Advisory Committee on the Marine Environment ICES CM 1998/ACME:8 Ref.: E
{)/O
REPORT OF THE
ICESIHELCOM BENTHOS TAXONOMIC WORKSHOP
Roskilde, Denmark 4-7 November 1997
This report is not to be quoted without prior consuItation with the General Secretary. The document is areport of an expert group under the auspices of the International Council for the Exploration of the Sea and does not necessarily represent the views of the CounciI.
International Council for the Exploration of the Sea Conseil International pour I'Exploration de la Mer
Pal:rgade 2-4 DK-1261 Copenhagen K Denmark TABLE OF CONTENTS
Section Page
INTRODUCTION 1 2 TERMS OF REFERENCE 1 3 PROGRAMME FOR THE WORKSHOP 1 3.1 Phyllodocidae 1 3.2 Nephlyidae, Nereidae, and Polynoidae 1 3.3 Porifera 1 3.4 Sampling Techniques and Preservalion Practices 1 3.5 Pholoidae, Cirratulidae, and Ampharetidae 2 3.6 Gastropoda: Hydrobia. and Potamopyrgos 2 3.7 Gastropoda: Tectibranch Opistobranchia 2 3.8 Bivalvia: The Cardium-complex 2 3.9 Polychaeta: Spionidae and Flabelligeridae 2 3.10 Amphipoda 3 3.11 Feedback and General Discussion on Future Workshops 3 3.12 Closing ofthe Workshop 3 4 REFERENCES 3 ANNEX 1: LIST OF WORKSHOP PARTICIPANTS 4 ANNEX 2: AGENDA 6
ANNEX 3: PHYLLODOCIDAE 7 • ANNEX 4: SELECTIVE LIST OF REFERENCES FOR POLYCHAETA FROM THE KATIEGAT AND THE BALTIC SEA 9 ANNEX 5: IDENTIFICATION OF SPONGES 10
ANNEX 6: FINAL NOTES 21 - ANNEX 7: CIRRATULIDAE FROM THE KATIEGAT, 0RESUND, AND BALTIC SEA 22 ANNEX 8: A TAXONOMIC SCHEME 33 ANNEX 9: MORPHOLOGICAL FEATURES 37 ANNEX 10: THE LUMBRINERIDAE OF THE KATIEGAT AND THE BALTIC SEA 38 ANNEX 11: THE SPIONIDAE OF THE KATIEGAT AND THE BALTIC SEA 39 1 INTRODUCTION
The ICESIHELCOM Benthos Taxonomie Workshop (WKBT) was the outcome of work initiated by the ICESIHELCOM Steering Group on Quality Assurance of Biological Measurements (SGQAB). An intercalibration exercise carried out by the Steering Group indicated that misidentification of species often occurs in the monitoring of benthie macrofauna. The aim of the Workshop was to minimize the possibility for species misidentification and to raise the general level of taxonomic knowledge among the persons who carry out the practical monitoring.
The Workshop was held at the National Environmental Research Institute (Roskilde, Denmark) from 4-7 November, under the chairmanship of J~1rgen N~rrevang Jensen who opened the Workshop and welcomed participants. Danny Eibye-Jacobsen made a short presentation on practical issues of the Workshop. Taxonomie experts from the Zoological Museum ofCopenhagen acted as Iecturers.
Twenty people (the maximum number) participated in the Workshop. The names and addresses of Workshop participants are listed in Annex I.
2 TERMS OF REFERENCE
The terms of reference for the ICESIHELCOM Benthos Taxonomie Worksho{? were to:
leES C.Res.1996/3:4
• a) identify benthie invertebrate species over which there is taxonomie disagreement and find a common identification; b) review rriäterial brought together by researchers and research assistants that has created taxonomie problems and determine ways to obtain a correct identification; e) review the development of 'in-house quality assurance manuals' for benthos studies; d) prepare the materials resulting from this Workshop as a laboratory report for publication.
3 PROGRAl\1~1E FOR THE WORKSHOP
The Workshop was a mixture of lectures on special taxonomie groups and exercises in species identification. In general, the Agenda for the Workshop, as presented in Annex 2, was followed.
3.1 Phyllodocidae
The first lecture on the polychaete family Phyllodocidae was given by Danny Eibye-Jacobsen, who started with an overview of the morphological features which are important for distinguishing the different species. A list of these are found in Annex 3. After the lecture, participants could examine preserved specimens ofthis family.
3.2 Nephtyidae, Nereidae, and Polynoidae
A lecture on the Nephtyidae, Nereidae, and Polynoidae families was also given by Danny Eibye-Jacobsen. The characteristic morphological features were presented and Rainer (1991) was recommended for use as the key for the genus Neptlzys. A selective list of references for Polychaeta from the Kattegat and the Baltie Sea are given in Annex 4. The lecture was followed by a practical session on preserved specimens.
3.3 Porife ra
ale Tendal gave a lecture on the phylum Porifera. An overview of the taxonomie problems with sponges was given as weil as a key to the speeies of sponges found in the Baltie Sea. The presentation by ale Tendal appears in Annex 5. The lecture was followed by a practical session on preserved specimens.
1998 WKBT Report 3.4 Sampling Techniques and Preservation Practices
A diseussion on sampling teehniques, preservation praetices, and interaetions between monitoring units and zoologieal museums were on the programme. A range of sampling gear (Van Veen, HAPS, Smith-McIntyre) is apparently used in the Kattegat/Belt Sea area. It was reeommended that sampling follow the ICES guidelines for soft bottom maerofauna (Rumohr, 1990).
Problems with sampling epifauna as weIl as the more searee infauna were diseussed. It was reeommended to use a dredge to supplement the traditional infauna sampling, although this gear is normally not quantitative. It was noted that a quantitative dredge has been developed in Holland (Bergman and Santbrink, 1994).
Different fixation teehniques were diseussed. Although fixation in formaldehyde is generaIly reeommended in the HELCOM and ICES guidelines, preservation in ethanol is preferred by several counties in Denmark for health reasons. However, the Zoologieal Museum in Copcnhagen wiIl only aceept material that has been fixed in formaldehyde.
The aspeet of quality assuranee (QA) was diseussed. Several laboratories have already practieed resorting 10-20 % of the sampies for quality assuranee purposes. This practice is recommended.
The eoIlection of referenee-speeimens was reeommended. It was further suggested that the Zoological Museum should make a list of rare speeies (with synonyms) for the internet.
The use of denaturated ethanol as a preservative is not recommended for health reasons if the sampies are to be used for histological purposes after preservation. The use of Rose Bengal as a coloration dye is not recommended as it may make • the identifieation of some species difficult.
It was generally agreed that identification only to the genus level is preferred to an incorrect species name.
3.5 Pholoidae, Cirratulidae, and Ampharetidae
A lecture on the families Pholoidae, Cirratulidae, and Ampharetidae was given by Mary E. Petersen. Taxonomie problems within these families were presented as weIl as keys for identifying species. These keys are attached as Annex 6. The lecture was followed by a practical session on preserved specimens.
