Check List of Species (Algae, Invertebrates and Vertebrates) Found in the Vicinity of the Island of Helgoland (North Sea, German Bight) — a Review of Recent Records J

HELGOLÄNDER MEERESUNTERSUCHUNGEN H elgoländer M eeresunters. 47, 1—34 (1993)

Check list of species (algae, invertebrates and vertebrates) found in the vicinity of the island of Helgoland (North Sea, German Bight) — a review of recent records J. Harms

Biologische Anstalt Helgoland (Meeresstation); D-27483 Helgoland, Federal Republic of G e rm a n y

ABSTRACT: The species lists presented for benthic macroalgae, invertebrate and vertebrate species presented are extracted from recent publications (from 1977 to date). The lists summarize the species composition of the intertidal and subtidal hard-bottom communities around Helgoland. Additional information is supplied for the species composition of the “Steingrand" and "Tiefe Rinne“. The lists do not claim completeness, but have been intended to provide a working platform for further listings and for comparison with data published earlier.

INTRODUCTION The island of Helgoland is situated some 50 km off the nearest mainland. The marine environment around Helgoland displays an offshore character and is for the major part of the year under the influence of North Sea water bodies (Martens, 1978). Fresh water inflows from the rivers Westerschelde, Maars, Rhine, Weser and Elbe, and the pollutants transport therein reach the area around Helgoland only in diluted form (Korringa, 1968; De Ruijter et al., 1987). Therefore, the area can be considered as relatively unpolluted compared to more inshore localities in the southern North Sea. Nevertheless, the environment around Helgoland is influenced by human activities, which has been proved by regular analyses of nutrients and phytoplankton biomass during the last 30 years on the Helgoland Reede (Radach & Berg, 1986; Radach & Bohle-Carbonell, 1990). The increase in nutrients and phytoplankton biomass results in an overall increase of benthic macrofauna biomass in the German Bight (Rachor, 1990), which is in good agreement with other organically enriched areas in the North Sea (Rachor, 1990 and literature cited therein). Rachor (1990) demonstrated that deep-dwelling and long-living species derive less benefit from these changes than animals which ( 1) live near the sedim ent surface; 2 () have an adaptive feeding behaviour; and (3) grow and reproduce quickly. Helgoland is the only natural hard-bottom substrate in the southern part of the North Sea. The rocky area around Helgoland covers about 40 km2, which is more or less identical with the area of the nature reserve (5138 ha). In the vicinity of Helgoland, an additional hard-bottom substrate occurs at ''Steingrund'', a glacial end moraine north-

Senckenbergische Bibiiothak | Frankfurt a. Main I 2 J. Harms Species list Helgoland 3 east of Helgoland. The species settling on these hard-bottom substrates can be consi dered as isolated. The next natural hard-bottom communities occur along the Norwegian and East English coast line. We know only little about recruitment and population dynamics of the species living in the intertidal and subtidal area around Helgoland. This \A8 * ^ V area is under particular pressure by human activities such as fishery and ship traffic, A a 9 \ * b 6 ^ few miles south of Helgoland there are major shipping routes to the harbours of Wilhelmshaven, Bremerhaven, Hamburg and to the Baltic Sea. The potential danger of these activities was demonstrated vividly on January 26th, 1993 when the Hudson Bay , a 299 m cargo ship, was in peril of stranding in the southwest intertidal area of the island of Helgoland. During the last 15 years, several investigations on community structure in the intertidal and sublittoral area around Helgoland have been performed (see literature dted in Table 1). The species lists presented here summarize data on benthic algae (Table 2), benthic invertebrates (Table 3) and acrania and vertebrates (Table 4) as supplied by these publications. Additional information is supplied by records on plank tonic polychaete larvae (Table 5), whose adults are not listed in Table 3. So far, only a list of benthic macroscopic algae settling around Helgoland exists (Kornmann & Sahling, 197?, 1978, 1983; Kornmann, 1986). The algae species recorded in recent publications comprise 2/3 of the species listed in the publications by Kornmann & Sahling (1977, 1983). The frequency of recordings might be a useful first indication of the abundance of each species. As far as possible, rare species and species with highly fluctuating abundance are marked. The relevant publications should be consulted for more detailed information about sampling methods, locations and literature used for species determina tions. As far as possible, sampling areas are given in Figure 1 for the intertidal and in Figure 2 for the subtidal area. The species lists were supplemented by observations (from 1980 until the present date) made by members of the Biologische Anstalt Helgoland. Most of these additionally listed species can be considered as rare. The literature used for taxonomic classification is listed in Table 6 . Fig. 1. Sampling area in the rocky intertidal zone of the island of Helgoland. For explanation of station numbers, see Table 1 The lists presented are a preliminary attempt to give an overview of the species « diversity of the hard-bottom community around Helgoland. The lists do not claim completeness; more detailed investigations with regular sampling programs are needed made by Berberich (1989) for the "Tiefe Rinne“, and by Kühne (1992) for the "Stein- for both intertidal and subtidal communities. For further completion of these lists in the grund". Berberich (1992) compared her data with earlier recordings made by Caspers near future, information on additional species, with reference material if possible, is (1939). For numerous species, changes in abundance could be recognized. At “Stein- especially requested from guest researchers and student courses which are held in great grund", almost complete extinctions of macroalgae compared to early investigations number at the Biologische Anstalt Helgoland each year. (Hagmeier, 1925) were recorded. As shown by Berberich (1989) and Kühne (1992), only The lists of the current species composition around Helgoland will form the basis dramatic changes in abundance, extinction of species or new records can be detected by for comparison with data sets published in earlier years (Dalla Torre, 1889; Heincke, direct comparison between new and old data sets. Comparisons can only be a provisional 1894a,b; 189?; Hartlaub, 1894; Kuckuck, 1894, 1897a, b, 1900, 1902; Michaelsen, 1897, means of assessment for the detection of changes in the species composition. In dealing 1923; Attems, 1897; Ehrenbaum, 1897; Weltner, 1897; Ascherson, 1900; Heydrich, 1900; with changes in species composition, one must take into account that both natural Sokolowsky, 1900, 1902, 1925; Schultz, 1902; Longchamps, 1904; Hartmeyer, 1908; phenomena and human activities are continously changing the living conditions for wild Borowsky, 1910; Grimpe, 1925; Hagmeier, 1925, 1930; Zimmermann, 1925; Klie, 1927; flora and fauna. Although it has become obvious that human impact is of greatest Hertling, 1930, 1932; Nienburg, 1930; Schlottke, 1932; Caspers, 1939, 1950, 1952). significance for the often dramatic changes observed in the environment during the last Additional information on the community structure around Helgoland is given by Stein et decades (Agger, 1989), it is unrealistic to dream about an unchangeable nature. Nature al. (1990), who summarized data sets between 1902 to 1912 for the Baltic and North Sea has been continuously transforming since its very beginning, and will keep on changing area. The comparison between new and old data sets is time consuming and is not the long after the last human has turned into a fossil (Reise, 1989). purpose of this publication. First attempts to quantify changes in species abundance were Detection of long-term trends in species abundance and species composition, J, Harms Species list Helgoland 5 4 combined with the recording of variables such as climatic effects, changes in sea level, long-term and wide-scale effects of exploitation, nature management, and pollution can clarify the reasons for changes in species composition, Helgoland is an ideal place for •E- such a monitoring project. It fulfills all the requirements needed for a successful monitor ing program (Reise, 1989): (1) Historical comparisons and paleologieal analysis provide interpretations of the present state; ( 2 ) an institution exists to guarantee uninterrupted •E: time series studies into the next decades; and (3) the hard-bottom substrate around the island of Helgoland has been designated as nature reserve since 1981, •E. Acknowledgements, This publication would not have been possible without the selfless help of my colleagues who gave helpful information and supported this work in great manner. My thanks are especially due to H.-G, Andres (Zoolog. Inst. u. Mus,, Univ. Hamburg), D. Barthel (IFM, Univ. Kiel), I, • K. Bartsch (BAH-Hamburg), A, Dzwillo (Zoolog. Inst. u. Mus., Univ. Hamburg), G, Hartmann-Schröder (Zoolog. Inst. u. Mus., Univ. Hamburg), R. Janssen (Senckenberg, Frankfurt), G. Jarms (Zoolog. Inst, •E u. Mus., Univ. Hamburg), P. Kornmann (BAH-Helgoland), U. Muhlenhardt-Siegel (Zoolog. Inst. u. Mus., Univ. Hamburg), P.-H. Sahling (BAH-Helgoland), P. Sundberg (Univ. of Göteborg), B. Wahl (BAH-Hamburg), C. Warnecke-Cremer (Zoolog. Inst, u. Mus., Univ. Hamburg) for taxonomic and • E, systematic advice and to R. Foster for correcting the English.

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British marine isopods, - Synopses Br. Fauna,3, 1-86, Hamburg, 110 pp, Newman, A. & Ross, A., 1976. Revision of the balanom orph barnacles; including a catalogue of the Smaldon, G., 1979. British coastal shrimps and prawns. — Synopses Br. Fauna.15, 1-126, species. Society of Naural History, San Diego,9, 1—107. Smock, U., 1991. Licht- und Elektronenmikroskopische Untersuchungen anLagotia spec. Otophora: Nielsen, C., 1989. Entoprocts. - Synopses Br. Fauna.41, 1-131. Heterotrichida. Dipl, Arb., Univ. Köln, 71 pp. Nienburg, W., 1930. Die Besiedlung des Felsstrandes und der Klippen von Helgoland. Teil 2: Die Sokolowsky, A., 1900. Beiträge zur Meeresfauna von Helgoland. XI. Die Amphipoden Helgolands, — Algen. — Wiss. M eeresunters, (Abt. Helgoland). 15, 1-15, Wiss. M eeresunters, (Abt. Helgoland). 4, 141—166. Nierstrasz, H. F. & Schuunnans Stekhoven, J. H., 1930, Anisopoda. - Tierw. Nord- u. Ostsee,10 (Xe), Sokolowsky, A., 1902. Beiträge zur Meeresfauna von Helgoland. XIII, Ober drei in der Nordsee bei 134-172. Helgoland gefundenen Caprellidea. - Wiss, Meeresunters. (Abt. Helgoland). 5, 1.1-16. Nordheim, H. von, 1983. Systemati.es and ecology ofProtodrilus helgolandicus sp. n,, an interstitial Sokolowsky, A., 1925. Nachträge zur Ampohipodenfauna Helgolands. — Wiss. Meeresunters. (Abt. polychaete (Protodrilidae) from subtidal sands off Helgoland, German Bight. - Zool. Ser.12, Helgoland). 16, 1-6. 171-177. Stein, U., Hudriede, W. St Rumohr, H., 1990. Historische Benthosdaten aus Nord- und Ostsee in den Nordheim, H. von, 1984. Life history of subtidal interstitial polychaetes of the families Polygordiidae, Jahren 1902—1912. — Mitt. zool. Mus. Univ. Kiel (Suppl.),3, 1—189. Protodrilidae, Nerillidae, Dinophilidae, and Diurodrilidae from Helgoland (North Sea), - Helgo Ten Broeke, A., 1929, Sipunculoidea und Echiuroidea. — Tierw. Dtl. 15, 156—168. länder Meeresunters. 38, 1—20. Thompson, T. E. & Brown, G. H., 1976, British opistobranch molluscs. — Synopses Br. Fauna,8 ,1-203. Nordheim, H. von, 1989. Six new species ofProtodrilus (Annelida, Polychaeta) from Europe and Ude, H., 1929. Oligochaeta. — Tierw. Dtl. 15, 1-132. New Zealand, with a concise presentation of the genus. — Zool. Ser.18, 245-268. Vervoort, W., 1946. Hydrozoa (Cl). A. Hydropolypen. - Fauna Ned,14, 1-336. Nordsieck, F., 1969. Die europäischen Meeresmuscheln. Fischer, Stuttgart, 253 pp. Weltner, W., 1897. Beiträge zur Meeresfauna von Helgoland. IX. Die Cirripedien Helgolands. — Nordsieck, F., 1982, Die europäischen Meeres-Gehäuseschnecken (Prosobranchia). Fischer, Stutt Wiss. M eeresunters. (Abt. Helgoland). 2, 437—447. gart, 539 pp. W estheide, W., 1990, Polychaetes: Interstitial families. — Synopses Br, Farma.44, 1—152, Pesta, O., 1932. Krebstiere oder Cmstacea. I, Ruderfüßler oder Copepoda 3. Unterordnung; Harpac W hitehead, P. J, P., Bauchot, M.-L,, Hureau, L.-C., Nielsen, J. & Totonese, E, (Eds.), 1984-1986. ticoida (1. und 2, Hälfte), - Tierw. DÜ. 24, 1-164. Fishes of the north-eastern Atiantic and the Mediterranean. Unesco, Paris, 1—3, 1473. Plate, S„ 1992. Untersuchungen zum zeitlichen Auftreten, zur Abundanz und Entwicklung mero- Zimmermann, W., 1925. Helgoländer Meeresalgen I-VI, Beiträge zur Morphologie, Physiologie und planktischer Polychaetenlarven im Gebiet Helgoland Reede; Ordnungen: Orbiniida, Spionida Oekologie der Algen. — Wiss. Meeresunters, (Abt. Helgoland). 1 6 ,1-25. und Capitellida. Diss., Ruhr-Univ,, Bochum, 235 pp. Platt, H.M. & Warwick, R. M., 1983. Free-living marine nematodes. Part I; British enoplids, - Synopses Br, Fauna.28, 1-307. Pleijel, F. & Dales, R. P., 1991. Polychaetes: British phyllodocoideans, typhloseolecoideans and tomopteroideans. - Synopses Br. Farma.45, 1-202. Prudhoe, S., 1982. British polyclad turbellarians. — Synopses Br. Fauna.26, 1-77, Radach, G, &¿ Berg, J., 1986. Trends in den Konzentrationen der Nährstoffe und des Phytoplanktons in der Helgoländer Bucht (Helgoland Reede Daten). -B er, Biol. Anst, Helgoland. 2, 1-65, Radach, G. & Bohle-Carbonell, M., 1990. Strukturuntersuchungen der meteorologischen, hydro graphischen, Nährstoff- und Phytoplankton-Langzeitreihen in der Deutschen Bucht bei Helgo land. - Ber. Biol. Anst, Helgoland, 7, 1—425, Rachor, E., 1990. Changes in sublittoral zoobenthos in the German Bight with regard to eutrophica tion. - Neth, J. Sea Res. 25, 209-214. J, H arm s Species list Helgoland 11

