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Chapter 42 – Cold-Water Corals 1 2 Convenor (lead author): Erik Cordes 3 Authors: Sophie Arnaud-Haond, Odd-Aksel Bergstad, Ana Paula da Costa Falcão, 4 Andre Freiwald, J. Murray Roberts 5 Commentators: Peter Harris (Group of Experts) 6 Group of experts: Patricio Bernal 7

8 Abstract 9

Cold-water corals occur on continental margins throughout the world’s oceans and create 10 habitat for a diverse group of associated fauna. Cold-water corals prefer irregular 11 topographies with enhanced current flow, including submarine canyons and seamounts, 12 and can form bioherms many kilometers in area and rising over 100 meters from the 13 seafloor. These structures support a distinct community that represents a significant high-14 biomass and high-diversity hotspot in deep waters. This community is ultimately reliant 15 on surface productivity as the base of the food web, but provides a significant ecosystem 16 service through remineralization and recycling of nutrients, which may then become 17 available to surface waters through mixing and upwelling events. Cold-water corals are 18 increasingly threatened as human activities, including fishing, mining, and oil extraction, 19 move into deeper waters. Global climate and ocean change also represent significant 20 threats to these communities, primarily through the rise in ocean temperature, ocean 21 acidification, expansion of oxygen minimum zones, and changes in surface primary 22 productivity. Conservation efforts have primarily focused on deep-water fisheries 23 activities, although significant efforts are underway to regulate mining and oil and gas 24 industry activities in these habitats. Scientific research on the continental margins and in 25 the deep sea has greatly increased our understanding of cold-water corals in recent years, 26 and new discoveries of deep-water communities continue each year. From a scientific 27 point of view, these types of focused surveys should precede the expansion of human 28 industrial activities into deeper, unexplored waters in order to avoid or minimize impact 29 on these remarkable and fragile habitats. 30

31 1. Inventory and Ecosystem Functions 32

33 Globally viewed, framework-constructing cold-water corals cover a wide range of depths 34 (39 - 2000 m) and latitude (70°N – 60°S). This chapter primarily focuses upon the 35 framework-constructing scleractinian corals, although the communities of gorgonian and 36 stylasterid corals that form highly significant structural habitat in the North Pacific and 37 elsewhere (e.g. Stone 2006) are also briefly discussed. The most representative cold-38 water, framework-building, scleractinian corals are Enallopsammia rostrata, 39 Goniocorella dumosa, Lophelia pertusa, Madrepora oculata, Oculina varicosa and 40