3.6 Gastropoda: 1I)"drobia, and Potamopyrgos
A lecture of the genera llydrobia and Potamopyrgos was presented by Bent Muus. An overview of the taxonomie and ecological aspeets of these genera was given. Copies of drawings of these speeies are attached as Annex 7. The leeture was foIlowed by a practical session on preserved speeimens. 3.7 Gastropoda: Tectibranch Opistobranchia ., A lecture on Tectibranch Opistobranchia was given by Tom Schi~dte. The taxonomie problems in this group were presented. A taxonomie scheme of the group is attached as Annex 8. The lecture was followed by a praetical session on prcserved specimens.
3.8 ßivah'ia: The Cardium-complex
A leeture on the Cardium-eomplex was given by Gotfred H~'pner Petersen. The taxonomie problems of this group were presented, foIlowed by a praetical session on preserved speeimens.
3.9 Polychaeta: Spionidae and Flabelligeridae
A lecture on Spionidae and FlabeIligeridae was given by Dannny Eibye-Jacobsen, including an overview of the morphologieal features which is important for the distinetion of the different speeies. A list of morphological features are found in Annex 9. The leeture was followed by a practical session on preserved specimens.
2 1998 WKBT Report 3.10 Amphipoda
A lecture on Amphipoda was given by Teunis Jansen who gave an overview of taxonomie features, which is a problem for this genus. The lecture was followed by a praetical session on preserved speeimens.
3.11 Feedback and General Discussion on Future Workshops
J0rgen N0rrevang Jensen introdueed the diseussion on the evaluation of the Workshop and suggestions for future workshops.
Several partieipants expressed that the idea behind taxonomie workshops is good. Several suggestions to improve the quality of future workshops were given. Suggestions included:
workshops should coneentrate on speeific phyla, families, or genera in order to have more time for detailed studies; speeimens of problematic genera/species should be distributed in order to insure a correct identification; partieipants should be encouraged to bring their own materials to future workshops; programmes should be sent in advanee announcing the topies and leetures on the different groups; regional/national workshops should be held every second year, alternating with international workshops held during off years; • participation in taxonomie workshops should be optional for institutes collecting monitoring data in the OSPAR area.
3.12 Closing ofthe Workshop
The Chairman thanked the staff of the Zoological Museum for their engaged teaehing and for arranging a very fruitful Workshop. The partieipants were acknowledged for their active participation and positive attitude. The Workshop was adjoumed.
4 REFERENCES
Bergman, M.J.N., and Santbrink, van, J.W. 1994. A new benthos dredge ('Triple-D') for quantitative sampling of in fauna species of low abundance. Netherlands Journal of Sea Research, 33: 129-133.
Rainier, S.F. 1991. The genus Nephtys (Polyehaeta: Phyllodocida) of Northern Europe: a review of species, including the deseription of N. puIch ra sp. n. and a key to the Nephtyidae. Helgoländer Meeresuntersuchungen, 45: 65-96.
Rumohr, H. 1990.· Soft bottom macrofauna: Collection and treatment of sampIes. ICES Techniques in Marine Environmental Sciences, No. 8.
1998 WKBT Report 3 ANNEX 1
LIST OF WORKSHOP PARTICIPANTS
Name Address Telephone Fax E-mail Ann-Britt Andersin Finnish Institute of Marine +45 6556 1813 +456556 1505 [email protected] Research P.O. Boks 33 FIN-00931 Helsinki Finland Grete Dinesen Bioconsult +458625 1811 +4586258173 gtd@bioconsulLdk Johannes Ewaldsvej 42-44 8230 Abyhpj Denmark Dan Evander Department of Animal +46 90 786 7703 +4690 786 7665 Ecology, Umea University 5-901 87 Umea, 5weden. Ute Fischer Niedersächsisches +494421947176 +494421947110 Landesamt für Ökologie Fliegerdeich 1 - D-26382 Wilhelmshaven, - Germany Rune Fredriksen Bioconsult, Johannes +4586251811 +4586258173 ruf@bioconsulLdk Ewaldsvej 42-44 8230 Abyh~~j Denmark Fritz Gosselck Institut für Angewandte +49 382O..t/61812 +4938204/61810 Ökologie GmbH Lindenweg 2 D-18184 Neu Broderstorf Germany Peter A. Göransson PAGHB +46 4226 1078 +46 4210 5044 Kustgatan 40b 5-25270 Raä 5weden J~~rgen Nprrevang Jensen National Environmental +45 4630 1200 +454630 1114 [email protected] (Chairman) Research Institute . Frederiksborgvej 399 P.O. Boks. 358 - ; DK-4000 Roskilde Denmark Kirsten Johansen Marin-lD +45 3325 5244 Halls Alle 12 1802 Frederiksberg C Denmark Heta Kankainen-Raunisto Finnish Institute of Marine +3589613941 +358961394494 Research P.O. Boks 33 FIN-00931, Helsinki, Finland Anika Buur Leth Fyns Amt +456556 1813 +45 6556 1505 0rb:rkvej 100 5220 Odense 50 Denmark
4 1998 WKBT Report Name Address Telephone Fax E-mail Lene Rueskov Nielsen Fyns Amt +4565561813 +45 6556 1505 0rba:kvej 100 5220 Odense S0 i Denmark Erik Skinh[lj Pedersen Marin-ID +45 3325 5244 Halls Alle 12 1802, Frederiksberg C Denmark Christine Peters Baltic Sea Research +493815197440 Institute Seestrasse 15 D-18119 Rostock- Wamemunde Germany Heye Rumohr Institut für Meereskunde, +494315973957 +49341 5973994 [email protected] Kiel University Düstembrooker Weg 20 D-24105 Kiel Germany
Doris Schiedek Baltic Sea Research +49881519701 +493815197440 doris.schiedek@io- Institute 205 wamemuende.de Seestrasse 15 - D-18119 Rostock- Warnemunde Germany Yolker Schroeren Landesamt für Natur und +494347704443 +494347704402 [email protected] Umhelt des Landes Schleswig-Holstein Hamburger Chaussee 25 - D-24220 Flintbek Germany
Susan Smith National Board of Fisheries +46 31 697825 +4631691109 Institute of Coastal Research Nya Yarvet S-42671 Yästra Frölunda Sweden
Jette Steinmeyer Storstrl'lms Amt +45 5482 3232 +4554823224 Parkvej 37 4800 Nykl)bing F Denmark -, - Wiebke Swartzbach Federal Environmental +49 30 8903 2028 +49 3089 032285 [email protected] Agency Bismarck Platz 1 D-14193 Berlin Germany
Tomasz Wandzel Intitute of Environmental +48 5820 17 28 +4858 204950 Protection ext 310 ul. Kollataja PO-81-33I, Gdynia Poland
1998 WKBT Report 5 ANNEX 2
AGENDA
Tuesday, October 4
10.00-11.00 Introduction, practical issues 11.00-12.30 Polychaeta: Phyllodocidae 12.30-13.30 Lunch 13.30-16.30 Polychaeta: Nephtyidae, Nereididae, Polynoidae
Wednesday, October 5
9.00-10.00 Porifera 10.00-12.30 General discussion on sampling techniques, preservation practices and interactions between monitoring units and zoological museums 12.30-13.30 Lunch 13.30-16:30 Polychaeta: Pholoidae, Cirratulidae, Ampharetidae
Thursday, October 6
9.00-11.00 Gastropoda: Hydrobia, Potamopyrgus 11.00-12.30 Gastropoda: tectibranch Opisthobranchia 12.30-13.30 Lunch 13.30-15.30 Bivalvia: the Cardium-complex 15.30-16.30 Polychaeta: Lumbrineridae
Friday, October 7
9.00-11.00 Polychaeta: Spionidae, Flabelligeridae 11.00-12.30 Amphipoda 12.30-13.30 Lunch 13.30-15.30 Feedback and general discussion on future workshops and the role of ICES
6 1998 WKBT Report ANNEX 3
PHYLLODOCIDAE
PIlyl/odoce character maclllata (Lilllle, (767) mllcosa 0rsted, 1843
ventral cirri ofmidbody rounded or with obtuse :wit~ acute point distally I point
midlateral rows on proximal 6-8 papillae (rarely 9) 9-11 papillae part ofproboscis with
dorsal tentacular cirri of segment 7-9 segment 9-11 sgments 2 and 3
prostomium anterior to eyes not particularly dark with dark pigment
segment 1 ventrally not particularly dark with dark pigment ("posterior lip")
tentacular cirri subdistally often with dark spot without dark spot
segments 3 and 4 dorsally with dark pigment only with middorsal spots 2
greenish-yellow pigment as middorsal spots, between dorsolaterally, at base of present 3 dark spots each parapodium ..