Table 1, literature used for the survey, and explanations of the symbols used in Tables 2-4 Table 2. Check list of benthic macroscopic algae settling around Helgoland as listed in recent publications (for a more complete overview see Kornmann & Sahling, 1977,1983). Due to methodical O rig in of d a ta A: K. Janke (1985, 1986) difficulties (diving) or because individuals were juveniles, individuals were not always determined B: Bock (1992) to species level; therefore, individuals marked as "sp." do not necessarily represent additional C: Gillandt (1979) species to the already listed members of the same genus. Synonyms used in one of the summarized D: Breiholz (1992) publications are given in brackets; • The Wva-group is now undergoing a taxonomic revision E: Hinz (1992) (Kornmann & Sahling, pers, comm.); o Taxonomic classification uncertain; for more information, see F: Kriiß (1988) Table 1 G: Schultze (1987); Schultze et al. (1990) H: Karez (1991) Chlorophyceae I: Haß (1992) Acrosiphonia arcta (Dillw.) J. Ag. A7, J2, R J: Harms & Anger (1983) Acrosiphonia centralis (Lyngb.) KjeUm. Ei, G¡, G2i R K: Kluijver (1991) Acrosiphonia sonderi (Kütz.) Komm. Ei, G2, R L: Müller (1990) Acrosiphonia sp. Gi, G2 M; Berberich (1989) Blidingia minima (Näg. ex Kütz.) Kylin A2 i A q, Bí—Bß, R N; Kühne (1992) Blidingia marginata (J. Ag.) P. Dang. Bi-B4, Bs, R O: Skaumal (1977) Blidingia sp. As P; Dahms (1990) Bryopsis hypnoides Lamour. Bí, D3, D4i H i , K7i Kg Q: Nordheim (1984) Bryopsis lyngbyei Homem. Ki.Kg R; M unda & M arkham (1982) Bryopsis plumosa (Huds.) Ag. o e 2, e 3 S; Smock (1991) Capsosiphon flavescens (C. Ag.) Setch. et Gardn, Bí, B3 and personal communications from members of the Biologische Anstalt Helgoland Chaetomorpha aerea (Gooden, ex Dillw.) Kütz. Di, D3 —D5 , Ei, E2 Chaetomorpha melagonium (Web. et Mohr) Kütz. B4i B7i D2i D3i D3i Gi, G2, Hi—H3i Kt—Ka, K7 i A, B, C, H3, L í , O, R: Samples from the eulittoral zone, R D, E, F, G, Hi_2 , I, J, K, L2, Q: Samples from the subtidal zone, Chaetomorpha tortuosa (Dillw.) Kütz. A31 A4 , Aß, A?, H2i H3i R Gj; Fouling species onLaminaría digitata, Cladophora albida (Huds.) Kütz. b 4 , k 2 G2; , Fouling species onLaminaría hyperborea Cladophora rupestris (L.) Kütz. A3i Aß—Ag, B4 -B 7, Ki, K2i R M: Species of the "Tiefe Rinne" Cladophora sericea (Huds.) Kütz. A3i A4 , Ae, Af, Ag, B2, B7i Gi, K4 , R N; Species of the "Steingrund" (gravel fields), Cladophora sp. h 3 N2: "Steingrund", additional species on nearby sandy bottoms. Codium fragile (Sur.) Hariot R, Ka Enteromorpha compressa (L.) Grev. B2, B3i Bg, R The symbols used correspond to the station descriptions given by the authors as follows: A; = Enteromorpha intestinalis (L.) Link R sandy bottom; A2 = gravel fields; A 3 = Myfflus-beds; A 4 = Littoiina-habitat; A 5 = Enteromorpha- Enteromorpha linza (L.) J. Ag. Ki, K2, K8i R zone; Ae = Fucus-zone; A7 = Priei (creek); A 8 = Laminaria-zone; Ag = species on the north west Enteromorpha prolifera (O, F. Müll.) J. Ag. K i , K2, R pier; Bt = station 1, 2, 3; B2 = station 5, 6 , 7, 8 , 9; B3 = station 10, 11, 12, 14; B4 = station 13, 15; B5 Enteromorpha sp. A2i As, Ag, Bí—B7i Di, Ei, E2, Es, Ga, J3 = station 16, 17, 18, 20; B6 = station 19; B7 = station 4, 21; Di = station 1, 5; D2 = station 2; D3 = Hormiscia neglecta Komm, B s station 3; D4 = station 4; D5 = station 6 ; Ei = station 1, 2, 3; E2 = station 4, 5, 6 ; E 3 = staion 7, 8 , 9; Hormiscia penicilliformis Roth (Fries) R Ei = station 10; E5 = station 11; Ht = south harbour; H2 = white Priei; H 3 = north-east intertidal; Monostroma grevillei (Thur.) Wittr. R Ii = station A; I2 = station B; I3 = station C; I4 = station D; I5 = station E; Jx = station A; 2 J = Monostroma undulatum Wittr. Ag, R station B; J3 = station C; Kt = community A;K 2 = community B; K3 = community C; K4 = Prasiola stipitata Suhr in Jessen Bi-B 4 community D;Kä = community E; K6 = communityF; K7 = community G;Ka = community H; N Rhizoclonium kochianum Kütz. g 2 = gravel fields; N 2 = additional species on nearby sandy bottoms, Rhizoclonium riparium (Roth) Harv. B2—B4i Bg Spongomorpha aeruginosa (L.) Hoek. B2, R 1 rare species (for more detailed information see Kühne, 1992) Ulothrix sp. Ag, Ei—E3, H3 i R 2 rare species (for more detailed information see Berberich, 1989) Ulva lactuca L. • A2 —A 4 , Ag—Ag, Bí—B7, Ki, Kg, K7, R 3 rare species (for more detailed information see Skaumal, 1977) Ulva curvata (Kütz.) De Toni • Ag, B4, J 2 rare species (observations by members of the Biologische Anstalt Helgoland) Ulva sp. • Dx, D3 -Ds, Et-Es, Gi, G2, Hi-H3, Ki, K2 * rare species considered by Heiber & Rachor (1989) * * endangered species (Rote Liste, D eutschland [Blab et al. 1984]) Phaeophyceae 1 high variation in species number from year to year Ascophyllum nodosum (L.) Le Joi, B3i K2i Ks Chorda tomentosa Lyngb. Ei, E2 Chorda filum (L.) Stackh. d 2, D 4 Chordaria Oagelliformis (O. F. Müll.) C. A g.’ R Cladostephus spongiosus (Huds.) C. Ag. B4 Desmarestia aculeata (L.) Lamour. D s, Ei , E2, Gi , G2p K1( K s 12 J. H arm s Species list Helgoland 13

Table 2 (Continued) Table 2 (Continued)