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DRAFT: NOT TO BE QUOTED OR CITED FOR PUBLICATION Solenosmilia variabilis. The most common and widespread of the large, structure-41 forming octocorals are found in the genera Corallium, Isidella, Paragorgia, 42 Paramuricea, and Primnoa. 43 Cold-water corals (CWC) occur in continental slope settings, on deep shelves and along 44 the flanks of oceanic banks and seamounts (Fig. 42.1). The majority of CWC occur 45 between the depths of 200 to 1000 m, with the bathymetric ranges becoming shallower 46 towards the poles (Roberts et al., 2009). The shallowest occurrences are associated with 47 the rocky slopes and sills of fjords (Wilson 1979). Continental slopes exhibit a variety of 48 specific topographic irregularities which attract cold-water coral larvae to settle. In many 49 parts of the world ocean, the shelf edge is incised by gullies and submarine canyons 50 (Harris and Whiteway, 2011; Harris et al., 2014). Some prominent examples are located 51 on the canyon-rich slope of the Gulf of Lion (Fabri et al., 2014), the Bay of Biscay under 52 French and Spanish jurisdiction (De Mol et al., 2011; Sánchez et al., in press), The 53 Gulley off the coast of Nova Scotia (Mortensen and Buhl-Mortensen 2005), and the Mid-54 Atlantic canyons off the United States (Brooke and Ross 2014). Narrow straits between 55 land masses may also provide suitable substrate, such as the Straits of Florida (Correa et 56 al., 2012), Gibraltar (De Mol et al., 2012), Sicily (Freiwald et al., 2009), and the Yucatan 57 (Hebbeln et al., 2014). Open-slope coral mounds are known from the Northeast Atlantic 58 along the Rockall and Porcupine Banks (Van der Land et al., 2014), the Southeast coast 59 of the United States (Stetson et al., 1962; Reed et al., 2006), the Gulf of Mexico (Reed et 60 al., 2006; Lunden et al., 2013), Southwestern Atlantic Ocean (Viana et al., 1998; Sumida 61 et al., 2004; Pires, 2007; Kitahara et al., 2009; Caranza et al., 2012), and off Mauritania 62 (Colman et al., 2005). These mounds are not randomly distributed over the slope but 63 show a strong affinity with distinct water mass boundaries passing along the slope 64 (Mienis et al., 2007, Arantes et al, 2009, White and Dorschel, 2010). Oceanic seamounts 65 represent another important cold-water coral-rich environment (see Chapter 52), such as 66 the Tasmanian seamounts off South Australia (Thresher et al., 2011), the seamount 67 speckled Chatham Rise (Tracey et al., 2011), and seamounts of the Mid-Atlantic Ridge 68 system (Mortensen et al., 2008). A compilation of cold-water coral occurrences is 69 displayed in Figure 42.2 based on the UNEP-WCMC database (Freiwald et al., 2005) and 70 more recent findings. 71 Deep-sea corals have been known since the first descriptions in the 18th century and the 72 first deep-water research expeditions of the 19th century (Roberts et al., 2006). The 73 presence of large reef structures in deep water was not appreciated until the first 74 submersibles were available in the late 20th century. More recently, based on these 75 occurrence records, the use of habitat modeling has led to the discovery of numerous 76 cold-water coral sites and habitats (Davies and Guinotte 2011, Yesson et al., 2012). As an 77 example, scleractinians were discovered on steep submarine cliffs after modelling 78 (Huvenne et al., 2011) and field observation in the Mediterranean (Naumann et al., 2013) 79 and the Bay of Biscay (De Mol et al., 2010; Reveillaud et al., 2008). Similarly, an 80 extensive screening of the Mediterranean revealed additional and more extensive coral 81 formations than anticipated hitherto (Freiwald et al., 2009). Habitat modelling has thus 82 far mostly been applied to a few of the most common species at a regional scale 83

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DRAFT: NOT TO BE QUOTED OR CITED FOR PUBLICATION (Rengstorf et al., 2013; Yesson et al., 2012) with coarse resolution (Ross and Howell, 84 2013). However, models are now being applied at finer resolution levels in order to guide 85 surveys with the visual tools of remotely-operated and manned submersibles (Georgian et 86 al., 2014). Additional fine-grained and broad-scale habitat modeling, specifically 87 incorporating the best available taxonomic identifications (Henry and Roberts, 2014) is 88 still needed to discover additional habitats, and to forecast the fate of CWC facing both 89 direct (fisheries) and indirect (environmental) impacts (Guinotte et al., 2006). 90 Cold-water coral formations support a highly diverse community, comprising faunal 91 biomass that is orders of magnitude above that of the surrounding seafloor (Mortensen et 92 al., 1995; Henry and Roberts, 2007; Cordes et al., 2008; Roberts et al., 2008). In addition 93 to this tightly-associated community, cold-water coral frameworks may also serve as 94 temporary but important spawning, nursery, breeding and feeding areas for a multitude of 95 fishes and invertebrates (Moberg and Folke, 1999; Fossa et al., 2002; Husebo et al., 2002; 96 Ross and Quattrini 2009; Baillon et al., 2012; Henry et al., 2013), and transient habitats 97 for diel vertical migrators (Davies et al., 2010). The ability to construct massive calcium 98 carbonate frameworks, which makes both shallow and deep-water coral reefs unique, 99 provides an important biogeochemical function in both the carbonate system (Doney et 100 al., 2009) and in calcium balance (Moberg and Folke, 1999). CWC skeletons also provide 101 an information function (sensu Groot et al., 2002) through their archiving of paleoclimate 102 signals (Adkins et al., 1998; Williams et al., 2006). Besides this, CWC ecosystems posses 103 an inherent aesthetic value demonstrated through countless films, photographs, and 104 paintings of reefs or reef organisms (Groot et al, 2002). 105 Cold-water corals and the communities they support rely on surface productivity as their 106 primary source of nutrition; either through the slow, relatively steady deposition of 107 marine snow, which may be enhanced by hydrographic mechanisms (e.g. Davies et al., 108 2009; Kiriakoulakis et al., 2007), or through more active transport provided by vertical 109 migrators (Mienis et al., 2012). As in shallow-water systems, corals and sponges of the 110 deep reefs recycle these nutrients and form both the structural and trophic foundation of 111 the ecosystem. In addition to these ties from shallow to deep water, the transport of 112 nutrients from deep to shallow water is accomplished both by the diel vertical migrations 113 of plankton and small fishes (Davies et al., 2010) as well as by periodic down- and 114 upwelling that can occur near some of the reefs (Mienis et al., 2007; Davies et al., 2009). 115 Although the mechanisms for deep-to-shallow water transport are well established, the 116 input of deep-water secondary productivity to shallow ecosystems remains unquantified. 117