I Note that in both species the ventral cirri ofthe midbody are longer than the neuropodium. It is important that a parapodium be removed to correctly judge this character, as all ventral cirri appear pointed in dorsal view. I• 2 As on more posterior segments, where such middorsal spots are found in both species. I 3 .- This charader is only useful with live material, as the greenish-yellow pigment always disappears when the animals are transferred from formalin to alcohol. .. These two spots are sometimes connected across the dorsum as a thin transverse band, but this is still easily distinguishable from the middorsal spot found in P. maculata.
1998 WKBT Report 7 Eteolle character flam (Fabricius, 1780) 1 [onga (Fabricius, 1780) 1
sides ofprostomium in convex straight or slightly concave dorsal view
slightly thicker than dorsal not thicker than dorsal ventral tentacular cirri tentacular cirri tentacular cirri about 4 times as large as about twice as large as dorsal cirri ofmidbody 2 neuropodium neuropodium
colour ofliving animals pink or orange white or yellow 3
1 Pleijel (1993: MIOS 8) noted that both species bearing these names in our area may be different from those originally described (both from Greenland). Material from Iceland and the Faroe Islands appears to show intermediate morphologies, so species clines may be involved. Until further work is done, I would suggest that both names can routinely be used for specimens from the Kattegat and the Baltic. . 2This is, in my opinion, the most reliable character. In order to correctly assess it, a parapodium I. must' be reinoved and laid flat, preferably in posterior view. Note that the classical character in the literature, dorsal cirri broader than long in E. flam and longer than broad in E. IOllga, is very unreliable aS these differences are only apparent on very large specimens; otherwise the dorsal cirri are about as long as wide in both species. . 3 Paradoxically, specimens ofE. tonga ?ften turn Y§Y dark following preservation.
EUlIlit!a character baltusiensis Bergström, 1917 sanguinea (0rsted, 1843)
conspicuously broader than slightly longer than broad, tip dorsal cirri ofmidbody 1 long, with a drawn-out tip not drawn out
about 4 times as large as about twice as large as dorsal cirri ofmidbody 1 neuropodium neuropodium
asymmetrical and sharply almost symmetrical and ventral cirri ofmidbody 1 pointed bluntly pointed prostomium often with dark pigment not particularly dark without or with only weakly with conspicuous transverse dorsum developed transverse segmental bands segmental bands
1 These characters only reliable on animals longer than 10 mm.
8 /998 WKBT Report ------
ANNEX 4
SELECTIVE LIST OF REFERENCES FOR POLYCIIAETA FROM TIIE KATTEGAT AND TIIE BALTIC-
Holthe, T. 1991. Identification of Annelida Polychaeta from northern European and adjacent Arctic waters. - GWlIleria 66: 1-30. [An excellent paper with a family by family listing ofrelevant literature. ]
Phyllodocidae: Pleijel, F. 1993. Polychaeta Phyllodocidae. - AIal'ille !llI'erlehrales ojSc.:al1llillal'ia 8: 1-159. [The bible.]
Nephtyidae: Rainer, S. F. 1991. The genus Neph(l's (Polychaeta: Phyllodocida) ofnorthern Europe: a review ofspecies, including the description ofN. pulchra sp. n. and a key to the Nephtyidae. lIelKolüllder Ivfeeresulllel'suclulI1gell 45: 65-96. [The bible.]
Polynoidae: Loshamn, -A.-A. 1980. En systematisk og dyregeografisk underokelse over skjellrygg-gruppen (Familie Aphroditidae sensu Fauvel 1923). -Masters' Thesis. University ofOslo. [In Norwegian, but with excellent figures and based on a large amount ofmaterial. Possibly available from: Zoologisk Institutt, Universitetet i Osol, Postboks 1050, Blindern, Oslo 3, Norway.] Tebble, N. & S. Chambers. 1982. Polychaetes from Scottish waters. A Guide to Identification. Part I. Family Polynoidae. - Royal Scottish Ivfuseum Sludies. Royal Scottish l'vfuseum, Edinburgh. 73 pp.
Lumbrineridae: Winsnes, I. M. 1980. En systematisk og dyregeografisk undersokelse over familiene Eunicidae _ og Lumbrineridae (Polychaeta) fra Norge. - Masters' Thesis. University ofOslo. [In Norwegian, but with excellent descrip~ions. Possibly available from: Zoologisk Institutt, Universitetet i Osol, Postboks 1050, Blindern, Oslo 3, Norway.] Frame, A. B. 1991. The lumbrinerids (Annelida: Polychaeta) collected in two northwestern Atlantic surveys with descriptions ofa new genus and two new specie?- j)roceedillgs 0/the Biological Socie(l' 0/WashillgtOIl 105: 185-218. [With good information on the latest swarm oflumbrinerid genera and the best description ofScole/oJ11ajragilis.]
Spionidae: Ramberg, 1. P. & T. A. Schram. 1982. A systematic review ofthe Oslofjord species ofPolydora Bosc and Pselldopolydora Czerniavsky, with some new biological and ecological data (Polychaeta: Spionidae). - Sarsia 68: 233-247. Sigvaldad6ttir, E. 1992. Redescription of Pl'iollospio hallYlIlellsis Laubier, 1966, ami re examination ofP. ockelmalllli Pleijel, 1985 (Polychaeta, Spionidae). - Oplwlia 35: 209 217. Sigvaldad6ttir, E. & A. S. Y. Mackie. 1993. Priollmpio sleells/1'lIpi, P. jallax and P. duhia (Polychaeta, Spionidae): re-evaluation ofidentity and status. - Sal'sia 78: 203-219.