Phaeophyceae Rhodophyceae D i, D3—DSl Ej, Gj, G21 Ki, K2i Ka, R Desmarestia viridis (O. F. Müll.) Lamour. Membranoptera alata (Huds.) Stackh. By, Di, Da, Ds, El, E2, Gi, G2, H2, Ku R G i, G2, R Ectocarpus fasciculatus Harv, Phycodrys rubens (L.) Batt. Di-Da, Ds, Ei, Ea, Hi, Ki, K2, K7 Ectocarpus siliculosus (Dillw.) Lyngb. R [Phycodryssinuosa Ectocarpus sp. A s, B2, D i , D 3i D 4 i E i , K x, K2, Kg (Good, et Woodw.) Kütz.) Elachista fucicola (Veli.) Aresch. B4, R Phyllophora pseudoceranoides Ej, Ea, Hlf Ki-K 3 , K7 Elachista sp. Kl r K2 (S. G. Gmel.) Newr. & A. R. A. Taylor Fucus serratus L. Aj—A4, A 6-A 9, B2| B4—Bg, R Phyllophora traillii Holm, ex Batt. Ea, K2 i K, Fucus spiralis var. platycarpus L, A 2i A3, Ag, B j—B7i R Phyllophora truncata (Pali.) Zinova Di—D3, D5i Ei—Eg, E 5 Fucus vesiculosus L. A 2—A4, Ag, Ag, B o B 4, Bg, R Phyllophora sp. Gi, G2 Fucus sp. Ag Plocamium cartilagineum (L.) Dixon B5, D2i D3, D5i Ej, Ea, Ht-H 3, Ku K2 Halidrys siliquosa (L.) Lyngb, As Plumaria elegans (Bonnern.) Schmitz B4, By, Di, D4, Ei, Gi, Ga, H t, K7 Laminaria digitata (Huds.) Lamour. Ae, A7, B4, D i—D3i D s, Elf Gi, G2, R Polyides rotundus (Huds.) Grev. Da. D5 , Ei, Ea, K2 Laminaria hyperborea (Gunn.) Fosl. Ei—Es, Ki, Ks Polysiphonia elongata (Huds.) Spreng. Ea, J 2 Laminaria saccharina (L.) Lamour. As, A7, D i, D3—D5, E i , Kj, K3, R Polysiphonia nigra (Huds.) Batt. * Kx,K2 Polysiphonia nigrescens (Huds.) Grev. Laminaria sp. E1-E4, H 3 Ei, Ea, Hi, H3, J3, K2, R Laminariocolax tomentosoides (Farl.) Kylin G i , G2 Polysiphonia urceolata (Lighti, ex Dillw.) Grev. B4, D2 -D 4i E1-E 5 , Gi, Ga, Ja, Kí-K3, Ka, R Polysiphonia violacea (Roth) Spreng. Leptonematella fasciculata (Reinke) Silva E i—E3i Es Aa, B4 Polysiphonia sp. Litosiphon filiformis (Reinke) Batt. G i , G2 A7 i D i—D3, Ds, Hi—Ha Porphyra leucosticta Thur. in Le Joi. Petalonia fascia (O. F. Müll.) O. Kuntze Ag, D i—Ds, Ex—Es, Ki, R R Porphyra linearis Grev. R Petalonia zosterifolia (Reinke) Kuntze R Porphyra purpurea (Roth) C. Ag. b 2 Pilayella littoralis (L.) Kjellm. R Porphyra umbilicalis (L.) J. Ag. A5 , Ag, Bí—B7, Ki, R Punctaria hiemalis Kylin* J 2 Porphyra sp. Di, D4, Hi Punctaria plantaginea (Roth.) Grev. Ei, Ki, K2, K s Rhodochorton floridulum (Dillw.) Nag. d 3 Ralfsia verrucosa (Aresch.) J. Ag. K i , R Rhodochorton purpureum (Lighti.) Rosenv. Ba, B3, B5 Scytosiphon lomentaria (Lyngb.) lin k Ja, R Rhodomela confervoides (Huds.) Silva R Sargassum muticum (Yendo) Fensholt M. Janke, Schilling (pers. comm.) Rhodomela virgata Kjellm. D4 Spongonema tomentosum (Huds.) Kütz. R Rhodomela sp. Ex—E5 , Gi, G 2 Sphacelaria caespitula Lyngb. K3 Trailliella intricata Batters D3i E3i Gi, Ga, H 1-H 3 Sphacelaria plumosa Lyngb. E i, Ki, Ka, K? Sphacelaria radicans (Dillw.) C. Ag, K! Red crust algae Dermatohthon pustulatum (Lamour.) Fosl. Gi, G2 Rhodophyceae Hildenbrandia rubra (Sommeri.) Menegh. Ei-E3, Gi, G2, Ki, IQ-K7, R Ahnfeltia plicata (Hudson) Fries B4 , D4 , Ds, Ei, R Melobesia membranacea (Esper) Lamour. Gi, Ga, H 1-H 3 Antithamnion plumula (Ellis) Thur. in Le Jolis d 4, k 7 Petrocelis hennedyi (Harv.) Batt. R Audouinella membranacea (Magnus) Papenfuss Gi, G2i Kj, Ka, K4, K7i Kb Phymatolithon laevigatum (Fosl.) Fosl. R Bangia atropurpurea (Roth) C. Ag. Bs, R Phymatolithon lenormandii R Brongniartella byssoides D 3 , E3, E4 (Aresch. in J. Ag.) Adey (Good, et Woodw.) Schmitz Phymatolithon polymorphum (L.) Fosl. R Callithamnion hookeri (Dillw.) S. F. Gray B4, B7i Da, D4i K i , K s Phymatolithon sp. Ki, Ks—K7 Ceramium deslongchampsn Chauv, in Duby D 5, Ei, Kj, K2, Kj, K b, R Rhodophysema elegans R Ceramium rubrum (Huds.) C. Ag. Ag—Ag, Bx—B4, B7i Dg—D5 , Ei, E2, Gi, G2, H x—H3, (Crouan frat, exJ. Ag.) Dixon J2, J3, Ki, K2i K7l R Cyanophyta Chondrus crispus Stackh, A 2 —A4, Ag—Ag, Ba, B4, Bg, B7, Dx, Ei, G2, R Spirulina sp. Hi Corallina officinalis L. 6 A2, A3, A -A 9i Bj, Et, E2, Gj, G2, H2, Kx, K3, R Arthospira sp. Hi Cystoclonium purpureum (Huds.) Batt. Di, Ei—E3i Eg, G2i R Delesseria sanguinea (Huds.) Lamour, D 1-D 5 , Ei-Es, Gi, Ga, Ki, K2 Diatomeen Dumontia incrassata (O. F. Müll.) Lamour. A4( Ag, A7, Ag, R Benthic diatoms Ei Furcellaria lumbricalis (Huds.) Lamour. R Goniotrichum alsidn (Zanard.) Howe J3 Lomentaria clavellosa (Turn.) Galil. Di, D 31 Ea, E3, E5i Hi, Ki, Ki Lomentaria orcadensis (Harv.) Coli, ex Taylor K6 J. H arm s 14 Species list Helgoland 15

Table 3, Check list of benthic protozoa and invertebrates settling around Helgoland as described in Table 3 (Continued) recent publications. Due to methodical difficulties (diving) or because individuals were juveniles, individuals were not always determined to species level; therefore, individuals marked as "sp," do not necessarily represent additional species to the already listed members of the same genus. Hydrozoa, Athecata Synonyms used in one of the summarized publications are given in brackets; o: Taxonomic Bougainvillia sp. classification uncertain; for more information see Table 1 Ii Clava multicornis (Forskàl, 1775) A2, A3 , Aß—Ag, Bj, B2, Bí, B5 , B7, H3 , Ii, Ji Corymorpha sp. 0 n 2 Coryne pusilla Gärtner, 1774 Rhizopoda, Foraminifera A2i Ae-Ag, Bí, B2, B4 -B 7 , Gi, G2, Hi—H 3 Coryne sarsii (Lovén, 1835) Ks Miliolinae sp. M Eudendrium rameum (Pallas, 1766) A8 i Ht-Ha Ciliata Eudendrium sp. Ii, I2 , Kj, IQ, KSl IQ, N Ascobius fauré-fremieti Fauré-Fremiet, 1932 S Hydractinia echinata (Fleming, 1828) n 2 Ascobius simplex (Dons, 1917) Hadzi gen. S Perigonimus repens (Wright, 1858) 0 J i Eufolliculina uhligi Mulisch & Patterson, 1983 S Perigonimus sp. Ii Lagotia obstétrica (T. S. Wright, 1862) Hadzi gen, S Sarsia tubulosa (M. Sars, 1835) Jarms (pers. comm.) Lagotia spirobis (Dons, 1912) Donsgen. S Sarsia sp. h Lagotia viridis T. S. Wright, 1858 S Tubularia indivisa Linnaeus, 1758 E2, Eg, IQ, K5 Lagotia sp. S Tubularia larynx EUis & Solander, 1786 Ds, E!-E4i Ha, I 1-I 5 , J 1-J 3 , K2i K4i Ks Metafolliculina andrewsi Hadzi, 1938 S Tubularia sp. N Parafolliculina violacea (Giard, 1888) Dons gen. S Zoothamnium sp. Harms (pers, observ.) Hydrozoa, Thecata Abietinaria abietina (Linnaeus, 1758) Ks, N Porifera Calycella syringa (Linnaeus, 1767) Hi, Ki, K2l IQ, Ks, N Acervochalina loosanoffi (Hartmann, 1958) IQ Calycella sp. Q Biemna variante (Bowerbank, 1866) E3, K 5 Campanularia sp, In I2 , M 1 5 Cliona celata Grant, 1826 E -E 4, IQ, K Campanulina hincksii Harüaub, 1897 H t,H 2 Cliona lobata Hancock, 1849 N2 Clytia hemisphaerica (Linnaeus, 1767) Hi-Ha, L, I4, Jx, J2, Kx, IQ, K5, IQ, N Cliona sp. N [Campanularia johnstoni (Alder, 1856), Dysidea fragilis (Montagu, 1818) Da Laomedea pelagica (Van Breemen, 1905)] Esperiopsis fucorum (Esper, 1974) E2—E4, K3 Diphasia rosacea (Linnaeus, 1758) Ex—E4, K j, K j , K4i IQ, N [Amphilectus fucorum (Esper, 1794)] [Nigellastrtim rosaceum (Oken, 1815)] Halichondria bowerbanki Burton, 1930 D¡, D4, E lf E2, K7 Dynamena pumila (Linnaeus, 1758) A2i A3i As—Ag, Bx—B7, Gi, G2 ,Hx—H3 ,Kx, K2i Ky [Halichondria coalita (Grant, 1825)] Gonothyraea loveni Allman, 1864 Jarms (pers comm.) Halichondria panicea (Pallas, 1766) A2, A 6-A g, Dx-D4, B4 -E 4, Gi, G2, Hx-H3, Hartlaubella gelatinosa (Pallas, 1766) Ag, As, Gi, G2, J 3 Kr-Kg, K7, Ka [Laomedea gelatinosa (Pallas, 1766), Haliclona oculata (Pallas, 1766) E3, E4, IQ Obelia gelantinosa (Pallas, 1766)] Haliclona rosea (Bowerbank, 1866) Klr K4, K 7 Halecium halecium (Linnaeus, 1758) IQ, N Haliclona urceolus (Rathke & Vahl, 1807) K 4 Hydrallmania falcata (Linnaeus, 1758) Ae, A7l Bs, Ex-E 3, E5i e j, H2 i H3 i K„ K j , IQ, N Halisarca dujardini Johnston, 1842 A7 -A 9l D3, D5, E2, G2, G2r H 1 -H 3 Kirchenpaueria pinnata (Linnaeus, 1758) D1( Ex, H3( Ki, K s Leucandra fistulosa [Johnston, 1842) ' 1 N 2 Lafoea dumosa (Fleming, 1820) H i , K j Leucandra johnstoni (Carter, 1871) Ka [Lafoea fruticosa (Sars, 1850)] [ieucozua johnstoni (Carter, 1871)] Laomedea calceolifera Hincks, 1871 N Leucosolenia botryoides (Ellis & Solander, 1786) A e-A g, Blt B3-B 5, Hx-H 3 [Laomedea conferta (Hartlaub, 1897)] Leucosolenia complicata (Montagu, 1812) A2, G2 Laomedea ßexuosa Alder, 1857 A2 —Ag, Bx—B7, Dx—Da, Ex, Es, Gx, G2i Hx, H3, Leucosolenia variabilis (Haeckel, 1870) . D i-D j, E .-E 4, I 4 —15, K2, Ks, K7, Ks Ix- I3 , J i—J3, Kx, K3 Mycale macilenta Topsent, 1924 K4 Myxilla incrustans (Johnston, 1842) E2, K2, K4, Ks Laomedea neglecta Alder, 1856 N Laomedea sp. d 3, n Oscarella lobularis (Schmidt, 1862) D1( D 3 -D 5i H 4, H2, IQ Obelia dichotoma (Linnaeus, 1758) B2r B3, B7i D3i D4i Gx, G2i H i , H2i J2i J 3, K i , Pleraplysilla minchini Topsent, 1905 IQ, K5 [Laomedea dichotoma (Linnaeus, 1758), Polymastia mammillaris (O. F. Müller, 1806) K4 IQ-Ka, N Laomedea longissima (PaUas, 1766), Scypha ciliata (Fabricius, 1780) A7, D 1-D 5 , Ex-E3, G2, Hx-H3, Ii-I4,Jt- J 3, K1( [Sycon cihatum (Fabricius, 1780)] K4, K7, Kb Obelia longissima (Pallas, 1766)] 5 Scypha coronata (EUis & Solander, 1786) ASl B4, G 2 Obelia geniculata (Linnaeus, 1758) A —A7, B4i B5i Dx, D3i D4i Ex—E3i Gx, G2i [Syconcoronatum (Ellis & Solander, 1786)] [Laomedea geniculata (Linnaeus, 1758)] H 1-H 3 , Ix, I3, Q, J1-J 3 , Kt, K3, IQ, N Suberites ficus J. E. Gray, 1806 Ei, Ka Opercularella lacerata (Johnston, 1847) IQ J. H arm s 16 Species list Helgoland 17

