruk, Andrey V. ; Hilario, Ana ; Kihara, Terue C. ; Ramirez-Llodra, Eva ; Sarrazin, Jozée ; Tunnicliffe, Verena ; Amon, Diva ; Baker, Maria C. ; Boschen‐Rose, Rachel E. ; Chen, Chong ; Cooper, Isabelle J. ; Copley, Jonathan T. ; Corbari, Laure ; Cordes, Erik E. ; Cuvelier, Daphne ; Duperron, Sébastien ; Du Preez, Cherisse ; Gollner, Sabine ; Horton, Tammy ; Hourdez, Stephane ; Krylova, Elena M. ; Linse, Katrin ; LokaBharathi, P. A. ; Marsh, Leigh ; Matabos, Marjolaine ; Mills, Susan W. ; Mullineaux, Lauren S. ; Rapp, Hans Tore ; Reid, William D. K. ; Rybakova, Elena Goroslavskaya ; Thomas, Tresa Remya A. ; Southgate, Samuel James ; Stöhr, Sabine ; Turner, Phillip J. ; Watanabe, Hiromi K. ; Yasuhara, Moriaki ; Bates, Amanda E.

Motivation Traits are increasingly being used to quantify global biodiversity patterns, with trait databases growing in size and number, across diverse taxa. Despite growing interest in a trait‐based approach to the biodiversity of the deep sea, where the impacts of human activities (including seabed mining) accelerate, there is no single repository for species traits for deep‐sea chemosynthesis‐based ecosystems, including hydrothermal vents. Using an international, collaborative approach, we have compiled the first global‐scale trait database for deep‐sea hydrothermal‐vent fauna – sFDvent (sDiv‐funded trait database for the Functional Diversity of vents). We formed a funded working group to select traits appropriate to: (a) capture the performance of vent species and their influence on ecosystem processes, and (b) compare trait‐based diversity in different ecosystems. Forty contributors, representing expertise across most known hydrothermal‐vent systems and taxa, scored species traits using online collaborative tools and shared workspaces. Here, we characterise the sFDvent database, describe our approach, and evaluate its scope. Finally, we compare the sFDvent database to similar databases from shallow‐marine and terrestrial ecosystems to highlight how the sFDvent database can inform cross‐ecosystem comparisons. We also make the sFDvent database publicly available online by assigning a persistent, unique DOI. Main types of variable contained Six hundred and forty‐six vent species names, associated location information (33 regions), and scores for 13 traits (in categories: community structure, generalist/specialist, geographic distribution, habitat use, life history, mobility, species associations, symbiont, and trophic structure). Contributor IDs, certainty scores, and references are also provided. Spatial location and grain Global coverage (grain size: ocean basin), spanning eight ocean basins, including vents on 12 mid‐ocean ridges and 6 back‐arc spreading centres. Time period and grain sFDvent includes information on deep‐sea vent species, and associated taxonomic updates, since they were first discovered in 1977. Time is not recorded. The database will be updated every 5 years. Major taxa and level of measurement Deep‐sea hydrothermal‐vent fauna with species‐level identification present or in progress. Software format .csv and MS Excel (.xlsx).
Volcanically hosted venting with indications of ultramafic influence at Aurora hydrothermal field on Gakkel Ridge

(Nature Communications, 2022-10-31) German, Christopher R ; Reeves, Eoghan P ; Türke, Andreas ; Diehl, Alexander ; Albers, Elmar ; Bach, Wolfgang ; Purser, Autun ; Ramalho, Sofia P ; Suman, Stefano ; Mertens, Christian ; Walter, Maren ; Ramirez-Llodra, Eva ; Schlindwein, Vera ; Bünz, Stefan ; Boetius, Antje

The Aurora hydrothermal system, Arctic Ocean, hosts active submarine venting within an extensive field of relict mineral deposits. Here we show the site is associated with a neovolcanic mound located within the Gakkel Ridge rift-valley floor, but deep-tow camera and sidescan surveys reveal the site to be ≥100 m across-unusually large for a volcanically hosted vent on a slow-spreading ridge and more comparable to tectonically hosted systems that require large time-integrated heat-fluxes to form. The hydrothermal plume emanating from Aurora exhibits much higher dissolved CH/Mn values than typical basalt-hosted hydrothermal systems and, instead, closely resembles those of high-temperature ultramafic-influenced vents at slow-spreading ridges. We hypothesize that deep-penetrating fluid circulation may have sustained the prolonged venting evident at the Aurora hydrothermal field with a hydrothermal convection cell that can access ultramafic lithologies underlying anomalously thin ocean crust at this ultraslow spreading ridge setting. Our findings have implications for ultra-slow ridge cooling, global marine mineral distributions, and the diversity of geologic settings that can host abiotic organic synthesis - pertinent to the search for life beyond Earth.
Hot vent beaneath an icy ocean: the Aurora vend field, Gakkel Ridge, revealed

(Oceanography Society, 2022-11-08) Ramirez-Llodra, Eva ; Argentino, Claudio ; Baker, Maria ; Boetius, Antje ; Costa, Carolina ; Dahle, Håkon ; Denny, Emily M. ; Dessandier, Pierre-Antoine ; Eilertsen, Mari H. ; Ferre, Benedicte ; German, Christopher R. ; Hand, Kevin ; Hilário, Ana ; Hislop, Lawrence ; Jamieson, John W. ; Kalnitchenko, Dimitri ; Mall, Achim ; Panieri, Giuliana ; Purser, Autun ; Ramalho, Sofia P. ; Reeves, Eoghan P. ; Rolley, Leighton ; Pereira, Samuel I. ; Ribeiro, Pedro A. ; Sert, Muhammed Fatih ; Steen, Ida H. ; Stetzler, Marie ; Stokke, Runar ; Victorero, Lissette ; Vulcano, Francesca ; Vågenes, Stig ; Waghorn, Kate Alyse ; Buenz, Stefan

Evidence of hydrothermal venting on the ultra-slow spreading Gakkel Ridge in the Central Arctic Ocean has been available since 2001, with first visual evidence of black smokers on the Aurora Vent Field obtained in 2014. But it was not until 2021 that the first ever remotely operated vehicle (ROV) dives to hydrothermal vents under permanent ice cover in the Arctic were conducted, enabling the collection of vent fluids, rocks, microbes, and fauna. In this paper, we present the methods employed for deep-sea ROV operations under drifting ice. We also provide the first description of the Aurora Vent Field, which includes three actively venting black smokers and diffuse flow on the Aurora mound at ~3,888 m depth on the southern part of the Gakkel Ridge (82.5°N). The biological communities are dominated by a new species of cocculinid limpet, two small gastropods, and a melitid amphipod. The ongoing analyses of Aurora Vent Field samples will contribute to positioning the Gakkel Ridge hydrothermal vents in the global biogeographic puzzle of hydrothermal vents.

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