Open Access in Geophysics Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Nonlinear Processes , 2 − , and 1 Open Access Open Access Open Access Open Access Open Access Open Access Open Access Open Access Open Access Open Access Open Access Open Access habitats ). In non- ) and how 6 . cm 40

Open Access = 18 Discussions Discussions Discussions Discussions Discussions Discussions Discussions Discussions Discussions Discussions Discussions Discussions

Climate < Lophelia pertusa, Sciences Sciences Dynamics Chemistry of the Past Solid Earth Techniques , V. Unnithan , SD Geoscientific Methods and and Physics and Atmospheric Atmospheric Atmospheric 1 2 Data Systems Geoscientific Earth System Earth System Measurement − Instrumentation Hydrology and Ocean Science Annales Biogeosciences The Cryosphere Natural Hazards Natural Hazards habitats (41.6 shrimp m and Earth System and Earth arborea Model Development Geophysicae . We conclude that CWC reef 2 , R. Tong 1 − , dead structure, coral rubble) shrimpm 2 < 3366 Open Access Open Access Open Access Open Access Open Access Open Access Open Access Open Access Open Access Open Access Open Access Open 3365 Access Open Access , L. Thomsen

habitats (24.4 shrimp m 2 Primnoa resedaeformis Climate Sciences Sciences Dynamics Chemistry of the Past Solid Earth Techniques Geoscientific Methods and and Physics and ), live Advances in Atmospheric Atmospheric Atmospheric Data Systems Geoscientific Geosciences 5 Earth System Earth System Primnoa resedaeformis Measurement . Instrumentation Hydrology and Ocean Science Biogeosciences , T. Schoening The Cryosphere Natural Hazards Natural Hazards 2 EGU Journal Logos (RGB) EGU Journal Logos and Earth System and Earth and pertusa = 35 Model Development ˚ 2 a et al., 2002; Freiwald et al., 2002; Roberts et al., 2006; Reveillaud , SD 2 − , J. Ontrup 1 ) and live 1 . = 26 In this study we quantified shrimp densities associated with key CWC habitat cate- We found shrimp abundances at the Røst reef to be on average an order of This discussion paper is/has been under review for the journal Biogeosciences (BG). Please refer to the corresponding final paper in BG if available. Jacobs University Bremen, Campus RingBielefeld 1, University, Faculty 28759 of Bremen, Technology, Biodata Germany Mining and Applied Neuroinformatics et al., 2008;(Weaver White et et al., al., 2004; 2012). Turley et Commonly al., identified 2007; as Levin local and biodiversity Sibuet, hotspots 2012) these biogenic habitats shrimp densitieshabitats were clearly supporthabitats greater on shrimp the Norwegian densities margin. than the surrounding non-biogenic 1 Introduction Cold-water coral (CWC) ecosystems have beenlast subjects decade of (Foss much study throughout the magnitude greater inshrimp biogenic densities reef were habitats observed(43 than shrimp in m in association non-biogenic withSD habitats. live Greatest may be surrounded andground, intermixed gravel/pebbles, steep with walls). non-biogenicacross To these habitats date, microhabitats (soft studies have been sediment, offauna more distribution. hard- distribution numerous than of those sessile investigating mobile fauna gories at the Røst reef, Norway,We by analysing also image investigated data shrimp collected distribution bythese patterns towed may video on vary sled. the with local habitat. scale ( Cold-water coral reefs areof highly diverse microhabitats. heterogeneous In ecosystemshabitats a comprising typical (live of European colonies a cold-waterParagorgia coral of range arborea reef locally various biogenic common coral such as Abstract Biogeosciences Discuss., 10, 3365–3396, 2013 www.biogeosciences-discuss.net/10/3365/2013/ doi:10.5194/bgd-10-3365-2013 © Author(s) 2013. CC Attribution 3.0 License. Microhabitat and shrimp abundance within a Norwegian cold-water coral A. Purser 1 2 Group, P.O. Box 100131, 33501 Bielefeld, Germany Received: 6 December 2012 – Accepted: 7Correspondence February to: 2013 A. – Purser Published: ([email protected]) 22 FebruaryPublished 2013 by Copernicus Publications on behalf of the European Geosciences Union. T. W. Nattkemper 5 25 15 20 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | and Pandalus propin- , ¨ offker et al., 2011) Primnoa resedaeformis ¨ uggeberg et al., 2010; Wagner colonies often comprising a num- 3368 3367 ´ opez Correa et al., 2004), with the complexity of ˚ a et al., 2005; Roberts et al., 2009) is commonly the key Pandalus borealis, Pandalus montagui (Hopkins and Nilssen, 1990; Jonsson et al., 2004; Buhl- Primnoa resedaeformis (Herrera et al., 2012; Tong et al., 2012). Growth morphologies of colonies stand more erect, often facing fan-like into the direction of , Fig. 1 (Foss Caridion gordoni and Aside from fish (Costello et al., 2005; Wheeler et al., 2005; S Methodologies for assessing CWC reef biodiversity have changed over the years. In addition to scleractinians, gorgonian are found within many European CWC Cold-water coral reefs are often associated with elevated seabed structural features and Mortensen, 2004b). How shrimp vary in abundance across and between typical shrimp are the mobileractinian fauna (Henry most and commonly Roberts, reported(Krieger 2007; in and Lessard-Pilon association Wing, et with al., 2002;vations CWC 2010) Buhl-Mortensen have scle- tended and and to gorgonian Mortensen, be corals from qualitative 2004a). Norwegian rather reefs Published than include quantitative. obser- Shrimpquus species reported Mortensen and Mortensen,and 2004b). scleractinian The or gorgonian relationship corals isultative between uncertain. commensals these The on hypothesis the shrimp that associated shrimps coral species are species fac- has been raised (Buhl-Mortensen Davis, 2005; Mortensena et al., lesser 2008; numberCWC Orejas of reef et habitats attempting (Jonsson al., et quantificationLessard-Pilon 2008; al., et 2004; of Purser al., Roberts mobile 2010; et et D’Onghia al., macrofauna al., et 2008; al., Le across 2009), 2011). Guilloux with various et al., 2010; sessile species abundances, mobile faunaapproach. are Analysis often of missed video ormarine data, under-sampled collected or by by video this Remote sledmore Operated success Vehicle can (ROV), than gather sub- grab sampling, datathe although on level video of the resolution taxonomical and distribution2012). illumination identification of The can possible mobile limit illumination (Purser macrofauna et requiredhaviour with (Trenkel al., to et 2009; collect al., Schoening video 2004).primarily et on At data al., occurrence CWC may and reef also distribution sites, of influence video sessile faunal species studies (Henry, be- to 2001; date Metaxas have and focused prevalent flow (Mortensen and Buhl-Mortensen, 2005) (Fig. 2). Early studies usingcampaigns dredge employing Van sampling Veen or (Jensen box2007; core and Henry sampling techniques Frederiksen, et (Henry al., 1992) and 2010). Roberts, gave Although way these to techniques are adept at collecting data on Paragorgia arborea sitional rates in variousloops areas above the of reef a structure (White, reef,with 2007; or the Wagner et entrapping coral al., structure suspended 2011). and Sediments materialhabitat the deposited for in surrounding meiofauna coral turbidity (Raes rubble and areas Vanreusel, provide 2006; a Bongiorni further et micro- al.,ecosystems. 