Table 3 (Continued) Table 3 (Continued)

Hydrozoa, Thecata Nemertini Plumularia setacea (Linnaeus, 1758) Dj, Ei—Ea, Ej, G u G 2, H 2i H 3i Ki—K4, N Oerstedia dorsalis (Abildgaard, 1806) A2i A3i Ag—Ag, Gj, G2, H 1-H 3 [Polyplumularia setacea (Linnaeus, 1758)] Oerstedia striata Sundberg, 1988 Sundberg (pers. comm.) Sertularella polyzonias (Linnaeus, 1758) Ka, K4, Ks Procephalothrix filiformis (Johnston, 1828-29) a 8, g 2 Sertularella rugosa (Linnaeus, 1758) Ei, G2, Hi, H2, K2i K4, Ka, N Tetrastemma candidum (Müller, 1774) A2i A3i Ay, Ag, H3 Sertularia cupressina (Linnaeus, 1758) A3, Aß—Ag, B5 , Dg, Ei—E3l H i—H3, Ki, K2, K4, N Tetrastemma helvolum Bürger, 1895 h 3 Sertularia tenera G. O, Sars, 1874 N Tetrastemma vermiculus (Quatrefages, 1846) A 3 , Ay, Ag, G2, H i—Hg Sertularia sp. II, M Mollusca, Polyplacophora Scyphozoa Lepidochitona cinerea (Unneaus, 1767) A3 —A4i Ag—Ag, Ha, K,, Kg Aureüa aurita (Linnaeus, 1746) Ae, Ag, D3, G2, Hi, Ij, Ji, J 2 Leptochiton asellus (Gmelin, 1791) A2, A3 , Ag—Ag, M Craterolophus tethys (Clark, 1863) Ae-Ag Mollusca, Gastropoda, Prosobranchia Anthozoa Acmaea virginea (Müller, 1776) * At--A4, Ag—Ag, Bg, G21 I31 I5 Ag Actinia equina (Linnaeus, 1758) Aporrhais pespelecani (Linnaeus, 1758)' M Actinothoe sphyrodeta (Gosse, 1853) e 2 Buccinum undatum Linnaeus, 1758 3 E3, E4i Ka, K4, Kg, Kf, M, N A , Ag—Ag, M, N Alcyonium digitatum Linnaeus, 1758 Calliostoma zizyphinum (Linnaeus, 1758) ' Ag Cerianthus lloydii Gosse, 1859 Kb, M, N [Calliostoma conuloide (Lamarck, 1822)] Diadumene cincta Stephenson, 1925 Ei—E4i Ka, K4-Ka Chrysallida obtusa (Brown, 1827) h 2 Edwardsia sp. M, N Chrysallida pellucida (Dillwyn, 1817) h 2 N Haliplanella lineata (Verrili, 1869) *' [Partulida spiralis (Montagu, 1803)] A2, A3, Ag—Ag, B4, B5 , By, D i—D3, Dg, Ei—E4, Gi, Metridium senile (Linnaeus, 1761) Crepidula fornicata (Linnaeus, 1758)' Harms (pers, observ.) G2, H !-H 3, I4, Ig, K —Ka, N Epitonium clathrus (Linnaeus, 1758) ’ M Metridium senile var. pallidus (Rawlinson, 1934) Goemann (pers, comm.) [Clathrus clathrus (Linnaeus, 1758)] Sagartia elegans (Dalyell, 1848) A2, Af, Ag, Di—D5i E!—Eg, Ilf I3l I4, Ki—Ki, Kg Gibbula cineraria (Linnaeus, 1758) A2 -A b, B1( Gi, G2i H i-H 3, Ky-Ky, N Sagartia troglodytes (Price, 1847) Ab, Ei, Ki-Ki, Ke, N Gibbula tumida (Montagu, 1803) M Sagartiogeton laceratus (Dalyell, 1848) Ay Helcion pellucidum (Linnaeus, 1758)’ K. Janke (unpubl.) Sagartiogeton undatus (O. F. Müller, 1788) A3, Ag—Ag, Di, D3—Dg, E¡—E3, Ii, I4, Kg [Patina pellucida (Linnaeus, 1758)] Urticina felina (Linnaeus, 1761) A2—A4, Ag—Ag, Dg, Ei—E5 , Ki, K2, Ki, Ky, N Hydrobia ulvae (Pennant, 1777) A3 , Ag—Ag, G1( Ga. M, N [TeaEa felina Stephenson, 1935] Lacuna pallidula (da Costa, 1778) A2 , A3 , Ag—Ag, Gi, G 2 Kamptozoa Lacuna vincta (Montagu, 1803) A2, A 3 , Ag—Ag, Bi, Ba. Bg, B7, G 1, G2| H i—H 3 Barentsia sp. Ii [Lacuna divaricata (Fabricius, 1780) non Linné] Lamellaria latens (Müller, 1776) K. Janke (unpubl.) Pedicellina cernua (Pallas, 1774) A 7 —Ag, H i—H 3 , Ii, I3 Pedicellina nutans Dalyell, 1848 Dx Lamellaria perspicua (Linnaeus, 1758) K. Janke (unpubl.) Littorina littorea (Linnaeus, 1758) Aj—Ag, B,—B7, Ha Nemathelminthes, Nematoda Littorina mariae Sacchi & Rastelli, 1966 Ag, B2 -B 7, Hj, Ha Oncholaimus sp. N Littorina neritoides (Linnaeus, 1758) ' K. Janke (unpubl.) Plathelminthes, Turbellaria Littorina obtusata (Linnaeus, 1758) A 2—Ag, Hg Littorina saxatilis (Olivi, 1792) Aa—Ag, Ag, B Ll Ba. B4—By Cryptocelis alba Lang, 1884 N Littorina saxatilis var. rudis (Maton, 1797) K. Janke (unpubl.) Stylochoplana agilis Lang, 1884 Ha, H3 Lunatia alderi (Forbes, 1838) M Turbellaria sp. Iii I41 Is [Natica alderi Forbes, 1838] Nemertini Lunatia nitida (Donovan, 1800) N N Amphiporus bioculatus McIntosh, 1873-74 G 1( G2 Lunatia catena (da Costa, 1778) N Amphiporus lactifloreus (Johnston, 1827-28) a 7 Nassarius (Hima) incrassatus (Strom, 1768) Cephalothrix rufifrons (Johnston, 1837) Ag, Ag Nucella lapillus (Linnaeus, 1758) ’ A6 p Ag Emplectonema gracile (Johnston, 1837) Ag Omalogyra atomus (Philippi, 1841) K. Janke (unpubl.) Bg, Gi, Ga. H2. Ha Lineus bilineatus (Renier, 1804) A3 , Ag—Ag Onoba aculeus (Gould, 1841) Ag~Ag Lineus longissimus (Gunnerus, 1770) Ia Onoba semicostata (Montagu, 1803) [Cingula striata (Montagu, 1803)] Lineus ruber (Muller, 1771) A2, Ag, Ab—Ag, Gi, G2, H3i I 4 Lineus sanguineus McIntosh, 1873 Sundberg (pers. comm.) Patella vulgata Linnaeus, 1758' K. Janke (unpubl.) Lineus viridis (Müller, 1774) Aa Pusillina inconspicua (Aider, 1844) Ag, J2| J 3 [Rissoa inconspicua Aider, 1844] Micura fasciolata Ehrenberg, 1831 A7 , G 2 J. H arm s 18 Species list Helgolandand 19

Table 3 (Continued) Table 3 (Continued}

Mollusca, Gastropoda, Prosobranchia Mollusca, Bivalvia Pusillina parva (da Costa, 1778) Ae, Ag, Bt-B7, Ei, Gi, G2, Hi—H3, I1( I4, I5, M, Abra alba (W. Wood, 1802) M , N [Rissoa parva (da Costa, 1778)] Kl—Ky Abra nitida (Müller, 1776) M, N Raphitoma linearis (Montagu, 1803) H i Acanthocardia echinata echinata M Skeneopsis planorbis (Fabricius, 1780) A 3, Ag, A 7 , Ag, B i, B4 —B7 (Linnaeus, 1758) Turritella communis Risso, 1826* Anger (pers. comm.) Aequipecten opercularis (Linnaeus, 1758) Anger (pers. comm.) Anomia ephippium Linnaeus, 1758 Ag Mollusca, Gastropoda, Opisthobranchia Arctica islandica (Linnaeus, 1767) M , N Acanthodoris pilosa (Müller, 1789) A 7 , Ag, Ki, K4 , Kg, K7 , N [Cyprina islandica (Linnaeus, 1767)] Acteon tornatilis (linnaeus, 1758) M Arenomya arenaria (Linnaeus, 1758) BS, N Aeolidia papillosa (Linnaeus, 1761) A7 i Aa, N [Mya arenaria (Linnaeus, 1758)] Antiopella cristata (Chije, 1841) K4 Cerastoderma edule edule (Linnaeus, 1758) N 2 [Janolus cristatus (Chiaje, 1841)] [Cardium edule (Linnaeus, 1758)] Antiopella sp. N Cerastoderma sp. g 2 Aplysia punctata Cuvier, 1803 # Schilling (pers. comm.) Chamelea striatula (da Costa, 1778) M, N Archidoris pseudoargus (Rapp, 1827) Aß—Ag, Kj—Kg [Venus striatula da Costa, 1778] Coryphella gracilis (Alder & Hancock, 1944) G2 Cochlodesma (Bontaea) praetenue M, N Coryphella pedata (Montagu, 1815) Ag (Pulteney, 1799) Corphella pellucida (Alder & Hancock, 1843) K4 Corbula (Varicorbula) gibba (Olivi, 1792)*2 N Coryphella verrucosa ruübranchialis A 7 , Ag Donax vittatus (da Costa, 1778) N (Johnston, 1832) Dosinia exoleta (Linnaeus, 1758) N 2 Coryphella sp. N Ensis siliqua (Linnaeus, 1758) N Cuthona concinna (Alder & Hancock, 1843) g 2, N Ensis sp. N2 Gari fervensis (Gmelin, 1791) M Cuthona foliata (Forbes & Goodsir, 1839) Hs. It, I3 Cuthona gymnota (Couthouy, 1838) N Goodallia triangularis (Montagu, 1803) N 2 Cylichna cylindracea (Pennant, 1777) M [Astarte triangularis (Montagu, 1803)] Hiatella arctica (Linnaeus, 1767) Ag—Ag, Gj, G2, Ji , M Dendronotus frondosus (Ascanius, 1774) Aa, K4l N [Saxicava arctica (Linnaeus, 1767)] Doris sp. N Hiatella rugosa (Linnaeus, 1767) Ag-Ag, N Doto coronata (Gmelin, 1791) A8, N [Saxicava rugosa (Pennant, 1777)o] Eubranchus exiguus (Alder & Hancock, 1848) H i. H3l Ii, la, N 2 Mactra stultorum (Linnaeus, 1758) N Eubranchus tricolor Forbes, 1838 K. Janke (unpubl,) [Mactra corallina (Montagu, 1808)] Eubranchus sp. N M odiolus aff. barbatus (Linnaeus, 1758) N Facelina auriculata (Muller, 1776) As Modiolus modiolus (Linnaeus, 1758) Ei, Gi, G2, M ,N Facelina auriculata coronata Ii Mysella bidentata (Montagu, 1803) M, N (Forbes & Goodsir, 1839) [Montacuta bidentata (Monatgu, 1803)] Facelina sp. J 21 J 3 Mya truncata Linnaeus, 1758 Gi. Ga, M Hermaea dendritica (Alder & Hancock, 1843) K. Janke (unpubl.) Mytilus edulis Linnaeus, 1758 A i—A9, Bj-By, D 2i Gi, G2i Hi—H3, L—15, J 1—J31 Jorunna tomentosa (Cuvier, 1804) K. Janke (unpubl.) Ki—K4, M Limacia clavigera (Müller, 1776) N Nucula nucleus (Linnaeus, 1758) G2,M Limapontia depressa Alder & Hancock, 1862 K. Janke (unpubl.) Nucula nitidosa Winckworth, 1930 M, N 2 Limapontia senestra (Quatrefages, 1844) K. Janke (unpubl.) [Nucula turgida Leckenby & Marschall, Onchidoris bilamellata (Linnaeus, 1767) Ga. Ii 1875¡ Nucula nitida Sowerby, 1833 Onchidoris muricata (Muller, 1776) Aï . Ag, Gi, G2 non Brocchi, 1814] Palio dubia (M. Sars, 1829) Ii Petricolaria pholadiformis (Lamarck, 1818) E1-E3 * 2 M, N Philine punctata (Adams, 1800) H 2 Phaxas pellucidus (Pennant, 1777) Polycera quadrilineata (Müller, 1776) A71 Ag, H2l Ka, K4, N [Cultellus pellucidus Pennant, 1777] Retusa obtusa (Montagu, 1803) K. Janke (unpubl.) Pholas dactylus Linnaeus, 1758 E2 Retusa truncatula (Bruguiére, 1792) As Spisula elliptica elliptica (Brown, 1827) M Spisula solida (Linnaeus, 1758) G2, N Tergipes tergipes (Forskâl, 1775) Ii. la Tritonia plebeia Johnston, 1828 N Spisula subtruncata (da Costa, 1778) M, N M Tritonia sp. N Tellimya ferruginosa (Montagu, 1808) [Montacuta ferruginosa (Montagu, 1808)] 20 J. H arm s Species list Helgoland 21