118 2. Features and Trends 119 120

All geoform structures mentioned share some environmental factors that facilitate coral 121 settlement and subsequent growth: provision of current-swept hard substrate, and often 122 topographically-guided hydrodynamic settings. It has been suggested that corals are 123 preferably confined to narrow seawater density (Sigma-theta) envelopes (Dullo et al., 124 2008) in which along slope larval dispersal propagation may be facilitated. Survival and 125 World Ocean Assessment © 2014 United Nations

DRAFT: NOT TO BE QUOTED OR CITED FOR PUBLICATION growth may be most closely associated with specific hydrodynamic settings including 126 tidal-driven internal-wave fronts hitting continental slopes and seamounts (Mienis et al., 127 2007; Henry et al., 2014), specific up- and downwelling currents affecting the summits of 128 shallow-water seamounts (Ramirez-Llodra et al., 2010), and tidal-driven downwelling 129 phenomena on inner shelf settings (Davies et al., 2009; Findlay et al., 2013). These 130 hydrographic transfer processes tend to concentrate or prolong the retention time of 131 nutrients and food that sustain the metabolic demands of the suspension-feeding 132 community. 133 Another perspective on the occurrence of coral habitat is a combined, biogeophysical and 134 hydrochemical analysis of the ambient seawater, a very recent endeavor in the still young 135 research history of cold-water coral systems (e.g., Findlay et al., 2014; Flögel et al., 2014; 136 Henry et al., 2014). The global habitat suitability study by Davies and Guinotte (2011) 137 conducted on the six major cold-water framework-building corals (Enallopsammia 138 rostrata, Goniocorella dumosa, Lophelia pertusa, Madrepora oculata, Oculina varicosa 139 and Solenosmilia variabilis) used the Maximum Entropy modelling approach 140 (MAXENT). This approach uses species-presence data, global bathymetry 30-arc second 141 grids (1 km2 resolution) and incorporates environmental data from several global data 142 bases. Viewed on such a global scale, these corals generally thrive in waters that: 1) are 143 supersaturated with respect to aragonite, 2) occur shallower than 1500 m water depth, 3) 144 contain dissolved oxygen concentrations of >4 ml l-1, 4) have a salinity range between 34 145 and 37 ppt, and 5) show a temperature range between 5 and 10°C. Laboratory 146 experiments have confirmed many of these ranges, with L. pertusa being the most 147 commonly studied species. Mediterranean L. pertusa and M. oculata colonies survived 148 and grew at 12oC for three weeks, with M. oculata showing a greater sensitivity to high 149 temperature (Naumann et al., 2014). Gulf of Mexico L. pertusa colonies survived and 150 grew at up to 12oC, but died when exposed to 14oC for 8 days (Lunden et al., submitted). 151 Studies of L. pertusa from west of Scotland demonstrated that this species can maintain 152 respiratory independence and even survive periods of reduced oxygen (Dodds et al., 153 2007). 154 However, some remarkable outliers to these trends exist in the Red Sea and the Gulf of 155 Mexico. The Red Sea represents the warmest and most saline deep-sea basin on Earth, 156 with temperatures >20°C throughout the water column and salinity in excess of 40 ppt. 157 Recent findings of typically deep-dwelling corals in these habitats shed new light on the 158 persistence of corals in deep waters (Roder et al., 2013; Qurban et al., 2014). Although 159 none of the coral species found in the Red Sea belong to the classical six (see above for 160 list), limited framework growth is recorded mainly by Eguchipsammia fistula under food- 161 and oxygen deprived conditions (1.02 – 2.04 ml l-1). Coral survival under such extreme 162 environmental conditions is probably at the expense of aerobic respiration and 163 calcification rates. The latter benefitted from high temperatures in combination with high 164 aragonite saturation values of 3.44-3.61 in the Red Sea (Roder et al., 2013). The cold-165 water coral communities in the northern Gulf of Mexico belong to the most intensively 166 studied sites in waters of the United States (e.g., Cordes et al., 2008). The major 167 framework-constructor is L. pertusa and almost all environmental variables such as 168