2010). Common species at EuropeanParagorgia reefs arborea include these species differ, with ber of branched arms draped across or close to the underlying substrate, whereas the reef morphology providing refuge2002; for mobile Costello organisms et such as al.,et fish 2004; al., (Husebø 2012) et Ross shrimp al., and andRoberts other Quattrini, crustaceans et 2007; (Reed al., et Ballion al., 2008; etnamic 1982; Le Kreiger al., flow and Guilloux 2012; may Wing, et 2002; D’Onghia be al., influenced 2010; by D’Onghia coral et structure, al., 2012). enhancing Local or hydrody- reducing local depo- volumes of refractory material,(Duineveld et or al., pulses 2004; ofal. Kiriakoulakis labile 2007; et Davies material et al., al., to 2005; 2009;et the Thiem Van Oevelen al., et benthic et 2011; al., al., ecosystem Duineveld 2009; 2006; R etsive Kiriakoulakis generations al., et of 2012). coral Sizableet reef al., structures growth 2007). (De can The mol, develop aragoniteat with 2002; skeleton of and succes- Dorschel these above et polyps the al., increasesfor 2005; seafloor habitat sessile complexity Wheeler (Rogers, filter both 1999). feeding organisms Dead (L coral polyps provide hard substrate the calcium carbonate skeletonsLophelia pertusa of scleractinian coralframework species. building species. In European waters, or found in locations where hydrodynamic conditions assist in the delivery of large develop over time with the growth of complex biogenic reef structures, formed from 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 | ˚ a pix- , with m sclerac- Primnoa 3 1080 × ∼ 1920 coverage on dead, Lophelia pertusa Lophelia pertusa and those too blurred for analysis, dis- 2 ¨ offker et al., 2011). It has been hypoth- m 25 . 1 3370 3369 . > cm or 2 also abundant in some regions of the reef complex m %) coverage recorded. Two image frames per second 75 . 25 0 ± ( < 2 m ˚ a and Skjoldal, 2010; S 1 polyp cup is 1 ∼ ), with change in CWC habitat category. Paragorgia arborea cm %. Prior to analysis, the image frames were checked for suitability, with those and 50 20 < Shrimp densities will differ by CWC habitat category. Distribution patterns of shrimp observed( at CWC reefs will differ on the local scale ∼ Lophelia pertusa A Sony HD video camera was used to film the seabed from an altitude of 1. 2. els into the web-based imageical annotation Labelling platform BIIGLE and (BioImageet Exploration) Indexing, al., Graph- (Ontrup 2009; et Bergmann etwas al., al., 2009; 2011). The Schoening seabedcovering et a overlap seabed al., of area successive of 2009; imagecarded. Purser frames Following this sorting stage, aImage set coverage of estimates 3534 image were framesof based remained a on for analysis. the assumption that the average diameter common at CWCtransect coral data reefs has on recentlybenthopelagic been the fauna used Norwegian quantification in (D’Onghia margin a et similar (see al., 2011). ecosystem Sect. in 2.3). thean Mediterranean Video average for area sled of were exported from the video camera and imported with a resolution of (Tong et al., 2012). 2.2 Video data Data was collecteddense from ridge three crests, video lessduring sled populated the flanks traverses and (Fig. ARKXXII/1a sparselyThese 3), “FS populated video cross-cutting Polarstern” intermediate transects the areas cruise were coral (Klages planned and to Thiede, provide data 2011) from of typical 2007. habitat categories tinian coral growth on aduring series the of Cenozoic seafloor (Damuth, crestsgions formed 1978). by of Between the these coral Traenadjupet landslide scleractinian rubbleexposed reef coral and crests framework areas (Wehrmann are et of re- arborea al., low 2009) density with the gorgonian corals the edge of the continental shelf, and is marked by vigorous granted by the Institute of Marine Research (IMR), Norway. The complex is situated on species of shrimp. 2 Methods 2.1 Location of study – theThe Røst Røst reef reef complex, Norway, oneet of the al., most 2002; extensive in Nordgulen Norwegianthe waters et focus (Foss al., of 2006; this Wehrmann study. Permission et to al., conduct 2009; video Tong surveys et of al., the 2013), reef was complex was areas of significant human activity,ration/production primarily (Foss commercial fishing andesised oil and that gas CWC explo- reefjuvenile ecosystems fish are species highly important (Husebøcould refuges et also for play al., a some 2002; role commercial Ballion in the et lifecycle al., of 2012) other mobile and commercial that fauna, these such as reefs some By testing these twofunctioning hypotheses, of the CWC aimble reef was use biodiversity to for hotspots, increaseis the our and particularly development understanding to prescient of given of provide that management the many information plans CWC of reefs for in possi- these European waters areas. are This located in latter aim ture, dead coral framework etc) haswe not investigate to these date patterns been in investigatedRøst distribution in across detail. reef reef In on habitat this study categoriescamera the found was Norwegian at analysed, the with margin. the Video following two data hypotheses collected tested: by ship towed video-sled CWC coral microhabitats (live scleractinian coral structure, live gorgonian coral struc- 5 5 25 15 20 10 20 15 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | (a), have a far Paragorgia (approx. 5– , therefore the 2 mm m , Live 5 1 ∼ ∼ Lophelia pertusa (c), dead structure (d), coral (Brown and Forsythe, 1974) used Lophelia pertusa Primnoa resedaeformis F % coverage of an image frame). Ad- 50 and > at the Røst reef (Tong et al., 2012, 2013). 3372 3371 habitat categories), dead coral (Dead structure measure) (Field, 2009). 