Table 3 (Continued) Table 3 (Continued)

Mollusca, Bivalvia Annelida, Polychaeta, Errantia Tellina (Moerella) donacina (Linnaeus, 1758) N 2 Harmothoe lunulata (Delle Chiaje, 1841) M [Angulus donacinus (Linnaeus, 1758)] Harmothoe sp. M Tellina (Fabulina) fabula Gmelin, 1791 M, N Kefersteinia cirrata (Keferstein, 1862) C [Tellina fabula Gronovius, 1781] Lepidonotus squamatus (Linnaeus, 1767) Ag, A7, C, K4, N Tellina (Angulus) tenuis teams da Costa, 1778 N Microphthalmus sczelkowii Mecznikovi, 1865 Ab, C Teredo navalis Linnaeus, 1758 A3 Nephtys caeca (Fabricius, 1780) M, N Thracia papyracea papyracea (Poli, 1791) M N ephtys cirrosa Ehlers, 1868 N [Thracia phaseolina (Lamarck, 1818}] N ephtys hombergii Savigny, 1818 I4, M, N Thracia villosiuscula (MacGillivray, 1827) n 2 Nephtys longosetosa 0rsted, 1843 N 2 Thyasira fíexuosa (Montagu, 1803) M Nereis (Hediste) diversicolor O. F. Müller, 1776 A2i Ag, I4, J 2, J3 Timoclea ovata (Pennant, 1777)*2 M Nereis elitoralis Eliason, 1962 M [Vanus ovata Pennant, 1777] N ereis longissima Johnston, 1840 M Venerupis senegalensis (Gmelin, 1791) Ai, As, N Nereis pelagica linnaeus, 1761 A3, Ag-Ag, By-B4l C, G^, G2, H i , H 2i Ii , Lt> Is, [Venerupispullastra (Montagu, 1803)] Ja,N C Zirfaea crispata (Linnaeus, 1758) k 4 Nereis (Neanthes) succinea Frey & Leuckart, 1847 Mollusca, Cephalopoda Nereis (Neanthes) virens Sars, 1835 A2, A3, Ag, C, M Alloteuthis media (Linnaeus, 1758) M Ophryotrocha gracilis Huth, 1934 Ag, C Loligo sp. M Ophryotrocha puerilis siberti (McIntosh, 1885) H2l J 1( J 2 Sepia officinalis Linnaeus, 1758 Goemann, M. Janke (pers. comm,) Pholoe m inuta (Fabricius, 1780) C, M, N M, N Sipunculidea Phyllodoce (Anaitides) groenlandica 0 rsted , 1843 Golfingia minuta (Keferstein, 1862) As, Ag Phyllodoce (Anaitides) lineata (Claparède, 1870) M Golfingia sp. g 2, n 2 Phyllodoce (Anaitides) maculata A i, A3, Ag—Ag, B2l B3, C, G i, G2| H* Ii, I3—Is, Echiuroidea (Linnaeus, 176?) M, N M, N Echiurus echiurus (Pallas, 1774) # * * * Phyllodoce (Anaitides) mucosa 0rsted, 1843 M Phyllodoce (Anaitides) rosea (McIntosh, 1877) M, N 2 Annelida, Polychaeta, Errantia [Anaitides subulifera Eliason, 1962] Ji-J3, M Aphrodita aculeata Linnaeus, 1761 M Phyllodoce sp. Pirakia punctifera (Grube, 1860) C Autolytus edwardsi Saint-Joseph, 1887 C G | G2 Autolytus prolifer (O, F. Müller, 1776) [Eumidia punctifera (Grube, I860)] A g-Ag, Bi-By, C, Gi, G2, H1- H 3i I1-I4, J lr J3, N Pisione remota (Southern, 1914) h 2, h Brania pusilla (Dujardin, 1839) C, h 2 Polynoe (Enipo) kinbergi Malmgren, 1865 M Eteone (Mysta) barbata Malmgren, 1865 N Eteone flava (Fabricius, 1780) Proceraea cornuta (Agassiz, 1862) G i, G2 N 2 Protodorvillea kefersteini (McIntosh, 1869) M, N Eteone longa (Fabricius, 1780) C, M, N C, M, N Eteone sp. Sphaerodorum flavum 0rsted, 1843 N 2 [Spaerodorum gracile (Rathke, 1843)] Eulalia bilineata (Johnston, 1840) N Spaerosyllis hystrix Claparède, 1863 C, M Eulalia expusilla Pleijel, 1987 Hi Sthenelais boa (Johnston, 1839) A8i C, G i [Eulalia pusilla 0rstedt, 1843] Sthenelais limicola (Ehlers, 1864) M, N2 Eulalia viridis (Linnaeus, 1767) A 2, A3, Ag—As, Bí, C, G i, G2, Ha, It—lg, Jf-J3, N Streptosyllis websteri Southern, 1914 G i, G2 Eumidia sanguinea 0rsted, 1843) A 2, A 6i C, G2, N Syllides articulocirratus Gillandt, 1979 A8i C Eusyllis blomstrandi Malmgren, 1867 H t- H 3, N Syllides longocirrata 0rsted, 1845 N Exogone naidina 0rsted, 1845 C Syllis gracilis Grube, 1840 C, N 2 Gattyana cirrosa (Pallas, 1766) Ag, C, M, N Typosyllis armillaris (O. F, Müller, 1776) Ag, Ag, C, G2 Glycera capitata 0rsted, 1843 M, N Typosyllis hyalina (Grube, 1863) C Glycera rouxii Audouin & Milne-Edwards, 1833 N 2 Glycinde nordmanni (Malmgren, 1865) M, N Annelida, Polychaeta, Sedentaria Glycinde sp. M Ampharete finmarchica (M. Sars, 1864) M, N Goniada maculata 0rstedt, 1843 M, N Amphitrite figulus (Dalyell, 1853) Ae, C, Gi, G 2 Goniadella bobretzkii (Annenkova, 1929) N Anobothrus gracilis (Malmgren, 1865) M [Sosane gracilis (Malmgren, 1865)] Harmothoe imbricata (Linnaeus, 1767) A7, Ii, I4 Aonides paucibranchiata Southern, 1914 M, N Harmothoe impar (Johnston, 1839) Ag—Ag, C, J f M, H 22 J. H arm s Species list Helgoland 23

Table 3 (Continued) Table 3 (Continued)

