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M A R I N E F A U N A OFF T H E C O A S T OF EAST ANGLIA J. R. ELLIS AND S. I. ROGERS The marine fauna of East Anglia has been relatively well documentcd over thc last Century, and faunal lists exist for many groups (Collings, 1933; Anon, 1934, 1937, 1938; Hamond, 1971; Hughes & Quinn, 1983). The marine ecology of several estuarine and inshore sites has also been studied (Mistakidis, 1951, 1957; Hamond, 1963; Millner et al„ 1977; George et al„ 1995), although the sub-tidal fauna has reeeived little attention (Covey, 1998). Hamond (1969) reported on the sublidal fauna off thc north coast of Norfolk and some studies of the North Sea have included a few stations off thc Suffolk coast (e.g. Davis, 1925; Dyer et al„ 1983). This lack of suitablc information is despite the fact that the coastal zone of Suffolk is used extcnsively for thc extraction of sand and gravel (Rogers, 1997), and supports locally important fisheries for demersal species, including flatfish, edible crab, Cancer pagurus, brown shrimp, Crangon crangon, and other organisms on the sea floor (Pawson & Robson, 1998a,b) (see Plate 3 & Cover). A Community is broadly defined as a group of organisms living in a defincd habitat. In the present study we use the term assemblage in prefercnce to Community as marine habitats are often patchy, a trawl of 30 minutes my pass over several habitat types and the catches themselves are affected by the gear used (Basford et al„ 1990). The aim of the current study was to describc and quantify the invertebrate and fish assemblages present off the coast of East Anglia. The Centre for Environment, Fisheries and Aquaculture Science (CEFAS) undertakes regulär fish surveys in order to provide abundancc indices of commercial fish stocks. Such surveys will also catch a ränge of invertebrates and, during 1998, this invertebrate by-catch has been quantified and the data for 20 stations along the coast of East Anglia recorded. Sampling Methods Data were collected on R.V. Corystes during a research cruise in August 1998 and a total of 20 stations (A-T, Fig. 1) off the coast of East Anglia were sampled. Station details are given in Table 1. All catches were made using a 4-m beam trawl, with 40 mm mesh, as described by Kaiser and Spencer (1994) and tows were of 30 minutes duration, resulting in an approximate sampling area of 15,000 m 2 . All fish were identified, counted and weighed and the total weight of the by-catch recorded. The invertebrate by-catch was completely sorted at nine stations, and randomly sub-sampled on thc remaining 11, duc to either a large catch or time constraints. Invertebrates were identified to the lowest taxon possible, and the biomass (wet weight) of all macro-epibcnthic species recorded. Macro-epibenthic invertebrates being thosc species which are large enough to be sampled by the gear and occur on thc sca floor and not within sediments. Non-colonial species were also enumerated. Catch data for both fish and invertebrates were subsequently raised to catches per hour. These data were used in the analysis of Community stmeture, which was performed using the PRIMER analytical package (Clarke & Warwick, 1994). © British Crown Copyright reserved
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53.0-
Norfolk
52.5 N
sand/gravel
Suffolk
V
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51.5-
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Figure 1: Map of study area showing trawl locations (A-T) and main sediment types. See Barne et al. (1998) for additional information regarding sediment types.
