Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | , 2 1,4 Discussions Biogeosciences , and T. J. Pitcher 1 in the OSPAR region and of , B. E. Narayanaswamy 2 1 ect the biological diversity and produc- ff , R. S. Santos 3 , F. Tempera 1 18952 18951 , C. Wienberg 1 erences will likely a ff , G. H. Taranto 1 in the Mediterranean Sea, which is much larger than previously 2 erent disciplines (Staudigel et al., 2010). Since small underwater fea- ff , G. Menezes 3 , K. Ø. Kvile 1 This discussion paper is/has been under review for the journal Biogeosciences (BG). Please refer to the corresponding final paper in BG if available. Centre of IMAR of the University of the Azores, DepartamentoSAMS, de Scottish Oceanografia Marine e Institute, Pescas Oban,MARUM, Argyll, Center , for UK Marine Environmental Sciences, University of Bremen, LeobenerFisheries Straße, Centre, University of British Columbia, Vancouver, Canada – Published: 20 December 2012 the needs of di tures may also play an important role in deep and high sea ecosystems (e.g., Koslow 1 Introduction Seamounts are ubiquitous underwaterally features in described the as world isolated(Menard, oceans elevations 1964; and greater International are than tradition- Hydrographic 1000ical m Organization, rationale in 2008). seems However, relief no to above2007) ecolog- support the and this the seafloor definition traditional has size been extensively limit modified (Pitcher in et the literature al., to 2007; better satisfy Wessel, in the region), 22 in25 the in Mediterranean the Sea North-East (9.2 Atlanticmapped % south in of both of all areas the seamounts are OSPAR in incharacteristics. have general in-situ the These very information. region) heterogeneous, di Seamounts and showing diversetion geophysical of resident and associated organisms. search, identifying examples ofseamount well-studied population in seamounts. the OurSea OSPAR is study area large showed (North-East with that aroundseamounts Atlantic) 1061 the occupy and and large in 202 seamount-like Mediterranean areasabout features, of 184 respectively. 000 Similarly, about km 1thought. 116 000 The km presence of seamountsthe in late the 19th North-East Century Atlantic butis has overall still been knowledge relatively known regarding poor. since seamount Only ecology 37 and seamounts geology in the OSPAR area (3.5 % of all seamounts This work aims at characterising theConvention seamount limits physiography (North-East and Atlantic biology Ocean) in and thepotential Mediterranean OSPAR Sea. abundance, We first location inferred secondly, and summarized the morphological existing characteristics biological, geological of and seamounts, oceanographic in-situ and re- Abstract Biogeosciences Discuss., 9, 18951–18992, 2012 www.biogeosciences-discuss.net/9/18951/2012/ doi:10.5194/bgd-9-18951-2012 © Author(s) 2012. CC Attribution 3.0 License. D. Hebbeln 1 and LARSyS Associated2 Laboratory, Universidade dos 9901-382 Ac¸ores, Horta, Portugal 3 28359 Bremen, Germany 4 Received: 23 November 2012 – Accepted: 30 November 2012 Correspondence to: T. Morato ([email protected]) Published by Copernicus Publications on behalf of the European Geosciences Union. Seamount physiography and biology in North-East Atlantic and Mediterranean Sea T. Morato 5 20 25 15 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | 2 ect pelagic ff cial OSPAR database (con- ffi 18954 18953 ) is an international legal instrument made of representatives of At the same time, the scientific community and international organizations recog- Recently, the importance of seamount ecosystems has been recognized by the The Oslo-Paris convention for the Protection of the North-East Atlantic (OSPAR; The total number of seamounts at a global scale has been estimated in several stud- moves long-lived commercially valuable fish species (Pitcher, 2010) that are extremely 2006; McClain, 2007; O’Hara, 2007; Shank, 2010). nized that human threats tosome seamount vulnerable ecosystems marine are growing ecosystems2009). fast (Pitcher and Therefore, et are some impacting al., actions 2010;sources have been Taranto (NEAFC, et taken 2011; al., to Santoscerns regulate 2012; et are the FAO, al., related use 2009; to of seamount 2010;building seamount fishing, organisms Morato especially re- (Williams et trawling et that al., al., physicallying 2010b). destroys 2010), communities reef- Major disturbs con- the by abundant sediment seamount filter re-suspension feed- (Clark et al., 2010) and selectively re- that draw marine scientists oflar, diverse recent backgrounds investigations into are seamount trying research. toand In reveal particu- evolutionary how geological, processes oceanographic, interact ecological altering to nutrient regimes, shape connectivity distinct amongdiversity distant communities (Genin populations on et and al., deep-sea seamounts 1986; habitat by Richer de Forges et al., 1987; Genin, 2004; Samadi et al., ducted in the region. scientific community, management(Stocks authorities, et al., industry 2012). Properties andand associated benthic with conservation life underwater (e.g., reliefs Hubbs, initiatives 1959; might2010a), Samadi a et influence al., global 2006; and Clark local etWhite al., circulation et 2010; (e.g., Morato al., Vastano et 2007) et al., and al.,Koppers enlighten and 1985; deep Watts, Roden, 2010; Earth 1987; Staudigel chemistry and and Clague, processes 2010; (Wessel, Wessel et 2007; al., 2010); all factors OSPAR area and about 59(2011) in estimated the the number Mediterranean of Sea.area seamount More greater as recently, than Kim 132 1000 m and (850 inthe Wessel greater height Mediterranean in than Sea. the 500 OSPAR However, m)datasets these while that estimates they were did consider based not no on seamounts consider global to the bathymetry occur most in recent multibeam bathymetry surveys con- the Kitchingman et al. (2007) dataset there may be at least 339 large seamounts in the http://www.ospar.org 125 000 to 25 millionet small seamounts al., or 2010; knolls Kimsuch greater estimates and than highlight 100 Wessel, m 2011; seamountsthe in as Yesson height seabed, et (Wessel one forming al., of one(Etnoyer et 2011). the of al., Despite most 2010). the this prevalent largest imprecision, set biomes