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Chemosymbiotic Species from the Gulf of Cadiz (NE Atlantic)

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Chemosymbiotic Species from the Gulf of Cadiz (NE Atlantic) EGU Journal Logos (RGB) Open Access Open Access Open Access Advances in Annales Nonlinear Processes Geosciences Geophysicae in Geophysics Open Access Open Access Natural Hazards Natural Hazards and Earth System and Earth System Sciences Sciences Discussions Open Access Open Access Atmospheric Atmospheric Chemistry Chemistry and Physics and Physics Discussions Open Access Open Access Atmospheric View metadata, citationAtmospheric and similar papers at core.ac.uk brought to you by CORE provided by Repositório Institucional da Universidade de Aveiro Measurement Measurement Techniques Techniques Discussions Open Access Biogeosciences, 10, 2569–2581, 2013 Open Access www.biogeosciences.net/10/2569/2013/ Biogeosciences doi:10.5194/bg-10-2569-2013 Biogeosciences Discussions © Author(s) 2013. CC Attribution 3.0 License. Open Access Open Access Climate Climate of the Past of the Past Discussions Chemosymbiotic species from the Gulf of Cadiz (NE Atlantic): Open Access Open Access distribution, life styles and nutritional patterns Earth System Earth System Dynamics Dynamics C. F. Rodrigues, A. Hilario,´ and M. R. Cunha Discussions Departamento de Biologia & CESAM – Universidade de Aveiro, Aveiro, Portugal Open Access Correspondence to: C. F. Rodrigues ([email protected]) Geoscientific Geoscientific Open Access Instrumentation Instrumentation Received: 28 November 2012 – Published in Biogeosciences Discuss.: 7 December 2012 Revised: 7 March 2013 – Accepted: 25 March 2013 – Published: 17 April 2013 Methods and Methods and Data Systems Data Systems Discussions Open Access Abstract. Previous work in the mud volcanoes from the Gulf 1 Introduction Open Access Geoscientific of Cadiz (South Iberian Margin) revealed a high number Geoscientific of chemosymbiotic species, namely bivalves and siboglinid Model Development polychaetes. In this study we give an overview of the distri- The finding thatModel hydrothermal Development vent mussels have a signifi- 13 Discussions bution and life styles of these species in the Gulf of Cadiz, cantly depleted δ C signature, very distinct from the values determine the role of autotrophic symbionts in the nutrition of other typical deep-sea invertebrates, led Rau and Hedges Open Access of selected species using stable isotope analyses (δ13C, δ15N (1979) to postulate that these mussels rely onOpen Access some non- and δ34S) and investigate the intra-specific variation of iso- photosynthetic food source.Hydrology Later this findingand was repeated Hydrology and tope signatures within and between study sites. During our for several cold seep invertebrates,Earth System and confirmed by the ob- Earth System studies, we identified twenty siboglinidae and nine bivalve servation that they host symbiotic chemoautotrophic bacteria chemosymbiotic species living in fifteen mud volcanoes. (Conway et al., 1994; CavanaughSciences et al., 1992). In this symbi- Sciences Solemyid bivalves and tubeworms of the genus Siboglinum otic relationship, the invertebrate host facilitates the access to Discussions Open Access are widespread in the study area, whereas other species inorganic carbon, oxygen and reduced sulphur-compounds,Open Access were found in a single mud volcano (e.g. “Bathymodiolus” and in return it uptakes the bacterial metabolic byproducts Ocean Science mauritanicus) or restricted to deeper mud volcanoes (e.g. or digests symbiont tissueOcean for itsScience nutrition (Fisher, 1990; Discussions Polybrachia sp., Lamelisabella denticulata). Species distri- MacAvoy et al., 2002). The carbon source for thiotrophic and bution suggests that different species may adjust their posi- methanotrophic bacteria is different, with the former using tion within the sediment according to their particular needs, pore water or near-bottom water dissolved inorganic carbon Open Access Open Access and to the intensity and variability of the chemical sub- (DIC) and the latter using CH4 (Fisher, 1990; Conway et al., strata supply. Tissue stable isotope signatures for selected 1994). Solid Earth The different carbon fixationSolid pathways Earth involve distinct species are in accordance with values found in other stud- Discussions ies, with thiotrophy as the dominant nutritional pathway, and isotopic fractionation (e.g. carbon, sulphur and nitrogen), with methanotrophy and mixotrophy emerging as secondary which makes stable isotope approaches particularly useful for elucidating the nutritional status of organisms in vent strategies. The heterogeneity in terms of nutrient sources (ex- Open Access Open Access pressed in the high variance of nitrogen and sulphur values) and seep environments (Conway et al., 1994; Van Dover and the ability to exploit different resources by the different and Fry, 1994). Carbon isotopic values have been used to The Cryosphere differentiate animalsThe with thiotrophicCryosphere symbionts from those species may explain the high diversity of chemosymbiotic Discussions species found in the Gulf of Cadiz. This study increases the with methanotrophic symbionts (Brooks et al., 1987; Kenni- knowledge on distributional patterns and resource partition- cutt et al., 1992), and to identify the source methane pool ing of chemosymbiotic species and highlights how trophic as either thermogenic or biogenic (Sassen et al., 1999). 