Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | , 6 , R. R. Kirby 5 , J. Kennedy 3,4 , A. Hosia 630 629 1 1 , A. Fischer 1,2 , and P. Tranter 1 , M. Blackett 1 , R. Stern 7 This discussion paper is/has beenData under (ESSD). review for Please the refer journal to the Earth corresponding System final Science paper in ESSD if available. Sir Alister Hardy Foundation for Ocean Science (SAHFOS), The Laboratory,School Citadel of Hill, Ocean and Earth Science, National Oceanography Centre, UniversityUniversity of Museum of Bergen, UniversityInstitute of of Bergen, Marine P.O. Box Research, 7800, P.O. 5020 BoxDepartment 1870, Bergen, of 5817 Norway Environment, Nordnes, Fisheries Bergen, and Norway Sealing Division, P.O. Box 1000Marine Station Institute, 1390, Plymouth University, DrakeInstitute Circus, for Plymouth Marine PL4 Resources 8AA, and UK Ecosystem Studies (IMARES), P.O. Box 68, 1970 AB Received: 26 February 2014 – Accepted: 26Correspondence June to: 2014 P. Licandro – ([email protected]) Published: 5 November 2014 Published by Copernicus Publications. Earth Syst. Sci. Data Discuss.,www.earth-syst-sci-data-discuss.net/7/629/2014/ 7, 629–667, 2014 doi:10.5194/essdd-7-629-2014 © Author(s) 2014. CC Attribution 3.0 License. K. Raab 1 Plymouth PL1 2PB, UK 2 Southampton, European Way, Southampton3 SO14 3ZH, UK 4 5 Iqaluit, Nunavut, XOA OHO,6 Canada 7 IJmuiden, the Netherlands Biogeography of jellyfish in theAtlantic, North by traditional and genomic methods P. Licandro Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | G.O. erent ff http://doi. ers significantly ff E Long. Jellyfish were ◦ W–5 ◦ erent data sets produced within the EU ff N Lat and 62 ◦ 632 631 ). erent types of data, key jellyfish taxa for the spring-summer period erent water masses and with the bathymetry. Higher densities were ff ff shore waters between 59–68 ff transatlantic cruise show that information on jellyfish diversity di All observations presented here can be downloaded from PANGAEA ( Combining di The identification by DNA barcoding of 23 jellyfish specimens collected during the Based on CPR data from 2009–2012, blooms of Cnidarians occurred in all seasons Using a net adapted to sample gelatinous zooplankton quantitatively, Cnidaria and Overall the data show high variability in jellyfish abundance and diversity, mainly in EUROBASIN cruises contributes tosequences increasing available the on still GenBank. very limited number of jellyfish were identified in thesouthern northern North North Atlantic Atlantic could regions.CPR be Key survey, inferred although species based this for on should the be Cnidariandata. confirmed central blooms further and identified by by comparison the with quantitative to the presence ofSars meropelagic medusae. Comparisons ofdepending net on the records sampling from gear the large utilised. scyphozoan Indeed, and the big hydrozoan species, trawlsctenophora mostly while are collect small only relatively hydrozoans caught and by early smaller stages nets. of across the whole North Atlantic basin.what Molecular was analysis previously revealed hypothesized, that, the in CPRand contrast is with holoplanktonic able hydrozoans to and detect scyphozoans. blooms of meroplanktonic 2012 were re-analysed withblooms the across the aim North of Atlantic identifying basin. the time andrelation location with of di Cnidarian generally recorded on the shelves, where populations tend to be more diversified due to date overview ofregion. the diversity and standing stocks of jellyfishCtenophora in were collected the in the North epipelagicinshore Atlantic layer and during o spring-summer 2010–2013, in also identified and countedequipment in in the samples same opportunisticallythe region collected Gulf and by at of two other Cadiz. coastal sampling Continuous stations Plankton in Recorder the (CPR) Bay samples of Biscay collected and in in 2009– Scientific debate onrelated whether to the a recent trueCnidaria increase rise and in Ctenophora. in Here their reports we abundance, describeprogram of di EUROBASIN, have jellyfish which outlined have the outbreaks been lack is assembled of with reliable the records aim of of presenting an up Abstract might be planktonic andshapes benthonic, and solitary sizes), their or fragility colonial, thatand can with the compromise a poor some key knowledge large morphological of range features their of taxonomy. di 2013). Within this debate, itdata has (Purcell, been 2009; recognised Brotz that et there al.,used is 2012; to Condon a describe et lack a al., of defined 2012). reliable planktoning “Jellyfish” jellyfish functional is to group, a i.e. the general gelatinous term two carnivoresbe phyla belong- extremely Cnidaria challenging, and due to Ctenophora. their The morphological identification complexity (Cnidaria of for those instance, groups can In recent years a globala increase general in consensus jellyfish on abundancescientific has community this been pointed matter widely out has debated,marine increasing but not frequencies and of been estuarine jellyfish achieved regionset outbreak yet. worldwide al., events 2005; While (e.g. in Kawahara Brodeur a et2012), al., et part 2006; some al., of Atrill studies 1999; et the suggested al., Mills,up- that 2007; and 2001; Licandro the downward Xian et oscillations rise al., characterising in 2010; their long-term Brotz jellyfish et periodicity abundance (Condon al., is et al., just a phase of pangaea.de/10.1594/PANGAEA.835732 1 Introduction 5 5 25 15 20 10 25 20 10 15 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | the G.O. ff Multiple Open- 2 cruise they were col- ) and are trapped between 2 erent sampling gears provides ff G.O. Sars cruise, Table 2 and Fig. 1). the Cumberland Peninsula, in the ff cruise and the transatlantic 2013- Arctic 634 633 Icelandic 100 cm long) and an internal mechanism contain- × ) is fundamental to collect intact specimens that would 1 − 50 cm tall erent jellyfish data sets produced within the EU program EU- × ff Cruise, the 2012- Meteor erent depths in the 0–1000 m layer using a standard 1 m ff erent taxonomists to ensure consistency and quality control of the data. cruise (Table 2 and Fig. 1). The same net was used to sample jellyfish o ff erent types of nets were used to collect jellyfish in several North Atlantic regions Even though the sampling methodology is not particularly suitable to quantitatively Jellyfish were also identified and counted in samples opportunistically collected with A “gentle” net, hereafter called the “jellynet”, was designed following the main spec- Here we describe di Conventional sampling methodologies are often inappropriate to quantify jellyfish ff on the back of the sampler. Once back at the laboratory, the internal mechanism is The Continuous Plankton Recordertowed at (CPR) the surface is (7 