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The Benthic Infauna of the North Sea: Species Distribution and Assemblages

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The Benthic Infauna of the North Sea: Species Distribution and Assemblages ICES]. mar. Sci.,49: 127-143. 1992 The benthic infauna of the North Sea: species distribution and assemblages A. Kiinitzer, D. Basford, J. A. Craeymeersch, J. M. Dewarumez, J. Dorjes, G. C.A. Duineveld, A. Eleftheriou, C. Heip, P. Herman, P. Kingston, U. Niermann, E. Rachor, H. Rumohr, and P. A. J. de Wilde D.. I. 1..~ Kiinitzer.. ~~.A.. Basford. , . Craevmeersch.~2~~~~~~~ ~. A.. Dewarumez.~~~~~~~~, 1. M.. Diiries.,~~, , Duineveld, G. C. A,, Eleftheriou,A., Heip, C., ~ermin,P., Kingston, P., ~,ermann.U., Rachor, H., Rumahr, H., and de Wilde, P. A. 1.1992. The benthic infauna ofthe North Sea: species distribution and assemblages.lCES I. mar. Sci., 49: 127-143 In 1986 participants of the Benthos Ecology Working Group of ICES conducted a synopticmapping ofthe infauna ofthe southernand central North Sea. Together witha maooine.. ofthe infauna ofthe northern North Sea bv Eleftheriou and Basford (1989) this nrnvides thedatabase for thedescrintionofthe benthicinfaunaofthe whole~orth~eain Assemblages were further separated by the 30,50 m and 100 m depth contour as well as by thesediment type. In addition to widely distributed species, cold water speciesdo not occur further south than thenorthernedgeoftheDogger Bank,whichcorrespands to the 50m depth contour. Warm water species were not found north of the 100m depth contour. Some species occur on all types of sediment but most are restricted to a special sediment and therefore these soecies are limited in their distribution. The factors struc- (tirlng ,p,..~:i clslnhufion\ 2nJ a,semhl.tge~ <:em ld h: rr.nlprrdtL.rc. thc ~rlrlurn;~ .,I d~ticrcnln.4tr.r nur\cr. c &:. :\llaiit~cu.dcr thc t\pr. ol cr.iimznl anJ the id01 <uppi) 12 rhc h~,ntlid\ tlr.cthcrtt,u, ,\ ,rn.l H.~riorJ. I) J Juurrl;jl .,ilhr \Idr#n< H~olocl..~l Association of the UK, 69: 123-143 Key words: benthos, North Sea, assemblages Received 21 March 1991; accepted 9 January 1992 A. KCnitrer: Alfred- Wegener-tnrtitul fir Polar- undMeertiforrchu,zg. Columbusstra&, D-2850 Brfmerhovm, Germany. Present address: Umwellburzdesaml, Bismarckplofz I, D- 1000 Berlin 33, Germany. D. Bosford: Marine Laboratory, P.O. Box 101, Victoria Road, Aberdeen AB9 808,Seotland. J. A. Croeymeersch: Delta Inslituul voor Hydrobiologisch Onderzoek, Vierstran1 28, NL-4401 EA Yerseke, The Netherlands. J. MM. Dewarumez: Stalion Marine, BP 41 Wimereux, F-62930, France. J. Ddrjes: lnslitut fir Meererhiologie und Meerergeologie "Senkenberg", Schleusenstr. 39A, D-2940 Wilhelmshaven, Grrmony. G. C. A. Duineveld & P. A. J. de Wilde: Nederlands Insriruur voor Onderroek der Zee, PO.Box 59. NL-1790 AB Den BurglTerel, The Nerherlonds. A. Elejiherioli: Marine Lnboralory, P.O. Box 101, Victoria Rood, AberdeenAB9 8DB. Seotland. Presenr address: Depcrtment ofBiology, U~iversilyof Crete, P.O. Box 1470, Heraklion 71110, Crere, Greece. C. Heip: Insticut voor Dierkunde. Ledegoncksrroar 35, 8-9000 Gent, Belgium. Present addrers: Della lnsliruul voor Hydrobiologirch Onderzoek. Vier.slraa1 28, NL-4401 EA Yerseke, The Netherlands. P. Herman: Delta Insti~uulvoor Hydrobiologisch Onderzoek, Yier~rroot28, NL-4401 EA Yerseke. The Netherlands. P, Kingston: Heriot- Walt Univer,~iry,Insrilule of Offshore Engine@*. Research Park Ricearton, Edinburgh EH14 4AS. Scotland U. Nirrmann: Biologiscke Anstan Helgoland, Notkestr. 31, 0-2000 Hamburg 52. Germany. E. Raehor Alfred-Wegener-lnstifuf fir Polar- und Meeresforschung, Columbuss~raJe,0.2850 Bremerhauen, Germany. H. Rumohr: Insfilur fsr Meereskurrde, Diisternhrooker Weg 20.0-2300 Kiel. Germany. Introduction describe and quantify areas with similar fish food. During the following 20 years several areas in the North Sea were At the beginning of this century Petersen (1914) devel- investigated for their benthic infanna which were classi- oped the concept of infaunal communities in order to fied into certain communities or assemblages. Most of 128 A. Kunitzer et al. Figure I. Stations sampled by participants of the North Sea Benthos Survey (NSBS) in 1986. Stations in the northern North Sea (Aberdeen) were sampled between 1980 and 1985. these surveys were carried out in coastal areas, not Sea from 60 to 100 m depth; and the northern North Sea further north than the Dogger Bank area. A review of the from 100 to 200 m depth. The assemblages of these zones infaunal assemblages of the North Sea was given by are further structured by sediment composition. Before Kingston and Rachor (1982), showing the low number of the North Sea Benthos Survey (NSBS) it was not possible benthic surveys in the central and northern North Sea. to confirm or reject Glemarkc's concept of the zones in the Investigations in the vicinity of oil platforms suggested North Sea. that infauna assemhlagesnorthofthe Dogger Bankmight Since in most earlier surveys sampling of bottom fauna be similar to those south of it. On the other hand, was done with