Trans. Suffolk Nat. Soc. 35
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MARINE FAUNA OFF THE COAST OF EAST ANGLIA
Table 1: Locations, dcpths and the catches (per 30 minutcs) of fish and invertebrates at 20 stations off ihe coast of East Anglia Station A B C D E F G H I J K L M N O P
Q R S T
Latitude (°N) 51° 51° 51° 51° 51° 51° 51° 51° 51° 51° 52° 52° 52° 52° 52° 52° 52° 52° 52° 52°
44-1' 43-1' 37-9' 30-4' 27-8' 251' 27-9' 18-2' 15-8' 55-5' 01-5' 04-4' 30-6' 21-8' 17-6' 08-5' 14-3' 15-7' 45-6' 491'
Longitude Depth (m) (°E) 01° 20-9' 12 01° 28-7' 9 01° 31-4' 10-5 01° 22-2' 6 01° 19-9' 13 01° 23-5' 13 01°4M' 13 01° 4 5 1 ' 37 01° 39-3' 50 01° 34-5' 8 01° 30 0' 11 02° 05-3' 18 01° 50-8' 15 01° 45-4' 16 01° 44 0' 18 01° 41-4' 6 01° 40-6' 8-5 01° 45-3' 18 01° 57-9' 17-5 01° 38 1' 16 5
Benthos Wt. of fish, Wt. of benthos, fully kg (% catch) kg (% catch) sampled • 5-7(85) 1-0(15) • 13-7 (77) 4-1 (23) • 7-5 (78) 2-1 (22) 9-3 (37) 16 0 (63) 11-5(9) 115-4 (91) 11-0(1) 730-7 (99) 7-3 (2) 289-4 (98) • 11-9(83) 2-5 (17) 20-5 (15) 120-7 (86) 11-1 (15) 62-0 (85) 4-1 (3) 143-8 (97) 27-3 (39) 42-4 (61) 10-9 (30) 25-6 (70) • 5-6 (74) 2-0 (26) 10-1 (39) 15-7(61) • 0-7 (20) 2-9 (80) • 9-9 (86) 1-6 (14) • 13-9(70) 6-1 (31) • 1-6(55) 1-3 (45) 17-8 (5) 368-0 (95) -
Data analysis The stations were elustered aecording to the biomass of invertebrates, using the Bray-Curtis similarity index on fourth-root transformcd data (which reduces the bias towards dominant species) and stations with a 50% similarity in their catches were identified from the resulting dendrogram. Species that contributed to the observed differences between groups of stations were identified using the similarity of percentage procedure SIMPER (Clarke & Warwick, 1994). The mean species composition, by biomass, for stations with similar catches was used to describc the assemblage structure and to determine the dominant fauna. Similar analyses were thcn undertaken for dcmersal fish and for fish and invertebrates combined.
Results Dominant Fauna Over 75 species were recorded during the survey (Table 2) and were dominated by fish (31 species), crustaceans (18 species) and echinoderms (9 species). The predominance of larger bodied organisms is a refiection of the gear selectivity of the beam trawl. The most frequently recorded species were the swimming crab Liocarcinus holsatus and common starfish Asterias rubens which were
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Table 2: Taxonomic list of fauna PORIFERA Haliclona oculata (Pallas) Porifcra findet.) CNIDARIA Nemerlesia anlennina (L.) Hydrozoa (indel.) Alcyonium digitatum L. Urlicina felina (L.) Metridium senile (L.) POLYCHAETA Aphrodita aculeata (L.) Sabellaria spinulosa Lcuckhart ECHIURA Maxmuelleria lankesteri (Hcrdman) CRUSTACEA Idolea linearis (L.) Pandalus montagui Leach Crangon allmani Kinahan Crangon crangon (L.) Pagurus bernhardus (L.) Pisidia longicomis (L.) Hyas araneus (L.) Inachus dorsellensis (Pennant) Macropodia roslrala (L.) Corysles cassivelaunus (Pennant) Alelecyclus rotundatus (Olivi) Cancer pagurus L. Carcinus maenas (L.) Liocarcinus depuralor (L.) Liocarcinus holsatus (Fabricius) Liocarcinus marmoreus (Leach) Necora puber (L.) Pilumnus hirtellus (L.) MOLLUSCA Crepidula fomicata (L.) Buccinum undatum L. Acanlhodoris pilosa (Abildgaard in Müller) Aequipeclen opercularis (L.) Bivalvia (indet.) Sepiola atlantica d'Orbigny Alloteuthis subulata (Lamarck) Loligo sp. BRYOZOA Alcyonidium diaphanum Lamouroux Flustra foliacea (L.)