of on habitats earth of with about 28.8 million km Mid Atlantic Ridge or in many exclusive economic zones (EEZs). For example, from al., 2010; Kim andare Wessel, unable 2011; to adequately Yessonglobal et detect al., small abundance and 2011). of deep Currently,proach seamounts peaks these would still and approaches be thus bear thebathymetric to large estimates profiles. locate uncertainties. However, of the seamounts the A smallbeam on more area bathymetry bathymetry of accurate prevents improved the grids ap- analyses oceanabundance originated of floor (Hillier seamount by explored and location, using shipboard Watts, morphologyworldwide multi- 2007). range and from Recent about 25 estimates 000 to of about the 140 000 number large features of and potentially seamounts from the Governments of 15has Contracting included Parties seamounts andfined in the these the European features Commission. list asabove OSPAR of undersea the threatened mountains surrounding whose and/or seafloorsulted summits declining (OSPAR, in 2008). rise habitats September The more and 2011)waters o than included de- of 1000 m 104 Norway, Sweden, seamountsThey Faroe in are, Islands, however, the still UK, underestimated High Ireland, since Seas many France, more and Spain seamounts territorial and are known Portugal. in the and defined elevations taller2007). than 1000 m as large seamounts (sensuies mainly Pitcher by running et mathematical al., algorithmssatellite on altimetry global and bathymetry grids acoustic inferred soundings from (e.g., Kitchingman and Lai, 2004; Wessel et et al., 2001), we considered features greater than 100 m in height as small seamounts 5 5 25 15 20 10 15 20 10 25 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | ect seamount ff erent bathymetry grids, the ff 18956 18955 the southern border of the Azores EE , where several studies have ff Mediterranean Sea as revealed by in-situ studies Atlantic) and Mediterranean Sea erent resolutions available for the OSPAR area (North-East Atlantic) and Mediter- ff This work aims at characterising the seamount physiography and biology in OSPAR However, not all seamounts share the same set of properties and threats (Rowden group located o contributed significantly to our current knowledge of seamounts. Thus, in this part of consulted literature: (1) knowledgeof scores, an used individual to seamount classifylogical based attributes; the on and degree a (2) of set actualof knowledge values, of attributes used key used to geological, to describe oceanographicpress) describe seamount and and attributes. individual in eco- The seamounts shown list in wasinformation Supplement adapted 3. regarding from In seamounts this Kville part not etstill of strictly al. belong the (in belonging study, to we to additionally thearound included the Northeast the OSPAR Canary, Atlantic. areas Madeira We but and refer Cape that specifically, Verde to archipelagos the and the seamounts Meteor located seamount Seamount characteristics in themining NE seamount related Atlantic information andof from Mediterranean the 245 were literature references determined and wereSea, online by consulted databases. including A for scientific total themunications publications, North (Supplement reports, Eastern 2). online Two Atlantic databases, main and and types Mediterranean of personal information com- were recorded from the outputs originated by eachoverlapped grid the were selection compared. ofresolution. When seamounts The parts final was of list undertaken the ofthen seamounts on compiled bathymetry in and the grids resulted the from layer OSPAR multiple areaology with sources. and used A the Mediterranean detailed is description Sea higher presented of was in the Supplement method- 1. 2.2 Seamount characteristics in the OSPAR area (North-East Atlantic) and pography with a set of parametricseamount-like thresholds features. to For determine this whether the study,ingman peaks we et represent applied al. a (2007) methodology andof developed adapted di by by Kitch- Morato etranean al. Sea (2008) (Fig. on 1). several bathymetric After grids applying the methodology to di highlights topographic local maxima or peaks and then compares the surrounding to- try or ship-based soundings)abundance. soundings This is generally key to implies any a attempt two-steps to seamount estimate isolation seamounts process that first build future seamount research and management. 2 Methods 2.1 Seamount abundance and distribution in the OSPAR area (North-East The ability to identify seamounts from bathymetric grids (derived from satellite altime- formation available. Secondly, weoceanographic summarized in-situ the research, pointing existing out biological, gapsdistinctive in geological features seamount and investigation, and potentially examplesachieve of a snapshot well-studied of seamounts. the In knowledgeAtlantic status this and of seamount way, Mediterranean we ecosystems areas, in hope which the to North-East may represent basic information on which to 2012). Therefore, a clear understandingsity, of production spatial and patterns geophysical ofand properties threats, management is biological actions diver- crucial (Morato for etgraphic, al., geological the and 2010b) success ecological and of processes for conservation associated the with understanding seamounts. of oceano- area, North-East Atlantic, and thedance, Mediterranean location Sea. We and first morphological inferredand potential characteristics Mediterranean abun- of waters seamounts applying in a the detection OSPAR model area on the latest bathymetric in- is an emerging issueecosystems in in the seamount future managementVan (Halfar Dover, that 2011). and may Fujita, seriously 2007; a Hein et al.,et 2010; al., He 2005; et Morato al., et 2011; al., 2010c; Clark et al., 2012; Taranto et al., 2012; Kvile et al., vulnerable to heavy fishing (Morato et al., 2006). Besides fisheries, deep-sea mining 5 5 25 15 20 10 25 15 20 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | 202; 284) 1061; = = = n 2.7). Ac- n cial statis- = ; SD 377) for the ffi 2 = 3.5) while Mediter- = 1004 km erent heights show a ff for OSPAR seamounts = ◦ 14.8 km (SD b ) and the average slope r a = ξ . The relationship between 104 (OSPAR o b ◦ 1.50 r = ± n 