Annelida, Polychaeta, Sedentaria Annelida, Polychaeta, Sedentaria Arenicola marina (Linnaeus, 1758) A^—A3 , C Polycirrus sp. M, N Capitella capitata (Fabricius, 1780) Ai—A3 , C, Jj, J 2 , M2 Polydora antennata Claparède, 1868 N Capitella giardi (Mesnil, 1897) A 2 , A3, C Polydora caeca (0rsted, 1843) N {Capitomastus giardi (Mesnil, 1897)] Polydora ciliata (Johnston, 1838) A 2—A4, Ag—Ag, Bj, B2-B s, C, Ej—E3, G i, G2, H|, Capitella minima (Langerhans, 1880) M [Capitomastus minimus (Langerhans, 1880)] H 3 , B, I31 •II- J 3 Polydora ligni Webster, 1879 C, Ji, J 2 Capitella sp. G2 Polydora pulchra Carazzi, 1895 M, N2 Caulleriella bioculata parva Gillandt, 1979 C Polydora quadrilobata Jacobi, 1883 g 2 Caulleriella caput-esocis (Saint-Joseph, 1894) c , h 3 Pomatoceros triqueter (linnaeus, 1758) Ag—Ag, B1-B4, C, D1-D5, B1-E3, Es, Gi, G2, Hj, Chaetozone setosa Malmgren, 1867 g 2, n 2 H2, Ii , I3—15, J i—J3, K5, Kg, M, N Chone duneri Malmgren, 1867 M C Chone infundibuliformis Kroyer, 1856 M, N Pygospio elegans Claparède, 1863 g 2 Cirratulidae sp. M Sabellaria alveolata (Linnaeus, 1767) Sabellaria spinulosa Leuckart, 1849 A8 i C, N Cirratulus cirratus (O, F. Muller, 1776) G2 Scalibregma inflatum Rathke, 1843 M, N Cirratulus sp. II Cossura longocirrata Webster & Benedict, 1887 N Scolelepis bonnieri (Mesnil, 1896) M, N A i—A3i A5i C Ctenodrilus serratus (Schmidt, 1857) M, Janke (pers. comm.) Scolelepis squamata (Müller, 1789) Diplocirrus glaucus (Malmgren, 1867) M Scoloplos armiger (O, F. Müller, 1776) M, N Dodecaceria concharum 0rsted, 1843 C Spio armata (Thulin, 1957) N [Zeppelina monostyla (Zeppelin, 1883), Spio filicornis (O. F, Müller, 1766) M, N Zeppelina mediopigmentata Gillandt, 1979] Spio goniocephala Thulin, 1957 N Spiophanes bombyx (Claparède, 1870) M, N Fabricia stellaris (Müller, 1774) A2 —Ag, B i—B7, C, Gi, G2, Hi—H3, Ii, J i [Fabricia sabella (Ehrenberg, 1836)] Spirorbis spirillum (linnaeus, 1758) A2, Ag—Ag, B,-., G i, G2, H i—H3 Flabelligera affinis M, Sars, 1829 C, N [Circeis spirrillum (linnaeus, 1758)] Hydroides norvegica Gunnerus, 1768 Harms (pers. observ.) Spirobis spirobis (linnaeus, 1758) A2, Ag—Ag, Bí, Bs, Bfi, C, Gi, G2, H1 Uj Hypania invalida (Grube, 1860) M Spirobis tridentatus (Levinsen, 1883) A2, A3, A5—Ag, S3, B5, C, Gi, G2, H 1-H 3 , Ii—I5 Di, Ds, Ki, K? Janua (Dexiospira) pagenstecheri A2 —A4 , Ae—Ag, B2i Bs, Bg, C, G t, G2l H1-H3, Spirobis sp. (Quatrefages, 1865) Terebellides stroemi M. Sars, 1835 M Ï1-I 4 [Spirobis pagenstecheri Quatrefages, 1865] Tharyx marioni (Saint-Joseph, 1894 M A 7, Ag, I Janua sp. Ki, K6 Tharyx multibranchiis (Grube, 1863) C Lanice conchilega (Pallas, 1766) N A i—A3, Ag—Ag, C, Ei, B2, G i , G 2i Ji, J 2, K i, K4, Thelepus cincinnatus (Fabricius, 1780) Kg, M, N Travisia forbesi Johnston, 1840 n 2 Lanice sp. M Annelida, Interstitial Polychaeta Lysilla loveni Malmgren, 1865 M Magelona alleni Wilson, 1958 M, N Dinophilus gyrociliatus O. Schmidt, 1857 Ji Q Magelona papillicornis F, Müller, 1858 N Diurodrilus minimus Remane, 1925 Nerilla antennata O. Schmidt, 1848 Q Malacoceros fuliginosus (Claparède, 1868) Ai, Aa, Ag, A7, C, H3, M Q Malacoceros vulgaris (Johnston, 1827) C, M Nerillidium gracile Remane, 1925 Myriochele oculata Zaks, 1923 M Nerillidium troglochaetoides Remane, 1925 Q Polygordius appendiculatus Fraipont, 1887 Q Nicolea venustula (Montagu, 1818) h Nicolea zostericola (0rsted, 1844) Polygordius lacteus (Schneider, 1868) Q Bí, C, Gi, G2, H!-H3i Ï1-I 5 , J3, M, N N Notomastus latericeus Sars, 1851 M Polygordius sp. Ophelia limacina (Rathke, 1843) M, N Protodriloides chaetifer (Remane, 1926) Q Ophelina acuminata 0rsted, 1843 M, N Protodriloides symbioticus (Giard, 1904) Q R Orbinia sertulata Savigny, 1820 N 2 Protodrilus adhaerens Jäger s ten, 1952 Owenia fusiformis Delle Chiaje, 1841 M, N Protodrilus ciliatus Jägersten, 1952 Q Pectinaria koreni Malmgren, 1865 M, N Protodrilus gracilis von Nordheim, 1989 Nordheim (1989) Pherusa ßabellata (Sars, 1872) C Protodrilus helgolandicus von Nordheim, 1983 Nordheim (1983) Pherusa plumosa (O. F, Müller, 1776) C, M Protodrilus oculifer Pierantoni, 1908 Q Poecilochaetus serpens Allen, 1904 M Protodrilus purpureus (Schneider, 1868) Q Polycirrus aurantiacus Grube, 1860 M Trilobodrilus axi Westheide, 1967 O Polycirrus medusa Grube, 1850 M, N Trilobodrilus heideri Remane, 1925 Q J, H arm s 24 Species list Helgoland 25 Table 3 (Continued) Table 3 (Continued)

Annelida, Oligochaeta Crustacea, Copepoda, Harpacticoida A2—A4, Ag Paranais litoralis (Müller, 1788) Parategastes sphaericus (Claus, 1863) P Annelida, Hirudinea Parathalestris harpactoides (Claus, 1863) P Callobdella nodulifera Malm, 1863 n 2 Rhizothrix minuta (Wilson, 1932) P Scutellidium hippolytes (Kroyer, 1863) P Arachnida Acari Thalestris longimana Claus, 1863 P h, I4 Acari sp. Tisbe furcata (Baird, 1837) G. Sahling (pers. comm.) Pantopoda Tisbe gracilis (T. Scott, 1895) G. Sahling (pers. comm.) Achelia echinata Hodge, 1864 h 2, h 3, n Zaus spinatus Goodsir, 1845 P Achelia hispida Hodge, 1864 Ag, Ag, Gi, G2 Crustacea, Cirripedia Anoplodactylus angulatus (Dohrn, 1881) A2i Aa, Gj, G2 Anoplodactylus petiolatus (Kroyer, 1884) G i.G a Balanus balanus (Linnaeus, 1758) Ag, Ag, D3, II, Jj, J2 Anoplodactylus pygmaeus (Hodge, 1864) Bl-B ?, Hr-Ha Balanus crenatus Bruguiére, 1789 A2, A3, Ag—Ag, Bj—B4, B„ Dj—Dj, Ej, E2i Gi, Nymphon gracile Leach, 1814 Ii. hi Ui Is G2i Hj, 1,- 14, J ,-J 3 Nymphon rubrum Hodge, 1865 A8, Aa, G,, G2, Hg, H 3i K i , Ka, K4 -K 7i N Balanus improvisus Darwin, 1854 Di. D2; D4, D 5 , 1,—I4, J ,—J3 Phoxichilidium femoratum (Rathke, 1799) A3, As, As, Ag, H 2 Elm inius m odestus Darwin, 1854 Aa—A?, Ag, Bj—By, Dlf D 3—D5, E,, E2, H,, Ii—Is, Phoxichilidium tubulariae Lebour, 1947 K. Janke (unpubl,) JlT J3 T Pycnogonum littorale (Strom, 1762) A3, A?, Ag, K4 Sacculina carcini Thompson, 1836 M. Janke (pers. comm.) Semibalanus balanoides (Linnaeus, 1767) A 2—A?, Ag, B ,—By, D j, D 2, Ds, N 2 Crustacea, Ostracoda [Balanus balanoides (Linnaeus, 1767)] Cyteromorpha aff. fuscata (Brady, 1869) O Peltogaster paguri Rathke, 1842 Scharte (pers. comm.) Elofsonella concinna (Jones, 1857)'3 O Verruca stroemia (O. F. Müller, 1776) Aß—Ag, B,, B4, D,, D2, D4p D s, E j, E2, G j, G2i H lt Eucytheridea punctillata (Brady, 1865)*3 O H2, Ij—Ig, Jj, J2, Kj, K4, K5, N Hemicythere villosa (G. O. Sars, 1866) O Hemicytherura cellulosa (Norman, 1865) O Crustacea, Mysidacea Heterocythereis albomaculata (Baird, 1838) O Gastrosaccus spinifer (Goes., 1864) M Hirschmannia viridis (O. F, Muller, 1785) O Mysis relicta Lovén, 1862 g 2 Paradoxostoma variabile (Baird, 1835) O Praunus inermis (Rathke, 1843) M Semicytherura nigrescens (Baird, 1838) O Schistomysis kervillei (G. O. Sars, 1885) M Semicytherura sella (G. O. Sars, 1866)*3 O [Paramysis kervillei (G. O. Sars, 1885) Crustacea, Copepoda, Harpacticoida Schistomysis spiritus (Norman, 1860 M Alteutha interrupta (Goodsir, 1845) P [Paramysis spiritus (Norman, I860)] Alteutha oblonga (Goodsir, 1845) P Crustacea, Amphipoda Ameiropsis aff. nobilis Sars, 1911 P M, N Amphiascoides nanus (Sars, 1906) P Ampelisca brevicornis (Costa, 1853) M, N Amphiascoides debilis (Giesbrecht, 1881) P Ampelisca diadema (Costa, 1853) Ampelisca spinipes Boeck, 1861 N Amphiascus minutus Claus,{ 1863) P Ampehsca tenuicornis Liljeborg, 1856 g 2, N Amphiascus parvus Sars, 1906 P N Amphiascus propinquus Sars, 1906 P Am pelisca sp. Am philochus m anudens Bate, 1862 M Canuella perplexa T. & A. Scott, 1893 P Amphilochus neapolitanus Della Valle, 1893 M Diarthrodes nobilis (Baird, 1845) P Amphilochus sp. N Ectinosoma melaniceps Boeck, 1864 P Amphithoe rubricata (Montagu, 1808) N Ectinosoma obtusum Sars, 1920 P A m phitoe sp. Longipedia minor T. & A. Scott, 1893 P Is Aora typica Krayer, 1845 B3, Bg, Hj—H3, N Halectinosoma gothiceps (Giesbrecht, 1881) P Apherusa bispinosa (Bate, 1857) A7, Ag, B3 , Gj, Ga, Hj—H 3 , Ïj—I4 Harpacticus uniremis Krayer, 1842 P Apherusa jurinei (Milne Edwards, 1830) Ag, Bj, B4, Bß, By, G j, Ga, H j—Hg Paraleptastacus brevicaudatus Wilson, 1932 P Paramphiascella fulvofasciata Apherusa ovalipes Norman & Scott, 1906 g 2 P N Rosenfeld & Coull, 1974 Argissa hamatipes (Norman, 1869) A tylus falcatus M etzger, 1871 N Paramphiascella vararensis (T. Scott, 1903) P Atylus swammerdami (Milne Edwards, 1830) H2, I2, 14, Is, M, N Paramphiascopis longirostris (Claus, 1863) P 26 J, H arm s Species list Helgoland 27 Table 3 (Continued) Table 3 (Continued)

Crustacea, Amphipoda Crustacea, Amphipoda Autonoe longipes (liljeborg, 1852) Ii, L, Is, M [Lembos longipes (Liljeborg, 1852)] Synchelidium tenuimanum Norman, 1895 n 2 [Synchelidium haplocheles (Sars, 1895)] Bathyporeia elegans Watkin, 1938 N 2 Bathyporeia guilliamsoniana (Bate, 1856) n 2 Unciola planipes Norman, 1867 M Bathyporeia gracilis Sars, 1891 n 2 Urothoe poseidonis Reibisch, 1905 N Bathyporeia pelagica (Bate, 1856) n 2 Crustacea, Cumacea Calliopius laeviusculus (Krayer, 1838) As, Ag—Ag, B2, Gi, G2, H 2i H3, I3 Bodotria scorpioides (Montagu, 1804) A7, Ag, H2, N Caprella linearis (Linnaeus, 1767) A7, A 8i G 1( G2i H 2i I4, N Diastylis bradyi Norman, 1879 M, N Caprella sp. Kr-Kg Diastylis cornuta (Boeck, 1864) N Cheirocratus sundevallii (Rathke, 1843) Ag, H3, L, M Diastylis laevis Norman, 1869 M N Corophium crassicorne Bruzelius, 1859 Diastylis rathkei (Krayer, 1841) M Corophium insidosum Crawford, 1937 A3, A 6—As, Bí—Bg, G lr G2i H j—H3| Ij—15, Jt , J 3 Diastylis rugosa Sars, 1865 M Corophium sextoni Crawford, 1937 N Diastyloides biplicata (Sars, 1865) N Corophium sp. M Eudorella truncatula (Bate, 1856) M Dexamine spinosa (Montagu, 1813) A 6- A b, B í , B2i G j, G 2, H j- H 3 Iphinoe trispinosa (Goodsir, 1843) M, N Dexamine thea Boeck, 1861 i i j A3, Ag—Ag, Bj, B3—B5 G i, G2 H , H 2, Ij—I3 Lamprops fasciata Sars, 1863 N Dyopedos porrectus Bate, 1857 M Pseudocuma similis Sars, 1900 M, N 2 Echinogammarus marinus (Leach, 1815) A 2i A3i A 5—A7i Ag, Bg, H lt H3 [Chaetogammarus marinus Leach, 1815; Crustacea, Tanaidacea

