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Methane Seep Ecosystem Functions and Services from a Recently Discovered Southern California Seep Benjamin M

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Methane Seep Ecosystem Functions and Services from a Recently Discovered Southern California Seep Benjamin M Marine Ecology. ISSN 0173-9565 ORIGINAL ARTICLE Methane seep ecosystem functions and services from a recently discovered southern California seep Benjamin M. Grupe1, Monika L. Krach1,*, Alexis L. Pasulka1,†, Jillian M. Maloney2,‡, Lisa A. Levin1 & Christina A. Frieder1,§ 1 Center for Marine Biodiversity and Conservation, Integrative Oceanography Division, Scripps Institution of Oceanography, University of California San Diego, La Jolla, CA, USA 2 Geosciences Research Division, Scripps Institution of Oceanography, University of California San Diego, La Jolla, CA, USA Keywords Abstract Authigenic carbonate; chemosynthetic production; Del Mar Methane Seep; The recent discovery of a methane seep with diverse microhabitats and abun- macrofauna; Sebastolobus; stable isotope dant groundfish in the San Diego Trough (1020 m) off the coast of Del Mar, ecology. California raised questions about the role of seep ecosystem functions and ser- vices in relation to continental margins. We used multicorer and ROV grab Correspondence samples and an ROV survey to characterize macrofaunal structure, diversity, Benjamin M. Grupe, University of San Diego, and trophic patterns in soft sediments and authigenic carbonates; seep micro- Department of Biology, 5998 Alcala Park, habitats and taxa observed; and the abundance and spatial patterns of fishery- San Diego, CA 92110. E-mail: [email protected] relevant species. Biogenic microhabitats near the Del Mar Seep included micro- bially precipitated carbonate boulders, bacterial mats, vesicomyid clam beds, *Present address: Farallones Marine frenulate and ampharetid beds, vestimentiferan tubeworm clumps, and fields of Sanctuary Association, San Francisco, CA, Bathysiphon filiformis tubes. Macrofaunal abundance increased and mean faunal USA d13 † C signatures decreased in multicorer samples nearer the seep, suggesting Present address: Division of Geological and that chemosynthetic production enhanced animal densities outside the seep Planetary Sciences, California Institute of center. Polychaetes dominated sediments, and ampharetids became especially Technology, Pasadena, CA, USA ‡Present address: Department of Geological abundant near microbial mats, while gastropods, hydroids, and sponges domi- Sciences, San Diego State University, San nated carbonate rocks. A wide range of stable isotopic signatures reflected the Diego, CA, USA diversity of microhabitats, and methane-derived carbon was the most prevalent § Present address: Department of Biological source of nutrition for several taxa, especially those associated with carbonates. Sciences, University of Southern California, Megafaunal species living near the seep included longspine thornyhead (Sebas- Los Angeles, CA, USA tolobus altivelis), Pacific dover sole (Microstomus pacificus), and lithodid crabs (Paralomis verrilli), which represent targets for demersal fisheries. Sebastolobus Accepted: 18 October 2014 À altivelis was especially abundant (6.5–8.2 fishÁ100 m 2) and appeared to aggre- doi: 10.1111/maec.12243 gate near the most active seep microhabitats. The Del Mar Methane Seep, like many others along the world’s continental margins, exhibits diverse ecosystem functions and enhances regional diversity. Seeps such as this one may also con- tribute ecosystem services if they provide habitat for fishery species, export production to support margin food webs, and serve as sinks for methane- derived carbon. (reviewed in Armstrong et al. 2012; Thurber et al. 2014). Introduction In particular, a myriad of human activities acutely affect The deep sea is popularly described as remote, alien, and deep continental margins. These areas host a diversity of disconnected from society, but this vast region covering habitats that contribute essential fisheries production, 64% of the earth’s surface contributes many ecosystem mineral and gas resources, and the regulation of gas and functions, as well as ecosystem services benefiting humans climate cycling, carbon sequestration, waste absorption Marine Ecology 36 (2015) 91–108 ª 2015 Blackwell Verlag GmbH 91 Ecosystem functions of methane seeps Grupe, Krach, Pasulka, Maloney, Levin & Frieder and ecological systems (Levin & Dayton 2009; Levin & community structure have previously been examined only Sibuet 2012). at the San Clemente seeps, which occur much deeper Continental margin ecosystems have ecological func- than the OMZ at 1800 m [Bernardino & Smith 2010; but tions including benthic production and trophic transfer, see references for other Northeast Pacific seeps at Monte- decomposition of organic matter and nutrient remineral- rey (Barry et al. 1996), Northern California (Levin et al. ization, and carbon sequestration (Levin & Sibuet 2012), 2003, 2010), and Oregon (Kulm et al. 1986; Sahling et al. and methane seeps may perform many of these functions. 