Trans. Suffolk Nat. Soc. 35
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ECHINODERMATA Crossaster papposus (L.) Henricia oculata (Pennant) Asterias rubens L. Ophiura albida Forbes Ophiura ophiura (L.) Ophiothrix fragilis Thomson Echinus esculentus L. Psammechinus miliaris P. L. S. Müller Echinocardium cordatum (Pennant) ASCIDIACEA Molgula sp. ELASMOBRANCHII Scyliorhinus canicula (L.) Galeorhinus galeus (L.) Mustelus asterias Cloquet Raja clavata L. Raja montagui Fowler TELEOSTEI Anguilla anguilla (L.) Spratlus spratlus (L.) Engraulis encrasicolus (L.) Gadus morhua L. Merlangius merlangus (L.) Trisopterus luscus (L.) Trisopterus minutus (L.) Ciliata mustela (L.) Trigla lucema L. Myoxocephalus scorpius (L.) Taurulus bubalis (Euphrasen) Agonus cataphractus (L.) Liparis liparis (L.) Mullus surmuletus L. Echiichthys vipera (Cuvier) Hyperoplus lanceolatus (Le Sauvage) Callionymus lyra L. Gobius niger L. Pomatoschistus minutus (Pallas) Glyptocephalus cynogtossus (L.) Limanda limanda (L.) Microstomus kitt (Walbaum) Platichthysßesus (L.) Pleuronectes plattesa L. Solea solea (L.) Buglossidium luteum (Risso)
MARINE FAUNA OFF THE COAST OF EAST ANGLIA
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r c c o r d c d at all stalions. T h e d o m i n a n t specics, b y b i o m a s s , were the b r y o z o a n Alcyonidium diaphanum a n d A. rubens. N u m c r i c a l l y , A. rubens, L. holsatus and pink s h r i m p Pandalus montagui dominated catchcs. The highest-ranking spccics, b y o c c u r r c n c c , b i o m a s s and n u m b e r s arc given in T a b l e 3.
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Macro-cpibenthic invertebratc assemblages Cluster analysis for invertebratc data alone indicated that thc twenty catches could be attributed to six groups (Table 4a), with the stations within each of these groups having a similarity of more than 50%. These six groups formed two broad categories, those predominantly in thc south of the region dominated by A. diaphanum and those in the outer Thames Estuary and off Suffolk dominated by more motile fauna. Station T had the most disparate invertebratc catch, and was dominated by A. rubens and A. diaphanum. Thc common sun star Crossaster papposus was also abundant and. although not recorded in the random sub-sample, the starfish Henricia oculata and common sca urchin Echinus esculentus occurred at this Station. Stations I and L were also dominated by A. diaphanum and A. rubens, and the "reef'-building polychaete Sabellaria spinulosa was also an important component. This group was relatively similar to stations F, G, J and K, which were in the outer limits of the Thames Estuary and characterised by the dominance of A. diaphanum. The two latter groups were primarily differentiated on the basis of higher catches of shore sea urchin Psammechinus miliaris and C. pagurus at stations I and L. The stations described above were distinet from the remaining sites on the basis of their larger catches of A. diaphanum, A. rubens, C. pagurus and hydroids. Stations off Margate (D, E) were mostly dominated by motile fauna, especially A. rubens, L. holsatus and sea mouse Aphrodita aculeata. Although A. diaphanum was a significant component of the assemblage, other sessile invertebrates, including hydroids, S. spinulosa and plumose anemone Metridium senile formed only 3-6% of the biomass. Stations off the coast of Suffolk between Aldeburgh to Lowestoft (M, N, O, Q, R) were also dominated by motile invertebrates, including A. rubens, shore crab Carcinus maenas, C. pagurus, P. miliaris and L. holsatus. The remaining stations (A, B, C, H, P, S) were dominated by A. rubens and A. diaphanum, with S. spinulosa and C. pagurus also important constituents. Demersal fish assemblages Cluster analysis of the demersal fish fauna resulted in the identification of three fish assemblages (Table 4b). Stations H and S