J. Mar. Biol. Ass. U.K. (2005), 85, 323^326 Printed in the United Kingdom

An enriched Chaetopterus tube mat biotope in the eastern English Channel

E.I.S.Rees*, M. Bergmann, M. Galanidi, H. Hinz, R. Shucksmith and M.J. Kaiser School of Ocean Sciences, University of Wales Bangor, Menai Bridge, Anglesey,Wales, LL59 5AB, UK. *Corresponding author, e-mail: [email protected]

Patches of a very dense tube mat biotope were found during ¢sh habitat studies in the eastern English Channel. At three locations in the lows between linear sand banks o¡ the French coast an un-described small Chaetopterus sp. occurred with small Lanice conchilega as an enriched sediment stabilizing biotope. This biotope was distinct though having similarities to other tide swept sub-tidal biotopes dominated by L. conchilega. Using cameras and side-scan sonar it was seen to overlay heterogeneous cobbles and shell hash with intermittent rippled sand veneer. The patchiness of this enriching biogenic feature contributed to the variability in trawl catches of ¢sh.

INTRODUCTION form of Chaetopterus has an easily collapsed muddy tube and is distinct from the widespread large parchment tube Classi¢cations of sub-tidal sedimentary biotopes from building species (Mary E. Petersen, personal communica- north-west European shelf seas (Connor et al., 1997, tion). Similar specimens of a small Chaetopterus, collected 2003) include several categories wherein tube worms are o¡ Brittany had previously been sent to Dr Petersen by so abundant that they form biogenic mats. Such mats have important ecological functions in trapping ¢ne sedi- Professor M. Glemarec. Taxonomic descriptions of the new species are in preparation by Dr Petersen. ment and in provision of food or shelter for small ¢sh (Auster et al., 1997). A worm tube dominated biotope, di¡ering from those previously recognized, is described here from the eastern English Channel. The character- MATERIALS AND METHODS izing species in this case was a small sized, but not yet Habitats were studied at a series of locations in the described species of Chaetopterus. It occurred together with English Channel, previously ¢shed by the Centre for very large numbers of small Lanice conchilega. The small Environment, Fisheries and Aquaculture Science

Figure 1. Photographs of the Chaetopterus tube mat biotope. Vertical views are at di¡erent enlargement scales. Image (A) 0.3m across, image (B) 0.04 m across.

Journal of the Marine Biological Association of the United Kingdom (2005) 324 E.I.S. Rees et al. Chaetopterus tube mat in the English Channel