m aping nominal routes high-speed depth) by (Richardson plankton ships et of sampler al.,proximately opportunity 50 2006). that along cm The their wide is usual CPR ship- ising composed a of spool an with externalthe two body plankton overlapping (ap- enter bands through of thethe silk mouth mesh filtering of (270 the silk µm CPR aperture).wound and (1.61 cm During up the a on covering tow, a silk. spool The located two in bands a of formalin-filled silk tank, are driven then by progressively a propeller situated tion, with the exception ofGulf the of Cadiz samples and collected in o by the di Bay of Biscay. The taxonomic identification was crosschecked 2.2 CPR data 2003) (quantitative data), Harstad (Nedreaas andtrawls Smedstad, (qualitative 1987) data) and macroplankton (Tables 1 and 3). catch jellyfish specimens, samplesof collected Cadiz during (i.e. 2010 IEOble by dataset, 3) Bongo Table were 3) nets analysedcomposition and in to of in the provide jellyfish the baseline Gulf populationsidentification Bay in information of the of on jellyfish southern Biscay the was, regions (i.e. whenever relative of possible, abundance AZTI undertaken the and dataset, immediately North Ta- after Atlantic. collec- The Cumberland Peninsula (Canada) in 2011 (i.e. other sampling gears (Table 3lected and at Fig. di 1). Duringing/Closing the Net and Environmental Sensing System (MOCNESS, Wiebe and Benfield, Sars i.e. the 2012- (Fig. 1 and Table 1). ifications of a Régentlections net, of which gelatinous has organismsmouth been (Braconnot, fitted with shown 1971). a to The 2 mdiameter be jellynet long and suitable tapered has 30 cm net for a in and quantitative length. a 1mesh m col- The large aperture. non-filtering net diameter mesh rigid The is cod-end200 knitted 14 jellynet m) cm polyester across in was with the a used whole nominal North to 800 Atlantic µm basin, collect during jellyfish three main in EUROBASIN Cruises, the epipelagic layer (0– 2 Data 2.1 Net data Di Slow towing speed (0.5–1 m s be otherwise badly damaged. ROBASIN, assembled with the aim ofand presenting abundance an of up North to date Atlanticthe overview jellyfish. of opportunity The the to use diversity discuss of the di ence limitation on of the each quality methodological of approach the and data. its influ- standing stocks and to evaluate the diversitywater of must their populations. be A filtered large(Purcell, volume to of 2009). sea- collect Silk planktonic or jellyfish, polyesterstramine which mesh mesh should are (traditionally be usually used preferablyages very used to or dispersed rather destroys collect than many plankton nylon delicate samples), or species which of gelatinous severely zooplankton dam- (Braconnot, 1971). 5 5 25 20 10 15 20 25 15 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | er ff C (10 min). ◦ erent standard ff er then processing ff C. ◦ ) and 10 µL of 10 % SDS and 1 − C for 10 min in a dry-block heater, ◦ C (1 min), followed by 40 or 50 cycles of ◦ erent North Atlantic routes (Fig. 1) were vi- ff 636 635 C (1 min) and a final extension of 72 ◦ erent Polymerase Chain Reaction (PCR) amplification strategies and C (1 min) and 72 C for 4 h. Each sample was then vortexed briefly and centrifuged at ◦ ff ◦ C (1 min), 51 ◦ The PCR products were visualised on a 1 % agarose gel and either purified using A third protocol was used to extract DNA from jellyfish material embedded in the silk. A number of di In one case, a 540-bp partial, mtDNA 16S rDNA sequence was amplified by PCR Protocol 2 consisted of washing the tissues samples in TE bu Protocol 1 followed the methodology developed by Kirby et al. (2006). Briefly, small The visual identification of cnidarian jellyfish tissue and/or nematocysts in CPR sam- Jellyfish DNA was extractedwhich from were about collected 80 ethanol-preserved during cnidarian the specimens, EUROBASIN cruises and identified on board or to remove primer-dimers. Purified(MWG PCR Biotech, products Germany, or were Source thention Bioscience, primers sequenced Nottingham, as UK) commercially sequencing using primers. thewas Alternatively amplifica- performed Sanger using sequencing BigDye of kit PCRreverse products (Applied primer Biosystems, for USA), amplification, withelectrophoresis either according of the to sequencing forward manufacturer products or instructions carried out and at capilliary Source Bioscience. 2.3.2 DNA extraction from net samples preserved in ethanol using the primers of Cunninghaminvolved and Buss an (1993) initial and Schroth denaturation94 et al. step (2002). of The PCR 94 Montage spin columns (Millipore) or treated with ExoSAPIT (Illustra, supplied by VWR) 30 µL. In this case, approximatelyand one total third environmental of DNA abased was protocol CPR extracted developed sample by from was Ripley it et cut, according al. washed (2008). to in a TEmarkers phenol-chloroform bu were used. the sample either withBiotechnologies, USA) the using Masterpure protocol BK total (tissue digestion DNA samples) step with and of anthe RNA 4–12 extended h Proteinase homogenisation extraction or of kit using tissues DNAzol (Epicentre scribed reagent procedure by (LifeTechnologies, with USA) the the applying manufacturers. optional DNA centrifugation pellets step were as de- then dissolved in a final volume of then recovered. All centrifugation steps were performed at 22 incubated at 55 12000 g for 15 s. Samplesvortexed for were 10 then s and heated centrifuged atferred at 105 12000 to g a for Micropure-EZ 3 min.a centrifugal The Microcon supernatant YM-30 filter CFD was device (Millipore then (CFD) Corp.) trans- carding and (Millipore centrifuged the Corp.) at Micropure-EZ 14000 inserted CFD, gtimes into the for 8 with sample min. 200 After µL retained dis- of in8 sterile min the water; and YM-30 the the was first final two washed wash washes three was were centrifuged centrifuged at at 14000 14000 g g for for 5 min. The retained DNA was protocols. pieces of tissue from individual specimensdividually (approximately into 1 mm 180 µL length) were of placed chelex1 in- M solution Dithiothreitol (Instagene (DTT), Matrix, 4 Biorad) µL together of with proteinase-K 6 (10 µL mg of mL classified as records of jellyfishmethods outbreak were events then (Licandro used et in al.,identify some 2010, CPR cnidarian Fig. samples blooming 1). where species. Genetic swarm events were recorded to 2.3 Genetic analysis of Jellyfish 2.3.1 DNA extraction from CPR samplesJellyfish preserved DNA in collected formaldehyde from CPR samples was extracted using three di 10 nautical miles. ples has been carriedsamples out collected routinely in since 2009–2012 1958.sually along Within the di re-analyzed project and EUROBASIN, those CPR fully covered in jellyfish tissue and nematocysts were unloaded, the spool is unrolled and the silk is cut in