2 Primnoa resedaeformis ω Lophelia pertusa Paragorgia arborea (b), live . Given the complex topography of many of the habitats investigated, Primnoa resedaeformis 2 − , Live Paragorgia arborea We focused our study on shrimp abundance on determining whether or not the abun- The high heterogeneity of habitat categories within CWC reefs on the Norwegian The ten habitat categories used in this study (see Sect. 2.3) are not of uniform spatial Due to the high reflectivity of the eyes of shrimp, spotting them on the image frames tat type as thebiogenic investigated habitat). factor Levene’s test (4 oftributed levels, and homogeneity live therefore showed the coral, the robust dead Brown-Forsythe’s for data coral, the to coral analysis. be A unequally rubble,the Bonferroni dis- non- post-hoc factor test levels was (habitatFor used categories) the to ANOVA shrimp determine and distribution betweenfects post-hoc size which differences tests was of were also the determined significant. statistical ( package SPSS 17.0 was used. Ef- dances observed in association witharborea live coral (Live and Dead gorgonian categories), coralWall, rubble soft or sediment non-biogenic and habitats hardground (Gravel/pebble, a categories) one-way differed ANOVA statistically. test To do with this shrimp we abundance ran as the dependant variable and habi- coverage within a particular reef,gorgonian and can coral vary species greatlyless in coverage spatial between coverage reefs. than The This variation in coveragelematic. can To address make this problem statistical weobserved analysis decided in to association of group with the video together various thecomparable transect habitats shrimp spatial into data abundances broader areas categories of prob- covering reef more for statistical analysis. margin has been2010; widely Tong et reported al., 2012), (Buhl-Mortensenscales with of et great cm changes or al., in m. In habitat 2010;categories this complexity are study Gheerardyn often associated we occurring with et aim over to different al., show shrimp statistically abundances. whether or not these habitat other features such as the small overhangs of larger pebbles or coral blocks. 2.4 Statistics 2.4.1 Habitat category and shrimp density The video frames used innumber this of analysis covered shrimps a logged seabedof area in shrimp of each m imagefurther was shrimp assumed individuals to may be have the been minimum obscured number from view by coral structure, or 10 pixels). The mid-pointfor between every the shrimp two in eyes everyof was image, each individual with marked within these into each mid-pointsto the image representing differentiate BIIGLE frame. the particular In system point the shrimp present location the species. study dataset Fig. no within 5 attempt the shows was BIIGLE made an system, with example a image number frame of from shrimp positions logged. live rubble (e), gravel/pebbles (f),ground wall (j). These (g), habitat dead categoriesfor gorgonian were the selected (h), Røst as soft they reef were (Wehrmann seabed all et (i) previously al., reported and 2009; Tong hard- etwas al., not 2012) a significant problemthe (Fig. majority 5). of As shrimps the video observedindividual sled were were filmed observed distinct the from bright seabed above. points from The separated above, eyes of from each each shrimp other by Each image frame wasmain inspected habitat category in was turn loggedditionally, in (that presence/absence the with of BIIGLE eachimage system. of was For logged. the each Ten habitat remaining image categories habitat were the categories used (Fig. within 4): each live 2.3 Habitat characterisation and shrimp labelling 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- cm 2 ω ), coral = 1678 8 ) than in . n 4 . = 7 = 19 , SD habitat categories 7 . 4+ 25, SD = 8 = , mean , mean ), so indicating whether there was = 779 cm n = 378 n 40 ∼ 3374 3373 (Brown-Forsythe robust equality of means test ) or non-biogenic habitat categories ( 5 . 26 . scale) habitat categories had an influence on shrimp = 2 2 = 0 ). The Bonferroni tests also showed that differences in 2 m ω 1 01 , SD . , 0 < 45 . 001 . p < 0 = 1 )( 8 . p < , 8 = 1 . , mean , SD 6 = 699 . n = 0 3530) = 503 , (3 biogenic habitat. in images containing predominantly liveimages coral containing ( mainly deadrubble coral ( structure ( mean shrimp abundances were alsoages containing significant predominantly at dead the coralrubble. structure same No and significant confidence those difference level containingimages was mainly between containing indicated coral predominantly im- coral between rubble shrimp habitat and densities those observed containing in mainly non- There was a significant effectF of habitat category on numbersemployed of as shrimp Levene’s in test each image, indicateddicates that the the data habitat to typeBonferroni be has post-hoc in-homogenous). a tests The large indicated high effect that on shrimp shrimp abundances abundance at were the significantly Røst higher reef. deviation of shrimp densitiesTable 1. observed The median in shrimp associationgiven densities and with in quartile Fig. each distribution 6. of ofas Of these these the the observations predominant is ten is category habitat given inof image categories in the frames. outlined “dead Although gorgonian in some habitat” images Sect.always contained and 2.3, less areas “soft than eight sediment” 50 categories, were % coverage present of by an these image was frame. 3.1 Shrimp densities and habitat category A total of 18 269main shrimp habitat were category manually present labelled in withinin each the image each 3534 was images identified image analysed. andprimarily The frame the number each logged. of of shrimps The the number habitat of categories, image along frames with identified the as mean containing average and standard 3 Results In addition to logging thethe main number of habitat additional category habitat visibleshrimp categories in present densities each in present an analysed in image imagewas was image frame, compared also frames counted. with The containing a 1,number Kruskal-Wallis of test, 2, locally to 3 available ( determine or abundance. whether or not increase in the individual to maintain a certain distancetial from randomness other was shrimps. tested, In this withpixels study Ripley’s-L (half complete values spa- computed the up smallest toany image a trends frame in maximum distribution axis, of (shrimp 540 clustering or spacing) over2.4.3 distances of up Habitat to complexity 40 and shrimp abundance scales. Image frames containingas the less method than requiresConfidence 20 envelopes a Shrimps (95 %) minimum were were number computedviations omitted of by from from means points the of this (shrimps) Monte-Carlo confidencethis sampling. test, envelope for De- instance) by a allows points valid rejection withina outcome. an of particular spatial image the scale (shrimp assumption (thedence locations spatial that envelope) scale in at the at the which point confidence distribution levelnot pattern deviates determined. these from The is the deviations test random confi- represent also indicates on a whether tight or clustering of shrimps or a tendency for a shrimp By analysing the distances betweenimage each frame (i.e. shrimp all and the allpendent other its shrimps on neighbours in distance in an was a image)provides particular determined a a positive/negative by association statistical applying de- measure Ripleys-L for (Ripley, 1976). the Ripleys-L distribution of point items over different spatial 2.4.2 Local scale shrimp distribution 5 5 25 15 20 10 15 20 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | , ) 2 4 − . cm 10 ) were = 19 2 habitat, , Fig. 7. < − 01 . (SD 0 shrimps m 2 − 20 p < , > however, shrimps Primnoa resedae- Paragorgia arborea 54 . cm individuals m ). It is unlikely that this , live 10 8 . 20 Lophelia pertusa > > = 7 (3) = 1052 habitat categories were not sig- H , SD 7 . 