Table 1. Stations in the eastern English Channel where the Table 2. Numerical abundance of the more common species in Chaetopterus tube mat biotope was found in August 2002. grab samples as N/0.1 m2. Columns A, B and C based on the ¢ve Positionsofthecamerasledgetows,andthelimitsoftrawling grab samples which had 420 Chaetopterus sp. in each. areas within which the 2 m and 4 m beam trawls and grabs were Similarity values are per cent contributions to the similarity of deployed. the grouped stations analysed by SIMPER after square-root transformation. Column D has median abundances from samples Station code 69 71 72 with 520 Chaetopterus but 450 Lanice conchilega. Column E has median abundances from samples with few or Camera tows none of either tube building species. Deployed Latitude 50846.85’N50839.78’N50833.62’N AB CDE Longitude 01833.16’E01831.00’E00832.74’E Median Maximum Similarity Recovered Latitude 50847.12’N50839.81’N50833.35’N Tube building and Longitude 01833.48’E01830.81’E00832.73’E tube attached or Distance towed km 0.71 0.57 0.50 occupying Depths m 19^20 26 16^17 Chaetopterus sp. 34 609 8 3 0 Beam trawl and grab area spread Lanice conchlega 416 2280 22 352 2 North-east limit Eumida sanguinea 101 178 10 53 0 Latitude 50847.99’N50839.67’N50834.90’N Apseudes latreilli 3 571 4 1 0 Longitude 01834.27’E01830.94’E01833.10’E Phtisca marina 17 90 4 6 0 South-west limit Infauna Latitude 50846.64’N50838.17’N50832.12’N Notomastus 40 129 6 10 0 Longitude 01832.88’E01830.33’E01832.28’E latericeus Nemertea 14 136 3 15 6 indet. Echinocyamus 31 113 5 6 0 pusillus (CEFAS) for stock assessment purposes. The primary aims Poecilochaetus 22 118 5 13 0 were to compare the diversity and consistency of ¢sh serpens catches with the benthic environments, and to help de¢ne Lagis koreni 21 119 3 20 2 the habitat requirements of three £at¢sh species (Hinz et Echinocardium 185100 al., 2003). At each site observations were made by digital cordatum side-scan sonar to indicate ground type mosaics prior to Cirratulidae indet. 20 29 5 9 0 deployment of sledge mounted underwater television and Mysella bidentata 341230 still cameras. The video system was a Rovtech Seacam Urothoe 728202 camera pointing obliquely forward and recording on a brevicornis Ensis arcuatus 322253 Sony digital 8 tape recorder. The still camera was a Sipuncula indet. 1 24 1 0 Photosea 1000 system arranged with the camera pointing Glycinde 614 11 directly downwards, with the lens 0.7 m above the bed, nordmanni and the strobe at 60 degrees to it. Photographs were Abra alba 114120 taken at 42 second intervals spread along each 30 minute Aonides 212 00 camera sledge tow. Distances covered by camera sledge pauchibranchiata and trawl tows were calculated from shot and hauling Mobile epifauna positions derived from Digital Global Positioning System Ophiura ophiura 557110 satellite navigation. Spread randomly along each of the Abludomelita 11 48 2 7 0 previous CEFAS trawl tow lines a 0.1m72,daygrabwas obtusata deployed four times, with extra samples sometimes being Polinices pulchellus 215 70 Asterias rubens 16 00 taken if the side-scan showed obviously di¡erent ground types at opposite ends of the tows. These macrobenthos samples were sieved over 1-mm mesh before preservation for later laboratory processing. Four tows of 5 min duration cruise on RV ‘Prince Madog’ in August 2002. The stations at a speed of two knots were made with a 2 m small mesh with the tube mat (Table 1) were on the French side of the beam trawl and four tows of 20 min duration at four knots eastern English Channel o¡shore from Boulogne and Le were made with a 4 m, beam trawl with a 82 mm cod-end Touquet. mesh. Both trawls had chain matrix. Material from the Photographs of the seabed with the Chaetopterus and trawls was sorted, enumerated and weighed at sea on a Lanice conchilega tube mat biotope are shown in Figure 1. Pols motion compensated balance accurate to 1g. The Di¡ering scales of enlargement from the original 35 mm camera tows and grabs were made prior to using the colour slides are used to illustrate the habitat and the trawls at each site. disposition of the Chaetopterus tubes. Of 83 interpretable photographs taken at the three localities, 32 showed the dense tube mat biotope with Chaetopterus. Assuming conti- RESULTS nuity between sequential ¢lm frames during a camera The Chaetopterus tubes were noted at three of the local- sledge tow, the tube mat patches seen were spread along ities in the eastern English Channel sampled during a an estimated combined distance of 0.38 km. As video