sections that correspond to circa 5 5 25 20 10 15 25 15 20 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | C. ◦ Mug- Nanomia and . A few stations 3 cruise, Stns. 255 cruise) and west- − ) 3 spp. represented 11 % − Arctic O. A 1 µL aliquot of the 2 spp. and Mertensidae were G.O. Sars , the siphonophores ) and digested for 4 h at 55 Muggiaea atlantica 1 − Beroe Hormiphora cruise) ctenophores made up 90–100 % cruise, St. 152- 638 637 er (10 mM Tris-Cl pH 7.9, 100 mM EDTA and ff Aglantha digitale of jellyfish tissue was placed into a 1.5 mL Eppen- 3 G.O. Sars Meteor 1 mm , and the ctenophores erent regions of the North Atlantic. Further analysis was ∼ ff cruise) Atlantic shelves exhibited elevated abundance with Arctic cruise, St. 162- (not shown). Generally only cnidarians were found in the samples (Ta- Dimophyes arctica Icelandic and In the 0–200 m layer, cnidarians tended to be generallyOverall 27 more cnidarians and abundant 5 than ctenophore taxa were identified and counted in North A 540-bp partial, mtDNA 16S rDNA sequence was then amplified by PCR using the Overall 23 cnidarian taxa were successfully sequenced and published on GenBank giaea kochi ble 5), except in March 2010 when the ctenophore cara the most common taxa in epipelagic waters across the3.1.2 northern North Atlantic Bongo region. data In shallow waters in theand Gulf of time. Cadiz, They jellyfish were distributionwith was relatively high highly more variable peaks in abundant due space in to early swarms spring of and the autumn siphonophores (Fig. 3a), of the total jellyfish abundance. Atlantic epipelagic waters (Table 4).northeast Jellyfish Atlantic, populations mainly were moreand due Leptomedusae. diversified to The in trachymedusa the the presence of meroplanktonic species of Antho- The data collected in epipelagic watersstanding in stocks 2011–2013 showed across high the variabilitydance in northern jellyfish (Fig. North 2a–c) Atlantic generally basin rangedlocated between (Fig. on 0.42 2). the and Total eastern jellyfish 12ern (i.e. ind. abun- (Stns. St. 100 m 1 3- anddensities 2- one order of magnitude greater (max. 246 ind.ctenophores 100 m (Fig. 2d–f), evenand though 315- in some stations (St.4- 3.1 Jellyfish abundance and diversity in3.1.1 epipelagic waters Jellynet data 3 Results performed by aligning DNAmarker sequences from with public databases Cnidaria using sequences Bioeditexported (Hall from et into the al., MEGA 1999). same 5.1 These DNA were (Katohjoining trimmed et methods and al., with 1995) a toconfidence Kimura-2 produce intervals. substitution phylogenies using model Neighbour and tested using 1000 bootstrap 2.3.3 DNA sequence analysis Sequence identity of CPRpublic repositories cnidarian and tissue to wasplankton private established net databases firstly samples of in by Cnidaria di comparison DNA to sequences taken from extracted DNA was then used as template in a PCR. primers of Cunningham and Buss (1993)file and described Schroth above. et PCR al. (2002) productsusing and were the Montage visualised thermal spin on pro- columns a (Millipore).commercially 1 % (MWG Purified agarose Biotech) PCR gel using products and the were purified amplification then primers sequenced as sequencing(Table primers. 9). chloroform protocol. Briefly, dorf tube containing 400 µL0.5 % cell SDS) lysis with bu 4 µLFollowing proteinase-K a solution phenol-chloroform (10 purification mg mL theing DNA NaCl was recovered and by precipitation EtOH us- and resuspended in 40 µL nanopure H shortly after collection. DNA extraction followed a standard SDS, Proteinase-K, phenol- 5 5 20 25 15 10 20 10 15 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | ) and Aglan- cruise, shore from ff G.O. Sars ) were mostly col- Clytia, Gilia, Muggiaea shore from spring to late autumn, ff erent sampling methodologies during ff erent sampling gears ff the Norwegian trench and in the Icelandic ) were consistently sampled by all the gears. sp. were recorded in summer on the eastern ff 640 639 Atolla, Pelagia, Praya, Vogtia Cyanea Beroe cucumis et al., 2003). The relatively small mouth opening character- , ff erent depths in the 0–1000 m layer during the ff shore across the whole North Atlantic basin (Fig. 6). Genetic were recorded inshore and o ff transatlantic cruise showed that only a few dominant species (e.g. Nanomia cara , Pelagia noctiluca G.O. Sars erent species of colonial siphonophores were swarming inshore and o The comparison of the data collected with di Species diversity was generally higher in the mesopelagic than in the epipelagic layer Jellyfish species typically distributed in cold-temperate and warm-water regions were ff ples collected with 300 andthe 700 µm best mesh compromise demonstrated that to the quantitatively latter size catch represents meso- and macroplanktonic gelatinous equipped with monofilament wovenspecies nylon, of often irremediably Cnidariapolyester damage and have many shown Ctenophora, delicate to(Braconnot, while better 1971; preserve softer Rasko the material delicateising body standard such plankton of nets as gelatinous (e.g. circalimits zooplankton silk 50 the cm volume or mouth of diameter seawater knitted in filteredtative Bongo and records and therefore of WP2 is jellyfish. nets) not Even though appropriatesuitable 200 to µm to provide mesh quanti- collect size might small be considered hydromedusae the (e.g. most Cornelius, 1995), comparisons of sam- 4 Discussion Sampling jellyfish is challengingpersed as or these unevenly organisms distributed are (Purcell, delicate 2009). and Conventional often nets, very which dis- are usually sons, inshore andanalysis o of jellyfish material collecteddrozoans from as CPR well samples as identified blooms ofdi of relatively small big hy- scyphomedusae (Table 8).summer to Among early the autumn first (Fig. 7). group, ian In the second group, blooms ofwhile the swarms holopelagic cnidar- of the meropelagic and western Atlantic shelf. and the Mocness (Tables 4 and 6). 