1998 WKBT Report 9 ANNEX 5
HANDOUT ON SPONGE IDENTIFICATION AND HALTIC SPONGES
Idcntification ofspongcs a) Taxonomical problems with sponges Sponges (Porifera) is one of the few large phylae (about 6000 li ving species) where many higher category taxa have not yet been finally outlined. Over the last 20 years orders have been erected, split and laid down again,and the same goes for families, not to speak of genera·and species. There are several reasons for this: 1) Many original descriptions of sponge species are short and in sufficient with todays eyes, and sometimes the original material has disappeared or is insufficiently labelIed, so there is doubt about the identity of the single specimens in relation to the des cription ("The type specimen problem") . 2) Historically, terminology has been different in different lan guages ahd as used by various "schools", making it difficult to compare descriptions especially of very similar species or speci mens .where small differences are important (llThe terminology pro blem) . 3) Ideas on species variation has changed several times over time, in some periods leading to unjustified splitting into too many spe eies, in others to unrealisting synonYmisation ("The splitter lumper problem)". 4) Character deficiency, viz. low number of and high variation in taxonomically usable characters, is a wellknown problem in many sponge taxa. It has lead to a search for new charaeter complexes within fields as reproduction, genetics and bioehemistry, so far with'some success in single cases, but difficult to carry through for a larger number of species ("The eharaeter problem"). 5) While most sponge species show only little variatiorr within the 4t same population or between different populations, other species may vary literally from one side of an individual to another, certainly between individuals and over both the geographical and bathymetri- cal range as weIl as with season and according to loeality faetors, for example exposure to waves or currents (llThe variation pro blem)".
b) The identification proeedure Three categories of information should be considered: 1) The locality, substrate, depth, and exposure to current and light. 2) The general outer morphological characteristics, at best in fresh condition, such as form, dimensions, presence of oscula, _su;:_~a_c.e, e()~~l~r! texture and smell "- ._ 10 /998 WKBT Report These characteristics are often difficult to express in more exact terms, but nevertheless the appearence of a species can be experienced as very characteristic, once you are familiar with the local fauna, because only one or a few species of the genus are represented. 3) Skeletal characteristics such as presence and geometric pattern of spongin and spicule fibers, and the number, dimensions and ar rangement of spicule types. Generally spoken, the organization of the skeleton, the position of the spicules in the skeleton, and the kinds of spi cules considered, together will lead to order, family and genus, while spicule dimensions are species characters. Skeletal organization is sometimes difficult to make out, and in a local fauna, one can often directly use the kinds of spi cules present and their dimensions as a good starting point for identification. Within some groups, for example Haliclona, where very few characters are at hand, it is, however, always necessary to get an idea of th~ skeletal arrangement.
. In many cases the ~nformation can be drawn from very small (at most, a couple of mm maximum dimension) pieces of the sponge taken from different parts of the sponge body and squeezed in water (ar alcohol) between slide and cover glass. Always start with the low power objective of the compound microscope to get an overview. This kind of preparation is fast to make, and is not permanent. The risk-is that skeletal arrangement is dif-ficult to make out because of distorting, and that small or rare spicule types are not discovered. A better idea of the skeleton can one get from handmade sec tions, which should"always be made from specimens which have been hardened instrong alcohol for at least some days. Use a razor bla de and cut as thin as possible. One section should be perpendicular to the surface. In case of cylindrical growthform both tranverse and longitudinal sections should be taken. A section made parallel to the surface, including dermal membrane mayaIso be useful. Such sections can be studied in water or alcohol at low power, and then trown away,~of they can be taken taken through adehydration and mounting procedure to give permanent slide preparations. Clean spicule preparations are necessary when thefi~e struc ture of the spicules is to be studied and when numerous measure ments are taken. A small piece of sponge tissue (or several small pieces originating from different parts of the sponge (in all, the size of a pea) should be boiled in a few"ml concentrated nitric acid). In the case of calcareous sponges chlorine or a similar ba sic bleaching agent can be used. After boiling (in the case of ni tric acid, when the acid besomes clear again or when no more brown fumes are emanated), the remaining acid is pipetted off, and the spicules are cleaned in destilled water (be careful not to loose too many spicules during the procedure). Then the spicules can be rinsed.in 96% alcohol. With a pipette a portion is spread on a sli de, the alcohol burned off (or let to dry), and the dry slide with spicules"mounted with a cover glass.
11 /998 WKBT Report . Having the necessary preparations at hand, either use a key (one is given herein, but a few are also found in same of the books referred to in the list of "local" literature), or try to find a matching spicule picture. When an idea of the species name has been reached, always read the description very carefully in order to control. The key does only give a limited degree of certainty. It is always a possibility that there are more species in the area than keyed out, either because they were not found at the time the key ~as constructed, or the'key was been made for a different geo graphical area. Be aware that because of numerous synonymization problems the same species may be mentioned under different generic or species names in different books.
"
A selection of literature covering the sponge fauna of Skagerrack, Kattegat, The Danish Belts and the Baltic sea.
Alander, H., 1942: Sponges fram the Swedish West-coast and adj acent . waters. Göteborg. 95 pp. Arndt, W., 1928: Porifera, Schwämme, Spongien. - Tierwelt Deutsch- . lands 4: 1 - 94. Arndt, W., 1935: Porifera. ~'Tierwelt der Nord- und Ostsee III a: , .1-140. Barthel, D. & O.S.Tendal (in. prep.): The Baltic spange fauna: " Ecology, taxonomic composition, species distribution, and annotated bibliography and key (working title) . Hansson, H.G., 1994: Sydskandinaviska marina flercelliga evertebra ter. - Länsstyrelsen i Göteborgs och Bohus Län 1994:15. 205 : pp. (Porifera, p. 2-11) Tendal, O.S., 1967: On the freshwater sponges of Denmark. , Videnskabelige Meddeleüser fra Dansk naturhistorisk Forening : 130: 173 - 178. Tendal, O.S., 1973: De danske farvandes boresvampe. - Flora og ,Fauna 79: 105 - 108. , Tendal, O.S._ & T. Brattegard, 1997: Phylum Porifera. Pp. 65-80 in .' 'Brattegard, T. & T. Holthe (eds.): Distribution of marine, benthic macroorganisms in Norway. - Research Report for DN 1997-1. Directorate for Nature Management, TrondheJ:m, -Norway. 409 pp. Veethaak, A.D., R.J.A. Cronie & R.W.M. van Soest, 1982: Ecology and distribution of two sympatric, closely related sponge species, Halichondria,panicea (PalIas, 1766) and H. bower banki Burton, 1930 (Porifera, Demospongiae), with remarks on their speciacion. - Bijdragen tot de Dierkunde 52: 82-102. Weerdt, .W. H. de, 1986: A systematic revision of the north-eastern Atlantic shallow-water Haplosclerida (Porifera, Demospon-' giae) , part II: Chalinidae. - Beaufortia 36: 81 -165.
/998 lVKBT Report 12 Baltic sponges
List of species recorded from the Danish belts to The Bottom Sea
Species in paranthesis have been recorded only once, and that in older time. Species marked with an asterisk have not been found in the area but they are expected to occur there.