had similar catches of lesserspotted dogfish Scyliorhinus canicula, dab Limanda limanda and lesser weever fish Echiichthys vipera. The remaining stations represented two relatively similar flatfish assemblages, one dominated by lemon sole Microstomus kitt, L. limanda and sole Solea solea, with S. canicula also an important species, and a second dominated by 5. solea, plaice Pleuronectes platessa and L. limanda. The former assemblage occurred at four stations (B, I, L, T), the latter at the remaining 13 stations. Demersal assemblages of fish and invertebrates A third analysis was undertaken for all fauna (i.e. fish and invertebrates), and five distinet groups of stations, with similarities of 45-50%, were identified (Table 4c). Stations P and S were dominated by motile fauna, notably A. rubens, S. canicula, C. pagurus, L. limanda and thomback ray Raja clavata. Sessile taxa included hydroids, S. spinulosa, and the bryozoans Flustra
Trans. Suffolk Nat. Soc. 35
(1999)
MARINE FAUNA OFF THE COAST OF EAST ANGLIA
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Suffolk Ne ural History, Vol. 35
52
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Trans. Suffolk Nat. Soc. 35 (1999)
MARINE FAUNA OFF THE COAST OF EAST ANGL1A
53
foliacea and A. diaphanum. The lattcr species accountcd for only 2-7% or ihc catch. Stations M, N, O, Q and R were also dominated by motilc fauna. including S. solea, A. rubens and R. clavata. Sessilc fauna, which accounlcd for only 3-3% of the catch, includcd A. diaphanum, hydroids. S. spinulosa, F. foliacea and dahlia anemone Urlicina felina. These stations were adjaccnt to stations P and S, but were distinet from this pair of stations duc to the greater abundance of S. solea, common hermit crab Pagurus bernhardus, pogge Agonus cataphractus and bib Trisopterus luscus. Stations I, L and T were dominated by A. diaphanum, A. rubens and S. canicula, and these stations occurred at the extremities of the study area. Stations D, E, F, G, J and K were also dominated by A. diaphanum, and the nudibranch Acanthodoris pilosa, which gra/.es on A. diaphanum, was frequently caught. A. rubens, L. holsatus and S. solea were also significanl components of the catch. These stations were situated off the coasts of northeast Kent and southeast Suffolk. The remaining stations (A, B, C, H) were dominated by fish, including R. clavata, P. platessa, S. solea and L. limanda, although A. diaphanum was still a relatively important componcnt of this assemblage. These stations, which were in close proximity to the previous group, were biologically differentiated by their lower biomass of A. diaphanum, A. rubens, long-legged spider crab Macropodia rostrata, S. solea and M. senile. Discussion This survey is one of few examining the composition and distribution of macro-epibenthic assemblages, as inferred from beam trawl catches, off the coast of East Anglia. Beam trawls are considered to be effective gears for catching and sampling macro-epifauna and demersal fish, and they allow a large area of the seabed to be sampled. They have, however, bcen considered inappropriate for some Community analyses, due to the poorly defined relationship between epifauna (i.e. those animals living on the sea-floor) and Sediment, and the arbitrary nature of catches (Duineveld et al., 1991). Other methods of identifying epibenthic communities, such as the visual inspection of the seabed from towed photographic equipment and SCUBA surveys, may be unsuitable for the turbid waters of the south-western North Sea. The currcnt results indicate that groundfish surveys can be useful tools in the initial determination of the different faunal assemblages that may occur in an area. The identification of epifaunal assemblages is generally morc difficult than the description of infaunal communities (i.e. those animals living within the sediments), where Sediment type is an