different gears, and surveys were spread Glemarkc (1973) developed a concept of three different over more than half a century, during which major zones of benthic assemblages along the European North changes occurred in the benthic fauna (Kroncke, 1990), Atlantic Continental shelf, reflectingdifferencesin annual the results of these surveys are not comparable. Partici- variation of temperaturein bottom waters. He divided the pants of the Benthos Ecology Working Group of ICES North Sea into three zones: the southern North Sea up to therefore decided to map the henthic fauna of the whole the northern edge of the Dogger Bank; the central North North Sea during the same period of the year and with Benthic infauna of the North Sea Figure 2. Classification of stations by TWINSPAN, using only species presence/absencedata. 1st dlshotomy depth < 70m depth > 70m 2nd dichotomy 1 2 3 4 depth mu* <30m 30-70m 70-loom mainly >lorn deposit coarser sediment finer sediment liner sediment 3rd dichotomy It 1 b 2a 26 3 a 3b 4a 4 b depth 30.70m 30-50m > loom depo~~ fine sand muldy finesand fine sedimenf coarse sedimem Figure 3. Scheme of TWINSPAN classification (species presence/absence data), showing the environmental parameters of the assemblages. Figure 4. Classification of stations by TWINSPAN, incorporating species abundances. standardized gear. This survey, the North Sea Benthos far, only the distribution of echinoderms (Ursin, 1960), Survey (NSBS), was carried out during April 1986. The polychaetes (Kirkegaard, 1969) and bivalves (Petersen, area north of 58"Ncouldnot be sampledduring theNSBS 1977) has been described for the central and southern but the benthic fauna of that area had been sampled North Sea. Almost nothing is known about the dis- directly between 1980 and 1985 by Eleftheriou and tribution of crustaceans, such as amphipods and Basford (1989) with the same gear, and these data have cumaceans. The above mentioned authors found that the been included in the analysis. The combined data of the fauna of the North Sea was a mixture of arctic-boreal NSBS and Eleftheriou and Basford (1989) provide infor- (North Atlantic-North Pacific elements) and lusitanean- mation on theinfauna of the whole North Sea and should boreal species (Tethys elements), the first being dominant act as a basis for assessing natural and anthropogenic north of the Dogger Bank, the latter being important changes in the benthos. south of it. This pattern of distribution was demonstrated Besides the classification of the infauna into assem- for echinoderms and bivalves, the distribution of which blages, the NSBS enables us to describe the distrihution was related to temperature and water bodies, but was less of single species within the whole North Sea area. So clear for polychaetes, which seemed to be related more to Benthic infauna of the North Sea 1st dichotomy depth < 70m depth > 70m 2nd dichotomy I I1 111 Iv depth manly <30m 30-70m <loom deposit CO-ser sediment finer sediment fins sedment coar~ersediment 3rd dlshotomy la I b 118 Ilb llla depth 30-5Om 50.70m 70.100m deposit muday hnesand line sand indicator specles ~ephf~rc ~onioesp ~uculao. Opnelfa D. MlnoSDio c Ophe!t* 0. PmfldowYia C E~~nocaroiumcPfiomcephaius h Ca!fieoassa s NBphrys ,. rnyaaiia so. fmgone n. Umdioep PMOne r EUdoieNa i /viabe8 c sprophaner b. Exogone v. POWCiCiS SP number of staions 52 19 40 61 46 41 12 2 number of species 2718 29f9 44i9 43i10 54f16 51113 44f12 np(H') 12f4 1415 14-5 28r5 24+10 25*7 23+5 1nam2 805 + 728 873+ 623 1995 + 1499 1093 f 686 1224 1 1233 2863i18M 1775f1114 bianass (g AFDW.m.9 9.5+9.9 4354.3 12.61-75 7616.5 7.4f7.0 3.5? 3.7 3.8 f 2.2 .. ....... southern North Sea --------- --- central North Sea --- .. ... .... northern North Sea --------- Figure 5. Scheme of TWINSPAN classification (species abundance data), showing the environmental and biotic parameters (meanis.d.)ofthc assemblages and the indicator species. the type of sediment. The present paper tries to combine 1988) where invited experts on certain taxonomic groups the main pattern of species distributions with the checked the identifications of species, the species distribution of infaunal assemblages in the North Sea. list contained a lot of synonyms (e.g. Paramphinome jeffreysi P. pulchella, Pectinaria uurrcoma = Amphictene auricoma). Spelling and synonyms were checked usmg the Materials and methods Marine Conservatton Societv soecies directow.,. a coded checklist of the marine fauna and flora of the British Isles Sampling and its surrounding seas (Howson, 1987). The directory includes most of the species recorded during the NSBS. Species which do not really belong to the macrobenthos and hence, were badly sampled (e.g. fishes, larval stages, and Basford (1989) between 1980 and 1985, always in epibenthic species) were removed. During a final work- spring and early summer. The grid is of 30 x 35 nm with shop on Texel in 1989 the identification of all species four stations at the angles of the rectangle and one in the (including those of the northern North Sea) was checked centre.
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