303). Predicted seamount area 15.9 km (SD 15.54 = 4.87 = = 132 (Kim and Wessel, 2011) for the = b r ϕ = ; SD ϕ 2 n to ◦ erent with a greater proportion of seamounts for Mediterranean Sea seamounts). However, ff 18958 18957 0.027, 0.98 ◦ 910 km ). Our estimates are much higher when com- ± ) area and in the Mediterranean Sea ( 2 = 2 = ϕ 1.91 b ), the height-radius ratio ( a 67; 6.3 %) of the large seamount-like features in the ± 394) for the Mediterranean Sea. Predicted seamount b 0.085 r = = = n r 5.59 ξ = ϕ 59 for the Mediterranean Sea. These discrepancies are due to = 994; 93.7 %) of the summits located between 1000 and 3000 m n = 0.034, n er much between both areas. Large seamounts in the OSPAR region ± ff erent regions ( ff 339 (Kitchingman et al., 2007) and 0.098 = = n r Atlantic) and Mediterranean Sea ξ ), did not di Since our estimates included the bathymetry grids with highest resolution currently Nevertheless, the application of our methodology to the new bathymetry grids Seamounts in OSPAR area (North-East Atlantic) are mostly deep (Fig. 4) with a ϕ ranean regions remain partially unresolveddata, and with are largely only based a on small satellite altimetry portion resulting from finer ship-borne acoustic surveys. The gion. However, our results are similaret to the al. number (2011) of who seamounts ran estimatedmated by a 1288 Yesson similar seamounts methodology greater onthe than the Mediterranean 1000 GEBCO m Sea. bathymetry in grid, height and in esti- the OSPAR areaavailable and they 180 are in athus step more reliable further than the in datasets theties previously published. still identification Nonetheless need large of to uncertain- be seamounts tackled. in The bathymetry the grids region, available for and the OSPAR and Mediter- about 0.08 seamountspared per with 1000 km previouslytics), available seamount numbers: OSPAR area, and the fact that we used the most recent available bathymetry grids available for the re- distributions are shown inlar Fig. for 6. the di Additional seamountand characteristics were also simi- note that most ofthe the slope angles features that are rangedslope heterogeneous angle from and as summit height demonstrated, for for seamounts in example, the by OSPAR area isdemonstrated presented in that Fig. 7. seamountsabout may seamounts be 0.08 common per features 1000 km in the OSPAR ( were estimated to have aranean mean Sea basal seamounts radius showed of acordingly, mean basal basal areas radius are of and similar Mediterranean for seamounts OSPAR ( seamounts ( ( height distributions are shownradius in of Fig. the 5. seamount Shapes base of ( seamounts, as characterized by large majority ( depth. Only a smallOSPAR portion area are ( shallow, havingserved summits in at the less Mediterranean than Seabeing were 500 di m shallower depth. than The 500the patterns m OSPAR ob- area depth and (Fig. themean Mediterranean 4). height Sea Other of were, characteristics the however, similar.OSPAR identified of For area seamounts example, seamounts and was the from 1400 calculated m as (SD 1316 m (SD relationship between the cumulativestrong counts linear of relationship seamounts (Fig. ofwith 3). di 1000 Nevertheless, m the in cumulative height countsgory fell below of are the probably the fitted somewhat features line, underestimated. indicating that the numbers for this cate- 3.1 Seamount abundance and distribution in the OSPAR area (North-East A total of 1061 potentialin large seamounts the (heights OSPAR greater area, thanranean 1000 while m) Sea 202 were (Fig. identified seamount-like 2). featuresble The were 1, accepted while identified outputs the in of finaldepth each the list of bathymetric of Mediter- grid seamounts the generated are summitadequately by shown identified and our in topographic algorithm average structures Ta- – slope with including heights – location, larger is than 1,000m, provided since in the Supplement 4. Our method referring exclusively to seamounts in the OSPAR area. 3 Results and discussion the study we will refer to seamounts sampled in the North-East Atlantic rather than 5 5 25 15 20 10 20 25 15 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | erences ff ˜ ao Valente ˜ ao de Castro Bank in the OSPAR area and of 2 cult to formulate. ffi ) than in the North-East Atlantic ◦ ect their average productivity. 20 ff = ϕ 18960 18959 in the Mediterranean Sea, which is larger than generally thought. 2 erences in these characteristics will most likely influence the biological ff ). Di ◦ Mediterranean Sea 13 = Most of the detailed information available regarding seamount ecology in the NE The 84 explored seamounts comprise mostly large features, with an average height In terms of seamount geology, the most commonly known parameters were summit Nevertheless, our estimates draw seamounts as important and relevant areas of ϕ Sea. Research on seamount fish communities has been conducted in various locations Atlantic and Mediterranean Seation (Fig. available 9) for is 56seamounts) % related and of to deep-water the benthic fishcorals smapled invertebrates (49 (42 % seamounts), (informa- % of demersal of theon (51 the sampled % 36 seamounts). sampled of of Demersal seamounts) the invertebrates the(cnidarians and have seamounts of sampled been the deep-water in sampled class the ,and excluding NE Ceriantharia) the Atlantic have sea been anemones and reported in on 11and the 33 and in orders Mediterranean 8 Actiniaria the Sea, seamounts in Mediterranean respectively. the Seamountin North-East Sea. fish Atlantic about communities 34 have seamounts been in studied the North-East Atlantic and about 9 in the Mediterranean of volcanic origin, butseamounts interestingly (11 hydrothermal % activity of studied hashas seamounts). only been Of explored been in these, reported detail, the withpotentially on Azorean reports adapted 9 Jo of to a metal unique rich ecosystem waters of et (Colac¸o protists al., and 2006). bacteria communities of these seamounts and will likely a of over 1800 m. The5425 m Great Meteor above the is(see sea Sect. the 3.3.6). bottom, largest For known being comparison,4810 seamount m. Mount higher and The Blanc, than reaches depths the of highest anyBank, the peak mountain Cape seamount’ in Verde; summits in Monteiro the et