4+ = 8 % lower than observed in association 2 − 60 Paragorgia arborea . Table 2 shows that on a scale of ∼ (80 %) habitat image frames also containing 2 3376 3375 − , live . Just under 10 % of dead scleractinian structure 2 − ) and non-biogenic habitat categories (mean shrimps shrimps m 5 . 20 = 2 > shrimps m habitat category had a significantly greater number of shrimps 20 1 Lophelia pertusa (Fig. 6). The statistically significant observation of higher shrimp ). Coral rubble regions of CWC ecosystems are commonly made , SD > > 8 2 . were observed, with higher percentages of live 45 Primnoa resedaeformis − . 2 − = 1 = 1 2 − , SD 6 . and the dead scleractinian habitat categories. Table 2 shows that within just over shrimps m = 0 2 20 Sufficient shrimp densities to allow Ripley’s L analysis ( − observed for shrimp in sub-littoral environments (Hewitt et al., 2005). may occupy, it is difficultthe to imagine great that difference hiding in shrimpin utilizing density association these observed could with account whenThe this for comparing non-biogenic habitat the habitat categories, and abundances thosei.e. those observed not hardground dependant observed outcrops, on in gravel/pebble CWC fieldsabundances reef live of and development, and shrimps walls, observed were dead – regionsseveral though coral shrimp with some habitats. the areas m of lowest walldensities habitat in had images densities of containingthat more increased than local one heterogeneity habitat may category be is a of further benefit indication to CWC shrimps, as has been Shrimp abundances were lower again inshrimps association with m the coral rubble habitatm (mean up of small, brokenThough pieces this of habitat coral, contains weathered a and eroded number and of lying small on structural the gaps seabed. and features shrimp predominantly live coral, with arecorded in mean such average image density frames. ofhabitat Of 25 the is reef shrimps perhaps habitats m surveyed thecoral in most this structure, study, physically though the live complex still reef was (Buhl-Mortensen very associated et angular with and al., shrimpwith providing 2010). abundances the many Dead live physical coral habitatdifference habitats niches, (mean can shrimps be m were explained actually by easier observer to distinguish bias against or dead coral shrimp structure behaviour, than against as live shrimp coral. eyes From the results presented heregian it CWC is reef habitat clear with thathave change shrimp a in densities habitat large vary category, and influence acrossis that a on clear the Norwe- type shrimp that of shrimp density. habitat From abundances can the are highest images in analysed regions in where this the study available it habitat is 4 Discussion Densities of shrimps innificantly images different containing from 2, each 3 other. or shrimps were more negatively associatedage with frames their neighbours than in in livewere dead coral observed coral habitat to im- habitat be frames. clusteredpresent. At in association scales with of all habitats where 3.3 Habitat complexity and shrimp abundance Frames containing present than those with only 1 habitat type present, 50 % of the> images classified as(62.5 being %) and predominantly live live shrimp densities of image frames contained shrimps tended to be moreshrimps closely in spaced these in non-biogenic habitatsto habitats, probably use the indicates the low that density same of habitatthan the clumping features few together (such shrimp intentionally. as present pebble are overhang or tending fissure inobserved a in wall) 267 rather imageassociation frames with (Table 2). live These highformis densities were only observed in The minimum, maximum,across each mean habitat category and isbour given median in scores Table distance 1. indicate The mean betweenliving a distances to individual coral generally nearest habitats tighter neigh- shrimps than arrangement of with shrimps dead coral in structure association or with coral rubble habitats. Though 3.2 Patterns of shrimp distribution 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 | ) and scale colonies Primnoa Primnoa Lophelia cm and dead living coral 10 and and < also exude mucus Aristeus antennatus Lophelia pertusa within and surrounding may account for the dif- Lophelia pertusa free from accumulating sed- Lophelia pertusa Paragorgia arborea – a possibility unsupported by Paragorgia arborea of cm above the seafloor, whereas Paragorgia arborea s Lophelia pertusa 3378 3377 Lophelia pertusa Aristaeomorpha foliacea Primnoa resedaeformis within the living reef structural area. Primnoa resedaeformis and is not clear from this study. It is possible that they play colonies may acquire their food supply from more turbid wa- rather than ) found in this current study within live coral habitats may indicate cm likely utilise different food sources, with were similar, though the number of frames containing either of these Lophelia pertusa Paragorgia arborea Parapenaeus longirostris and dead scleractinian structure habitats reported here, the feeding hypothesis The differences in colony morphology and polyp spacing of Shrimp densities observed in association with The observation that shrimp at the Røst reef preferentially occupy regions of living trapped by the corals as a food source, then some minor difference in the food value the data here. those of ferences in shrimpcorals. abundances Possibly observed the in moreshrimps frames to flexible containing rest gorgonian predominantly on, coralsdition living or allow to a the a more gorgonians, greateras suitable scleractinians set surface a such of area cleaning as refuges agent for Wild for and et evasion potentially al., of 2008; as predators. Purser In a ad- et food al., gathering 2010). mechanism Should (Reitner, CWC 2005; shrimp be utilising material en- ters found in closerobserved on proximity both to gorgonian corals the arewithin seafloor. the the result Possibly mucus of the secreted shrimp by elevated feedingter the on shrimp column, corals material (Patton, densities trapped as 1972), such ratherParagorgia arborea than a material strategy in would the wa- likely have led shrimp to preferentially occupy tat (the gorgonian coral)favourable as position an with elevating respect substrate toand on the vertical prevalent which extent flow to reached conditions, by rest as coloniesMortensen the and of and these growth attain Buhl-Mortensen two morphs a gorgonian (2005) species more indicated isresedaeformis that very different. tending to stand nearered vertically, facing by the into stronger, the less prevalentPrimnoa turbid current, currents resedaeformis capturing found food 10 deliv- species resedaeformis coral species as the predominantble habitat densities would category not were be few expected if (Table 1). shrimps were Such simply compara- utilising the underlying habi- from within the reef itself, and a 10x greater abundance of the commercially significant CWC habitat, the complete absence of some shrimp species (e.g. a useful role in keepingiment, the a branches of process whichand Purser, can 2011). lead No attempt tovideo was tissue made