Class CALCAREA
Leucosolenia botryoides (Ellis & Solander, 1786) Leucosolenia complicata (Montagu, 1818)
(Clathrina coriacea (Montagu, 1818» (Guancha blanca (Miklucho-Maclay, 1868»
(Sycon ciliatum (Fabricius, 1780»
Class DEMOSPONGIAE
Polymastia robusta (Bowerbank, 1861)* Polymastia mammillaris (Müller, 1806)
(Pseudosuberites sulphureus (Bowerbank, 1866» (Terpios fugax (Duchassaing & Michelotti, 1864»
Suberites ficus (Johnston, 1842) (Cliona celata Grant, 1826) (Cliona vastifica Hancock, 1849) (Mycale lobata (Bowerbank, 1866) Haliclona urceolus (Rathke & Vahl, 1806) Haliclona rosea (Bowerbank, 1866)
(HaliclonaJ Acervochalina limbata (Montagu, 1818) Halichondria panicea (PalIas, 1766) Halichondria bowerbanki Burton, 1930*
Halisarca dujardini Johnston, 1842 % 5Y1ttU " D~ ~ ..k....A Ephydatia fluviatilis (Linne, 1758) .' Spongilla lacustris (Linne, ~758)
1998 WKBT Report 13 o •
Key to the species of sponges found in the Baltic
1a Without spicules Ha~isarca dujardini 1b With spicules 2 2a Spicules calcitic; at least some of them triradiates 3 2b Spicules siliceous; are rod-shaped 7 3a Form as a little bush of thinwalled tubes; at the open end the tubes may have a corona of spicules, but that is always shorter than the tube diameter 4 3b (Form as a little thick-walled urn with an oscular corona of spicules longer than diameter of the osculum ~. Sycon ci~iatum) 4a The spicules comprises a rod-shaped type S 4b (No rod-shaped type present 6) Sa Mostly triradiates; if there are tetraradiates the fourth ray is rudimentary Leucoso~enia complicata Sb tetraradiates common and their fourth ray is weIl developed .. ,~ ...... '.' Leucosolenia botryoides ,., 6a (triradiates regular Clathria coriacea) 6b (trir~~iates sagittal : Guancha blanca) 7a Spicules of the main skeleton are styles or tylostyles 8 7b Spicules of the main skeleton are oxea lS
8a Free living 10 8b (Boring galleries in calcareous material , 9)
9a (With two types of microscleres C~iona vastifica) 9b (no microscleres C~iona celata) 10a with long papillae on the upper surface 11 lOb Without papillae on the upper surface 12 11a Body surface hispid ("hairy"), different from the more smooth • papillae surface Polymastia~ammi~laris ~ 11b Body and papillar surface similar, almost smooth ~, ...... '...... Po~ymastia robusta 12a Skeleton spicules styles; microscleres chelae ...... ~ '. . • .... .' Myca~e ~ obaca 12b Skeleton spicules tylostyles 13 13a Body lumpy, with rod-shaped microscleres Suberices ficus - 13b (Body thinly encrusting, no microscleres 14) 14a (Dermal membrane strongly developed provided with numerous spicules, sponge interior massive Pseudosuberites su~phureus) 14b (Dermal membrane thin, without spicules, sponge interior ge~atinous Terpios fugaxJ
.'."
1998 WKBT Report 14 15a No fibers of spongin and spicules; spicules lying in confusion ·...... 16 15b spongin and fibers form pronounced fibers arranged in a network · " 17 16a Dermal skeleton a regular network of bundles of spicules ... · Ha~ichond.ria panicea 16b dermal skeleton a rather open and irregular network of spicule bundles and single lying spicules .... Ha~ichond.ria bowerbanki 17a Fibers very spongin-rich; sponge elastic Acervocha~ina lirnbata 17b Spongin mostly only at the ends of spicules 18 18a No gemmules; lives at marine salinity 19 18b With gemmules; lives in freshwater or at very low salinity... · " 20 19a Body of tubular elements, on short stalk " Ha~iclona urceolus 19b Body cushionshaped, encrusting ~ Ha~iclona rosea
20a Gemmule spicules spiny rods Spongilla lacustris 20b Gemmule spicules amphidiscs Ephydatia fluviatilis
1998 WKBT Report 15 o~ H.d.
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~ P, i ~ c.J I ~.s. <..ox.~ :Dermal rnun ~ b r-of'l-f. skdt' E:.on ({ra M1 ?Ja ri. man : P~(JboJl.j MlIj. ß" \\. Il 19 S"! ) c 18 /998 WKBT Report Spi c..u!-e s Coxea) u~ceo{us /998 WKBT Report 19 Gemmu (~ Sp\cu I H G\'\cl tl"n Q" bod~ spicul u .s k~(d:o,., spicur4~ (O)(('Cl) c 20 1998 WKBT Report ANNEX 6 FINAL NOTES If you believe that a specimen resembles a species but "something is wrong" there are several ways to deal with this situation. PllO/O€! halliea will be llsed as an example. A) VOll believe it probably ~ PllO/Ot.! ha/liea, but important structures are missing in order to be certain. PllO/O€! ?ha/liea B) You be1ieve it is not Photo€! haltiea, you don't know which species it is, but ofthe species YOll know, it most c10sely resembles Photo€! ha/liea. . Photo€! cf ha/liea [cf stands for confer, i.e., look there to see something resembling this] Ph%e near ha/liea C) Too many characters are missing and you don't feel very certain about Ph%e ha/liea at all. e-j~,/' Ph%e sp. 0) Too many characters are missing and you have no idea whether it is P/l0/OI! ha/liea. Oon't glless llnless you have very gooef sllpplementary information. Ph%e sp. Finally, if anyone would like to contact me er OIe Tendal, our e-mail adresses are [email protected] and [email protected] and our mail address is: Zoologisk Musuem, Universitetsparken 15, DK-2100 Copenhagen 0, Oenmark. I thank all of you for your active participation - it has been a pleasure to arrange this _. workshop. Danny Eibye-Jacobsen /998 WKBT Report 21 ANNEX 7 CIRRATULIDAE FROM TUE KATTEGAT, 0RESUND, AND ßALTIC Kcys and notcs for idcntifying spccics 2 November 1997 - Not to be cited without written permission from the author. Zoological Museum, University ofCopenhagen, Universitetsparken 15, DK-2100 Copenhagen 10, Denmark Tel +45-3532 10 17 - Fa'< +45-35 32 10 10 - E-mail: [email protected] Cirratulidae from the Kattcgat, 0resund, and Baltic: Key to genera I. T\vo grooved tentacles 2 .Two groups ofgrooved tentacles Cirratulus (part) (p.3) 2. Chaetae exclusively cap~llaries Aplte!oclzaeta (p. I) Chaetae include capillaries + variously sculptured acicular chaetae (aciculars) 3 3. .Aciculars distally spooned; grooved tentacles arising laterally Dodecaceria (p. 4) .Acic~lars not distally spooned; grooved tentacles arising dorsally 4 4. .Aciculars small, distinctly bidentate, usually present in neuropodia (Le., ventrally) from , chaeti'ger 2 or 3, appearing later in notopodia Caulleriella (p.2) Aciculars not as above, fIrst present in middle or posterior segments 5 5. Aciculars slender, similar in width to capillaries, fIrst recognizable in posterior segments (arise through a transition from capillaries), distally irregular or with knobby tips, not altemating regularly with capillaries : Tltaryx (p. 5) Aciculars broader, mostly with tips entire, altemating regularly with capillaries 6 6.;Anterior region broad, posterior region narrow and taillike; cuticle ofter: tessellate; aciculars not very conspicuous, usually flat against