range Alps, Western al., from isAvraham 2008) et a Europe “only” to al., few 2554 meters 1983), m showing (Jo (Alfeo Seamount, an Ionian average depth Sea; Ben- of about 600 m. Most seamounts are 71 km in theranean North-East Sea Atlantic) and andbetween 560 average m the summit in depth twofor the (625 areas. m seamounts North-East The in in Atlantic) the average( the Mediter- did slope Mediterranean not of Sea show the ( major studied di seamounts was higher average proximity of a neighbouring seamount (79 km in the Mediterranean Sea and those from the North-Eastthe Atlantic, Mediterranean since Sea the distance and to about the 1100 km shelf for averaged 61km the for North-East Atlantic. However, the the Azores region, and the Mediterranean Sea’s Eastern and Westerndepth, basins. height and ageSea. (Fig. 9), In both general, in(2 the million seamounts yr North-East in studied Atlantic average) and than inwhich the those may the Mediterranean in have the Mediterranean a North-Eastexample, direct age Sea Atlantic implication of (39 million are the in yr seamount in much theally, may seamounts average), be biological younger in linked diversity the to of the Mediterranean presence seamount Sea of since, are endemisms. much for Addition- less geographically isolated than From our database, ain total the of Mediterranean Sea 37 (Fig.poses seamounts 8) or in have directly the been sampledinformation OSPAR mapped for area regarding bathymetrically biological and 25 or (Supplement 22 seamounts geologicalSect. 5). seamounts pur- not Additionally, 2.2). strictly we The belonging included Greenland to areas basin, the the where OSPAR areas seamount Trough, (see the research Bay has of Biscay, been the Western conducted Iberian include Margin, the Overall, seamounts from theselarge regions ranges showed in a size, widemorphological depth, variety characteristics slope of demonstrates angles characteristics oncethe and with again same areas that set of of not properties, their all making bases. seamounts generalisations Such share di 3.2 diversity in Seamount research in the OSPAR area (North-East Atlantic) and preclude their detection by the algorithms.as Hence, our an results approximate still list have to of be OSPAR considered area and Mediterranean Seathe seamounts. seabed occupying aabout total 183 area 856 km of about 1 115 925 km coarse resolution of such grids results in the blurring of smaller seafloor elevations that 5 5 25 15 20 10 25 15 20 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | 18962 18961 , 2009) has shed further light on both oceanographic and eco- ff Conservation organizations have recently been engaged in seamount research in In the Mediterranean Sea, seamount research has traditionally been focused on Ever since, other seamounts such as the Josephine and the Great Meteor seamount Prince Albert I of Monaco was one pioneer of the marine science in general and Stony corals (order ) are the most common deep-sea corals (reported both the North-East Atlanticthe and knowledge Mediterranean, of which some2008, features has 2010). in augmented These terms considerably findings oftected promoted Area habitat the (MPA) and in creation the species of Mediterranean diversity the Sea (OCEANA, first (see Sect. seamount 3.3.7). Marine Pro- regarding seamount ecology reflect a growing(e.g., interest for Ettoumi this et field in al., theand 2010; Mediterranean the Bo Eastern Mediterranean et regions, al., excludingremain 2011; few largely features, Sevastou unknown. seamount et ecosystems al., 2011). In the Cape Verde al, 2010b; Giacomello and Menezes, 2011). geology with manymapped. Western In particular, Mediterranean seamountsterms seamounts in of being the volcanology and Tyrrhenian bathymetrically mineralogy SeaCompared (e.g., are well- Trua to et relatively al., geological well-studied 2002; exploration,Sea Dekov in and has early Savelli, been 2004). ecological sparse: only researchPerrone, a 1982; in few Strusi publications the et are al., Mediterranean available 1985; from Galil the and last Zibrowius, century 1998). (e.g., However, recent studies (Christiansen and Wol logical aspects of twostrated Northeast the importance Atlantic seamounts ofthe multidisciplinary (Seine seamounts research studies. and was This Sedlo) recentlywhich multidisciplinary extended and in was approach demon- Condor temporary of seamount closed (see Sect. to 3.3.3), fishing for longer term research purposes (Morato et Nellen, 2004; Nellen and Ruseler, 2004; Tempera et al., 2012). The OASIS project zonation studies (e.g., Rogers, 1994; Uiblein et al., 1999; Fock et al., 2002; Martin and 1970s, when German scientistsHesthagen, led 1970). the way with the Great Meteorcomplex expeditions have (e.g., been subjecttrastingly, many to other features immense have been ecological bathymetricallyreceived mapped research no but ecological to (see our research Sect. knowledge (e.g.,1984). the 3.3.6). Vigo Since Con- and the Porto seamounts, transitionstudies Mougenot regarding into et seamount al., the ecology. Biological 21stand studies century, have Great continued a Meteor in seamounts, boost the with Azorean occurred a in focus on the fisheries, number plankton of and benthic biotope charts and identifiedseamounts the in major 1896 (Porteiro, topographical 2009).the features, The area diversity including was and the abundance remarkableence of Princess (Carpine-Lancre of species Alice and found seamounts in Saldanha,Century in 1992). (Buchanan, the Although 1886), North-East the little pres- Atlantic information on has their been ecology known was since published until the the late 19th tipatharia) have also beencomprised arthropods, found mainly on decapod manysea crustaceans, spiders seamounts. but (class Other Pycnogonida), also intenselyoderms, mollusks, other in both studied crustaceans particular gastropods taxa and sea and urchinsand bivalves, (Class bryozoans and Echinoidea). have echin- also Taxa such been as sampled , from annelids many of theseamount seamounts. research in particular.paigns Between in 1885 the and Azores 1911 area he commanded of 13 the cam- North East Atlantic, producing new bathymetric along the Iberian marginpending on (e.g., the Menezes depth et of(bathydemersal the al., or