3.4 Jellyfish blooms as identified byBased the on CPR CPR deployments from 2009 to 2012, jellyfish blooms occurred in all sea- early stages of ctenophora were only caught by the smaller nets, such as the Jellynet Conversely, relatively large species (e.g. lected by big trawls (Table 7), while small hydrozoans (e.g. 3.3 Jellyfish diversity: comparison of di Thirty-seven species/genera of jellyfishble were 6), identified while thirty-two inand taxa the Harstad were Mocness trawls counted (Table samples 7). in (Ta- samples collected by thethe Macroplankton tha digitale evenly distributed across the water column or mainly concentrated at(Fig. the 5), with surface. the highest numberand of Labrador species seas. being recorded below 400 m in the Irminger 3.2 Jellyfish abundance and diversity in3.2.1 the 0–1000 m layer Mocness data The data collected at di show that in early Maymesopelagic 2013 the layer bulk (200–1000 of m the depth)Sea jellyfish o population (Fig. was 4). concentrated On in the the contrary, in the Irminger and Labrador Seas, jellyfish were more shown). recorded in the Bay of Biscayregion (Table 5). jellyfish Their are densities less inshould May abundant 2010 be than suggest further in that verified. in the this Bay of Cadiz (Fig. 3b), even though this and 63 % of the total jellyfish standing stock respectively at Stns. P-01 and G-01 (not 5 5 20 25 15 10 20 25 15 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | 800 and ≤ G.O. Sars ers a unique ff and other hydro- erent information ff Nanomia cara recorded by the CPR spp. and other Agal- ) and early stages of and ered. ff P. noctiluca, 6 m mouth opening and 3 cm ≥ erent nets during the Halistemma ff , erent sampling gears used during the Pelagia noctiluca spp. were the epipelagic species most ff , due to the large mesh size and large Dimophyes arctica , 642 641 Clytia, Gilia, Muggiaea shore waters the whole year round. Genetic erent species of hydrozoan siphonophores) Beroe ff ff erent water masses and with the bathymetry. The erent sampling gears provide di ff ff erent types of jellyfish communities. ff and di Muggiaea atlantica spp. and shore. Based on the CPR, ff Aglantha digitale sp.), which are characterised by the alternation of a benthic P. noctiluca Atolla, Pelagia, Praya, Vogtia Mertensiidae Cyanea transatlantic cruise, even if their abundance di Ireland in October 2007, were confirmed by net tows (see Fig. 2 in Licandro et al., Indeed, the re-analysis of CPR samples collected in recent years showed that jelly- The CPR Survey is the monitoring programme that covers the greatest spatial (tens Overall, the hydrozoans The use of modern technology, in particular of remotely operated vehicles equipped In agreement with previous studies (Hosia et al., 2008; Purcell, 2009 and references The data collected in epipelagic waters by the jellynet in the northern North Atlantic ff lantic regions. Those observations, in particular the high abundance of small hydrozoan fish blooms can occuranalysis in of coastal CPR andscyphomedusa o cnidarian material indicatespolyp that stage meroplanktonic andtonic jellyfish a species (e.g. pelagic (e.g. the medusa,swarm tend both to inshore and bloomzoan o over siphonophores the including shelf,matidae are while among holoplank- the main swarming species in the central and southern North At- scyphozoans. Outbreaks of the scyphomedusa o 2010 comparing CPR swarms eventsthat and the records CPR from can Doyle et providewhich al., reliable jellyfish 2008), information prefer suggesting to to help bloom. clarify the regions and periods in to thousands kilometres)plankton and at the temporal surface across (monthly theon whole plankton to North is Atlantic multidecadal) in usually regionsopportunity available scales, where to (Richardson no sampling document information et jellyfish al., swarms,tances 2006). which of are It ten-hundreds events therefore usually of o scale occurring kilometres methods over of dis- (e.g. data Brodeur collectionpreviously et are hypothesized needed al., (Atrill (Purcell, 2008) 2009). et and Inis al., contrast for able 2007; to with which Gibbons detect what large- blooms and was of Richardson, meroplanktonic 2009), as well the as CPR of holoplanktonic hydrozoans and erences therein). Though, as visualtowards observations species from of a detectable ship size andmethodologies or relatively from cannot simple a provide taxonomic pier reliable identification, are information these of biased on jellyfish the populations abundance throughout and the composition oceans. of relatively big scyphomedusae and other gelatinous plankton (Purcell, 2009 and ref- shore-based surveys have been used to provide semi-quantitative/qualitative estimates the ctenophores frequently recorded in thepresence northern of those North key Atlantic taxaG.O. region Sars was during detected by spring-summer. di The with underwater cameras andlect video-systems, in situ has information proven onet to gelatinous al., plankton, be 2008; particularly very Stemmann in et valuable deeptherefore al., waters to unlikely 2008). (e.g. to col- Nevertheless, Lindsay be video employed systems for are standard still jellyfish quite costly, monitoring. Ocean-surface and volume filtered. Small hydrozoansctenophora (e.g. were only caughtmesh by size in the this smaller study). netsprograms Therefore sampling are (i.e. gear set 1 should up m be to mouth carefully monitor considered opening di when and populations were less diverse inwhere Arctic more waters meropelagic than medusae on are the North-eastern present. Atlantic shelf, therein), a comparison oftransatlantic cruise records confirms collected that di withon di jellyfish populations.mesh Indeed, size the in big thisspecies trawls study) such (i.e. mostly as collected relatively large scyphozoan and hydrozoan 1997, 1999). regions, showed high variability inerally jellyfish observed standing on stocks, the withalso eastern varied, higher mainly and densities in western gen- relation North with di Atlantic shelves. Jellyfish diversity zooplankton, whilst limiting damage for jellyfish soft tissues (Braconnot, 1971; Buecher, 5 5 25 20 10 15 25 20 10 15 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | , 2014. (Cnidaria, , 2014. , 2014. erent planktonic ff (Cydippid Ctenophore) in Abylopsis tetragona and 10.1594/PANGAEA.830244 10.1594/PANGAEA.828647 10.1594/PANGAEA.829714 , 2014. 644 643 Pleurobrachia rhodopis , 2014a. , 2014b. Chelophyes appendiculata We wish to acknowledge the Instituto Español de Oceanografía (IEO) 10.1594/PANGAEA.828648 10.1594/PANGAEA.830265 10.1594/PANGAEA.830263 Overall, records of jellyfish swarms reported by the CPR, can help to identify North zan, H. W., Bogeberg, M.,Haddock, Purcell, S. J. H. E., Decker, D., M.Arthur, Malej, B., J. A., Uye, M., Parry, S., G. Madin, andoscillations, D., L. Graham, P. Eriksen, P., Natl. Brodeur, W. Acad. E., R. M.: Sci. D., Quiñones, Recurrent USA, J., 110, jellyfish Acha, 1000–1005, blooms M., 2013. are Harvey, M., a consequence of global Shuterland, K. R., Robinson, K.Malej, L., A., Dawson, Mianzan, M. N., H.gelatinous Decker, zooplankton M. W., in B., Uye, the Mills, S., world’s C. oceans, Gelcich, E., Bioscience, Purcell, S., 62, J. 160–169, and E., 2012. Madin, L. P.: Questioning the rise of in May 2012, doi: populations: trends in large marine ecosystems, Hydrobiologia, 690, 3–20,the 2012. Bay of Villefranche-sur-Mer (Northwestern Mediterranean)time from series, three Ann. di I. Oceanogr. Paris, 72, 173–184, 1997. Siphonophora) in the Bay of Villefranche,307, northwestern 1999. Mediterranean, J. Sea Res., 41, 295– and Hunt Jr., G. L.:shifts, The Progr. rise Oceanogr., and 