body Cirratulus (part) (p. 3) ~~ - Body spindle-shaped or of similar width throughout; aciculars in posterior segments usually conspicuous, forming complete or partial einetures, often. prntruding at right angles to body Cltaetozone (p. 3) Remarks There are presently 10 recognized genera ofcirratulids (Blake 1996), but 4 of these: Cirrifor mia, Monticellina, Protocirrineris, and Timarete are not known to occur in the area under consideration. Caulleriella? serrata Eliason, 1962 will not key out with the above key, but as it is still only known from a few specimens this is not likely to cause any problems. - Kcy to spccies ofApltelocltaeta Blake, 1991 1. Cryptofaunal, in Laminaria holdfasts or other protected substrata. Small species with , large eggs and brood protection Apltelocltaeta sp. Infaunal species, in sand or mud 2 22 1998 WKBT Report f... • ~ , ~ J. " 2. Body short and spindle-shaped, posterior segments distinct, often zigzag in dorsal view; hennaphroditic, viviparous A. vivipara (Christie, 1984) Body long, segments relatively short anteriorly and posteriorly, often beadlike in middle ...... ; :; : Aphelochaeta spp. Remarks Aphelochaeta was erected by Blake (1991) for bitentaculate cirratulids with exclusively capillary chaetae, as the type species of Tharyx was found to have slender, "knobby-tipped" aciculars in the most posterior segments (Blake 1991). At least 4 species ofAphelochaeta are present in Danish waters, and perhaps more in the Kattegat-Baltic area. One of these appears to be Tharyx marioni sensu Gibbs (1971), another is T. marioni sensu Hartmann-Schröder (1971, 1996) and many other authors, but neither seem to be this species. The identity of T. meintoshi (Southern, 1914) sensu Kirkegaard (1996) is uncertain. The presence ofonly capillary chaetae suggests that Aphelochaeta is the eorrect genus, but the identity ofMcIntosh's material (by Southem 1914 referred to Cirratllllls) and also the identity of Cirratu11iS filiformis Kefcrstein, 1862 (= Aphelochaeta) is presently not clear. A. vz\,'ipara was transferred to Aphelochaeta from Tharyx by Hartmann-Schröder (1996). In Danish waters I'have only seen the species in material from the Limfjord. ~) Apheloclzaeta sp. from eryptofaunal habitats such as Laminaria holdfasts is an unnamed hennaphroditic species with brood protection in the burrow and will be described by me (M. E. Petersen, in prep.). T/iaryx mliitibranchiis [sie] (Grube, 1863) was reported from Helgoland by Hart mann-Schröder (1971) and Gillandt (1979); Hartmann-Schröder (1996) later referred it to Aphelochaeta mllltibranchis. The speeies is not the one described by Grube as HeterocirnlS mliltibranchis Grube, 1863. Grube's type matenal has been examined and will be described separately. The speeies has not been seen in northem European waters. Key to species of Calil/eriel/a Chamberlin, 1919 1. Cryptofaunal in Laminaria holdfasts; pygiditim entire; live animals bright yellow ...... : C parva Gillandt, 1979 Infaunal in soft sediments; pygidium with two lobes; live animals olive green with golden sheen C bioculata (Keferstein, 1862) Remarks Calilleriella pan'a was originally described as C bioclliata pan'a Gillandt, 1979, and raised to a full species by Hartmann-Schröder (1996). The species may be identical with C fragilis (Leidy, 1854), described from Rhode Island, eastem USA, and cornmon along the NE American east coast (M. E. Petersen in Hartmann-Schräder 1996). The species is henna phroditic and has brood protection in the burrow (petersen 1994); it may occur together with Aphelochaeta 'sp. (see above). The above are the only two species of Cau/lerie/la presently known from Danish waters. Caulleriella caputesoeis (Saint-Joseph, 1888) is not a Calilleriella; the report by Gillandt (1979) is Cirratulus incertus (M. E. Petersen, unpublished). Calilleriella alata (Southern, 1914) is a more southem species not present in Danish waters. Cau/lerie/la? serrata Eliason, 1962 is not a true CalliIeriella; its generic placement is presently uncertain.. /998 WKBT Report 23 Cltaetozone l\1almgren, 1867 Cha,etozone setosa l\lalmgren, 1867 I Chaetozone setosa has been widely interpreted and the name.has been used for a large assortment ofspecies from all parts ofthe world. The true C. setosa is a northern species and probably not present in the area underconsideration. At least two species of Chaetozone are easily recognizable, but more are undoubtedly present. A lectotype of C. setosa has been selected from Malmgren's material from Spitsbergen, and a redescription of the species is in preparation (11. E. Petersen & S. F. Garcia-Martin, in prep.). For the present, the following designations are suggested. Key to some Kattegat-Baltic speciesof Cltaetozone I. Mud-bottom species; branchiae' I arising imrnediately posterior und slightly lateral to dorsal tentaeles; posterior segments with nearly complete cinctures, appearing zigzag in :'dorsal vie\v ; Cltaetozone cf. setosa •Sand-bottom species; branchiae I arising level with or anterior und lateral to dorsal .tentaeles; posterior segments with incomplete einetures, not particularly zigzag in dorsal .vie\v : : Clzaetozone sp. F group Remarks In both species the prostomium is pointed, but it is relatively longer und a bit more slender in Clzaetozolle sp. F group, where the first branchiae (branchiae 1) arise either level with the base of the,tentaculophores or slightly anterior to these; the "tops" of the tentaculophores are ofteh level with chaetiger 2. Aciculars of posterior segments are relatively short, and the meinbrunous "webbing" between posterior chaetae is not pronounced. .: In Clzaetozol,e cf. setosa, the fIrSt -branchiae are nearly in line with the tentaculophores, which are relatively short and weIl separated (elose together in sp. F group), and the aciculars of posterior segments are relatively long, often with those of right and left sides crossing dorsally where aciculars are best developed. Webbing between aciculars of posterior segments is often prominent. Key to species of Cirratulus Lamarck, 1818 1. ,Two tentacles, no eyes, cuticle tessellate C. caudatus Levinsen, 1893 ,Two groups oftentacles, 2 or more red eyes (more in larger individuals) .:.::; .. " C. incertus l\lcIntosh, 1923 Remarks Cirratulus caudatus is a fairly large infaunal species; it is not uncommon, but is rarely taken as complete specimens. Although the original description implies that there were two groups of tentaeles, none of the syntypes have more than two tentacles, which is also the case with more recently collected material (M. E. Petersen, unpubI. observations). :: Cirratulus. incertus is a small cryptofaunal species, usually found in sheHs bored by .. species of the boring sponge Cliona or in dead barnacle tests or moHuse shells in Laminaria holdfasts. It reproduces asexually by fragmentation (petersen 1991); regenerating specimens are'~ommon in late summer and earlyfall, but can be found )rear round. Sexual reproduction occ'urs in late sUf!lI11er; male and fe"?