seamount, bathypelagic) 2006, or the more dominant 2009; shallow species Pakhorukov, species are 2008). (demersal typical or De- deep-sea benthopelagic). fish on 27 of the abovementioned seamounts), but gorgonians and black corals (order An- within the OSPAR area, in particular around the Azorean and Meteor seamounts and 5 5 15 20 10 25 15 20 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | spp.), culties ffi (Tempera 2 ) were observed Caryophyllia 40 km in diameter) (Ja- ∼ pertusa W) ◦ W) 18964 18963 ◦ W) and Rosemary Bank Seamounts 70 km in diameter). Recent work undertaken on ◦ ∼ N, 29.04 ◦ N, 9.13 erent benthic invertebrate communities inhabiting this ff ◦ N, 11.1 ◦ W) ◦ N, 10.15 ◦ erent habitats. In addition, starfish, brittle stars, octocorals, hydrozoans and ff (59.25 Mediterranean Sea as revealed by in-situ studies Anton Dohrn and Rosemary Bank seamounts were sampled in detail in 2005 and about 26 km in lengthing and from 7.4 180 km to wide more at than the 1000 1000 m, m and depth an contour, estimated with planar depths area rang- of 432 km Area of Conservation (JNCC, 2011).cup On corals, sessile the holuthurians summit and of anemonesploughmarks, Rosemary were whilst Bank present on at fauna the the such edge terraced as ing of slopes stony the a iceberg corals, high gorgonians, diversity spongesepi-fauna of found and fauna at anemones. was the Ophiuroids found base includ- were of the Rosemary dominant Bank (Howell3.3.3 et al., 2007, Condor 2010). Seamount (38.55 Condor seamount is located about 17 km to the southwest of Faial Island (Azores), is addition cold scleractinian deep-watersupporting corals large assemblages (e.g., of sponges,and gorgonians, cup other corals fauna ( mobile on fauna the with summit noticeable(Narayanaswamy et edge changes al., (JNCC, in 2006). Anton 2011). species Dohrn dominance Echinoids has with recently were been increasing the proposed depths as dominant a Special associated with sampling steep topography.of Anton Dohrn deep-water has fishing been and howeverimagery the numerous (Jacobs, focus 2006). trawl-marks were observed on side-scan2006 sonar (Narayanaswamy et al., 2006;ing Howell place et in al., 2009 2007),covered (Stewart with by et additional al., a sampling 2009). tak- largelarge The number areas summit of of of sandy/gravel substrate Anton supporting valves Dohrn encrusting and was serpulids found and to sponges. plates be In interrupted by Rosemary Bank has found a numberslide of scars small and parasitic cones slopes (Jacobs,to that 2006) as an are well terraced area as (Howe commonlydeep-waters et of termed al., the the North-East 2006). “cradle Atlantic Boththe were of seamounts seamount sampled and belong deep-sea by banks Victorian biology” were scientists. (Gage, rarely However, sampled 2001), in as those the days because of the di elongate dome-shaped seamount ( cobs, 2006). The flankshills/parasitic cones are on devoid the of eastern and canyons northwestern and sides. gullies. Rosemary Bank Instead, is there a more are a few 3.3.2 Anton Dohrn (57.45 The Anton Dohrn andNE Rosemary Atlantic, Bank about seamounts 120 km areAnton northwest located of Dohrn in the the is UKto Rockall mainland a Trough, and a classically belong to relatively circular the flat UK’s shaped EEZ. completely seamount sediment with covered steep summit flanks ( that lead sponges, bryozoans and serpulid polychaetessummit cover large and proportions flanks, of with the depth-wise seamount or changes muddy in species substrate composition. within Patchesrange these of of mats, sandy di as wellbivalves as inhabit occasional this rocky seamount. The outcrops, Vesterisbankenthe create fauna rapid a is input likely of highly energy dependent from on upper waters during the short Arctic summer. in the OSPAR area forture which was detailed information visited is bydescribe available. This the researchers substrate large, during composition isolated and the fea- 3000 bathymetry 1980s of m this and high seamount, feature early whichan with is 1990s. a almost two Detailed nearly year-round peaks studies ice-cover, (Hempelgations it et in is al., the not 1991; Arctic. Cherkis straightforwarda However, one to thorough et expedition description al., conduct in of 1994). biological 1990 theseamount, With investi- di (Henrich from et the al., top 1992) of the provided summit to the deepest flanks. Bioherms constructed by 3.3.1 Vesterisbanken (73.52 The Vesterisbanken Seamount in the Greenland Basin is the northernmost seamount 3.3 Seamount characteristics in the OSPAR area (North-East Atlantic) and 5 5 25 15 20 10 25 15 20 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Madrepora and 16 %) (Wienberg et al., 2012). > Lophelia pertusa W) ◦ 18966 18965 N, 11.95 ◦ 67 %) and soft sediments ( > (Wienberg et al., 2012). 20 cm in diameter) thickets, namely of the species However, dead but relatively fresh-looking framework was often observed to be The south-western top of Coral Patch seamount is dominated by two major substrate Condor seamount has been an important fishing ground for bottom longline and Condor seamount hosts a peculiar multi-scale dynamic oceanographic conditions < was also found on the seamount (Wienberg et al., 2012). ( oculata entangled in lost fishing linesfishing indicating activities. that Overall, the clear seamount evidence is forplaces likely anthropogenic highly impact at frequented was by the found atfishing top several lines of that got Coral entangled withcoral Patch rocky frameworks. boulders seamount To a and minor that fractured degree pavement also mainly outcrops litter or comprising (small-sized unidentified the plastic objects) remnants of The macrofauna on Coralganisms Patch with seamount a comprises clear benthicoccurrences dominance to as of epibenthic well suspension-feeders, living which asall, or- were their the observed skeletal macrofauna as remnants shows live and (e.g., low shell diversity hash, which and coralwhich were abundance debris). the frequently except seamount Over- observed. for derived its cidarid Scleractinian name, echinoids deep-water are corals, rare and from just comprise isolated small-sized Seine seamount and Portugal.elongated seamount, It about exhibits 120 Coral kmits long Patch south-western and top is reaching 70 a kmSeismic a and sub-elliptical wide water bathymetric (D’Oriano WSW-ENE- depth data et show ofdistinct that al., about coalescent Coral 2010) 560 Patch volcanic m is cones with (Wienberg a developedemplaced composite on et on structure its al., a with top 2012). nine pre-existing (D’Oriano sedimentary et structural al., high 2010) (Zitellini that et are types: al., 2009). hard substrate ( (Menezes et al., 2011). 