Gammarus marinus Leach, 1815] Tanais dulongii (Audouin, 1826) b 5 Elasmopus rapax Costa, 1853 B j, B 7 Crustacea, Isopoda Gammarella fucicola (Leach, 1814) h 3 GammareUus angulosus (Rathke, 1843) g 2 Idotea baltica (Pallas, 1772) U Gammarellus homari (Fabricius, 1779) Ae, Gj, G2i Hj Idotea chelipes (Pallas, 1766) l 2 Gammaropsis maculata (Johnston, 1828) M, N 2 Idotea emarginata (Fabricius, 1793) l 2 Gammaropsis nitida (Sthnpson, 1853) N Idotea granulosa Rathke, 1843 A2j A3 , A5—Ag, Bj—B7, Gj, G2i Hj—H3i Lj, L 2 Gammarus locusta (Linnaeus, 1758) H 3 Idotea linearis (linnaeus, 1767) I* Gammarus salinus Spooner, 1947 K. Janke (unpubl.) Idotea neglecta Sars, 1899 L2 Hippomedon denticulatus (Bate, 1857) N Idotea pelagica Leach, 1815 Lj, L2 Hyale prevostii (Milne Edwards, 1830) Aa, A3, A5, Ag, B j—B7| H j Idotea sp. I3 I5 [Hyale nilssonii Rathke, 1843] Jaera albifrons Leach, 1814 A2i A3 , A 5 —Ag, B3, B5 —B7i Ha Jassa falcata (Montagu, 1808) À 2, A3, Ag—Ag, B j—Bg, B7i G j, G 2i H j—H3i Ij—I5, Janira maculosa Leach, 1814 G j, G2i H j, M, N Ji~J3, N Janiridae sp. N Jassa marmorata Holmes, 1905 Bj-B5, B7i Gj, G2, H j-H 3 Ligia oceanica (Linnaeus, 1767) Ag, Bj—Bg Lembos websteri Bate, 1857 n 2 Limnoria lignorum (Rathke, 1799) A3 , II, Jj Megaluropus agilis Hoek, 1889 M, N Crustacea, Decapoda, Natantia Melita palmata (Montagu, 1804) A3 , A e—Ag, H3 Athanas nitescens (Leach, 1814) 7 Metopa alderi (Bate, 1857) N a Crangon allmanni Kinahan, 1857 Al, A7, M, N Microprotopus maculatus Norman, 186? Hj, Na Crangon crangon (Linnaeus, 1758) Gj , J j, m , n Orchomene nanus (Kreyer, 1846) Gj, G2, Hj, Na Parapleustes bicuspis (Krayer, 1838) N Eualus occultus (Lebour, 1936) a 7 Pariambus typicus (Krayer, 1845) M Eualus pusiolus (Krayer, 1844) N Perioculodes longimanus N Pandalina brevirostris (Rathke, 1843) a 7, n (Bate & Westwood, 1868) Pandalus montagui Leach, 1814 # N Pandalus sp. 3 5 Photis longicaudata (Bate & Westwood, 1863) M, N 2 Ii. I -I Photis reinhardi Krayer, 1842 N Thoralus cranchii (Leach, 1817) M, N Phoxocephalus holbolh (Krayer, 1842) M, N Crustacea, Decapoda, Reptantia Pontocrates arenarius (Bate, 1858) N Homarus gammarus (linnaeus, 1758), 1837*, *** K j.It, Stenothoe marina (Bate, 1857) Hj, H 2i M Stenothoe monoculoides (Montagu, 1813) Bj, B3, B7, Hj, Ha Crustacea, Decapoda, Anomura Stenula rubrovittata (Sars, 1883) n 2 Callianassa aff. subterranea Montagu, 1808’ ’ ’ M Synchelidium haplocheles (Grube, 1864) M Galathea intermedia Lilljeborg, 1851 * * * M, N 28 J. H arm s Species list Helgoland 29 Table 3 (Continued) Table 3 (Continued)

Crustacea, Decapoda, Anomura Bryozoa, Cyclostomata Galathea squamifera Leach, 1814 Ae—As, Gi, G2, Ii, I31 I4 Plagioecia sarniensis (Norman, 1864) Galathea strigosa (Linnaeus, 176?) G 2, N Ea. E 5 A3, Ae—Ae, H 2, H3, IQ—Kg, M, N Tubulipora liliacea (Pallas, 1766) K j-K o Pagurus bernhardus (Linnaeus, 1758) Tubulipora sp. Pagurus pubescens (Kroyer, 1838)*, ’ * Scharte (pers. comm,) G j. G j, Ij, I4, Is Pisidia longicornis (Pennant, 1777) Ag, I2, N Bryozoa, Cheilostomata, Ascophora

