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Cold-Water Corals in a Changing Ocean

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Cold-Water Corals in a Changing Ocean Available online at www.sciencedirect.com ScienceDirect § Cold-water corals in a changing ocean 1,2,3 4 J Murray Roberts and Stephen D Cairns Although known since Antiquity, corals in deep, cold waters on continental shelves, slopes, canyons, seamounts and remained largely unappreciated and overlooked by the ridge systems across the globe [2]. The habitats engineered scientific community until the 1990s. The last two decades by cold-water corals vary from ‘coral gardens’, frequently have seen a dramatic increase in our understanding of these structured by bamboo or gorgonian corals, to the large deep- diverse ecosystems alongside growing realisation that many water framework reefsusuallyconstructedbyone ortwo ofa cold-water coral habitats had been degraded by bottom small group of scleractinian corals (Oculina varicosa, Madre- trawling and are threatened by ocean warming and pora oculata, Lophelia pertusa, Solenosmilia variabilis, Gonio- acidification. This paper discusses recent improvements in our corella dumosa, Enallopsammia profunda and Bathelia understanding of cold-water coral ecology, taxonomy and candida). In all cases the coral skeletons provide complex biodiversity following a variety of advances from the application three-dimensional structural habitat used by many other of predictive mapping to the use of molecular phylogenetic species as refuge or as a place from which to feed. approaches. Since many cold-water coral ecosystems occur in deep-waters beyond national jurisdiction, conservation Research on cold-water corals grew rapidly from the mid- management measures are being developed through the 1990s onwards with international symposia beginning in United Nations and related conventions. All such management 2000. This was first driven by technological advances measures require not only international agreement but also coupled with a growing need to understand ecosystems monitoring and enforcement to ensure their success. on deep continental margin and slope settings as offshore Addresses hydrocarbon exploration and deep-water fisheries 1 Centre for Marine Biodiversity & Biotechnology, School of Life expanded into these areas. The huge strides made in Sciences, Heriot-Watt University, Edinburgh, Scotland, EH14 4AS, acoustic seabed mapping using multibeam echosounders United Kingdom 2 have revealed a previously unknown density of cold- Scottish Association for Marine Science, Oban, PA37 1QA, United Kingdom water coral reefs and coral carbonate mounds [3,4]. Both 3 Center for Marine Science, University of North Carolina Wilmington, support extensive long-lived biogenic reefs that, through 601 S. College Road, Wilmington, NC, 28403-5928, United States the dynamic processes of growth and (bio)erosion, can 4 Department of Invertebrate Zoology, National Museum of Natural accumulate tens of metres above the surrounding seafloor, History, Smithsonian Institution, Washington, DC 20560, United States locally alter sedimentary patterns and provide niches for a Corresponding author: Roberts, J Murray ([email protected]) diverse community [5]. In the NE Atlantic, cold-water coral reef frameworks structured by Lophelia pertusa developed during interglacials, but were absent during Current Opinion in Environmental Sustainability 2014, 7:118–126 glacial climate periods indicating a very close coupling This review comes from a themed issue on Aquatic and marine with global climate [6 ]. The substantial structures pro- systems duced by coral carbonate mounds are formed by Georgios Tsounis Bernhard Riegl Edited by and sequences of interglacial coral reef framework overlain with glacial deposits (Figure 1). In other regions, cold- water coral research is growing rapidly with examples Received 30 June 2013; Accepted 12 January 2014 including studies of the solitary scleractinian Desmophyl- lum dianthus in Chilean fjords [7] and of colonial scler- actinians in Mediterranean canyons [8]. 1877-3435/$ – see front matter, # 2014 The Authors. Published by Elsevier B.V. All rights reserved. In parallel with the work to map and characterise cold- http://dx.doi.org/10.1016/j.cosust.2014.01.004 water coral habitats has been a growing understanding of the environmental factors that control their distribution. Cold-water corals are frequently associated with certain Introduction temperatures and salinities, with evidence that L. pertusa in the NE Atlantic is found in a particular density layer There are more coral species in waters over 50 m deep than À3 (sigma-theta 27.35–27.65 kg m ) perhaps because larvae there are on shallow, tropical coral reefs [1]. Set apart from disperse laterally along this horizon [9]. However, at local tropical corals by the cooler waters