3.3.4 Coral Patch seamount (34.93 Coral Patch is a still largely unexplored seamount, located roughly half-way between of the fish species and the time scales required for recoveries to earlier abundances dence from outside source areas appear to be important determinants for the status handline fishing (Giacomello andnever Menezes, employed 2011). may The help factin explain that a why bottom relatively the trawling good seamounting, was state still recreational (Tempera hosts fishing, et benthictant whale al., communities ancillary watching 2012). activities and Research in the activities, scuba-divingMenezes, 2011). Condor big-game activities Intense seamount handline fish- but are and to longlinetant also a fisheries reductions impor- lesser have, however, in extent caused (Giacomello the impor- characteristics, and abundance their of degree many of demersal residency fish on species. the Species Condor life seamount, history and their depen- mesopelagic organisms and the trappingto of lateral advection the increase latter the above2011). carbon the Preliminary supply seamount, to results in the of addition Condorgeneral the seamount low study et (Colac¸o availability of al., of food sediment for2011). microbial benthic assemblages consumers at revealed the a Condor summit (Bongiorni, including enhanced mixing,structures upwelling-downwelling over processes the and seamount, that closedmachnikov make et circulation it al, distinct from 2011).deep-water the It coral surrounding gardens ocean hosts (Bash- and habitatsBraga-Henriques deep-sea of et al., conservation aggregations 2011). importance, (Temperatrophic et The constituted levels Condor al., with by mesopelagic seamount 2012; organisms foodand being chain the the is link benthic composed between and the of epipelagic benthopelagic five realm organisms. Predation of epipelagic organisms by ther described by Temperamoratorium, et established al. in (2012). 2010, The allowingimprove seamount the the installation is understanding of under of a aet seamount scientific temporary al., observatory ecological fishing 2010b). to structure This andwith moratorium the functioning stakeholders. will (Morato last until 2014, when its future will be discussed et al., 2012). The main geomorphologic characteristics of the seamounts were fur- 5 5 25 15 20 10 25 15 20 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | erences in fish abun- erent from that of the ff ff erent species having been ¨ uller, 2002; Bartsch, 2008). ff cult to distinguish if such dif- ffi erent fishing pressures. A study ff erent invertebrate phyla have been ff W) , 2009). This scientific initiative assessed ◦ ff W) ◦ 18968 18967 ects” or to di ff N, 28.50 ◦ N, 14.38 ◦ stegui et al., 2009; Martin and Christiansen, 2009) but a ı ´ ect“ was observed on the microbial community structure and biomass ff ect on the fluxes of organic matter, with higher concentration of particulate ff ˜ no et al., 2000). Low local productivity might explain why the demersal fauna of The Great Meteor is among the few seamounts in the North-eastern Atlantic where The majority of the cruises that have visited the Great Meteor sampled the demer- The OASIS project did not find any evidence for constant high productivity at the ping of vertically migrating organisms abovea the seamount resource summit, which for might the provide 2004; demersal Schnack-Schiel community and (e.g., Henning,column Fock 2004). et that There al., were might 2002; alsomann retain observations Nellen and larvae, of and Mohn, a plankton Ruseler, 2002).production Taylor However, or the above nutrients levels the of seamount around primarysurrounding seem the open and not ocean, secondary seamount to though (zooplankton) be (Beck- Mouri temporary significantly variation di exists (Martinthis and seamount Nellen, is 2004; relatively poor. Fish species from the Great Meteor Seamount showed tal slope and neighbouring seamountsPerhaps surprisingly, (Piepenburg these and M studies foundGuyot to the be invertebrate relatively fauna poor, on both the in terms Great of Meteor abundancesecosystem and dynamics species have diversity. been addressed. Several studies have indicated the trap- though some geological investigationsWatts were et conducted al., in 1975; the Wendt et 1960s al., and 1976; 1970s Stackelberg (e.g., etsal al., invertebrate 1979). fauna. Aregarding total this of attribute. 16 Mostonomic references studies groups, have are been such qualitative, incorporated asfas, and copepods in 2007). often or the Nevertheless, focus some gastropods SEEF on quantitative (Georgeboth specific studies abundances and have tax- and Schminke, been diversity 2002; conducted, of Go- comparing invertebrate megafauna with the adjacent continen- The Great Meteor seamount, situatedseamounts South in of the the Azores, North isthe East one Atlantic, sea of rising the surface. largest from It known Atlantic about (Kville is 4500 et m al., also depth 2012). to the Its history 270 most m of below research thoroughly is studied dominated by seamount biological sampling, in the North-East 3.3.6 Great Meteor Guyot (30.00 two seamounts studied (Ar “seamount e et(Mendonc¸a al., 2012). Vilasated et an al. e (2009) furtherorganic suggested matter that above the the Seine seamounttute summit cre- an than important at food other source similardone for depths. to the This confirm benthic might community, this. consti- but further studies need to be et al. (2009) higheradjacent fish island abundances slopes were of observed similarferences at depths. were However, Seine it’s related seamount di to comparedconducted “seamount to by e Christiansen etdance al. at (2009) Seine seamount. found no significant di oceanographic, geochemical and biological aspectsof of Madeira Seine seamount Island), (Northeastern whichperiod. The were substrate visited