2002; Levin et al. 2010)]. Methane seeps play a role in global biogeochemical While the deep sea performs a host of ecosystem func- cycling and elemental transformation of carbon, sulfur, tions, here we focused specifically on habitat heterogene- and nitrogen (Hinrichs & Boetius 2002; Dekas et al. ity and biodiversity, trophic support for the benthic 2009; Boetius & Wenzhofer€ 2013). The anaerobic oxida- ecosystem and habitat for demersal megafauna. As the tion of methane (AOM) and associated precipitation of Del Mar Seep was discovered in 2012, another of our carbonate at cold seeps constitute a major carbon sink in objectives was to provide an initial characterization of its sediments, introducing a mechanism for benthic biogeo- habitats and taxa, and we suggest that this site may be chemical processes to influence potential greenhouse gas representative of other regional seeps within the OMZ. In sources (Ritger et al. 1987; Reeburgh 2007). The seep regards to seep ecosystem functions, we specifically biota acts as a methane filter that prevents methane hypothesized that: stored in gas hydrates and the deep biosphere from freely 1 Biogenic habitats associated with the seep would con- entering the hydrosphere and atmosphere; as much as tribute complex microhabitats and host distinct species 20–80% of methane carbon may be converted into ben- assemblages; thic biomass and carbonate, depending upon fluid flow 2 Macrofaunal density in sediments would increase with rates (Boetius & Wenzhofer€ 2013). proximity to the seep; The microbial biogeochemical processes that depend on 3 Species richness would be greater in sediments near reduced compounds (methane, sulfide, and hydrogen) at the seep periphery than those further away. The OMZ seeps create chemosynthetic primary production that sus- has depressed diversity, as few species are adapted to tains heterotrophic and symbiont-bearing fauna endemic the stressful physicochemical conditions, but the in-situ to cold seeps, as well as background consumers that may microbial production and microhabitat diversity at the aggregate at these productive benthic ecosystems (Levin seep might support more types of species than typical 2005; Sellanes et al. 2008). At a regional scale, seeps are background sediments (e.g. Levin et al. 2010); unique ecosystems that add physical, chemical, and biolog- 4 Methane-derived carbon (MDC) would be detectable ical habitat heterogeneity to continental margins (Cordes in the benthic food web near sources of seeping fluids, et al. 2010). High beta diversity at seeps and surrounding and isotopically light d13C signatures characteristic of regions can enhance the overall species richness, ecosystem chemosynthesis would be evident in sediment macro- function, and biological regulation that can occur on con- fauna but would decline with increased distance from tinental margins (Danovaro et al. 2008; Levin et al. 2010, the seep; 2012). Examples of non-endemic seep fauna utilizing the 5 Densities of demersal megafauna (Sebastolobus altivelis habitat offered by methane seeps include egg-laying sites and Microstomus pacificus) would be relatively high at the and nurseries for benthic octopuses and elasmobranchs methane seep compared with the surrounding seafloor. (Treude et al. 2011; Drazen et al. 2003), sponge-garden Moreover, we hypothesized these demersal fish would be refugia for macro-fauna (Thurber et al. 2010), and struc- more likely to occur specifically with three-dimensionally tural habitat for predatory fish and crabs (Patagonian complex seep microhabitats (carbonate outcrops, rubble, toothfish, Sellanes et al. 2008, 2012; orange roughy, Bowden clam beds) than over flat sediments. et al. 2013; sablefish, B. M. Grupe & L. A. Levin, personal observations; lithodid decapods, Niemann et al. 2013). Materials and Methods In this investigation, we document a suite of ecosystem functions associated with a methane seep recently discov- Study area ered in southern California. The Del Mar Methane Seep occurs at the lower boundary of the oxygen minimum This study was conducted at the Del Mar Methane Seep zone (OMZ), and with overlying water already hypoxic, (32°54.250 N, 117°46.940 W) at a depth of 1020 m in the benthic fauna and sediments are jointly affected by sharp northern portion of the San Diego Trough, approximately gradients in oxygen, sulfide, and organic carbon (Treude 50 km west of the Scripps Institution of Oceanography 2012). Despite knowledge of several methane seeps off (San Diego, California). This recently discovered seep is southern California, the macrofaunal biology and situated on a pop-up structure within a series of 92 Marine Ecology 36 (Suppl. 1) (2015) 91–108 ª 2015 Blackwell Verlag GmbH Grupe, Krach, Pasulka,
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