important discerning factor. The distribution of epibenthic organisms (i.e. those animals living above the sediments) is known to be affected by depth and Sediment composition (Basford et al., 1990) and, in the southwestern North Sea, salinity, turbidity and water movemenls are also important abiotic factors. Two distinet epifaunal assemblages have been reported in the northern North Sca: a sponge-F. foliacea-Hyas coarctatus assemblage and an A. rubens-Astropecten irregularis-Brissopsis lyrifer assemblage on coarsc and fine sediments respectively (Basford et al., 1990). Neither of these assemblages was evident in the more southern waters of our study area. Detcrmining bona fide communities or assemblages from trawl catches is complicated further by the
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sampling of an unknown number and variety of habitats Over a tow distancc of 1-5-2-0 km. Nevertheless, the groups of stations identified above provide thc first stcp in determining the broad ränge of faunal assemblages that occur in the study area. Distinct groups of stations, which were assumed to represent particular assemblages, were observed in each analysis. Analysis of the invertebrate catches resulted in thc identification of six assemblages, whereas demersal fish appeared to be less site-speeifie and only three assemblages were observed. That the larger and more motile fish communities appeared to be less habitatspeeifie was as expected. The third analysis, which was based on the biomass of both invertebrates and fish, may provide a better representation of the variety and faunal composition of demersal assemblages in the area. The fish assemblages reflected only a few of the differences in the invertebrate assemblages, for example stations at the outer limits of the study area (I, L, T) were distinct from other stations in both analyses. The majority of the stations in the coastal waters of the Thames Estuary and Suffolk had a relatively homogeneous fish assemblage and were not significantly different between sites with differing invertebrate assemblages. S. solea and other flatfish dominated the fish communities in this area, although two species of elasmobranch (R. clavata and S. canicula) were also important in terms of biomass. Thc ctenostome bryozoan A. diaphanum (often referred to as A. gelatinosum in early works - see Thorpe & Winston (1984, 1986) for taxonomic revision) was a key species in determining assemblage types, and accounted for 63-7% of the total faunal biomass. By contrast, large-bodied taxa such as crustaceans, fish and echinoderms constituted 5-0, 9-5 and 18-4% respectively of the total biomass. Despite the abundance of A. diaphanum in the Outer Thames Estuary and areas of the North Sea, including the Dogger Bank (Dyer et al., 1983), its biology and role in the ecosystem are poorly known and future research on the communities within these Alcyonidiumdominated habitats is needed. Acknowledgments We thank the scientists, captain and crew of the R.V. Corystes for their assistance at sea and J. Dann for his assistance with Computing. This work was funded by the Ministry of Agriculture, Fisheries and Food under MOU ' A' and the UK Department of the Environment, Transport and the Regions as a contribution to its co-ordinated programme of marine research for the North East Atlantic. References Anon. (1934). The crustacea of Suffolk. Part 1: crabs, lobsters and shnmps. Trans. Suffolk Nat. Soc., 2: 265-271. Anon. (1937). The Bryozoa of Suffolk. Trans. Suffolk Nat. Soc., 3: 254-256. Anon. (1938). The Mollusca of Suffolk. Trans. Suffolk Nat. Soc., 4: 2-22. Bamc, J. Hâ&#x20AC;&#x17E; Robson, C. F., Kaznowska, S. S., Doody, J. P., Davidson, N. C.& Buck, A. L., eds. (1998). Coasts and Seas of the United Kingdom. Region 7 South-east England: Lowestoft to Dungeness. Peterborough: Joint Nature Conscrvation Committee.