of inproviding the habitat total Seine both seamount seven for is times sessileburrowing a megabenthos infauna over mix such (Bett a et as of al., sponges during soft 2004). andidentified sediment A three-year corals total for and as of the hard 11 well seamount. di rock, as ersal In addition, and important benthopelagic studies fishobserved were community (Christiansen made with about et about the 50 al., dem- di 2009; Menezes et al., 2009). According to Menezes A seamount for which our knowledge hasSeine increased seamount, rapidly situated over near the the last fewthis Madeira years Islands. is seamount Virtually the nothing until wasStudy) known the project about in initiation 2002 of (Christiansen and the Wol OASIS (OceAnic Seamounts: an Integrated 3.3.5 Seine seamount (33.76 5 5 25 15 20 10 25 15 20 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | in the 2 E), and Emile ◦ N, 1.80 ◦ E) ◦ erent taxa were observed. Emile Bau- ff N, 32.67 ◦ 18970 18969 W), Ausias March (38.73 ◦ E) seamounts ◦ N, 2.85 ◦ N, 2.33 ◦ , much larger than previously thought. The 84 seamounts in the NE Atlantic 2 nities with the African coast, Madeira, and the Canary Islands (Uiblein et al., ffi Baudot (38.70 The seamounts in the Mallorca Channel had not been biologically researched until 184 000 km and Mediterranean from which we have found in-situ information are in general large 4 Conclusions Our study revealed that the seamount populationranean in the Sea OSPAR area is and inarea large the Mediter- and with about around 202vious 1061 in estimates seamount-like the features since Mediterranean inSimilarly, we these Sea. the topographic used These features whole occupy the values aOSPAR OSPAR latest area. large were In area bathymetry the higher of Mediterranean about than grids Sea, 1 seamounts pre- 116 available occupy 000 an km for estimated area the of about region. a nearly untouched areaindicated of relatively seamount high research. meiofaunal diversity Thesity with preliminary with a results increasing decrease of in depthand abundance this (Sevastou diversity and study were et diver- relatively al., low 2011). (Narayanaswamy, 2012). However, macrofaunal abundance gested recent hydrothermal activityprovided (Varnavas details about et both al., substrateatosthenes, 1988). composition a and Later, the platform a geological built drilling history1998). up of program There the mainly Er- was by onlycentury, limestone but one on the biological continental samples investigation were crustand on Zibrowius, described (Robertson, 1998). as this Various epifaunal “surprisingly seamount taxaof diverse” were during the for found seamount, attached the the including to region stony last the corals (Galil hardrecently and substrate been sponges. revisited The by Eratosthenes scientists Seamount investigating meiofaunal has and macrofaunal diversity, The Eratosthenes seamount, locatedscribed in the from Eastern geological Mediterranean surveysdefine Sea, that the was used regional first bathymetric, topography de- seismic1984). (e.g., and Early Ben-Avraham descriptions magnetic et of data hydrothermal al., manganese to 1983; deposits from Tanner the and seamount Williams, sug- 3.3.8 Eratosthenes Seamount (33.67 of encrusting red algae,anemones which and were often were also coveredSes frequently by observed. sponges. Olives The Deep-sea is summit corals, of largelymollusks. the sea covered deeper A by large sandy numbermany sediment of are of and demersal commercial fish inhabited interest.and species Emilie by at Baudot was crustaceans has Ausias also been and March a reported, damagework much from among by used OCEANA bottom fishing which these ground, trawling has threesigned been seamounts Marine along observed. Protected Areas with Based by the on the Bell the Spanish Guyot government were (Madina, recently 2011). de- depth, while Emile Baudot and150 Ausias m March and are 125 m, shallower respectively. seamounts, with depths of the sampling carried out by the2008). conservation During organization OCEANA intense in sampling 2007 (OCEANA, dot around 200 and di Ausias March both reach within the photic zone, allowing the development Three seamounts in theconservationists working Channel to of preserve the Mallorca biodiversityet have of al. the recently Mediterranean (2003, received Sea. 2004) Acosta theOlives, have Mont attention Ausias described of March the andMediterranean Emile geomorphology features Baudot. of are Compared located these to in Atlantic features,Olives, relatively seamounts, called and shallow these water Mont Ses (300–650 Ausias m Marcwhile depth). are the Ses both more flat-topped south-easterly andof located pinnacles, probably Emile and of Baudot might continental be is of origin, surrounded volcanic by origin. a The summit large of number Ses Olives, reaches 300 m 1999). 3.3.7 Ses Olives (36.52 strong a 5 5 20 25 15 10 25 15 20 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | th ´ erie Es- , 2011. ´ erie Estudos 1/2012, Horta, ect the biological diversity ff erent ages and slopes. Moreover, ff ˜ oz, A., Herranz, P., Urgeles, R., Palomo, C., 18972 18971 doi:10.1371/journal.pone.0016357 erences will likely a ff . ´ opez-Martinez, J., Mun This research has received funding from the European Community 7 stegui, J., Mendonc¸a, A., Vilas, J. C., Espino, M., Polo, I., Montero, M. F., and Martins, seamount ecosystems, Final Project Report,Portugal, Arquivos do 92–99, DOP,S 2011. Ionian Sea, eastern Mediterranean, Mar. Geophys. Res., 5, 389–404,thos: 1983. Seine and Sedlo SeamountsCentre, (NE DEEPSEAS Atlantic), Group, Data Southampton, report UK, 2004. for the National Oceanography Gasparini, G., Misic, C., Povero,acteristics P., Pusceddu, of A., Schroeder, the K.,Tyrrhenian Mesophotic and Sea), Bavestrello, PLoS Megabenthic G.: ONE, Char- Assemblages 6, e16357, of the Vercellidor Seamount seamount, (North in CONDOR Observatory for long-term study and monitoring of Azorean Retention potential of the seamount-induced circulation, Ocean Dynam., 52, 194–204, 2002. 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32 800 grid; http://www.gebco.net/ . . orth orth East Atlantic 000000 Mediterranean Mediterranean data/ http://www.emodnet-hydrography. grid; grid; 5 is the Azores grid; 6 is