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DRAFT: NOT TO BE QUOTED OR CITED FOR PUBLICATION temperature, salinity and aragonite saturation reflect the ranges known from Atlantic 169 Lophelia sites (Davies et al., 2010; Lunden et al., 2013). However, dissolved oxygen 170 values appear to be low, 2.7–2.8 ml l−1 are typically observed (Davies et al., 2010) and 171 values as low as 1.5 ml l−1 have been recorded adjacent to coral mounds (Georgian et al., 172 2014). Coral nubbins from these Gulf of Mexico populations survived and grew in the lab 173 at oxygen levels as low as 2.9 ml l-1, but eight-day incubations at lower oxygen 174 concentrations (1.5 ml l-1) caused complete mortality, suggesting that these conditions are 175 short-lived in situ (Lunden et al., submitted). Similarly low oxygenation levels were 176 found in the newly discovered Lophelia-Enallopsammia coral mounds in the Campeche 177 Bank coral mound province, in the southern Gulf of Mexico (Hebbeln et al., 2014). It is 178 possible that the low oxygen concentrations of the Gulf of Mexico result in lower growth 179 rates observed for L. pertusa on natural (Brooke and Young 2009) and man-made 180 substrata (Larcom et al., 2014), although this remains to be examined empirically. 181 There have been numerous recent advances in our knowledge of the oceanographic 182 variables describing coral habitat in the deep sea. However, knowledge gaps still remain 183 when up-scaling from local to regional to global scales. Furthermore, limited capacity to 184 carry out long-term in situ measurements with benthic landers and cabled observatories 185 persists. This knowledge is of utmost importance to understand the consequences of 186 already perceptible environmental change, such as ocean acidification, spread of oxygen 187 minimum zones, and rising temperatures on deep-sea ecosystems. 188

189 3. Major Pressures Linked to the Trends 190 191

Numerous anthropogenic threats to cold-water coral communities exist, including 192 fisheries, oil exploration and extraction, and mining, as well as global ocean change 193 including warming and acidification. The function of cold-water corals as habitat, feeding 194 grounds and nurseries for many fishes including deep-sea fisheries targets emerged 195 simultaneously with concerns as to the impact of fisheries on these ecosystems (Costello 196 et al., 2005; Grehan et al., 2005; Hourigan 2009; Maynou and Cartes 2012). Physical 197 impacts from both trawl fisheries and long-lining, now being conducted as deep as 1500-198 2000 m, are visible in the North Atlantic and Norwegian Seas (Roberts et al., 2000; Fossa 199 et al., 2002; Hall-Spencer et al., 2002), on the Australian seamounts (Koslow et al., 200 2001), off the coast of New Zealand (Probert et al., 1997), and Southwestern Atlantic 201 slope (Kitahara, 2009). Trawl fisheries have the most severe impacts, by the removal of 202 large volumes of organisms and of the cold-water coral framework from the seafloor and 203 the concomitant destruction of the habitat. These impacts have also been the most 204 recognized in terms of management efforts, thus far. 205 Installation of oil and gas offshore facilities and drilling activities (see Chapter 21) have a 206 great potential to adversely affect cold-water coral communities. The potential impact 207 should be higher in areas where much of the available substrate is from authigenic 208 carbonates related to natural oil and gas seepage, such as the Gulf of Mexico (Cordes et 209 al., 2008) and the Norwegian Margin (Hovland 2005). The most glaring example of oil 210 World Ocean Assessment © 2014 United Nations