Porcellana platycheles Pennant, 1777 L, Is Cellepora pumicosa (Pallas, 1766) K4 Crustacea, Decapoda, Brachyura Celleporella hyalina (Linnaeus, 1767) As, B4, Bs , B7l Ej, G j, G2i H1-H3, Ij, L(, Is , Kj-Kg Chorizopora brongniartii (Audouin, 1826) Cancer pagurus Linnaeus, 1758 Ag—Ag, Ii, I4, K 1( Kg—K7, Ivl, N Kj, Kg Cribrilina punctata (Hassall, 1841) A a, Kj, K4-K6 Carcinus maenas (Linnaeus, 1758) A j—Ag, B j—B 7, G2, H j—H3, Ij—14, J J—J31 Kj, K j , Kb, M Cryptosula pallasiana (Moll, 1803) A2, Ag-Ag, Bi-Bs, D i-D s, Ej, Gj, G2, Hj, H3, Ii-I5, Kj-Kg, Kg Corystes cassivelaunus (Pennant, 1777) n 2 Escharella immersa (Fleming, 1828) Ag, Hj, Kj-Kg Hyas araneus (Linnaeus, 1758) Ag—Ag, H2, J j , K j , K4—K g , M , N Escharella vadolosa (Johnston, 1838) Liocarcinus holsatus (Fabricius, 1798) A7, Ab, M, N Ag, Gj, G2i K j, Ks Escharella sp. [Macropipus holsatus (Fabricius, 1798)] I4 Microporella ciliata (Pallas, 1766) Liocarcinus pusillus (Leach, 1815) * * * N Kj, Ks , Kg Schizomavella linearis (Hassall, 1841) liocarcinus sp. M Kj, K, Schizoporella errata (Waters, 1878) Macropodia rostrata (Linnaeus, 1761) N , Kj, Kg, Kg Ag Smittina landsborovii (Johnston, 1847) Macropodia sp. M Ks Maja squinado (Herbst, 1788)* N Bryozoa, Cheilostomata, Anascan Goemann (pers. comm.) Necora p uber (linnaeus, 1767)* Aetea truncata (Landsborough, 1852) H i Pilumnus hirtellus (Linnaeus, 1761) Ag-Ag, Gj, Ga, Hi, H2i I j , I3-I5, N Bicellariella ciliata (Linnaeus, 1758) k 4 Pinnotheres pisum (Linnaeus, 1767) N Bugula ñabellata (Thompson in Gray, 1848) Ki, N Thia scutellata (Fabricius, 1793)* ’ * M , N Bugula plumosa (Pallas, 1766) E2, H j , IQ, K 7 Insecta Bugula stolonifera Ryland, 1960 G 2 Bugula turbinata Alder, 1857 d 2 Anurida maritima (Guérin, 1836) As, Ag, B2, Ba, Bß Callopora aurita (Hincks, 1877) G j , G o, H j Clunio marinus (Haliday, 1855) A 2, A j, Ag, Ag, Bj—B j, H j , H 3 , I 3 Callopora dumerilii (Audouin, 1826) H, Petrobius brevistylis Carpenter, 1913 A 5 Callopora lineata (linnaeus, 1767) Ag-Ag, Bj, Bs, Ej-Es, G ,, G2i H j, H 2i Ij, I3-Is, Phoronidea K j , K4-K j Phoronis mülleri Selys-Longchamps, 1903 H errm ann (1980) Conopeum reticulum (Linnaeus, 1767) A 2i A 7i Ag, K3-K 7i N i Phoronis sp. M, N Elektra pilosa (Linnaeus, 1767) A 2i A g - A g , B j —B 5 , Bj, D j, D 3i D 4i E j—E3i E5, Gj, Bryozoa, Ctenostomata G 2, H1-H3, I j —lg, J1-J3, Kj-Ks, N Elektra monostachys (Busk, 1854) K ,, K g Alcyonidium diaphanum (Hudson, 1762) K2,K 4 Flustra foliacea (Linnaeus, 1758) D 5 , E 2—E 4i O í , Ka, K 4, K s , N Alcyonidium gelatinosum (Linnaeus, 1761) A2, Ag—Ag, Gj, G a , N Membranipora membranacea (Linnaeus, 1767) A 7, A g , D j, Da, D s , G j, G 2, Kj, K3 [Alcyonidium polyoum (Hassal, 1841)] Membraniporella nitida (Johnston, 1838) G j, Ga, H j, K4i Ks Alcyonidium mamillatum Alder, 1857 K4, Ks Scruparia chelata (Linnaeus, 1758) H i Alcyonidium mytili Dalyell, 1848 H j, Ha, Ka, K ,, Ke Scrupocellaria reptans (Linnaeus, 1767) Bs, H1-H3, Is Bowerbankia gracilis Leidy, 1855 A s , B j-B 7, H j , H a Scrupocellaria scruposa (Linnaeus, 1758) B51 Ha, G j, G 2. H j —H 3 , K 4, K7 Bowerbankia imbricata (Adams, 1798) Ag—Ag, Jj—J3 Bowerbankia pustulosa (Ellis & Solander, 1786) Ag—Ag Echinodermata, Asteroidea Bowerbankia sp. Gj, Ga, I1-I4, KSl K8 Asterias rubens Linnaeus, 1758 A 2 i A 3 i Ag—Ag, G j, G 2, I4, K j —K7i M, N Flustrellidra hispida (Fabricius, 1780) Ag—Ag, B5, G j, H 2 Astropecten irregularis (Pennant, 1777)* " G o e m a n n (pers, c o m m .) Nolella sp. k 4 Solaster papposus (Linnaeus, 1768)*, *** Kb Walkeria uva (Linnaeus, 1758) A 2, Ag—Ag, Jj—J3 [Crossaster papposus (Linnaeus, 1768)] Bryozoa, Cyclostomata Echinodermata, Ophiuroidea Crisia aculeata Hassal, 1841 k 4 Acrocnida brachiata (Montagu, 1804) * *, * ’ * M, N2 Crisia denticulata (Lamark, 1816) k 2, k 4 Amphipholis squamata Aa, A 3i Ag-A8, H 2, Ha. Kj, IQ, Kg, M, N Crisia eburnea (Linnaeus, 1758) B5 , D 3 , Ds, Ej—E3i Gj, Ga, Ha, H3, K2, K4, N (Delle Chiaje, 1829)**, *** Crisidia cornuta (Linnaeus, 1758) h 2 Amphiura filiformis (O. F. Müller, 1776)*2 M, N 2 Disporella hispida (Fleming, 1828) E j , Ea, H 2, K4-K 6 Amphiuridae sp. M J, H arm s 30 Species list Helgoland 31 Table 3 (Continued) Table 4. Check list of Chordata found around the island of Helgoland as described in recent publications, for more detailed information see Table 1 Echinodermata, Ophiuroidea Ophiothrix fragilis (Abildgaard, 1789) h , Kj, Ki, N Acrania M, N Ophiura albida {Forbes, 1841) Branchiostoma lanceolatum Pallas, 1774 M, N Ophiura ophiura (Linaeus, 1758) N [Ophiura textura (Linnaeus 1758)] Chondrichthyes Scyliorhinus canicula (Linnaeus, 1758)* Goemann (pers, comm.) Echinodermata, Echinoidea Galeorhinus galeus (linnaeus, 1758) Goemann, M. Janke (pers. comm.) M, N Echinocardium cordatum (Pennant, 1777) Lamna nasus (Bonnaterre, 1788) * M. Janke (pers. comm.) Echinocyamus pusillus (O, F. Müller, 1776) * * M, N 2 Mustelus mustelus (linnaeus, 1758) * Goemann (pers. comm.) Echinus esculentus linnaeus, 1758* *, * * * M, N, Ki, K4-K , Brissopsis, lyrifera (Forbes, 1841) M Osteichthyes Clupeiformes Psammechinus miliaris (Gmelin, 1778)** Ag—Ag, H2, J2, J3, M, N Alosa alosa (Linnaeus, 1758) * * * Goemann, M. Janke (pers. comm.) Spatangus purpureus (O. F. Müller, 1776)*, * * * Goemann (pers. comm.) Clupea harengus Linnaeus, 1758 M Sprattus sprattus (Linnaeus, 1758) F, M Echinodermata, Holothurioidea Salmo salar Linnaeus, 1758*, *** M. Janke (pers. comm.) Holothurioidea sp. M Anguilliformes Tunicata Anguilla anguilla (linnaeus, 1758) F Ascidiella aspersa (Miiller, 1776) D ,, D4, G i, G a, Ij—13, Ji—■J31 Ks Ascidiella scraba (Muller, 1776) Ht , K4 Beloniformes Botrylloides leachi (Savigny, 1816) Ag-A g, Di, D3, D4, Ei—E5, G2, H 2, I j , I4, J2, Belone belone (Linnaeus, 1761) Goemann, M. Janke (pers. comm.) K i-K 4, K7 Syngnathiformes Botryllus schlosseri (Pallas, 1766) A2i Ag—Ag, Bj, B3, D i—D4, G i, G2, H i—Hs, Ij-Ij, Entelurus aequoreus (linnaeus, 1758) F JJ31 K7, Kg Syngnathus acus Linnaeus, 1758 n 2 Ciona intestinalis (Linnaeus, 1767) # B31 D j—Dj, H i, II—15, J l —J3, Kg Syngnathus rostellatus Nilson, 1855 M. Janke (pers. comm.) Clavelina lepadiformis (Müller, 1776) A-?—As , Bj, D i-Ds, G t, G2, H 1( Ilf I3, K7 Syngnathus typhle Linnaeus, 1758 M. Janke (pera, comm.) Dendrodoa grossularia (van Beneden, 1846) G2, Hi, K4, K6 Didemnum candidum Savigny, 1816 D j-D j , Et-Eg, Gj, G2, Hj-Ha, Ii-Ij, Klr K.,, K, Gadiformes [Didemnum maculosum Berrii, 1950)] Ciliata mustela (linnaeus, 1758) F possibly D. helgolandicum Michaelsen, Ciliata septentrionalis (Collett, 1875) F 1921, only the larvae can be distinguished Gadus morhua linnaeus, 1758 F, K,, M, Hz Didemnum sp. Ag—Ag Melanogrammus aeglefinus (Linnaeus, 1758) M Diplosoma listerianum (Milne-Edwards, 1841) D 4, H i [Gadus aeglefinus Linnaeus, 1758] Diplosoma migrans (Menker & Ax, 1970) I2 I4 Merlangius merlangus (linnaeus, 1758) F, M, N2 Molgula citrina Alder & Hancock, 1848 Ag-A g, D i, D 2, D j , G2, H i , H 3i I1( I3- I 5, J ^ , K2 [Gadus m erlangus Linnaeus, 1758] Molgula complanata Alder & Hancock, 1870 H 1-H 3 Molva molva (linnaeus, 1758) M. Janke (pera, comm.) Polyclinum aurantium Milne-Edwards, 1841 D i, D j, Ka Pollachius pollachius (linnaeus, 1758) Goemann, M. Janke (pera, comm.) Sidnyum turbinatum Savigny, 1816 A2i Ag-Ag, D i—D3, D j , E i —E4 i G i , G 2i H i —H3 i Pollachius virens (Linnaeus, 1758) F K j , K2, K, Raniceps raninus (Linnaeus, 1758) F Styela coriacea (Alder & Hancock, 1848) h 2 Rhinonemus cimbrius (linnaeus, 1766)* F Styela partita (Stimpson, 1852) B i , H 3 Trisopterus luscus (Linnaeus, 1758) F, M Trisopterus minutus (Linnaeus, 1758) F, M [Gadus capelanus Lacepède, 1800] Perciformes, Percoidei Ammodytes marinus Raitt, 1934 F Ammodytes tobianus Linnaeus, 1758 F. N 2 Ctenolabrus rupestris (Linnaeus, 1758) F, Ki—Kg, N Echiichtys vipera (Cuvier, 1829) F [Trachinus vipera Cuvier, 1829] Hyperoplus lanceolatus (Le Sauvage, 1824) F, N 2 Labrus bergylta Ascanius, 1767* F Mullus surmuletus Linnaeus, 1758 Goemann, M, Janke (pera, comm.) Trachurus trachurus (Linnaeus, 1758) f , n 2 32 J, H arm s Species list Helgoland 33 Table 4 (Continued) Table 5. Polychaete larvae, whose adults are not listed in Table 3, found in plankton samples from the Helgoland Reede by Husemann (1992) and Plate (1992), S: Husemann; T; Plates * species which Perciformes, Scombroidei live normally on rocky surfaces, shell grounds or coarse sands Scomber scombrus Linnaeus, 1758 M Perciformes, Gobioidei Chaetopterus variopedatus (Renier, 1804)* T Gobiusculus flavescens (Fabricius, 1779) F Harmothoe longisetis (Grube, 1863) S Pomatoschistus microps (Krayer, 1838) F Heteromastus filiformis (Claparède, 1864) T Pomatoschistus minutus (Pallas, 1770) F, M Laonice cirrata (M. Sars, 1851) T Pomatoschistus pictus (Malm, 1865) F Magelona filiformis Wilson, 1959 T Ophiodromus flexuosus (Delle Chiaje, 1822) S Perciformes, Callionymoidei Pectinaria (Amphictene) auricoma (O. F. Müller, 1776) S F Callionymus lyra Linnaeus, 1758 Polydora flava Claparède, 1870* T Callionymus reticulatus Valenciennes, 1837 N Polydora hermaphroditica Hannerz, 1956 T Callionymus sp. M Prionospio ciliifera Wirén, 1883* T Perciformes, Blennioidei Pseudomystides limbata (Saint-Joseph, 1888}* S Pholis gunnellus (Linnaeus, 1758) F, Kj, K2, K4 i Ke, N Sige fusigera Malmgren, 1865* S Zoarces viviparus (Linnaeus, 1758) F, Ke Spio martinensis Mesnil, 1896 T Perciformes, Mugiloidei Chelon labrosus (Risso, 1826) Goemann (pers, comm.) Scorpaeniformes

Agonus cataphractus (Linnaeus, 1758) F, M, N 2 Cyclopterus lumpus Linnaeus, 1758 F, G2 Eutrigla gurnardus (Linnaeus, 1758) M [Triglia gurnardus Linnaeus, 1758] Uparis montagui (Donovan, 1804) F Liparis liparis (Linnaeus, 1766) Ga, N Myoxocephalus scorpius (Linnaeus, 1758) F, Ki, K3 ---K7 , M, H Taurulus bubalis (Euphrasen, 1786) F, Ki, K4 -K 6 Trigla lucerna Linnaeus, 1758 M Gasterosteiformes Gasterosteus aculeatus Linnaeus, 1758* * * Goemann (pers. comm.) Spinachia spinachia (Linnaeus, 1758) F Pleuronectiformes

Arnoglossus laterna (Walbaum, 1792) F, N 2 Buglossidium luteum (Risso, 1810) F, N Hippoglossoides platessoides (Bloch, 1787) M. Janke (pers. comm.) Limanda limanda (Linnaeus, 1758) M, N 2 Microstomus kitt (Walbaum, 1792) M, N Phrynorhombus norvegicus (Günther, 1862) F, N Platichthys ñesus (Linnaeus, 1758) n 2 Pleuronectes platessa Linnaeus, 1758 k 5, m , n 2 Psetta maxima (Linnaeus, 1758) F Scophthalmus rhombus (Linnaeus, 1758)* F Solea vulgaris Quensel, 1806 F, M. N 2 [? Pleuronectes solea Linnaeus, 1758, Common synonym:Solea soiea] Mammalia, Cetacea Phocoena phocoena (Linnaeus, 1758)*, * ’ * Goemann (pers. comm.) Mammalia, Pinnipedia Phoca vitulina Linnaeus, 1758* * * Goemann (pers, comm.) Halichoerus grypus (Fabricius, 1791)* Goemann (pers. comm.) 34 J. H arm s

Table 6 . Literature used for taxonomic classification. Additional advice for special taxonomic groups given by the authors mentioned in the list

Algae Kornmann & Sahling (1977, 1983)-, Kornmann & Sahling (pers. comm.) Ciliata Hadzi (1951) Porifera Arndt (1934); A. Dzwillo (pers. comm.); D. Barthel (pers. comm.) Hydrozoa Vervoort (1946); G. Jarm s (pers. comm.) Scyphozoa Krumbach (1928) Anthozoa M anuel (1988) Kamptozoa Nielsen (1989) Nematoda Platt & Warwick (1983) Turbellaria Prudhoe (1982) Nemertini Gibson (1982); P. Sundberg (pers. comm.) Polyplacophora Jones & Baxter (1987) Prosobranchia Graham (1988); Nordsieck (1982); R. Janssen (pers, comm,) Ophistobranchia Thompson & Brown (1976); R. Janssen (pers. comm.) Bivalvia Nordsieck (1969); R. Janssen (pers, comm.) Cephalopoda Jaeckel (1926); C. Warneke-Cremer (pers, comm,) Sipunculoidea Gibbs (1977) Echiuroidea Ten Broeke (1929) Polychaeta Hartmann-Schröder (1971); George & H artm ann-Schröder (1985); West heide (1990); Pleijel & Dales (1991); G, H artm ann-Schröder (pers. comm. Oligochaeta Ude (1929) Hirudinea Johansson (1929) Pantopoda King (1974) Ostracoda Athersuch et al. (1989) Harpacticoidea Pesta (1932) Cirripedia Newmann & Ross (1976) Mysidacea Gordan (1957); U. M ühlenhardt-Siegel (pers. comm.) Amphipoda Schellenberg (1942); Lincoln (1979); Barnard & B arnard (1983); Barnard & Karaman (1991); H.-G, Andres (pers. comm.) Cumacea Jones (1976) Tanaidacea Nierstrasz & Schuurmens Stekhoven (1930) Isopoda Naylor (1972) Decapoda Schellenberg (1928); Smaldon (1979); Ingle (1983) Burukovskii (1986) Insecta Cheng (1976) Phoronida Emig (1979) Bryozoa Ryland & Hayward (1977); Hayward & Ryland (1979); H ayward (1985); Hayward & Ryland (1985) Echinodermata Ludwig (1900); Lieberkind (1928); I. Bartsch (pers. comm.) Tunicata Millar (1970); M enker & Ax (1970) Pisces H ureau & Monod (1973); W hitehead et al. (1984); E. W ahl (pers. comm,) Cetacea Freud (1933a) Pinnipedia Freud (1993b)