0310019 = bathymetry 50 grid; 8 is the . 1 Arctic grid; 2 is the N . 2

. 18984 18983 http://www.ipgp.fr/rech/lgm/MOMAR editerranean 2 products/gridded https://jetstream.gsi.ie/iwdds/index.html outh Western Iberian Margin and orn¸ etLourenc¸o al. (1999). available at: SWIM project. It bringsecuted together between 19 multibeam 2000 bathymetry andmately surveys 2006 180 ex- 000 and km covering a total area of approxi- The resolution of the2.5-km Polarstereographic grid to has 2-km. been TheJakobsson increased compilation et from of al. (2008). IBCAO Version 2.0 is described in controlled ship depthpoints soundings guided with interpolation by between satellite-derived sounding gravity data. data and Data Network (EMODNET).0.25 arc-minutes The using EGB the wassources best built available using at bathymetry a a from variety resolution a of of variety methods. of ological Survey of Ireland; Interactiveable Web at: Data Delivery System avail- eu/content/content.asp?menu (Table 1 for details) (Table

grid; 7 is the M land grid; 4 is S 77 MOMAR GIS Database v2 (08 October 2010) with a resolution of 90 m 11 Bathymetric dataset compilation funded by the ESF Euro Margins Number seamounts 4 EMODNET Gridded Bathymetry (EGB). European Marine Observation id Atlantic Ridge Number of seamounts identified by the bathymetry grids used in this study. Outlines of the bathymetry layers used to detect seamounts in the OSPAR area (North

Figure 1. Outlines of the bathymetry layers used to detect seamounts (North East in Atlantic) and the the Mediterranean OSPAR Sea area grid; 3 is the Ire the M grid 9 is the GEBCO and

3 4 5 6 7 8 9 1 2 AzoresMid Atlantic Ridge 104 Includes multibeam mapping and global grid data. Available from South West Iberia Margin Arctic 244GEBCO/All 610 IBCAO Grid v. 2.0. Includes the A recent years global of 30 arc-second multibeam grid mapping. largely generated by combining quality- Mediterranean 202 MED 250 mMED 800 m 151 51 As in North East Atlantic As in GEBCO/All AreaOSPARNorth of East Atlantic 1061 Bathymetry Grid Ireland 11 Digital terrain model (DTM) derived from swath bathymetry from the Ge- Fig. 1. East Atlantic) andis the the Mediterranean Ireland Sea. grid;Atlantic 1 4 Ridge Arctic is grid; grid; South 7the 2 is Western GEBCO the is Iberian grid Mediterranean Margin the (Table 1 250 grid; North for grid; details). 5 8 East is is Atlantic the the grid; Mediterranean Azores 800 3 grid; grid; 6 and is 9 the is Mid Table 1.

) 34

. ) dark dots Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper( | grey dots grey = n Sea (

33 . The relationships can be s 3500 = = 0.98 for the OSPAR area

2 3000 in the OSPAR (n=in the area 0.98 for the OSPAR area (dark dots) = (black dots) (black for for identified seamount

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Figure 1061) and in the Mediterranean Sea (n= 202) grids. H ( v

4 5 6 1 2 3 2 3 4 5 6 1 Fig. 3. and expressed as Fig. 2. 1061) and in thegrids. Mediterranean Shaded Sea layer ( shows the limits of the OSPAR area. Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper |

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Figure in the features

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2 3 4 5 1 5 6 2 3 4 1

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2 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussionr Paper |

37 + 0.80; + 1800 h ) in the h

1600 1400 4000 0.62. 1200 = ) 2 2 r 1000 Area (km Area 800 lope angle (φ) and seamount height ( Mediterranean Sea Mediterranean 0.80; + 600

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30 25 20 15 10 Frequency 2000 Height (m) Height 18990 18989 1800 1600 1400 like features. Bin size features. like is 100 m. - 1200 1000 ) 2 1000 . The expressed . The be asrelationship 0.003 φ can = Area (km Area 800 OSPAR area OSPAR Relationship between sea . 600 7 0 400

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0 0

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Figure (right) seamount Mediterranean 4 5 2 3 1 Fig. 7. OSPAR area. The relationship can be expressed as Mediterranean (right) seamount-like features. Bin size is 100 m. Fig. 6.

5 3 4 1 2 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper |

) 40 70

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39 that have 60

50 NE Atlantic (dark in the 40 30 erent attributes in the NE Atlantic (dark grey) and Number of sampled seamountssampledof Number ff 20 18992 18991 North North East Atlantic and Mediterranean Sea

. 10 ology or oceanography data (for list of seamounts, see sampled sampled for different attributes (grey) Shaded layer shows the limits layer shows the OSPAR of Shaded area.

a ). 5

0 Se Age Height Volume Endemism Taylor cap Taylor Elongation Basin depth Basin Macrophytes Cephalopods Demersal fish Demersal Summit depth Summit Summitshape type Substrate formingcorals - . Location of seamounts in the Mineral deposits Mineral Demersal invert. Demersal 8 breathing visitors breathing Sponge meadows Sponge Mediterranean Mediterranean Cold water Corals water Cold - Slope of seamount ofSlope prod. Zooplankton Nutrient upwelling Nutrient Enhanced org.mat. Enhanced . Number of seamounts Phytoplanktonprod. Aggregating DW fish DW Aggregating Air 9

Topographic blockage Topographic Visiting large pelagics large Visiting Hydrothermally active Hydrothermally Habitat

Chemosynthetic comm. Chemosynthetic Seamount attributes Seamount Number of seamounts sampled for di

Figure Figure been sampled for geology, bi Material Supplementary

Location of seamounts in the North East Atlantic and Mediterranean Sea that have been in the in the

and and Figure Figure

3 4 5 6 2 1 in the Mediterranean Sea (grey). Fig. 9. sampled for geology, biology orShaded oceanography layer data shows (for the list limits of of seamounts, the see OSPAR area. Supplement 5). Fig. 8.

4 2 3 1