data, here damage and to over potentially differentiatecoral different time shrimp habitats. species species if Such from could local unchecked the be spatialhas (Larsson utilising partitioning been the of reported live the coral reeftrawl at survey and ecosystem tropical comparing by dead species coral shrimp abundances species dicated reefs within that and (Hoeksema this outside may and of also CWCrable be Fransen, reef likely densities 2011). zones at of in- CWC A the reefs. recent shrimp D’Onghia et species al. (2010) report compa- Given the differences inpertusa shrimp densities observed inseems association the with more likely, living with thecoral physical structure structure of habitats the not live to being the hugely living different. Whether the shrimp are of benefit that some shrimp species mayserved play in a tropical role reefs as (Zann,coral cleaners 1987). associated at At shrimp CWC tropical reefs, individuals reefs, ascus have studies has and indicated of been zooplankton a stomach (Patton, ob- diet 1994). contents(i.e. consisting The of a of regular tendency shrimp both to spacing coral maintainof on mu- some less the distance than from 10 that neighbouring shrimps individual over shrimps distances requirethat a the minimum regularity of surrounding spacing space observed is for a feeding, function or of the perhaps underlying coral morphology. coral has previously beenfilter reported feeders as (Buhl-Mortensen a etthe likely al., Mediterranean behavioural 2010), trend (D’Onghia and in et reportedfollowing al., mobile for direct 2011). CWC some sampling Elevated reef shrimp of shrimpstructure species associate concentrations from in fauna observed the from Gulf within of Mexico (Cordes et al., 2008) led the authors to hypothesise 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 | , Primnoa , dead coral Paragorgia arborea Primnoa resedaeformis and than in frames where the less Primnoa resedaeformis , live ), so without further study the conclusion that 3380 3379 ¨ offker et al., 2011). From the shrimp densities coral was in dominance. The spaced, branched spp. colonies – although not a strictly quantita- Paragorgia arborea –8) analysed in this study was far less than frames = 365 structure, perhaps limiting this habitat in usefulness n ( = 5 Primnoa resedaeformis n in western Atlantic waters (Metaxas and Davis, 2005; in the Gulf of Alaska (Kreiger and Wing, 2002). Although Primnoa Paragorgia arborea cm 3 < , live The research leading to these results has received funding from the Eu- Paragorgia arborea in association with Lophelia pertusa 1 ARKXXII/1a cruise are thanked for their assistance, particularly B. Lessmann structure may be the most appealing coral habitat for smaller shrimp Lophelia pertusa − Polarstern Primnoa resedaeformis Lophelia pertusa Despite the absence of species level distribution data and some variation in seabed Should protection be the overriding advantage offered by living coral habitats over Previous studies have investigated megafauna associated with of FS for assistance with preparation of the video data. reported here it wouldresult seem in likely that a the dropavailability loss for in of many local fish CWC species. reef shrimp ecosystems numbers, would andAcknowledgement. likely therefore result inropean Community’s a Seventh Framework reduction Programme (FP7/2007-2013)project, in under grant the prey HERMIONE agreement no.a 226354) CORAMM group with collaboration. further We acknowledgepermission funding the to Institute received investigate of the from Marine Røst Research, Statoil. reef Norway complex. This for The work captain, crew is and onboard scientific party in association with thegian non-biogenic margin habitats seafloor of (gravel/pebbles, wall, thestudy, hardground). it investigated Though would region not seem of likely addressed the thaterwise in the Norwe- would this loss have of a CWC knock-onwith reefs effect the via on loss anthropogenic local activity of fish or structural stocks oth- fish in habitat addition (S to that associated coverage of the varioussented habitat here that categories shrimp investigated, densities can itreefs vary with is significantly across changes clear Norwegian in margin from habitat CWC tribution the category, also and data change that with pre- the changefor local in shrimp scale habitat. is patterns The also significance in very(live of shrimp clear, biogenic dis- with reef shrimp habitats densitiesstructure, coral observed rubble) in to association be with in these excess of an order of magnitude greater than observed to larger shrimp individualsther and of species. the gorgonian The species numbercontaining ( of live image frameshigher containing ei- densities ofproached shrimp tentatively. are associated with the gorgonian corals should be ap- densely packed morphologies of the gorgonians may offer more useful refuge niches than the more the surrounding dead reefresedaeformis areas, then the complexityspecies and and malleability individuals. of It the hasas a been refuge hypothesised by that shrimp thisfrom gorgonian the observations species reported is used intically the significant current differences study in it shrimpgonian is species, abundances not it between possible is the to quite twodegree observe possible investigated in statis- the gor- frames video dominated data by spatially under complex sampled shrimps to a greater from within the branchestive of estimate of shrimpassociated density, their with rough the figure living isAtlantic. gorgonians comparable in with this shrimp densities current study, on the eastern side of the ported on the difficulties involvedgorgonian in species, quantifying by shrimp either densities suction ining the sampling those vicinity fauna collected of from whilst these coralBuhl-Mortensen sampling branches and whole or by coral Mortensen count- branches.viduals (2005) From colony report video taken shrimprespectively. by These densities ROV, density of estimates 4.8 arestudy, far and and lower made 3.1 than from indi- thoseGulf reported video of Alaska, in data Krieger the recorded and current by Wing (2002) a reported more “hundreds intrusive of shrimp methodology. eyes” In reflected the could also explain the minor(Fig. differences 6). in shrimp densities observed by coral species and Buhl-Mortensen and Mortensen, 2005). Buhl-Mortensen and Mortensen (2005) re- of the mucus itself or the entrapped material found on scleractinians and gorgonians 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 Lophelia pertusa Lophelia pertusa reefs in Norway: experi- (Gunnerus, 1763), J. Nat. Lophelia ¨ alv, T., Jonsson, L., Bett, B. J., van Weer- Primnoa resedaeformis 3382 3381 () at the Mingulay Reef complex, Limnol. , 2012. (L., 1758) and ¨ uggeberg, A., Dullo, W. C., and Freiwald, A.: Growth and erosion Lophelia pertusa Paragorgia arborea 10.1371/journal.pone.0044509 ˚ ˚ ˚ a, J. H., Lindberg, B., and Christensen, O.: Mapping of a, J. H. and Skjoldal, H. R.: Conservation of cold-water coral reefs in Norway, in: Handbook