-ale epitokes with numerous long capillaries swim up off 24 1998 lVKBT Report • t" f, ~,' .t' \. .. the bottom and spaw-n in the free water mass, after which the epitokes die. The species is not very common in pure mud but may be present if there are dead shells. The epitokes were described by Stephenson (1950b, as C. cirratus). In both this and other species of Cirratulus, the dorsal part ofthe peristomium forms a transverse fold resembling a kind offorehead. C cirratlls (O.F: l\IüiIer,' '1776) (type locality Sondmor County,\V Norway), is a Iarge, black-eyed, infaunal species common in British waters but never observed in the Kattegat to the Baltic; reports of C. cirratus from these areas are referable to C. incertus. C. cirratus never reproduces asexually and does not become epitokous when sexually mature. Its biology has been studied in detail by Olive (1970, 1971, 1973); eggs are laid in an irregular jelly mass and development is direct (Stephenson 1950a). A color photo of a female from Devon, S\V England, can be seen at: http://www.aki.ku.d.k/zmuc/inv/staftlmep2.htm Cirratulus sp. is a small oculate species that has been found in masses ofPomatoceros tubes off Tjämö, western Sweden. The species resembles C. incertus but differs from it in being yellow with red eyes. To date no yellow C. incertus have ever been seen, even though numerous live specimens have been observed. Cirratlilus sp. is small and only known from a few specimens; its status is unclear and it may be undescribed (M.E. Petersen, unpubl.). Any additional material would be most welcome. Genus Dodecaceria Orsted, 1843 Syn.: Zeppelina Vaillant, 1890 Dodecaceria conc/rarum 0rsted, 1843. ?Syn.: Dodecaceriafimbriata (Verrill, 1879) Syn.: Dodecaceria calilleryi Dehorne, 1933 Not Dodecaceria conchamm sensu Gibson (see below) Remarks Only Dodecaceria concharllm is likely to be encountered in the Kattegat-Baltic area. The species is common in dead shells that have been bored by species ofthe boring sponge Cliona or in dead bamacle tests or bivalve shells in Laminaria holdfasts. It also occurs in encrusting coralline algae ("Lithothamnion " spp., now referred to other genera), but this is not a common habitat in Danish waters. The species reproduces asexually by schizogamy or fragmentation in I' late summer and fall, and single middle segments or anterior or posterior ends may be found I regenerating the missing portions. The original segmentes) are usually recognizable by their darker color, the regenerated portions being paler. The species is weIl illustrated by Dehorne (1933, as D. calilleryi). Gibson & Clark (1976) and Gibson (1978, 1979, 1981, 1996) have provided much useful information on this and other species ofDodecaceria, but unfortunately Gibson chose to use (also in 1996, see below) incorrect younger names (fIrst D. calilleryi, later D.fimbriata) [or the type species of the genus, D. concharom, which is the only species present at either of 0rsted's type localities (the 0resund off Hellebrek and the Kattegat off Jutland between Frederikshavn and Skagen) or in any Danish waters. D. jimbriata (Verrill, 1879) is an eastern North American species described in the genus Heterocirrus; it is either a junior synonym of the true D. concharum or a distinct but very similar North American species (see George & Petersen 1991 for a review of the main synonyms); its name has no place in th~ taxonomy ofEuropean Dodecaceria. D. cQulleryi Dehorne, 1933 was erected for an asexually reproducing species of Dodecaceria from northem France. The species is identical with the true D. conchanlm, but 1998 WKBT Report 25 ------_._------ Dehorne did not know that only one species was present in Danish waters. The name D. caulleryi is common in textbooks in connection with illustrations of asexual reproduction by schizogemisis (multiple fission). Zeppelilla Vaillant, 1890, is a genus earlier referred to the Ctenodrilidae and believed to reproduce only asexually; George & Petersen (1991) pointed out that most species of Zeppelina were referable to asexual regenerates orjuveniles ofDodecaceria spp.. D. concltarum has a life cycle similar to that of Cirratu/us incertus. with asexual reproduction by fragmentation and sexual epitokes with gametes shed into the free water masse The life span of the species was by Gibson & Clark (1976) estimated to be about 3 years or more. The parthenogenetic species referred to by Gibson and by Hartmann-Schröder (1971, 1996) as D. concllantm is a distinct species (D. ater (Quatrefages, 1866» and not present in Danish waters, where the salinity appears to be too low for it (Gibson 1996). " Tlzaryx \Vebster & Benedict, 1887 Tltary.'(; killariensis (Southern, 1914) Remarlcs Tlzaryx killariellsis is the most common - perhaps the only - true Tharyx in Danish waters. It is common in the \Vadden Sea and is the T. marioni of Farke (1979). The species-is weIl illustrated.in Southern (1914). It was transferred to Tharyx by Blake (1991); in Hartmann Schröder (1971) and Kirkegaard (1996) it is referred to Caulleriella. ,The name T. marioni has been applied to numerous species from many parts ofthe world (see remarks under Aphe/ochaeta); the true identity ofthe species is under investigation (M.E. Petersen, in prep.); it does not appear to occur in the Kattegat-Baltic area. Tltaryx viJ:lpara Christie, 1984 was by Hartrnann-Schröder (1996) referred to Aphelo clzaeta (see comments under this). References Blake, J. A. 1991. Revision of some genera and species of Cirratulidae (polychaeta) from the western North Atlantic. - In M. E. Petersen & J. B. Kirkegaard (eds): Systematics, Biology and Morphology of World .' Polychaeta, Proceedings of the 2nd International Polychaete Conference, Copenhagen, 1986.0phelia Suppl. 5: 17-30. Blake, J. A. 1996. 