Journal of the Marine Biological Association of the United Kingdom (2005) Chaetopterus tube mat in the English Channel E.I.S. Rees et al. 325 image resolution was lower than from the stills camera, only about 0.7^1.2 times their aperture diameter, but some the Chaetopterus tubes could not always be reliably distin- more exposed tubes were seen lying collapsed on the guished from those of other species. However, the open sediment surface. apertures of tubes of Chaetopterus could often be seen on The numbers of individuals per 0.1m72 of the more video monitors as small de¢ned dark spots in comparison abundant taxa in the ¢ve grab samples that each contained with the ragged and paler appearance of the fringed tops at least 20 individual Chaetopterus worms are shown in of the L. conchilega tubes. The underwater television Table 2. This arbitrary cut o¡ was chosen to coincide with (UWTV) showed a dense tube biotope apparently with a break in the ranked abundance sequence. For compar- Chaetopterus and L. conchilega for 41min of a combined ison, the table also shows median abundances of the same total of 108 min when the sledge was on bottom at the species in samples with more than 50 Lanice conchilega but three localities. For a further 39 min the video showed few Chaetopterus. Median abundances in samples from the sparse to moderately dense tubes which appeared to be same trawl tow locations with very few tubes of either L. conchlilega. This equates to 0.45 km over the ground for species are also shown. This indicates how sparse the the dense tube mat. The di¡erence from the still camera benthos was without the in£uence of the worm tubes. For estimate was due to ¢lm running out before the UWTV the whole 14 grabs, high variance to mean ratios of 460 in was recovered. On the enlarged still photographs, the the counts of Chaetopterus and 914 for L. conchilega indicated lengths of the shadows implies that the tubes of the small that both species had strongly aggregated distributions. Chaetopterus species often protruded from the sediment by Video observations con¢rmed this patchiness. The grabs were, however, spread over the length of the previous Table 3. Mean wet weights (grams) of fauna caught in 2 m CEFAS trawls and not targeted at where the shorter and 4 m beam trawls, at the three stations where the tube mat camera tows were taken. Of the ¢ve grabs, only one seems was found, based on a total of 12 tows with each gear type. to have coincided with the very dense Chaetopterus tube mat Species omitted where mean 510 g in 2 m small mesh trawls and as seen on the photographs and video. 525gin4mtrawls. Table 3 lists the mean wet weights, from the beam trawls, of the more abundant taxa by di¡erent life-form 2 m small 4m¢sh groups. Some of these, such as the attached epifauna, mesh trawl mainly came from habitats other than the tube mat. The trawls would have swept several habitats, including Attached epifauna patches of exposed cobbles and shells. On average the 4 m Alcyonidium diaphanum 60 1248 trawl picked up 29 kg of stones and 5 kg of shells per Alcyonium digitatum 836 20 min tow and 3 kg of stones and 2 kg of shells were in Nemertesia antennina 36 479 Hydralmania falcata 11 5 min tows of the 2 m small mesh trawl. Hydrozoa indet. 581 The super¢cial sediment collected by grab from Porifera indet. 291 amongst the Chaetopterus tubes was mainly cohesive Mytilus edulis 190 muddy sand (median phi diameter 3.04^3.09). Loss on Mobile epifauna and infauna ignition was 2.3^3.1%. In the patches where the Asterias rubens 1261 6748 tubes were most dense the video showed black patches Liocarcinus holsatus 321 849 around burrows and black coiled worm casts indicating Hinia reticulata 87 organic enrichment and anoxia under the sediment Ophiura ophiura 65 surface. Crangon crangon 63 Maia squinado 45 36 Pagurus bernhardus 37 28 DISCUSSION Diogenes pugilator 32 Necora puber 21 82 In this part of the eastern English Channel there is a Spisula elliptica 14 complex of linear sand banks aligned with the tidal Echinocardium cordatum 11 currents, with a range of more heterogeneous seabed Psammechinus miliaris 230 types in the lows between the sand banks. These include Cancer pagurus 180 cobbles, relict shell beds, and exposures of the underlying Spatangus purpureus 63 bedrocks (chalk and Kimmeridge clay), together with Liocarcinus depurator 32 rippled sand sheets (James et al., 2002). The water here is Ophiura albida 26 fairly turbid with moderately strong tides (2 knots on mean Squid and ¢sh Sepia o⁄cinalis 322 5069 spring tides) (Lee & Ramster, 1981).The bed surface adja- Pleuronectes platessa 31 11953 cent to the tube mat patches had asymmetrical sand Platichthys £esus 2240 ripples, with cobbles sometimes showing through the Limanda limanda 482 sand, or there was a lag composed mainly of shelly hash. Solea solea 73 294 However, within the tube mat patches the surface Microstomas kitt 117 appeared smooth with only slight mounding. The sedi- Scophthalmus rhombus 39 ment samples taken by grab indicated that the tube mat Callionymus lyra 136 557 had trapped a ¢ner and more cohesive muddy sand than Raja clavata 939 would be predicted from the strength of the tidal currents. Trigla lucerna 181 From the presence of Urticina sp. which attach to solid Trisopterus minutus 42 Agonus cataphractus 11 25 objects, the tube mat probably formed only a super¢cial ¢ne sediment layer.

Journal of the Marine Biological Association of the United Kingdom (2005) 326 E.I.S. Rees et al. Chaetopterus tube mat in the English Channel