DRAFT: NOT TO BE QUOTED OR CITED FOR PUBLICATION and gas industry impacts in the deep sea is the Deepwater Horizon disaster in 2010 in the 211 Gulf of Mexico. Material, conclusively linked to the spill, was discovered on octocoral 212 colonies (primarily Paramuricea biscaya) approximately 11 km away from the site of the 213 drilling rig (White et al., 2012). These colonies suffered tissue loss and many have 214 continued to decline in health since the spill (Hsing et al., 2013). Subsequent surveys 215 detected at least two additional sites, extending the impacts to 26 km from the site of the 216 well, and from 1370 m to 1950 m water depth (Fisher et al., 2014). Additional oil and gas 217 related impacts may have gone undetected in the past, particularly in deep-water areas 218 that are poorly explored and surveyed prior to industrial activity. Offshore energy 219 industry activity in the form of wind and wave energy is increasing as well (see Chapter 220 22), and physical structure placed on the seafloor could have an impact on cold-water 221 corals if the appropriate surveys are not completed prior to installation. 222 Global climate change is affecting every community type on Earth, and its effects are 223 already being felt in the deep sea. Ocean warming has been recorded in numerous deep-224 water habitats, but is particularly significant in marginal seas, which are home to many of 225 the world’s cold-water coral. In particular, there is evidence that the Mediterranean has 226 warmed by at least 0.1°C between 1950 and 2000 (Rixen et al., 2005). Cold-water corals 227 are highly sensitive to warming waters because of their clear upper thermal limits, and 228 the temperature excursions around this general upward trend are likely to be much higher. 229 Ocean acidification is an even more pervasive threat. Continued additions of CO2 into the 230 atmosphere exacerbate the problem as the oceans absorb approximately 30% of the CO2 231 from the atmosphere. Because the carbonate saturation state in seawater is temperature-232 dependent, it is much lower in cold waters and therefore cold-water corals lie much 233 closer to the saturation horizon (the depth below which the saturation state is below 1 and 234 carbonate minerals will dissolve) than shallow water corals. Solitary corals of the South 235 Pacific are already facing saturation states below 1 (Thresher et al., 2011), and small reef 236 frameworks constructed by Solenosmilia variabilis on Northeast Atlantic seamounts 237 grow in periodically undersaturated waters (Henry and Roberts 2014; Henry et al., 2014). 238 The Lophelia reefs of the Gulf of Mexico lie very close to the saturation horizon, at a 239 minimum saturation state of approximately 1.2 (Lunden et al., 2013). 240 Other possible effects of global climate change include deoxygenation and changes in 241 sea-surface productivity. Declines in oxygen availability are primarily linked to 242 increasing water temperature, but also to synergistic effects of pollution and agricultural 243 runoff, which are most significant in shallow water. However, because some cold-water 244 corals live at oxygen-minimum zone depths, even small changes in oxygen concentration 245 could be significant. Because cold-water corals live below the photic zone and rely on 246 primary productivity transferred from the surface waters to depth for their nutrition, 247 changes in surface productivity could have significant negative impacts. In particular, the 248 increased stratification of surface waters above the thermocline will lead to decreased 249 productivity in high latitude spring-bloom and upwelling ecosystems. This includes the 250 North Atlantic where the most extensive examples of the known cold-water coral reefs 251 exist. 252