a, J. H., Mortensen, P. B., and Furevik, D. M.: The deep-water coral Norwegian waters: distribution and fishery impacts, Hydrobiologia, 471, 1–12,ences 2002. and survey methods,and in: Roberts, Cold-Water J. Corals M., and Springer, Ecosystems, Berlin, edited 359–391, by: 2005. Freiwald, A. of Marine Fisheries ConservationSquires, and D., Tait, Management, M., edited and Williams, by: M., Grafton, Oxford R. University Q., Press, US, Hilborn, 215–240, R., 2010. Watmough, T., and Witbaard,water R.: coral Spatial reefs of and the tidalSer., Mingulay variation 444, Reef Complex 97–115, in (Outer 2012. food Hebrides, Scotland), supply Mar. to Ecol. Prog. shallow cold- a cold-water coral277, 13–23, community 2004. (Galicia Bank, NW Spain), Mar. Ecol. Prog. Ser., of a cold-waterPleistocene coral and covered Holocene, carbonate Earth Planet. mound Sci. in Lett., 233, the 33–44, Northeast 2005. Atlantic during the Late mentation, Mar. Geol., 28, 1–36, 1978. Roberts, J. M.: Downwellingto and the deep-water cold-water bottomOceanogr., currents 54, 620–629, as 2009. food supply mechanisms Maiorano, P.: Distribution andtowed behaviour cameras of in the deep-sea Santa benthopelagic443, Maria 95–110, fauna di 2011. Leuca observed cold-water using coral province, Mar. Ecol. Prog.paring Ser., deep-sea fishMaria di fauna Leuca betweendoi: cold-water coral coral and province non-coral (Mediterranean “megahabitats” Sea), in PLoS ONE, the 7, Santa e44509, ing, T. C. E., decoral Hass, reefs H., as Roberts, fish J.by: habitat M., Freiwald, in and A. Allen, the and D.: NE Roberts, Role Atlantic, J. of in: M., cold-water Springer, Cold-Water Berlin, Corals 771–805, and 2005. Ecosystems, edited of deep-water coral banksMediterranean Sea, on Deep-Sea the Res. Pt. abundance I, and 57, 397–411, size 2010. structure of the megafauna in the 50, 2010. and Fisher, C. R.:777–787, Coral 2008. communities of the deep Gulf of Mexico, Deep-Sea Res. Pt. I, 55, Huvenne, V., Ivanov, M., Swennen,the R., Porcupine Basin, and southwest Henriet, of J. Ireland, P.: Mar. Large Geol., deep-water 188, coral 193–231, 2002. banks in gonian Hist., 38, 1233–1247, 2004a. 61, 2004b. B., Gheerardyn, H., King,heterogeneity N. and J., biodiversity and on the Raes, deep M.: ocean Biological margins, structures Mar. Ecol.-Evol. as Persp., a 31, source 21– of habitat scleractinian corals promote higher biodiversitycontinental in margins, Biol. deep-sea Conserv., meiofaunal 143, assemblages 1687–1700, along 2010. equality of several means, Technometrics, 16, 129–132, 1974. semblages from two shelf stations west of Svalbard, Mar. Biol. Res., 7, 525–539, 2011. for fish larvae, Front. Ecol. Environ., 10, 351–356, 2012. Foss Foss Foss Field, A.: Discovering Statistics Using SPSS, SAGE Publications, London, UK, 822 pp. 2009. Duineveld, G. C. A., Jeffreys, R. M., Lavaleye, M. S. S., Davies, A. J., Bergman, M. J. N., Duineveld, G. C. A., Lavaleye, M. S. S., and Berghuis, E. M.: Particle flux and food supply to Dorschel, B., Hebbeln, D., R D’Onghia, G., Maiorano, P., Carlucci, R., Capezzuto, F., Carluccio, A., and Sion, L.: Com- Davies, A. J., Duineveld, G. C. A., Lavaleye, M. S. S., Bergman, M. J. N., van Haren, H., and D’Onghia, G, Indennidate, A, Giove, A., Savini, A, Capezzuto, F., Sion, L., Vertino, A., and Damuth, J. E.: Echo character of Norwegian-Greenland Sea: relationship to quaternary sedi- Costello, M., McCrea, M., Freiwald, A., Lund D’Onghia, G., Mairorano, P.,Sion, L., Giove, F., Capezzuto, F., Carlucci, R., and Tursi, A.: Effects Cordes, E. E., McGinley, M. P., Podowski, E. L., Becker, E. L., Lessard-Pilon, S., Viada, S. T., De Mol, B., Van Rensbergen, P., Pillen, S., Van Herreweghe, K., Van Rooij, D., McDonnell, A., Buhl-Mortensen, L., Vanrusel, A., Gooday, A. J., Levin, L. A., Priede, I. G., Mortensen, P.- Buhl-Mortensen, L. and Mortensen, P. B.: Crustaceans associated with theBuhl-Mortensen, deep-water L. gor- and Mortensen, P. B.: Symbiosis in deep-water corals, Symbiosis, 37, 33– Brown, M. B. and Forsythe, A. B.: The small sample behaviour of some statistics which test the Bongiorni, L., Mea, M., Gambi, C., Pusceddu, A., Taviani, M., and Danovaro, R.: Deep-water Bergmann, M., Langwald, N., Klages, M., Ontrup, J., and Nattkemper, T. W.: Megafaunal as- Ballion, S., Hamel, J.-F., Wareham, V. E., and Mercier, A.: Deep cold-water corals as nurseries References 5 5 30 25 20 15 30 10 25 20 15 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | and spp.) Acesta Pandalus : cleaning Primnoa on the Swedish Lophelia pertusa Gastroptychus formosus at Semporna, eastern Sabah, , Mol. Ecol., TS221, 6053–6067, Lophelia pertusa ¨ alv, T.: Distributional patterns of macro- and , Mar. Biol., 147, 775–788, 2005. colonies in the Gulf of Mexico, Deep-Sea Res. 3384 3383 Paragorgia arborea Heliofungia actiniformis ¨ offker, M. K., and Olu, K.: ˚ a, J. H., Furevik, D. M., and Jørgensen, S. B.: Distribution and Lophelia pertusa Paragorgia arborea and , 2012. (Scleractinaria) on the Faroe shelf, Sarsia, 77, 53–69, 1992. ¨ uhnerbach, V., Lindberg, B., Wilson, J. B., and Campbell, J.: The Sula Reef (Bivalvia: Limidae) with new data on its shell ultrastructure, in: Cold-Water Corals ) in Balsfjord, northern Norway: I. Abundance, mortality, and growth, 1979–1983, J. 10.1111/mec.12074 corals in Northeast Channel,1390, off 2005. Nova Scotia, Canada, J. Mar. Biol.Primnoa Assoc. resedaeformis UK, 85, 1381– Annu. Rev. Marine. Sci., 4, 79–112, 2012. excavata and Ecosystems, edited by: Freiwald, A. and Roberts, J. M., Springer, Berlin, 173–205, 2004. hydrocarbon exploration drilling site in119–129, the 2008. Træna Deep, Norwegian Sea, Environ. Geol., 56, composition associated with Pt. II, 57, 1882–1890, 2010. Lophelia pertusa megafauna associated with awest reef coast, of Mar. the Ecol. cold-water Prog. Ser., coral 284, 163–171, 2004. efficiency from natural sediments and drill cuttings, Mar. Poll. Bull.,cold-water corals 62, in 1159–1168, the 2011. North Atlantic, J. Mar. Biol. Assoc. UK, 90, 1363–1369, 2010. borealis Cons. Int. Explor. Mer., 47, 148–166, 1990. abundance of fish in deep-sea coral habitats, Hydrobiologica, 471, 91–99, 2002. in 2007 (ARK-XXII/1a-c) BerichteMarine Research), zur Alfred Polar- Wegener und2011. Institute Meeresforschung for (Reports Polar and on Marine Polar Research, and Bremerhaven, in the Gulf of Alaska, Hydrobiologica, 471, 83–90, 2002. cohabitating in the mushroomCoral coral Reefs, 30, 519, 2011. deep-water coral/mound systems of the96, NW 159–170, European 2007. Continental Margin, Int. J. Earth Sci., NE Atlantic, Deep-Sea Res. Pt. 1, 54, 654–672, 2007. ties off western Scotland, Coral Reefs, 29, 427–436, 2010. tion in the widespreaddoi: deep-sea coral structure for maintaining beta diversity, Ecology, 86, 1619–1626, 2005. and nitrogen isotopes of two deep-waterimplications corals from for the their North-East nutrition, Atlantic: in: initialand Cold-Water results Corals Roberts, and and J. Ecosystems, M., edited Springer, by: Berlin, Freiwald, 715–729, A. 2005. Biol. Assoc. UK, 81, 163–164, 2001. cold-water