77, fall 103–111, of 2008. jellyfish in the Bering Sea in relation to climate regime Atlantic collected in August and September 2011, doi: a more gelatinous future for the North Sea, Limnol.des Oceanogr., et 52, Doliolides 480–485, I. 2007. Hydrologie et écologie des Salpides,for Vie a Milieu, substantial 22, increase 257–286, in 1971. climate gelatinous change, zooplankton Fish. in Oceanogr., 8, the 296–306, Bering 1999. Sea, with possible links to pled during the G.O. Sars Cruise in May 2013, doi:10.1594/PANGAEA.828646, 2014. Cnidaria and Ctenophora taxa in the North Atlantic, doi: doi: doi: sampled during the M87/1 cruise in April 2012, doi: with the information collected in the Bay of Biscay andAtlantic Gulf regions of more Cadiz. impactedronmental by blooming change events and/or and anthropogenic helpcurrence. pressure to can discern whether explain envi- increasing jellyfish oc- siphonophores in coastal regions, while they are yet to be confirmed, are in agreement Condon, R. H., Duarte, C. M., Pitt, K. A., Robinson, K. L., Lucas, C. H., Shuterland, K. R., Mian- Condon, R. H., Graham, W. M., Duarte, C. M., Pitt, K. A., Lucas, C. H., Haddock, S. H. D., Licandro, P.and Raab, K.: Abundance of Cnidaria and Ctenophora in the Icelandic Sea sampled Buecher, E.: Distribution and abundance of Buecher, E.: Appearance of Brotz, L., Cheung, W. W. L., Kleisner, K., Pakhomov, E., and Pauly, D.: Increasing jellyfish Brodeur, R. D., Decker, M. B., Ciannelli, L., Purcell, J. E., Bond, N. A., Stabeno, P. J., Acuna, E., Brodeur, R. D., Mills, C. E., Overland, J. E., Walters, G. E., and Schumacher, J. D.: Evidence Licandro, P. and Kennedy, J.: Abundance of Cnidaria and Ctenophora in the North Western Braconnot, J. C.: Contribution à l’étude biologique et écologique des Tuniciers pélagiques Salpi- Licandro, P. and Hosia, A.: Abundance of Cnidaria and Ctenophora in the North Atlantic sam- Licandro, P., Blackett, M., Hosia, A., Kennedy, J., and Raab, K.: Presence and absenceReferences of Attrill, M., Wright, J., and Edwards, M.: Climate-related increases in jellyfish frequency suggest Licandro, P.: Abundance of CnidariaLicandro, and P. and Ctenophora Blackett, in M.: Abundance the of Bay Cnidaria of and Ctenophora Biscay in (May the 2010), North Atlantic Licandro, P.: Abundance of Cnidaria and Ctenophora in the Gulf of Cadiz 2010, EUROBASIN cruises, for facilitating injellyfish any specimens. possible This manner work theEuropean was collection Union). supported and by identification the of project EURO-BASIN (Ref. 264933, 7FP, Source Data References from Cádiz and AZTI Technaliaples from in San the Sebastian Gulf (Spain) ofC. for Cadiz González the and from collection in of IEO plankton the andC. sam- Bay to Vilas of Unai from Biscay. Cotano Thanks from IEODNA are AZTI. barcode. in particularly Thanks We due Cádiz, also are to to Spain, particularly M. L. for grateful Llope Prieto to providing and from the jellyfish CSIC heads and specimens of preserved mission and in scientists ethanol on for board the Acknowledgements. 5 5 30 25 20 15 10 25 20 10 15 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | in Irish communities, J. , 2010. 10.1186/1471-2148-2-1 Pelagia noctiluca Emiliania huxleyi 10.1098/rsbl.2010.0150 645 646 sp., BMC Evol. Biol., 2, doi: Aurelia (Scyphozoa: Rhizostomeae) in East Asian waters, Mar. Ecol.- , 2006. Nemopilema nomurai erent hydrographic regimes along the northern portion of the Mid-Atlantic Ridge, Deep- , K. A., Sommer, F. A., Hamner, W. M., and Cross, K. M.: Collection and culture tech- ff ff 10.1098/rsbl.2006.0530 conditions?, Hydrobiologia, 451, 55–68, 2001. 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Introduc- 5 5 30 15 20 25 10 20 15 10 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | (µm) diameter (m) JellynetJellynet 800MocnessHarstad trawl 1 Macroplankton trawl 800 180 30 3000 000 1 1 20 6 erent North Atlantic regions. ff 267, 272, 273,281, 274, 290, 292,305, 299, 307, 315,330, 324, 332, 333,340 338, 159, 160, 160bis, 161, 162, 163,166, 165, 167, 168,170, 169, 171 101, 102, 104,106, 105, 107, 108, 109, 111, 115, 116,118, 117, 120, 121,123, 122, 124, 125,127 126, E 152, 154, 155, 157, ◦ E 1, 2, 3 Jellynet 800 1 W 1, 2, 3, 4 Jellynet 800 1 ◦ ◦ W–5 ◦ WW T-01, P-01, G-01 58, 67, 68, 69 Bongo net Bongo net 200 200 0.4 0.4 W–1 ◦ ◦ ◦ 648 647 depth (m) (start, LT) depth (m) N 46 N 62–68 N 2 ◦ ◦ ◦ NN 6 5 EEE 150E 150E 150 16:14E 150 16:35 150 15 01:58 Apr 150 2012 15 02:22 Apr 2012 16 06:07 Apr 2012 165 16 06:34 Apr 2012 165 16 Apr 2012 165 16 Apr 2012 165 165 165 WWW 200W 200W 200 07:45W 200 08:13W 200 17:27 9W 200 Apr 2012 17:58 9W 200 Apr 2012 05:37 9W 200 Apr 2012 1350 06:07 9W 200 Apr 2012 1350 10 18:04 AprW 200 2012 1350 10 18:35 AprW 200 2012 1350 10 16:14 AprW 200 1350 2012 10 16:41 AprW 400 1350 2012 12 05:54 AprW 400 1350 2012 12 06:25 Apr 200 1350 2012 13 11:29 Apr 200 1300 2012 13 02:30 Apr 1300 2012 13 04:47 Apr 1300 2012 14 05:17 Apr 1300 2012 14 Apr 1300 2012 14 AprW 1300 2012W 1300 W 400 1300 W 200W 200 03:34W 200 05:03W 200 19 05:33 AprW 400 2012 19 17:26 AprW 200 2012 19 17:55 AprW 200 1350 2012 20 03:14 AprW 200 1350 2012 20 05:18 AprW 200 1350 2012 23 05:50 AprW 200 1350 2012 23 17:32 AprW 200 1350 2012 23 18:00 AprW 400 1300 2012 23 17:48 Apr 200 1300 2012 23 18:18 Apr 200 1300 2012 28 01:58 Apr 1300 2012 28 05:07 Apr 1300 2012 29 05:38 Apr 1350 2012 29 Apr 1350 2012 29 Apr 1350 2012 1350 1350 WWW 150W 200 200 17:44 200 11:40 22 Aug 13:33 2011 25 Aug 15:45 2011 12 Sep 2011 150 22 Sep 2011 200 334 200 ◦ ◦ 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0.01 0.00 0.00 0.00 0.00 0.00 0.06 0.00 0.01 0.01 0.01 0.01 0.04 0.08 ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ 59.99 59.99 59.99 59.99 59.99 30.00 30.11 29.98 29.98 30.16 30.11 30.05 30.05 30.03 30.03 30.04 30.00 30.01 59.97 59.97 59.98 59.98 59.98 45.18 91.23 50.82 36.00 ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ NN 10 N 10 N 10 10 NN 10 N 11 N 11 N 11 N 2 N 2 N 2 N 2 N 2 N 2 N 2 N 2 NN 1 N 1 N 1 N 1 N 1 N 1 11 N 11 N 11 N 11 N 11 N 2 N 2 N 2 N 2 N 2 N 10 10 NN 10 N 10 10 N 65 NN 65 N 62 68 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 30.00 30.00 30.00 30.00 30.01 29.95 29.99 29.99 50.00 49.99 50.01 50.01 50.04 50.01 50.01 50.01 20.00 20.00 20.01 20.01 20.01 20.01 30.00 29.99 29.99 30.14 30.33 50.00 50.00 50.00 50.00 50.00 29.99 29.99 29.99 29.99 29.99 08.43 75.95 08.48 04.00 ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ cruise cruise 111 61 1 61 1 61 1 61 1 61 1 61 2 61 2 61 2 62 2 62 2 62 2 62 2 62 2 62 3 62 3 62 3 60 3 60 3 60 3 60 1 60 1 60 1 61 1 61 1 61 2 61 2 61 2 62 2 62 2 62 1 62 1 62 1 61 1 61 1 61 61 61 1 66 234 65 67 63 Meteor Station LatitudeArctic Longitude Sampling Time Date Bottom . List of stations in which jellyfish were collected using the Jellynet. Main sampling in- Sampling gears used to collect jellyfish records in di cruise 3–20 May 2013 Bergen–Reykjavik–Nuuk 59–68 cruise 15–25 May 2012 Iceland 241, 246, 248, 255, cruise 9–29 Apr 2012 North of Scotland 60–62 cruise 22 Aug–22 Sep 2011 Cumberland Peninsula 63–67 G.O. Sars IEOAZTI Mar–Nov 2010 May 2010 Gulf of Cadiz Bay of Biscay 36 45 DatasetArctic Meteor DatesIcelandic Area Lat Long Stations Gear Mesh size Mouth formation is also indicated. Licandroand and Kennedy Blackett (2014), (2014), Licandro Licandro and and Raab Hosia (2014), (2014), Licandro Licandro et al. (2014) Table 2. Table 1. Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | erent collection gears. Main ff 650 649 depth (m) (start, LT) depth (m) E 200 22:30 3 May 2013 212 WWW 200W 200W 200 15:45W 200 08:40W 200 5 19:50W 200 May 2013 6 12:00W 200 May 2013 7 23:01W 200 May 2013 2912 8 11:10W 200 May 2013 3200 8 09:20W 200 May 2013 3693 9 11:50W 200 May 2013 1783 10 May 02:30W 2013 200 11 NA May 00:40W 2013 200 1498 12 May 06:40 1756 2013 200 14 May 19:25 1376 2013 200 15 May 16:25 1098 2013 15 May 19:35 2013 224 16 May 14:50 2013 315 17 May 1439 2013 20 May 2829 2013 2860 1100 WWW 400W 200W 400 04:45W 200 21:29W 200 16 May 01:36W 2012 200 16 May 22:22W 2012 200 17 May 23:32 1018 W 2012 200 17 May 15:24W 2012 200 217 18 May 17:57W 2013 200 450 19 May 19:57W 2012 200 990 19 May 14:08W 2012 200 698 19 May 22:59 1271 W 2012 200 20 May 04:10W 2012 200 963 21 May 23:51W 2012 200 805 22 May 22:49 1540 W 2012 400 22 May 02:28 1082 W 2012 200 23 May 02:18W 2012 200 261 24 May 03:57W 2012 200 537 25 May 02:07 1125 W 2012 200 26 May 19:36 2012 200 210 26 May 00:17 1079 2012 200 26 May 02:54 2012 990 27 May 15:42 2012 990 27 May 20:35 2012 896 27 May 1253 2012 27 May 2012 707 620 463 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 4.23 8.61 5.90 ◦ depths (m) (start, LT) ◦ ◦ 51.29 25.04 41.12 24.14 54.45 29.71 10.16 21.56 10.18 20.62 31.04 27.64 11.59 58.86 59.73 47.73 56.66 52.16 14.88 15.32 15.21 34.41 05.66 35.04 17.33 40.52 07.97 54.72 29.99 29.99 04.65 47.22 13.97 37.37 50.49 ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ E 0 : 25 : 50 : 100 18:50 3 May 2013 E 0 : 25 : 50 : 100 : 200 : 400 : 600 : 800 : 1000 19:01 4 May 2013 WW 290–310 0–700WW 0–1028W 500W 0–1000W 40–70W 30–38 400–420WW 0–1000W 120–150W 460W 250W 490–500 0–1000W 17:24W 695–705W 510–520 15:45W 320–330 01:58 05 May 2013 W 630–660W 170–200 05 11:44 May 2013 18:39 07 May 380W 2013 10:49 0–1000 10:52 08 13:08 May 07 2013 May 2013 09 11:35 May 2013 06:24 10 10 May May 2013 2013 11 May 2013 15 May 13:25 2013 16:31 15:16 20:31 22:14 15 16 May May 2013 2013 19:05 15 May 2013 16 20:55 May 2013 16 May 2013 23:40 17 May 2013 14:13 17 May 2013 17 May 2013 16:55 20 May 15:33 2013 20 May 20 2013 May 2013 WW 200 : 400 : 600 :W 50 800 : : 100 1000 : 200 :W 0 400 : : 25 600 : : 50/200 800 :W 0 : 400 : 1000 : 25 600 : : 50 800 :W 0 : 100 : 1000 : 25 200/400 : : 50 600 : 0W : 100 : 800 : 25 : 200 : 100 : 50 400 :W 0 : 100 : 600 : 25 : 200 : 800 : 50 : 04:22 400 : 1000W 0 : 100 : 600 : 25 : 05:59 200 : 06:27 800 : 50 : 300 : 1000 08:31W 0 100 : : 25 200/ : 05:12 50 : 06:18W 0 100 : : 25 200 : : 50 7 400/600 : May 50 : 100 : 2013 800 : 9 100 : 10 200 8 : May 1000 May : 200 May 2013 400/800 2013 : 2013 : 400 1000 : 11 600 5 May : May 800 2013 10:02 2013 : 1000 16 12:37 May 2013 03:22 18:34 17 May 2013 18:33 15 May 2013 20 May 2013 15 May 2013 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 NNN 5 0 N 3 N 3 N 7 N 8 N 9 N 12 N 15 18 NN 24 N 25 N 28 N 32 N 36 46 N 28 NN 25 N 26 N 23 N 18 16 NN 16 N 16 N 13 N 12 N 13 11 NN 13 N 14 N 19 N 21 N 21 23 NN 23 N 24 N 25 24 4.23 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 54.6 3.49 8.65 9.44 ◦ 33.39 ◦ ◦ ◦ ◦ 48.44 54.43 53.58 51.16 57.89 39.26 56.20 24.08 37.58 54.33 55.85 16.67 25.60 48.78 58.74 23.12 29.94 34.73 51.29 59.06 38.86 45.32 29.88 32.21 11.62 41.45 53.51 39.50 ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ 3.33 3.28 8.94 1.22 ◦ ◦ ◦ ◦ 25.00 45.86 40.10 40.30 29.59 33.80 47.65 29.98 18.37 32.05 32.71 31.13 22.83 20.36 50.23 35.06 44.11 00.11 44.83 29.91 14.79 21.49 21.73 00.11 39.98 52.11 07.23 58.09 58.09 03.38 43.05 51.57 17.02 38.81 ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ cruise NN 0 N 0 N 2 N 3 N 7 N 9 N 12 12 NN 15 N 25 N 27 N 28 N 32 33 NN 33 N 36 N 36 N 37 N 46 46 N 46 N 5 NN 1 N 0 N 2 N 7 N 9 N 12 N 15 N 25 N 28 N 32 N 36 46 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 8.4 ◦ 9.30 4.08 0.77 3.33 1.39 8.86 ◦ ◦ ◦ ◦ ◦ ◦ 15.82 39.70 50.63 43.66 36.33 40.12 11.49 29.41 56.56 54.55 50.58 49.10 51.58 51.36 51.37 38.80 40.64 43.89 25.00 40.72 39.52 33.83 32.09 52.75 56.90 54.61 46.97 ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ 152155 62 157159 65 65 160 65 66 161162163 67 67 165166 68 68 167 63 168 63 169 62 170 61 171 60 59 241 64 246248 65 255267 66 67 272 66 273 68 274 67 281 67 290 67 292 66 299 66 305 65 307 63 315 63 324 63 324 62 330 62 332 63 333 62 338 62 340 63 63 cruise 160bis 66 G.O. Sars Station LatitudeIcelandic Longitude Sampling Time Date Bottom Macroplankton trawl 101102104 65 105 65 106 65 107 65 108 66 109 67 111 67 115 67 116 68 117 63 118 63 120 62 121 61 122 61 123 61 124 60 125 60 126 60 127 59 59 59 Station LatitudeG.O. Sars Mocness Longitude152 Sampling154 62 64 Time Date 155157160 65 161 65 162 66 163 67 167 67 168 68 169 63 170 62 171 61 60 59 List of stations in which jellyfish were collected using di Continued. sampling information is also indicated.dro Licandro et al. (2014a, b), (2014). Licandro and Hosia (2014), Lican- Table 3. Table 2. Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | er- ff E N ◦ ◦ + + + + + + + + W–1 ◦ N 60–62 W 2 ◦ ◦ ∗ ∗ + + + + ++ + + + + + + 241–340 1–3 Icelandic Meteor 62–68 11–28 Day/Night Day/Night Icelandic Sea North of Scotland 16–25 May 2012 9–29 Apr 2012 E N ◦ ◦ + + + + + + + + + + W–5 ◦ cruise N 62–68 W 18 ◦ ◦ + + + + depths (m) (start, LT) G.O. Sars 652 651 WW 220 206 12:30 18:51 22 May 2010 23 May 2010 WW 29W 22 16W 21 20:05 03:28 01:18 19:22 4 Mar 2010 6 Mar 2010 7 Mar 2010 26 Jul 2010 WW 208 209 11:43 02:34 24 May 2010 24 May 2010 WW 16W 40W 27 28 21:34 21:24 03:00 27 02:18 Jul 2010 9 Nov 2010 11 Nov 2010 12 Nov 2010 0 0 0 0 0 0 0 0 0 0 0 0 N 60–63 W 36–24 ◦ ◦ + + + + 171 166–170 152–165 Day Day/Night Day/Night 0.96 2.52 59 46 ◦ ◦ 15.15 15.04 44.72 15.18 16.44 29.76 16.44 32.46 36.78 18.06 ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ 20 May 2013 14–17 May 2013 3–12 May 2013 Labrador Sea Irminger Sea Norwegian/Icelandic Seas taxon found only in samples collected at 0–400 m depth. Licandro NN 5 5 NN 5 5 NN 6 N 6 6 NN 6 N 6 N 6 N 6 6 N W 0 0 0 0 0 0 0 0 0 0 0 0 = ◦ ◦ ∗ + + + 45 45 Arctic ◦ ◦ 8.76 8.52 ◦ ◦ 45.02 14.97 39.96 24.72 22.26 44.70 22.26 43.08 ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ spp. larva 69 45 Station LatitudeIEO dataset Longitude