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DRAFT: NOT TO BE QUOTED OR CITED FOR PUBLICATION Through in situ habitat characterization as well as by experimental approaches, it has 253 become clear that acidification and the expansion of oxygen minimum zones, together 254 with rising temperatures, will affect the average metabolism and physiology of most 255 scleractinians (Gori et al., 2013; Lartaud et al., 2014; McCulloch et al., 2012; Naumann et 256 al., 2013; Tittensor et al., 2010). However, whether such changes will result in range 257 shifts and massive extinctions, or if species possess the resources to cope with variations 258 through phenotypic plasticity or adaptive genetic changes, is still largely unknown. The 259 solitary coral Desmophyllum dianthus and colonial scleractinian Dendrophyllia cornigera 260 have shown resistance to high temperature in aquaria (Naumann et al., 2013). The L. 261 pertusa colonies from the North Atlantic and Mediterranean have shown the ability to 262 acclimatize to ocean acidification in long-term experiments (Form and Reibesell, 2012; 263 Maier et al., 2012). In other experiments, certain genotypes of L. pertusa from the Gulf of 264 Mexico were able to calcify at saturation states as low as 1.0, suggesting a possible 265 genetic basis to their sensitivity to ocean acidification (Lunden et al., submitted). 266 However, to date no long-term studies combining acidification with temperature stress 267 have been produced and long-term effects on skeletal structure are unknown. As the 268 alteration of the oceans resulting from global climate change continues into the future, 269 cold-water corals will be among the first threatened. However, they also seem to be 270 resilient to some of these processes, and may hold some of the answers for coral survival 271 in future global climate-change scenarios. Regardless, the projected shoaling of the 272 aragonite saturation horizon (Orr et al., 2005) threatens the future integrity of deep-water 273 scleractinian reef structures world-wide (Guinotte et al., 2006). 274 The ability to keep up with the pace of ocean change and disperse into a new 275 environment or to recolonize depleted areas depends on the capacity for mid- or long-276 distance dispersal. This capacity has been demonstrated for L. pertusa by isotope 277 reconstruction and genetic analysis (Henry et al., 2014), supporting the hypothesis of a 278 post-glacial recolonization of the Atlantic by refugees in the Mediterranean (De Mol et 279 al., 2005; De Mol et al., 2002; Frank et al., 2009). Overall, L. pertusa shows a pattern of 280 relative homogeneity within regions (e.g. the North Atlantic), and modest but significant 281 differentiation among regions, both for the Western Atlantic (e.g. Gulf of Mexico vs. 282 Southeast United States vs. North Atlantic; Morrison et al., 2011), as well as along 283 Eastern Atlantic margins from the Bay of Biscay to Iceland for both L. pertusa and M. 284 oculata (Becheler, 2013). Previous studies on the Eastern Atlantic margin had shown less 285 extensive connectivity, possibly reflecting the peculiar position of fjord populations in 286 Sweden and Norway (Le Goff-Vitry et al., 2004). Preliminary studies on D. dianthus 287 suggest a lack of barrier to large-scale dispersal (Addamo et al., 2012), although 288 bathymetric barriers to gene flow are evident (Miller et al 2011). Bathymetric barriers to 289 dispersal are also apparent in the phylogenetic community structure of deep-water 290 octocoral assemblages in the Gulf of Mexico (Quattrini et al., 2014). Altogether, the 291 present state of knowledge of genetic connectivity of deep-water corals suggests that 292 there the potential exists for some species to disperse and colonize across large distances 293 in response to major environmental changes. However, a great need for these studies be 294 conducted at a finer spatial scale with specific genetic markers remains (e.g. Dahl et al., 295

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DRAFT: NOT TO BE QUOTED OR CITED FOR PUBLICATION 2012) in order to ascertain the scale of genetic connectivity. In addition, the level of 296 differentiation among ocean basins requires regional scale conservation strategies. 297