coral mounds and adjacent off-mound habitat in the bathyal Porcupine Seabight, 2001. ture of harpacticoid copepodsSeabight associated (North-East with Atlantic), cold-water Helgoland coral Mar. Res., substrates 64, in 53–62, the 2010. Porcupine complex, Norwegian Shelf, Facies, 47, 179–200, 2002. ´ opez Correa, M., Freiwald, A., Hall-Spencer, J., Taviani, M.: Distribution and habitats of Mortensen, P.B. and Buhl-Mortensen, L.: Morphology and growth of the deep-water gorgonians Metaxas, A. and Davis, J.: Megafauna associated with assemblages of deep-water gorgonian L Levin, L. A., and Sibuet, M.: Understanding continental margin biodiversity: a new imperative, Lessard-Pilon, S. A., Podowski, E. L., Cordes, E. E., and Fisher, C. R.: Megafauna community Lepland, A. and Mortensen, P.: Barite and barium in sediments and coral skeletons around the Jensen, A. and Frederiksen, R.: The fauna associatedJonsson, with L. the G., bank-forming Nilsson, P. deepwater G., coral Floruta, F., and Lund Larsson, A. I. and Purser, A.: Sedimentation on the cold-waterLe coral Guilloux, E., Hall-Spencer, J. M., S Husebø, A., Nøttestad, L., Foss Krieger, K. J. and Wing, B. L.: Megafauna associations with deepwater corals ( Hopkins, C. C. E. and Nilssen, E. M.: Population biology of the deep-water prawn ( Klages, M. and Thiede, J.: The expedition of the research vessel “Polarstern” to the Arctic Hoeksema, B. W. and Fransen, C. H. J. M.: Space partitioning by symbiotic shrimp species Kiriakoulakis, K., Freiwald, A., Fisher, E., and Wolff, G.: Organic matter quality and supply to Hewitt, J. E., Thrush, S. F., Halliday, J., and Duffy, C.: The importance of small-scale habitat Henry, L. A., Davies, A. J., and Roberts, J. M.:Herrera, Beta S., diversity Shank, of T. cold-water M., coral and reef communi- Sanchez, J. A.: Spatial and temporal patterns of genetic varia- Kiriakoulakis, K., Fisher, E., Wolff, G. A., Freiwald, A., Grehan, A., and Roberts, J. M.: Lipids Henry, L. A. and Roberts, J. M.: Biodiversity and ecological composition of macrobenthos on Henry, L. A.: Hydroids associated with deep-sea corals in the boreal north-west Atlantic, J. Mar. Gordon, J. D. M.: Deep-water fisheries at the Atlantic Frontier, Cont. Shelf. Res., 21, 987–1003, Gheerardyn, H., De Troch, M., Vincx, M., and Vanreusel, A.: Diversity and community struc- Freiwald, A., H 5 5 30 25 20 30 15 25 10 20 15 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Oculina – a first Lophelia Leptogorgia virgulata Madrepora oculata , 2012. (LINNAEUS 1758) and other deep-water reef- 3386 3385 (Scleractinia) zooplankton capture rates, J. Exp. Mar. , 2012. ˚ ¨ a, J. H., and Berntsen, J.: Food supply mechanisms for cold- alv, T., Ontrup, J., and Nattkemper, T.: Use of machine-learning Lophelia pertusa 10.1371/journal.pone.0043534 Lophelia pertusa ¨ ogel, S., Dullo, W. C., Hissmann, K., and Freiwald, A.: Water mass charac- coral reefs, B. Mar. Sci., 32, 761–786, 1982. 10.1371/journal.pone.0038179 pertusa using multiscale terrain variables,tween in: Disciplines, edited Earth by: System Lohmann Science:J., G., and Bridging Grosfeld Unnithan the K., V., Wolf-Gladrow Gaps Springer, D., Be- Berlin, Wegner A., 113–118, Notholt 2013. water corals along a continental shelf edge, J. Marineof Syst., deep-water 60, 207–219, gorgonian 2006. coralsPLoS in ONE, relation 7, to e43534, seabed doi: topography on the Norwegian margin, Nattkemper, T. W.:densities Semi-automated at image the analysisdoi: for Arctic the deep-sea assessment observatory of HAUSGARTEN,coral megafaunal reefs PLoS off Ireland, ONE, Deep-Sea Res. 7, Pt. I, 58, e38179, 818–825, 2011. teristics and sill dynamics inNorway, Mar. a Geol., subpolar 282, cold-water 5–12, coral 2010. reef setting at Stjernsund,proach northern for Visual Bioimagealization, Database Barcelona, Mining, Spain, IV 2009. 09 – Intl. Conference on Information Visu- 255–266, 1976. reusel, A., Olu-Le Roy,Bay K., of and Biscay, NE Henriet, Atlantic, J. Facies, 54, P.: The 317–331, 2008. distribution of scleractinian corals in the forming corals and impacts from human activities, Int. Rev. Hydrobiol.,southeastern 84, United 315–406, States, 1999. Deep-Sea Res. Pt. I, 54, 975–1007, 2007. areal and trophic relationships of decapodsvaricosa associated with shallow- and deep-water geobiological approach, in: Cold-Water CoralsRoberts, and J. Ecosystems, M., edited Springer, Berlin, by: 731–744, Freiwald, A. 2005. and Geology of Deep-Sea Coral Habitats, Cambridge, UK, 2009. munities associated withSeabight sediment-clogged (NE Atlantic), cold-water Deep-Sea Res. coral Pt. framework II, 53, in 1880–1894, 2006. the Porcupine megafuanal communities andNorth evidence East for Atlantic, coral Facies, 54, carbonate 297–316, 2008. mounds on the Hatton Bank, algorithms for the automatedProg. detection Ser., of 397, cold-water 241–251, coral 2009. habitats: a pilot study. Mar.food concentration Ecol. on Biol. Ecol., 395, 55–62, 2010. cold-water coral ecosystems, Science, 312, 543–547, 2006. SPP) from the , Australia, B. Mar. Sci., 55, 193–211, 1994. ploring of images from theIEEE, Arctic Bremen, deep-sea 2009. observatory HAUSGARTEN, Proc. OCEANS’09, (Lamark), B. Mar. Sci., 22, 419–431, 1972. a potential World Heritage Site based on natural criteria, Geological Survey of Norway, 2006. gonian corals in the NortheastFishing, Channel, edited by: Nova Barnes, Scotia, P.W. in: and41, Benthic Thomas, 369–382, Habitats J. 2005. P.,American and the Fisheries Society Effects Symposium, of ¨ ¨ uggeberg, A., Fl offker, M., Sloman, K. A., and Hall-Spencer, J. M.: In situ observations of fish associated with Tong, R., Purser, A., and Unnithan, V.: predicting habitat suitability for cold-water coral Tong, R., Purser, A., Unnithan, V., and Guinan, J.: Multivariate statistical analysis of distribution Thiem, Ø., Ravagnan, E., Foss Schoening, T., Bergmann, M., Ontrup, J., Taylor, J., Dannheim,S J., Gutt, J., Purser, A., R Schoening, T., Ehnert, N., Ontrup, J., and Nattkemper, T. W.: BIIGLE Tools – A Web 2.0 Ap- Ross, S. W. and Quattrini, A. M.: The fish fauna associated with deep coral banks off the Ripley, B. D.: The second-order analysis ofReveillaud, stationary point J., processes, Freiwald, J. A., Appl. Van Probab., 13, Rooij, D., Le Guilloux, E., Altuna, A., Fourbert, A., Van- Rogers, A. D.: The biology of Reitner, J.: Calcifying extracellular mucus substances (EMS) of Reed, J. K., Gore, R. H., Scotto, L. E., and Wilsom, K. A.: Community composition, structure, Roberts, J. M., Wheeler, A., Freiwald, A., and Cairns, S.: Cold-Water Corals. The Biology and Raes, M. and Vanreusel, A.: Microhabitat type determines the composition of nematode com- Roberts, J. M., Henry, L. A., Long, D., and Hartley, J. P.: Cold-water frameworks, Purser, A., Larsson, A. I., Thomsen, L., and Van Oevelen, D.: The influence of flow velocity and Roberts, J. M., Wheeler, A. J., and Freiwald, A.: Reefs of the deep: the biology and geology of Purser, A., Bergmann, M., Lund Patton, W. K.: Distribution and ecology of associated with branching corals ( Patton, W. K.: Studies on the symbionts of the gorgonian coral, Ontrup, J., Ritter, H., Scholz, S. W., and Wagner, R.: BIIGLE – Web 2.0 enabled labeling and ex- Nordgulen, O., Bargel, T. H., Longva, O., and Ottesen, D.: A preliminary study of Lofoten as Mortensen, P. B., Buhl-Mortensen, L., and Gordon, D. C.: Effects of fisheries on deepwater gor- 5 5 30 25 20 15 10 30 25 20 15 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | , 2009. (mm) (mm) ¨ alv, T.: Particulate organic matter ´ 10.5194/bg-6-663-2009 evas, S., Rochet, M. J., and D. Tracey, M.: 3388 3387 (mm) (mm) neighbour neighbour distance distance minimum maximum ¨ ottner, S., Naumann, M., Hoffmann, F., and Rapp, H. T.: neighbour neighbour nearest nearest ¨ alv, T., Guihen, D., Kiriakoulakis, K., Lavaleye, M., and Duin- 2 − of m frames nearest nearest distance distance Shrimp densities observed across the various habitat categories. Mean and median Organic matter release by coldMar. water Ecol. corals Prog. and Ser., its 372, implication 67–75, for 2008. fauna-microbe interaction, 405, 1987. tinental margin, Int. J. Earth Sci. 96, 1–9, 2007. eveld, G.: Cold-water coralfor carbon ecosystem cycling, Mar. (Tisler Ecol. Reef, Prog. Ser., Norwegian 465, Shelf) 11–23, 2012. may be a hotsport mounds on the NW European margin, Int. J. Earth Sci., 96, 37–56, 2007. mineralization and carbonate preservation inNorwegian modern shelf, cold-water Biogeosciences, coral 6, reef 663–680, sediments doi: on the ecosystem research on Europe’s deep-ocean margins, Oceanography, 17, 132–143, 2004. cold-water coral community as aweb hot analysis spot for from carbon2009. Rockall cycling Bank on continental (northeast margins: Atlantic), a food- Limnol. Oceanogr.,fluxes 54, and hydrodynamics 1829–1844, at the Tisler cold-water coral reef, J. Mar. Sys., 85, 19–29, 2011. ocean acidification destroy cold-water ecosystems?, Coral Reefs, 26, 445–448, 2007. Availability of deep-water fish trawling andcle cisual (ROV), Mar. observation Ecol. from Prog. a Ser., remotely 284, operated 293–303, vehi- 2004. Habitat Number ShrimpsLive SD LopheliaLive ParagorgiaLive Primnoa Mean 365Dead structure 8Coral rubble SDGravel/Pebbles 24.41 779 5Wall 43.00 Median 1330 18.64Hardground 699 35.58 8.70 41.60 11.08 Mean 0.28 10.92 26.06 1.45 7.80 30 5.81 0.93 SD 10.57 6.22 2.52 15.27 318 10.07 0.67 1.5 Mean 6.23 11.97 7.92 7.13 1.95 1.94 4.65 13.54 SD 7.20 8.02 13.64 3.38 3.62 4.15 4.38 8.52 0 6.67 0 4.71 2.29 11.04 24.45 10.74 12.73 6.08 24.95 2.26 11.42 29.79 5.41 1.21 13.99 21.00 0 20.94 12.57 4.01 7.41 8.65 10.55 10.96 2.10 4.50 19.35 4.43 10.67 10.68 11.30 11.50 20.65 Zann, L. P.: A review of macrosymbiosis in the coral reef ecosystem, Int. J. Parasitol. 17, 399– Wild, C., Mayr, C., Wehrmann, L., Sch White, M., Wolff, G. A., Lund White, M.: Benthic dynamics at the carbonate mound regions of the Porcupine Sea Bight con- Wheeler, A. J., Beyer, A., and Freiwald, A.: Morphology and environment of deep-water coral Table 1. distance between individual shrimpsalso observed given. in Mean associationshown. minimum with and each maximum habitat category distances are between nearest neighbours are also Wehrmann, L. M., Knab, N. J., Pirlet, H., Unnithan, V., Wild, C., and Ferdelman, T. G.: Carbon Weaver, P.P.E., Billett, D. S. M., Boetius, A., Danovaro, R., Freiwald, A., and Sibuet, M.: Hotspot Wagner, H., Purser, A., Thomsen, L., Jesus, C. C., and Lund Van Oevelen, D., Duineveld, G., Lavaleye, M., Mienis, F., Soetaert, K., and Heip, C. H. R.: The Turley, C. M., Roberts, J. M., and Guinotte, J. M.: Corals in deep-water: will the unseen hand of Trenkel, V. M., Chris Francis, R. I. C., Lorance, P.,Mah 5 25 20 15 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | % live 50 > dead struc- cm 10 > 50% The downstream > (b) cm and 10 scale scale < cm shrimps shrimps pattern Typical “cauliflower” growth form 10 of each “cauliflower” is made up of > (a) cm 25 cm 10 ∼ scale scale < Primnoa resedaeformis 3390 3389 cm scale) shrimp distribution patterns observed in of shrimp shrimp regularly regularly with no , live 40 < 2 20 − m > shrimp habitat clustering clustering spaced spaced spatial shrimps. The number of frames taken above the 20 > shrimps present is indicated. Insufficient shrimps were observed in colony at the Røst reef, Norway. 2 − 20 , the remainder the dead skeletal structure of previous polyp generations. Paragorgia arborea > , live of shrimp frame Frames Frames with Frames with Frames with Frames with Frames frames m Table summarizing the local ( Lophelia pertusa Lophelia pertusa Main Habitat Number Mean SD NumberLive LopheliaLive % ParagorgiaLive Primnoa 365Dead structure 8 24.41 % 779 5 43.00 18.64 35.58 8.70 41.60 187 26.06 7.80 5 % 51.3 4 71 62.5 20.3 80.0 9.1 % 40.0 55.6 50.0 42.6 60.0 % 31.0 75.0 75.7 60.0 % 25.0 8.6 10.0 20.0 14.4 0 7.1 0 12.9 0 at the Røst reef, withside live polyps of facing the into the “cauliflower” prevalent reef current structure. direction. The upper Fig. 1. living Image courtesy IFM-GEOMAR JAGO team. Table 2. image frames containing Lophelia pertusa association with the other habitat categories investigated to allow computation of Ripley’s L. ture categories with Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | (b) colonies at the Røst reef, Nor- structure or on the walls of coral blocks. 3392 3391 Paragorgia arborea and Lophelia pertusa . A large salmon coloured colony in the middle distance, a pink , the orange branching gorgonian in the lower centre of the image. P. arborea Primnoa resedaeformis P. resedaeformis Bathymetric map showing the location of the Røst reef on the Norwegian margin, and Typical (a) the location of the three video transects analysed in this study. Fig. 3. colony in top rightits and polyps a retracted. salmon coloured colony in bottom right. The bottom right colony has Three colonies of way. Often located in cracks in the Fig. 2. Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | dead scle- Paragorgia (d) live (b) coral and a number , dead gorgonian (right of (h) (left of image), wall, Lophelia pertusa (g) Lophelia pertusa Live (a) 3394 3393 Primnoa resedaeformis gravel/pebbles, (f) live (c) hardground. coral, a smaller region of live (j) coral rubble, (e) soft seabed and (left and right of image), (i) Habitat categories used in this study. Example frame extracted from the video sled data within the BIIGLE system. A number Primnoa resedaeformis Fig. 5. of slightly blurred shrimplive eyes have been labelledof shrimp in within green. and The around image the shows coral structures. predominantly Fig. 4. ractinian structure, image), arborea Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | 3396 3395 Boxplot showing shrimp densities observed in image frames with differing numbers of Boxplot showing observed median shrimp densities, quartiles and outliers observed in Fig. 7. habitat categories present. Fig. 6. association with each of the investigatedDead main habitat gorgonian categories. or No frames soft contained sedimentfigure. primarily habitats, and these categories are therefore absent from the