Sampling Time Date AZTI dataset Bongo net 586768 45 43 45 Bongo net TF-01 36 GD-02SP-01 36 TF-01 36 36 SP-01GD-01 36 SP-01 36 GD-02 36 36 Continued. Jellynet dataset. List of jellyfish taxa collected in epipelagic waters (0–200 m) in di spp. spp. spp. CNIDARIA Obelia Family Physophoridae Lensia Muggiaea atlantica Chuniphyes multidentata Pantachogon haeckeli Pantachogon Order NARCOMEDUSAE Family Aeginidae Aeginopsis laurentii Order LEPTOTHECATA Family Phialellidae Phialella quadrata Family Mitrocomidae Cosmetira pilosella Mitrocomella polydiademata Family Tiarannidae Modeeria rotunda Family Tiaropsidae Tiaropsis multicirrata Family Campanulariidae Clytia islandica Clytia Order SIPHONOPHORA Suborder Physonectae Physonectae Family Agalmatidae Agalma elegans Nanomia cara Physophora hydrostatica Suborder Calycophorae Family Diphyidae Dimophyes arctica Lensia achilles Lensia conoidea Family Clausophyidae Aglantha digitale Family Rhopalonematidae Order TRACHYMEDUSAE HYDROZOA Date 22 Aug–22 Sep 2011 Latitude 63–67 North Atlantic region Cumberland shelf StationsCruise 1–4 LongitudeTime 62–68 Day/Night Table 4. ent North Atlantic regions. et al. (2014). Table 3. Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | E N ◦ ◦ + + + + + + W–1 ◦ N 60–62 W 2 ◦ ◦ + ++ + + 241–340 1–3 Icelandic Meteor 62–68 11–28 Day/Night Day/Night Icelandic Sea North of Scotland 16–25 May 2012 9–29 Apr 2012 N ◦ W + + + ◦ E N ◦ ◦ + W–5 ◦ N 43–45 W 5 ◦ cruise + ++ + + + + + + + + ++ ++ + + + + ◦ N 62–68 W 18 ◦ ◦ +++ + + + G.O. Sars 654 653 N 60–63 W 36–24 ◦ ◦ + + + + 171 166–170 152–165 Day Day/Night Day/Night 59 46 larva 20 May 2013 14–17 May 2013 3–12 May 2013 Labrador Sea Irminger Sea Norwegian/Icelandic Seas spp. spp. spp. spp. CTENOPHORA CNIDARIA N W ◦ ◦ + + Order Cydippida Family Pleurobrachiidae Hormiphora North Atlantic regionLatitude Gulf of CadizHYDROZOA BayOrder of TRACHYMEDUSAE Biscay Family Geryoniidae Liriope tetraphylla 36 LongitudeFamily Rhopalonematidae Aglaura hemistoma 6 Aglantha digitale Order LEPTOTHECATA Family Lovenellidae Eucheilota paradoxica Family Campanulariidae Clytia hemisphaerica Clytia Obelia Order SIPHONOPHORAE Suborder Physonectae Physonectae Family Agalmatidae Agalma elegans Suborder Calycophorae Family Abylidae Abylopsis tetragona Bassia bassensis Family Diphyidae Chelophyes appendiculata Eudoxoides spiralis Lensia conoidea Muggiaea atlantica Muggiaea kochi Order ANTHOATHECATA Family Coryniidae Corynidae Maximum sampling depth (m)TimeMonth 16–40 206–220 Mar, Jul, Nov 2010 Day/Night May 2010 Day/Night Arctic spp. incerta sedis larva Continued. Bongonet dataset. List of jellyfish taxa collected in epipelagic waters (0–200 m or 0 m- spp. spp. CTENOPHORA North Atlantic region Cumberland shelf LatitudeTimeDateOrder ANTHOATHECATA 63–67 22 Aug–22 Sep 2011 Day/Night Cruise Longitude 62–68 StationsFamily Corymorphidae 1–4 Euphysa aurata Aplanulata Plotocnide borealis Family Rathkeidae Rathkea octopunctata Lizzia blondina Family Pandeidae Amphinema rugosum Family Zancleidae Zanclea Order Cydippida Cydippida Family Mertensiidae Mertensia ovum Mertensiidae Order Beroida Family Beroidae Beroe cucumis Beroe gracilis Beroe Bolinopsis infundibulum bottom) in 2010, in the Gulf of Cadiz and Bay of Biscay. Licandro et al. (2014). Table 5. Table 4. Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | E E N ◦ N ◦ ◦ ◦ + + + + + + + + + + + + + + W–5 W–5 ◦ ◦ cruise cruise N 62–68 N 62–68 W 18 W 18 ◦ ◦ ◦ ◦ ++ + + + + G.O. Sars G.O. Sars 656 655 N 60–63 N 60–63 W 36–24 W 36–24 ◦ ◦ ◦ ◦ +++ + + ++++++ + + + + + + + + + ++ ++ + + + + + + + + + +++++ + + + + + + + + ++ + + + spp. spp. spp. spp. spp. spp. spp. CTENOPHORA CNIDARIA , Mocness dataset. List of jellyfish taxa collected in the 0–1000 m layer, in Family Euplokamidae Euplokamis Order Lobata Family Bolinopsidae Bolinopsis infundibulum Order Beroida Family Beroidae Beroe abyssicola Beroe cucumis Order Cydippida Unidentified Cydippid Family Mertensiidae Mertensia ovum Mertensiidae LatitudeLongitude 59 46 North Atlantic regionStationsCruise Labrador Sea Irminger SeaLensia Norwegian/Icelandic achilles Seas Lensia conoidea Lensia hunter Muggiaea bargmannea 171Family Clausophyidae Chuniphyes multidentata Crystallophyes amygdalina Heteropyramis crystallina 166–170Family Sphaeronectidae Sphaeronectes Order ANTHOATHECATA Family Hydractiniidae Hydractinia areolata Family Tubulariidae 152–165 Hybocodon SCYPHOZOA Family Atollidae Atolla parva Atolla wyvillei Family Periphyllidae Periphylla periphylla TimeDate 20 May 2013 Day 14–17 May 2013 Day/Night 3–12 May 2013 Day/Night Order NARCOMEDUSAE Family Aeginidae Aeginura grimaldii Family Cuninidae Solmissus incisa Order LEPTOTHECATA Family Mitrocomidae Halopsis ocellata Rhopalonematidae Mitrocomella polydiademata Family Tiarannidae Chromatonema rubrum Obelia Order SIPHONOPHORAE Suborder Physonectae Family Agalmatidae Marrus orthocanna Nanomia cara Halicreas minimum Halicreatidae Family Rhopalonematidae Aglantha digitale Crossota rufobrunnea Pantachogon haeckeli Sminthea arctica Family Campanulariidae Clytia islandica Suborder Calycophorae Family Hippopodiidae Vogtia serrata Family Diphyidae Dimophyes arctica Gilia reticulata North Atlantic regionStationsCruise Latitude Labrador Sea Irminger Sea Norwegian/Icelandic Seas HYDROZOA Order TRACHYMEDUSAE Family Halicreatidae 171Botrynema brucei 59 166–170 152–165 Longitude 46 TimeDate 20 May 2013 Day 14–17 May 2013 Day/Night 3–12 May 2013 Day/Night G.O. Sars Continued. erent North Atlantic regions. Licandro et al. (2014). ff Table 6. Table 6. di Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | N N W W ◦ ◦ ◦ ◦ + + + + + cruise cruise N 65–68 N 65–68 W 15–01 W 15–01 ◦ ◦ ◦ ◦ + + + + + + + + +++ + + + ++ + G.O. Sars G.O. Sars 658 657 N 60–63 N 60–63 W 36–25 W 36–25 ◦ ◦ ◦ ◦ +++++ + + + + ++ + + + + + + + + + + +++ + + + + ++ + + + + + spp. eni ff spp. erent North Atlantic regions. Licandro et al. (2014). ff spp. sp. CTENOPHORA , Harstad and Macroplankton dataset. List of jellyfish taxa collected in the CNIDARIA Order Cydippida Family Mertensiidae Mertensia ovum Family Hippopodiidae Vogtia glabra Vogtia serrata Family Diphyidae Dimophyes arctica Lensia conoidea Nectodamas diomedeae Family Clausophyidae Chuniphyes multidentata Order ANTHOATHECATA Family Bythotiaridae Bythotiara murrayi SCYPHOZOA Family Atollidae Atolla chuni Atolla parva Atolla vanhoe Atolla wyvillei Atolla Family Periphyllidae Periphylla periphylla Family Pelagiidae Pelagia noctiluca Order Beroida Family Beroidae Beroe abyssicola Beroe cucumis Beroe gracilis Beroe North Atlantic regionStationsCruise Labrador Sea Irminger Sea Norwegian/Icelandic Seas 125–127 115–124 101–111 Latitude 59 Longitude 46 TimeDate 20 May 2013 Day 15–17 May 2013 Day/Night 5–11 May 2013 Day/Night Longitude 46 North Atlantic regionStationsCruise Labrador SeaTime Irminger SeaDate Norwegian/Icelandic Seas HYDROZOA Order TRACHYMEDUSAE 125–127Family Halicreatidae Halicreas minimum 115–124 20 May 2013 Day 15–17 May 2013 101–111 Day/Night 5–11 May 2013 Day/Night Latitude 59 Halitrephes maasi Halicreatidae Family Rhopalonematidae Aglantha digitale Colobonema sericeum Crossota rufobrunnea Rhopalonematidae Family Cuninidae Solmissus incisa Order LEPTOTHECATA Family Laodiceidae Ptychogena lactea Family Tiarannidae Chromatonema rubrum Order SIPHONOPHORAE Suborder Physonectae Family Agalmatidae Marrus orthocanna Nanomia cara Pantachogon haeckeli Order NARCOMEDUSAE Family Aeginidae Aeginura grimaldii Modeeria rotunda Suborder Calycophorae Family Prayinae Praya dubia G.O. Sars Continued. Table 7. Table 7. 