298 4. Implications for Services to Ecosystems and Humanity 299 300

Impacts on cold-water corals and the structures they form would have significant 301 implications for the functioning of the surrounding deep sea and wider oceanic 302 ecosystems. The linkages from shallow to deep water, and back again, implicate cold-303 water corals as key components of the broader oceanic ecosystem. The physical 304 structures created by cold-water corals supports fisheries through the direct provision of 305 habitat or nursery grounds, which is likely to lead to increases in commercially 306 significant fish populations. This is likely to be most important where cold-water corals 307 are known to be highly abundant, such as on the North Atlantic and Australian and New 308 Zealand seamounts. 309 The ecosystem services provided go beyond the direct provision of substrate and shelter. 310 The complex habitat formed by cold-water corals increases the heterogeneity of the 311 continental margin, promoting higher beta diversity (Cordes et al., 2010). As in other 312 ecosystems (e.g. Tilmann et al., 1997), increased diversity most promotes higher levels of 313 ecosystem function. These ecosystem services may be important in relatively 314 oligotrophic regions such as the Gulf of Mexico and the Mediterranean where they may 315 support elevated levels of secondary productivity that may be transported back to the 316 surface. Recent findings from reefs off Norway demonstrated their significant role in 317 carbon cycling, raising additional concerns as to the impact of their disappearance on 318 global biochemical cycles (White et al., 2012). 319 Cold-water corals may also hold genetic resources that may provide services to humanity, 320 either directly or through their function as biodiversity hotspots in the deep sea (Arrieta et 321 al., 2010). Taxa such as cnidarians, sponges, and molluscs have been shown to harbor the 322 highest abundance of natural marine products of interest for biotechnology development 323 (Molinski et al., 2009; Rocha et al., 2011). As an example, the anti-AIDS drug AZT was 324 developed from an extract of a sponge from a shallow Caribbean reef (de la Calle, 2009). 325 At least half, and likely far more, of the diversity of corals and sponges lies in deep, cold 326 waters (Cairns, 2007), and therefore, these species have the highest potential for new 327 discoveries. With this potential comes the management concern, especially as some of the 328 potential genetic resources (See also chapter 29) harbored within the genomes of cold-329 water corals and sponges lie beyond national jurisdiction (Bruckner, 2002; de la Calle, 330 2009). 331 332 333 334 335

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5. Conservation Responses 336 337

Raised awareness of the susceptibility of cold-water coral communities to impacts of 338 human activities has, in recent decades, resulted in national and international actions to 339 protect cold-water corals and facilitate recovery of coral areas adversely affected in the 340 past. In some areas where significant damage was documented, e.g. along the continental 341 shelf off Norway (Fossa et al., 2002) and on seamounts in New Zealand and Australia 342 (Koslow et al., 2001), national legislation was introduced and specific management 343 measures were implemented. A growing number of reserves or protected areas within 344 national jurisdiction in the Atlantic and North Pacific have followed, and in some 345 countries, e.g. Norway, it is illegal to deliberately fish in coral areas even if the area is not 346 formally closed as a reserve. Since the mid-2000s a series of United Nations General 347 Assembly (UNGA) resolutions (e.g. 61/105, 64/72, 65/38) on sustainable fisheries have 348 called for a number of measures, including the implementation of the International 349 Guidelines for the Management of Deep-Sea Fisheries in the High Seas (FAO, 2009), and 350 action to avoid significant adverse impacts of fisheries on vulnerable marine ecosystems 351 such as coral areas. These resolutions refer specifically to areas beyond national 352 jurisdiction. Such actions run in parallel with efforts to create networks of marine 353 protected areas within national jurisdiction, partly motivated by the need to protect corals. 354 355 In response to the measures called for by the General Assembly, seamounts and 356 continental slope habitats with a documented or assumed coral presence have now been 357 set aside as marine reserves or fisheries closures within national jurisdiction of Australia, 358 New Zealand, the United States and in the North and Southeast Atlantic. In the Northeast 359 Atlantic, substantial areas have been protected within national jurisdiction in Norway, 360 Iceland, and EU member States. Beyond national jurisdiction, two RFMOs (Northwest 361 Atlantic Fisheries Organization, Northeast Atlantic Fisheries Commission) have closed a 362 range of seamounts and seabed areas to bottom fishing. These RFMOs also restricted 363 fishing to a limited agreed set of subareas, i.e. the “existing fishing areas”, and created 364 strict rules and impact assessment requirements. These measures are intended to protect 365 known areas with significant concentrations of cold-water coral but also essentially 366 reduce the incentive to explore areas outside these areas. Similar rules apply in the 367 Southeast Atlantic, implemented by the Southeast Atlantic Fisheries Organization 368 (SEAFO) which closed selected ridge sections and seamounts to fishing, and restricted 369 fisheries to certain subareas. In the Mediterranean, the General Fisheries Commission for 370 the Mediterranean (GFCM) implemented fisheries restriction zones in specific coral sites. 371 372 The Convention for the Conservation of Antarctic Marine Living Resources (CCAMLR, 373 http://www.ccamlr.org/) banned bottom trawl fishing within the CCAMLR convention 374 area. Bottom fishing regulations and area closures aim to prevent adverse impacts on 375 bottom-associated vulnerable marine communities. Marine protected areas in this area are 376 being considered but have thus far not been established. 377 378