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Basford, D., Eleftheriou, A. & Raffaelli, D. (1990). The infauna and epifauna of the northem North Sea. Netherlands Journal of Sea Research, 25: 165-173. Clarke, K. R. & Warwick, R. M. (1994). Change in marine communiiies: An approach to Statistical analysis and interpretation. Plymouth Marine Laboratory. Collings, D. W. (1933). The fishes of Suffolk. Trans. Suffolk Nat. Soc., 2: 104-133. Covey, R. (1998). Chapter 6: Eastem England (Bridlington to Folkestonc) (MNCR Sector 6). In Marine Nature Conservation Review: Benthic marine ecosystems of Great Britain and the north-east Atlantic (K. Hiscock, ed.). Peterborough: Joint Nature Conservation Committce. Davis, F. M. (1923). Quantitative studies on the fauna of the sea boltom No. 1 - Preliminary investigation of the Dogger Bank. Ministry of Agriculture and Fisheries, Fishery Investigations, Series II, 6(2); 54pp. Duincveld. G. C. A„ Künitzer, A„ Niermann, U„ de Wilde, P. A. W. J. & Gray, J. S. (1991). The macrobenthos of the North Sea. Netherlands Journal of Sea Research, 28: 53-65. Dyer, M. F., Fry, W. G„ Fry, P. D. & Cranmer. G. J. (1983). Benthic regions within the North Sea. Journal of the Marine Biological Association of the United Kingdom, 63: 683-693. George, J. D., Chimonides, P. J. Evans, N. J. & Muir, A. I. (1995). Fluctuations in the macrobenthos of a shallow-water cobble habitat off North Norfolk. In Biology and ecology of shallow coastal waters. Proceedings of the 28th European Marine Biology Symposium, Crete, September 1993 (A. Eleftheriou, A. D. Ansell & C. J. Smith, eds.). Hamond, R. (1963). A preliminary report on the marine fauna of the north Norfolk coast. Transactions of the Norfolk and Norwich Naturalists' Society, 20: 2-31. Hamond, R. (1969). On the Norfolk marine area, and the offshore stations worked within it. Transactions of the Norfolk and Norwich Naturalists' Society, 21: 209-228. Hamond, R. (1971). The leptostracan, euphausiid, stomatopod and decapod crustacea of Norfolk. Transactions of the Norfolk and Norwich Naturalists' Society, 22: 90-112. Hughes, R. G. & Quinn, P. (1983). Additions to the Suffolk marine fauna list. Trans. Suffolk Nat. Soc., 19: 346-357. Kaiser, M. J. & Spencer, B. E. (1994). Fish seavenging behaviour in recently trawled areas. Marine Ecology Progress Series, 112: 41-49. Miliner, R. S.; Dickson, R. R. & Rolfe, M. S. (1977). Physical and biological studies of a dredging ground off the east coast of England. ICES CM 1977/E:48; l l p p . Mistakidis, M. N. (1951). Quantitative studies of the bottom fauna of Essex oyster grounds. Fishery Investigations, Series II, 17(6): 1-47. Mistakidis, M. N. (1957). The biology of Pandalus montagui Leach. Fishery Investigations, Series II, 21(4): 1-52.
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Pawson, M. G. & Robson, C. F. (1998a). Chaptcr 5.5 Exploited sca-bcd species. In Coasts and Seas of the United Kingdom. Region 7 South Easi England: Lowestoft to Dungeness (J. H. Bamc, C. F. Robson, S. S. Kaznowska, J. P. Doody, N. C. Davidson & A. L. Buck, cds.). Petcrborough: Joint Naturc Conservation Commiltce. Pawson, M. G. & Robson, C. F. (1998b). Chaplcr 5.7 Fish: Exploited sea fish. Ibid, 121-125. Rogers, S. 1. (1997). A revicw of closed arcas in the United Kingdom Exklusive Economic Zone. Scientific Series, Technical Report, CEFAS (Lowestoft), 106: 20pp. Thorpe, J. P. & Winston, J. E. (1984). On the identity of Alcyomdium gelatinosum (Linnaeus, 1761) (Bryozoa, Ctenostomata). Journal of Natural History, 18: 853-860. Thorpe, J. P. & Winston, J. E. (1986). On the identity of Alcyomdium diaphanum Lamouroux, 1813 (Bryozoa, Ctenostomata). Journal of Natural History, 20: 845-848.
J. R. Ellis & S. 1. Rogers CEFAS Lowestoft Laboratory Pakefield Road Lowestoft Suffolk NR33 OHT
Trans. Suffolk Nat. Soc. 35
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Plate 3: Catch from CEFAS Research Vessel Corystes (p. 4 & p. 45). Large fish is a Cod, Gadus morhua, others include Sand eels, Ammodytes marinus, Gurnard, Aspitrigla cuculus, Dab, Limandci limanda and Herring, Clupea harengus.