0–1000 m layer, in di Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Scyphozoa Hydrozoa Hydrozoa Scyphozoa Hydrozoa Scyphozoa Hydrozoa Scyphozoa Scyphozoa Scyphozoa Scyphozoa Scyphozoa Scyphozoa Scyphozoa Scyphozoa Scyphozoa Scyphozoa Scyphozoa Scyphozoa Scyphozoa Hydrozoa Hydrozoa ∗ sample identified by visual sp. Hydrozoa = spp. Hydrozoa ∗ sp.sp. Scyphozoa Scyphozoa sp. Scyphozoa sp. Scyphozoa number 16S Pelagia noctiluca Pelagia noctiluca Pelagia noctiluca Pelagia noctiluca Pelagia noctiluca Pelagia noctiluca Cyanea Cyanea Pelagia noctiluca Aglantha digitale Agalma elegans Halistemma Apolemia Cyanea Cyanea Pelagia noctiluca Halistemma rubrum Muggiaea atlantica Pelagia noctiluca Diphyes dispar Pelagia noctiluca Apolemia uvaria Pelagia noctiluca Pelagia noctiluca Pelagia noctiluca Pelagia noctiluca Pelagia noctiluca Pelagia noctiluca NE Atlantic KJ866185 NE AtlanticNE Atlantic KJ866187 KJ866197 NE AtlanticNE AtlanticNE AtlanticNE Atlantic KJ866203 KJ866186 KJ866204 KJ866206 NE AtlanticNE AtlanticNE Atlantic KJ866200 NE Atlantic KJ866193 KJ866201 NE AtlanticNE Atlantic KJ866199 KJ866202 KJ866194 NW AtlanticNW AtlanticNW Atlantic KJ866189 KJ866190 KJ866191 NW Atlantic KJ866188 North Atlantic KJ866195 North Atlantic KJ866184 North Atlantic KJ866183 660 659 E Aug 2006 E Aug 2006 WW Mar Sep 2007 2007 Agalmatidae Hydrozoa W Sep 2007 W Oct 2007 W Oct 2007 W Oct 2007 W Oct 2007 W Nov 2007 W Dec 2007 W Jul 2008 W Jul 2008 W Jul 2008 W Aug 2008 W Aug 2008 W Nov 2008 W Nov 2008 W Dec 2008 W Dec 2009 W Dec 2009 W Mar 2010 WW JulW Jul Nov 2010 2010 2010 WW Oct Jan 2011 2012 W AugW Oct 2012 2012 0 0 sp. NE Atlantic KJ866196 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 spp. NW Atlantic KJ866192 90 48 66 80 57 01 59 59 59 59 60 12 55 41 45 18 03 10 47 04 48 34 22 07 08 03 28 14 08 ◦ ◦ spp. NE Atlantic KJ866198 ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ sp. NE Atlantic KJ866205 erent North Atlantic regions. Taxa identifiedHYDROZOA Order TRACHYMEDUSAE Family Halicreatidae Botrynema brucei Family Rhopalonematidae RegionCrossota rufobrunnea Pantachogon haeckelii Pantachogon Genbank accession Sminthea arctica Order NARCOMEDUSAE Family Aeginidae Aeginura grimaldii Family Cuninidae Solmissus Order LEPTOTHECATA Suborder Conica Family Laodiceidae Ptychogena lactea Family Mitrocomidae Mitrocomella polydiademata Suborder Proboscoida Family Campanulariidae Clytia islandica Order SIPHONOPHORAE Suborder Physonectae Family Agalmatidae Halistemma rubrum Marrus orthocanna Nanomia cara Nanomia cara Suborder Calycophorae Family Hippopodiidae Vogtia glabra Family Diphyidae Dimophyes arctica Gilia reticulata Lensia achilles Lensia conoidea Lensia Muggiaea bargmannea Family Clausophyidae Chuniphyes multidentata Heteropyramis crystallina Heteropyramis ff N 1 N 2 NN 41 18 N 25 N 4 N 16 N 16 N 16 N 1 N 1 N 18 N 15 N 18 N 25 N 25 N 4 N 4 N 21 N 25 N 6 N 29 N 51 NN 51 5 NN 9 66 N 38 N 4 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 05 18 83 63 50 37 48 48 48 29 31 72 90 70 14 16 45 60 47 10 01 46 59 00 50 12 03 51 57 ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ ◦ DNA sequences (mt16S rDNA) identified from cnidarian taxa collected during the Identity of cnidarian tissues collected from CPR samples and identified based upon CPR tows Latitude Longitude Month Year Taxa identified Class 330M 58 330M 58 535ZB438BB 49 45 438BC 43 3030PR 49 460W 54 460W 54 460W 54 707A 58 708A 58 464W 54 464W 54 464W 54 80FA 54 80FA 54 571SA 45 571SA 45 83FA 54 465BC 47 748V 60 468BC 45 349EA 45 349EA342PR 46 48 488BA373EB 49 42 499BD 42 364PR 49 Table 9. project EUROBASIN in di Table 8. mt16S rDNA analysis. Sampling information are also indicated. inspection. ! 14

Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | ! 15 and a ! cruise ( 10 Arctic ! ! ) and relative propor- 3 0 − ). Licandro and Blackett f teor!cruise!(B!and!E)!and!G.O.!Sars! Arc@c$cruise$ Meteor$cruise$ and c -10 cruise ( -20 -30 CPR$routes$ G.O. Sars 662 661 Icelandic$cruise$ Gulf$of$Cadiz$(IEO)$ -40 ) and e ! and -50 b erent jellyfish datasets presented in this study. ff $cruise$ -60 cruise ( ! Sars ( -70 Meteor Bay$of$Biscay$(AZTI)$ G.O.$ Jellynet!datasets.!Total!jellyfish!abundance!and!relative!proportion!of!Cnidaria!and!Ctenophora!in! ! and -80 Locations of the di Jellynet datasets. Total jellyfish abundance (individuals 100 m 60 50 40 70 ! cruise!(C!and!F).! ! Figure"2. the!stations!sampled!during!the!Arctic!cruise!(A!and!D),!Icelandic!and!Me Icelandic ), cations!of!the!different!jellyfish!datasets!presented!in!this!study. Figure 2. tion of Cnidaria and Ctenophora counts in the stations sampled during the d Figure 1. (2014), Licandro and Hosia (2014), Licandro and Kennedy (2014), Licandro and Raab (2014). Lo .! ! ! Figure"1 Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | ! ! samples 16 = ) in the stations 3 − . Licandro (2014a, b) erent depths in the 0–1000 m layer. (b) ff 664 663 and in the Bay of Biscay ! (a) ( .!Bongonet!datasets.!Total!jellyfish!abundance!in!the!stations!sampled!in!the!Gulf!of!Cadiz!(A)!and! Bongonet datasets. Total jellyfish abundance (individuals 100 m Mocness dataset. Abundance of jellyfish at di ! Figure"3 in!the!Bay!of!Biscay!(B).! ! preserved in formalin, not yet analyzed. Figure 3. Please note the shallower depths in Stns. 152 and 167. St. 155 is not shown. M Figure 4. sampled in the Gulf of Cadiz Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | ! 19 ! samples = ! 2012. > erent depths in the 0–1000 m ff 666 665 ! .!Jellyfish!swarms!recorded!by!the!Continuous!Plankton!Recorder!in!2009 Figure'6 Mocness dataset. Number of jellyfish taxa found at di Jellyfish swarms recorded by the Continuous Plankton Recorder in 2009–2012. ! ! Figure 6. layer. Please note the shallower depths inpreserved Stns. in 152 formalin, and not 167. St. yet 155 analyzed. is not shown. M Figure 5. Discussion Paper | Discussion Paper | Discussion Paper | Discussion Paper | 667 Jellyfish blooming species identified by genetic analysis from jellyfish material col- Figure 7. lected in CPR samples.also The shown. mean frequency of jellyfish presence recorded in 2000–2009 is