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DRAFT: NOT TO BE QUOTED OR CITED FOR PUBLICATION Currently little information exists to assess the impacts on target or by-catch species by 379 deep-sea fishing on seamounts in the Indian Ocean. The Southern Indian Ocean Deepsea 380 Fishers Association declared a number of seamounts in the Southern Indian Ocean as 381 voluntary closed areas to fishing. The ratification in 2012 of the Southern Indian Ocean 382 Fisheries Agreement (SIOFA), a new RFMO for the region, may lead to better 383 documentation and regulation of seamount fisheries. 384 385 In the North Pacific, the United States designated Habitat Areas of Particular Concern 386 (HAPCs) that contain Essential Fish Habitat (EFH) and closed subareas of the shelf and 387 upper slope from California to the Aleutian Islands to bottom trawling. Canada also has a 388 strategy to develop and implement further measures. In areas beyond national jurisdiction 389 in the North and South Pacific, respectively, the North Pacific Fisheries Commission 390 (NPFC) and the South Pacific Fisheries Management Organization (SPRFMO) 391 introduced measures similar to those adopted by the Atlantic RFMOs. 392 393 In the United States within national jurisdiction in the Gulf of Mexico, mitigation areas 394 are established around mapped seafloor seismic anomalies that often coincide with 395 hardgrounds that may support cold-water coral communities. Although these measures 396 may prevent most direct impacts from infrastructure, the persistent threat of accidental 397 loss of gear and catastrophic oil spills remains a concern. 398 399 A continued challenge is to assess the effectiveness of current and new protective 400 measures and to develop management in areas that need greater attention, e.g. the 401 Caribbean and areas for which no RFMOs exist. The fisheries sector is often perceived as 402 representing the major threat to cold-water corals, but a growing challenge is to avoid 403 adverse impacts from other industries moving into areas containing known coral habitats, 404 e.g. mining, oil and gas industries, and renewable energy industries operating under 405 different management regimes. 406 407

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White, M., Wolff, G.A., Lundalv, T., et al., (2012). Cold-water coral ecosystem (Tisler 754 Reef, Norwegian Shelf) may be a hotspot for carbon cycling. Marine Ecology 755 Progress Series 465, 11-23. 756

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768 769 Fig. 42.1. Examples of dense cold-water Lophelia pertusa reef frameworks, including 770 provision of fish habitat. a and b from 400-500 m depth in the Viosca Knolls region of 771 the Gulf of Mexico. c and d from 150 to 400 m depth on the Logachev coral carbonate 772 mounds on the Rockall Bank in the Northeast Atlantic. 773 774 775

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DRAFT: NOT TO BE QUOTED OR CITED FOR PUBLICATION 776 777 778

779 780 Fig. 42.2. Global distribution of the major framework-forming cold-water corals (source 781 Freiwald et al., 2005 and more recent published data, n = 7213 entries). 782 783

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