15 fuelling varies on spatial scales with direct implications to Chemoautotrophs tend to have lower δ N values than het- 34 seep assemblages and potentially to the biodiversity of con- erotrophs or marine phytoplankton, and the δ S values of tinental margin. animals hosting thiotrophic bacteria are depleted relative to animals with methanotrophic symbionts and to heterotrophs Published by Copernicus Publications on behalf of the European Geosciences Union. 2570 C. F. Rodrigues et al.: Chemosymbiotic species from the Gulf of Cadiz (Brooks et al., 1987). In fact, values of δ34S below 5 ‰ can Siboglinid polychaetes are represented in the Gulf of be used to infer a thiotrophic mode of nutrition (Vetter and Cadiz by a high diversity of frenulates (Hilario´ et al., 2010). Fry, 1998) reflecting the isotopic signature of the sulphide Frenulata, the most specious clade of siboglinids, include a source (Fisher, 1995). Stable isotope signatures can therefore wide range of body sizes and tube morphologies. They live in provide information on food resource use and partitioning, a variety of reducing environments, including shelf and slope both inter- and intraspecifically (Levesque et al., 2003). sediments, cold seeps, hydrothermal vents and organic falls, Previous work in the Gulf of Cadiz mud volcanoes (South with their tube spanning oxic–anoxic boundaries (Hilario´ et Iberian Margin) found a high number of chemosymbiotic al., 2011). The anterior end of the tube remains at the sedi- species, namely bivalves and siboglinid polychaetes (Ro- ment surface or extends into the oxygenated bottom water, drigues et al., 2008; Hilario´ and Cunha, 2008; Hilario´ et al., where oxygen is absorbed by a gill-like structure. Hydro- 2010; Oliver et al., 2011), and studies on the faunal assem- gen sulphide, or methane in the case of Siboglinum poseidoni blages in the area show high biodiversity and high variabil- that harbours methane-oxidising endosymbionts (Schmaljo- ity in structure, composition and density, which are likely to hann and Flugel,¨ 1987), is transported across the posterior result from a combination of biogeographic, historical and tube and body wall into the trophosome, a specialized tissue environmental factors (Cunha et al., 2013). Especially rel- where the endosymbiotic bacteria are housed (reviewed by evant for the distribution of chemosymbiotic species is the Hilario´ et al., 2011). In addition to the nutrition provided by variability in the fluid sources and transport mechanisms de- their symbionts, species of the genus Siboglinum can take up termining the intensity of the fluxes and the geochemistry of and metabolise dissolved organic compounds at a rate suf- the porewater and sediments. ficient to sustain respiration but not enough for growth and The Gulf of Cadiz is an extensive cold seepage area in- reproduction (Southward and Southward, 1981). cluding over 40 mud volcanoes at depths between 200 and Due to its characteristics (biogeographical location, large 4000 m (Pinheiro et al., 2003; Van Rensbergen et al., 2005). number of seeps, seep regime variability), the Gulf of Cadiz The molecular and isotopic composition of the hydrocarbon provides a unique setting for the study of chemosynthesis- gases present in the different mud volcanoes reveal a pre- based assemblages. The high diversity of chemosymbiotic dominance of thermogenic sources with varying degrees of species found in the Gulf of Cadiz led us to hypothesize thermal maturity (Mazurenko et al., 2002, 2003; Stadnit- that (1) at small spatial scales different morphological and skaia et al., 2006; Hensen et al., 2007; Nuzzo et al. 2008) life style adaptations allow the co-occurrence of species and and, in some cases, partial recycling of thermogenic methane trophic niche partitioning, and (2) trophic diversity under dif- and other hydrocarbons mediated by anaerobic oxidation of ferential geochemical conditions results in high biodiversity methane (AOM)-related methanogenic archaea in the shal- at local and regional scales. Herein, we determine the trophic low sediments (Nuzzo et al., 2009). Differences in both fluid range of chemosymbiotic species using stable isotope analy- geochemistry and composition of the microbial assemblages ses (δ13C, δ15N and δ34S)
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