they inhabit, cold-water scales cold-water corals occurrence is closely associated to corals are amongst the mostsignificant ecosystem engineers § This is an open-access article distributed under the terms of the Creative Commons Attribution-NonCommercial-No Derivative Works License, which permits non-commercial use, distribution, and reproduction in any medium, provided the original author and source are credited. Current Opinion in Environmental Sustainability 2014, 7:118–126 www.sciencedirect.com Cold-water corals Roberts and Cairns 119 Figure 1 Erosion of mound exposing Glacial - hard substrata (stones or Interglacial dead coral framework). Mound Cycle 10 - 1000 ka E a ial rly interglac Re-initiation of mound growth: coral colonisation Existing and new reef framework Initiation of mound Rec sediment ru growth: corals colonise development. itm Few epifauna e n elevated hard substrata. Gr t ow Sediment th infilled Biogenic rubble Living coral Ne al Reef w in laci framework terg Cycle Interglacial Lowest diversity Coral 0.01 - 1 ka n o i rubble s o e r c Highest diversity e ) n io a b (B r tu is Exposed D dead coral Mound ‘retirement’: Abundant epifauna no coral growth, glacial Mound development: deposits including dropstones. coral framework entraps mobile sediment. Glacial Interglacial Current Opinion in Environmental Sustainability Schematic illustration showing: (1) outer circle; cyclic stages of carbonate mound growth from initiation, development, ‘retirement’ and re-colonisation; (2) inner circle; smaller scale cycle of reef microhabitats, succession and faunal diversity. Figure redrawn from [5]. sites where fresh labile food material is transported Ecological Niche Factor Analysis [13] a number of studies rapidly from the surface [10], often in settings where have been carried out both at regional and global scales. seabed topography and water flow interact to promote The most recent work uses the Maximum Entropy enhanced food supply [11]. However, without a suitable approach with examples including scleractinian corals hard surface for larval attachment it is unlikely that cold- on seamounts [14] and deep-water octocorals [15 ]. Inter- water corals will be able to settle and flourish. estingly oceanic carbonate chemistry emerges as an important explanatory variable for both groups with the With the development of predictive habitat modelling aragonitic scleractinian corals and calcitic octocorals approaches and greater availability of regional and global related closely to aragonite and calcite saturation states datasets of key environmental parameters (e.g. tempera- respectively. As the progressive acidification of the oceans ture, dissolved O2, and nutrients) work to develop pre- continues there is great concern that large areas of the dictive models of cold-water coral occurrence has begun. global ocean will become undersaturated as the calcium Following the first studies published in 2005 [12] using carbonate saturation horizons shallow [16,17]. www.sciencedirect.com Current Opinion in Environmental Sustainability 2014, 7:118–126 120 Aquatic and marine systems CWC taxonomy and associated biodiversity circulation patterns [21]. McFadden et al. [22,23 ] have Cold-water corals are members of the Orders Scleractinia, reviewed the status of molecular phylogenetic analysis Zoanthidea, the Subclass Octocorallia and the Family among the deep octocorals, acknowledging that mito- Stylasteridae (Table 1). Understanding their global distri- chondrial gene sequences have limited value in this bution remains very limited by the lack of information group, advocating the use of sequencing selected nuclear from many deep-sea areas of the world. Where sufficient genes, complete transcriptomes, and using secondary information exists it has only recently become possible to structure, and gene order. The basic issue of identification produce summaries of global species diversity. For based on morphology of deep-water corals has been aided example, the azooxanthellate Scleractinia show three by the publication of a colour-illustrated key (with glos- high diversity centres (Figure 2), although recent unpub- sary) to the 120 deep-water scleractinian coral genera lished work by Kitahara and Cairns suggest that the New [24 ], and an illustrated glossary to all scanning electron Caledonian region and adjacent northern Norfolk Ridge microscopy characters used in the identification of the may be the overall centre of deep-sea coral biodiversity. Stylasteridae [25 ]. Finally, Stolarski et al. [26 ] have In parallel with work on other taxa, molecular approaches presented the latest phylogeny of the higher taxa of to phylogeny are now starting to help unravel patterns in Scleractinia, with heavy emphasis on deep-water taxa, the relationships between cold-water corals. For example, showing a third
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