Lanice conchilega was more widespread and the open variability noted in trawl tows made for ¢sh population topped Chaetopterustubesoccurredmainly whereL.conchilega monitoring (Hinz et al., 2003). Knowledge of where these was also particularly abundant. On some enlarged photo- biogenic features occur should be an important factor in graphs, the Chaetopterus tubes appeared to be the more abun- ecosystem based ¢sheries management and interpretation dant, but this was not so in any of the grab samples. of stock sampling trawl tows. Individuals of Chaetopterus may each have several tube open- ings in comparison with the single fringed top usual for We are grateful to the Master and crew of RV ‘Prince Madog’ L. conchilega which might account for L. conchilega being the for their skill in deploying and towing the range of gear used in moreabundant whenwormsratherthantubeswerecounted. this project, to Ben Powell and Ray Wilton for assistance with Under the system of biotope classi¢cations used for operating the side-scan sonar. Dr Mary Petersen advised on the the European Nature Information System (EUNIS) identity of the small Chaetopterus sp. Kerry Howell gave pre pub- (undated) there is a sub-tidal, L. conchilega dominated lication access to revisions of sub-tidal parts of the Joint Nature Conservation Committee Marine Habitat Classi¢cation. Clear- biotope, occurring in tide swept sand (EUNIS code ance to operate inside territorial waters was given by the A4.123). The Chaetopterus tube mat has obvious similarities Government of France. to this but seems su⁄ciently distinct to be considered sepa- rate at level 5 in the hierarchy. Based on side-scan sonar and video evidence, the wider REFERENCES extent of the individual trawl tow tracks previously used Auster, P., Malatesta, R. & Donaldson, C., 1997. Distributional for ¢sh stock assessment would have run over several responses to small-scale habitat variability by early juvenile di¡erent sedimentary habitats. These would have included silver hake, Merluccius biliearis. Environmental Biology of Fishes, rippled sands, with cobbles and shell hash both intermit- 50, 195^200. tently exposed at the bed surface as well as the enriched Connor, D.W., Allen, J.H., Golding, N., Lieberknecht, L.M., tube mat. Such super¢cial deposits, held together by Northern, K.O. & Reke, J.B., 2003. The National Marine worm tube mats, are liable to wash away when particular Habitat Classi¢cation of Britain and Ireland, Version 0.303. Peterborough: JNCC. www.ince.gov.uk/marine/biotopes/ cohorts of key species die out, so the tube mat patches default.htm could be ephemeral. This may explain why the biotope Connor, D.W., Dalkin, M.J., Hill, T.O., Holt, R.H.F. & has not been reported before in a comparatively well Sanderson, W.G., 1997. Marine Conservation Review. Marine studied inshore sea area. biotopes classi¢cation for Britain and Ireland. Vo l . 2. Sublittoral In north-west European shelf seas there are several biotopes. Version 97.06, Report 230. Peterborough: Joint Nature examples of biogenic habitats where key species create Conservation Committee. niches for rich and diverse ranges of co-occurring species. EUNISEuropean Nature Information System. Published by In some cases these biogenic habitats even appear as ‘oases’ European Environment Agency. www.eunis.eea.eu.int/ of higher biodiversity amongst wider areas of ‘semi- habitats-code-browser desert’, impoverished through sand instability or scour. Hinz, H., Kaiser, M.J., Bergmann, M., Rogers, S.I. & For example, a tube mat found on the edge of Nephrops Armstrong, M.J., 2003. Ecological relevance of temporal stability in regional ¢sh catches. Journal of Fish Biology, 63, norvegicus ¢shing grounds in the western Irish Sea had a 1219^1234. super-abundance of small bivalves and crustaceans James, J.W.C. et al., 2002. GEOSYNTH: a synthesis of the amongst a very dense mat of ampharetid worm tubes geology and sediments of the Dover Strait and its hinterland. (Rees & Holme, 1988). Biologically determined benthic British Geological Survey, Keyworth, Nottingham. Commissioned habitats seem to be disproportionately productive and Report CR/02/078. biodiverse in comparison with those at locations overwhel- Kaiser, M.J., Cheney, K., Spence, F.E., Edwards, D.B. & mingly in£uenced by physical sedimentary processes. In Radford, K., 1999. Fishing e¡ects in northeast Atlantic shelf the Chaetopterus biotope described here, nine taxa had seas: patterns in ¢shing e¡ort, diversity and community struc- maximum abundances equivalent to over 1000/m72.For ture. VII. The e¡ects of trawling disturbance on the fauna L. conchilega the maximum was equivalent to over 22,000/ associated with the tubeheads of serpulid worms. Fisheries Research, 40, 195^205. 72 Chaetopterus 72 m and for sp. over 6000/m . The existence Lee, A.J. & Ramster, J.W., eds. 1981. Atlas of the seas around the of such locally enriched habitats has implications for the British Isles, 5 pp.+75 sheets. London: Ministry of Agriculture, meso-scale distribution of several commercial ¢sh species Fisheries and Food. including plaice Pleuronectes platessa (Kaiser et al., 1999) Rees, E.I.S. & Holme, N.A., 1988. A local enriched macro- and silver hake Merluccius bilinearis (Auster et al., 1997). In benthos association in the Irish Sea. In Commission of the the eastern English Channel the plaice biomass was European Communities, COST 647 Workshop, La Corruna, highest in trawl tows with tube mat somewhere in the October 1986, pp. 173^174. same general location. Spatial and temporal patchiness associated with biogenic features will in£uence the Submitted 26 April 2004. Accepted 8 December 2004.

Journal of the Marine Biological Association of the United Kingdom (2005)