8. Family Cirratulidae Ryckholdt, 1851. Including arevision of the genera and species from the eastern North Pacific. - In J. A. Blake, B. Hilbig, & P. H. Scott (eds.): Taxonomie Atlas of the Benthic Fauna of the Santa Mafia Basin and the Western Santa Barbara Charme!. Volume 6 - The Annelida Part 3, Polychaeta: Orbiniidae to Cossuridae. Santa Barbara Museum ofNatural History, Santa Barbara, California: Pp. 263-384, figs. 8.1-8.47. , Christie, G. 1984. A new species of Tharyx (polychaeta: Ciratulidae) from the estuaries in north-east England. Sarsia 69: 69-73. Christie, G. 1985. A comparative study of the reproductive cycles of three Northumberland populations of Chaeto::one setosa (Polychaete: Cirratulidae). - Journal of the Marine Biological Association of the U. K. 65: 239-254. ' . Dehorne, A. 1933. La schizometamerie et les segmentes tetragemmes de Dodecaceria caulleryi. n. sp. - Bulletin Biologique de la France et Belgique 67: 298-326, pis. 10-12,6 figs. . . Eliason, A. 1962. Die Polychaeten der Skagerak-Expedition 1933. - Zoologiska Bidrag frän Uppsala 33: 207 293. Farke, H. 1979. Population dynamies, reproduction and early development of Thal'}~Y; mariom' (Polychaeta, Cirratulidae) on tidal flats of the Gennan Bight. - Veröffentlichen aus dem Institut für Meeresforschung in Bremerhaven 18: 69-99. 26 1998 WKnT Report George, J. D. & M. E. Petersen. 1991. Validity ofthe genus Zeppelina VaiIlant (polychaeta: Ctenodrilidae). -ln M. E. Petersen & 1. B. Kirkegaard (eds): Systematics, Biology and Morphology of World Polychaeta, Proceedings of the 2nd International Polychaete Conference, Copenhagen, 1986. Ophelia Supp!. 5: 89-100. Gibbs, P. E. 1971. A comparative study of reproductive cycles in four polychaete species belonging to the family Cirratulidae. - Journal of the Marine Biological ASsociation of the U. K. 51: 745-769. Gibson, P. H. 1978. Systematics of Dodecaceria (Annelida: Polychaeta) and its relation to the reproduction of the species. - Zoological Journal of the Linnean Society 63: 275-287. Gibson, P. H. 1979. The specific status of two cirratulid polychaetes, Dodecaceriafimbriata and D. caulleryi, compared by their morphology and methods ofreproduction. - Canadian Journal ofZoology 57: 1443-1451. Gibson, P. H. 1931. Gametogenesis in the cirratulid polychaetes Dodecaceria concharum and D. caulleryi. _ Journal ofZoology, London 193: 355-370. . Gibson, P. H. 1996. Distribution of the cirratulid polychaetes. Dodecaceria fimbriata, D. concharum and D. diceria in European waters between latitutes 48°N and 70oN. - Journal ofthe Marine Biological Association ofthe U. K. 76: 625-635. Gibson, P. H. & R. B. Clark. 1976. Reproduction in Dodecaceria cau/leryi (polychaeta: Cirratulidae). - Journal ofthe Marine Biological Association ofthe U. K. 56: 649-674. Gillandt, L. 1979. Zur Systematik, Autökologie und Biologie der Polychaeten des Helgoländer Felslitorals. - Mitteilungen aus dem Hamburgischen Zoolgischen Museum und Institut 76: 19-73. . Hartmann·Schröder, G. 1971. Annelida, Borstenwürmer, Polychaeta. - Die Tierwelt Deutschlands 58: 1-594. Hartmann-Schröder, G. 1996. Annelida, Borstenwürmer, Polychaeta. 2., neubearbeitete Auflage. - Die Tierwelt Deutschlands 58: 1-648. Kirkegaard, J. B. 1996. Havborsteorme II [Marine Polychaetes]. - Danrnarks Fauna 86: 1-451. Olive, P. J. W. 1970. Reproduction of a Northumberland population of the polychaete Cirratulus cirratus. Marine Biology 5: 259-273. Olive, P. J:W. 1971. Ovary structure and oogenesis in Cirratulus cirratus (polychaeta: Cirratulidae). - Marine Biology 8: 243-259. Olive, P. J.' W. 1973. The regulation of ovary function in Cirratulus cirratus (Polychaeta). - General and Comparative Endocrinology 20: 1-15. Petersen, M. E. 1991. A review of asexual reproduction in the Cirratulidae, with a redescription of Cirratulus gayheadius (Hartman, 1965), new combination, and ~mendation or reinstatement ofsome cirratulid genera. - Bulletin ofMarine Science 48: 592 (Abstract). Petersen, M. E. 1994. Hermaphroditic cirratulids (Annelida, Polychaeta) from Danish waters, with notes on . early development, description of a new species of Aphelochaeta Blake and a review of hermaphroditism among the Cirratulidae. - In J.-C. Dauvin, L. Laubier & D. J. Reish (eds.): Actes de la 4eme Conference Internationale de Polychetes. Memoires du Museum National d'Histoire Naturelle (Zoologie) 162. 634 (Abstract). Southern, R. 1914. Archiannelida and Polychaeta. Clare Island Survey Part 47. - Proceedings of the Royal Irish Academy 31: 1-160, pIs 1-15. Stephenson, W.,.l950a. The development of Cirratulus cirratus (O.F. Müller). Report of the Dove Marine Laboratory for 1943, sero III, No. 11: 7-20. Stephenson, W. 1950b. An epitokous cirratulid occurring in the Cullercoats tanks. Report of the Dove Marine Laboratory for 1948, sero III, No. 11: 21-30. - .- 1998 WKBT Report 27 Pholoidae from the Kattegat, 0resund, and BaUic Keys and notes for identifying species Mary E. Petersen 2 November 1997 - Not to be cited without written permission from the author. Zoological Museum, University ofCopenhagen, Universitetsparken 15, DK-2100 Copenhagen 0, Denmark Tel +45-35 32 10 17 - Fax: +45-35 32 10 10 - E-mail: [email protected] Spccics ofPholoe Johnston, 1839 from the Kattegat, Oresund and BaItic " Species included, type locality, principal synonyms: P/~oloe assimilis Orstcd, 1845 (Drobak, Oslo Fjord) I Syn.: Pholoe minuta ofauthors. [Not Fabricius, 1780] Pholoe baltica Orsted, 1843 (0resund and Kattegat) Syn.: Pholoe minuta ofmost N European authors. [Not Fabricius, 1780] :. Syn.: Pholoe tuberculata Southem, 1914 rvv Ireland) : Syn.: Pholoe inornata ofChambers 1985. [Not Johnston, 1839] 'Pholoe inornata Johnston, 1839. Type species. (NE England) Syn.: Pholoe minuta ofmany N European authors. [Not Fabricius, 1780] Pholoe pallida Chambers, 1985 (NE England) , Syn.: Pholoe cf. anoculata ofChristie 1982. [Not Hartman, 1965] • Key to species ofPltoloe Johnston, 1839 I.. \Vith eyes. Scales with or without pigment. FaciaI tubereie variable 2 Without eyes. Scales without pigment. Facial tubereie prominent, with papillae at base' ...... pallida , 2.' Scales with pigment. Elytral papillae evenly tapering or slender and capitate but not ,; moniliform. Facial tubereie inconspicuous Neuropodia without stylodes (papillae) on distal part 3 Scales without pigment. Elytral papilläe moniliform. Facial tubereie prominent, often as large as median antenna. Prostomium with dark pigment between eyes. Neuropodia with stylodes on distal part :::: : baltica 3. Prostomium with conspicuous black eyes, usually without other pigment. Elytra evenly pigmented brownish or blackish"often pale in deep water, covering dorsum completely, usually lying flat against dorsum. Elytral papillae marginal, slender, short and slightly capitate anteriorly, elongated and tapering posteriorly. Tentacular cirri smooth or irregular, never with distinct papillae ~: assimilis _ Prostomium sometimes with small amount of black pigment between eyes. Elytra often. strongly pigmented, with dark spots" usually not covering dorsum completely, leaving narrow middorsal strip bare, often ruffled and not lying flat against dorsum. Elytral papillae evenly tapering, never capitate, marginal laterally, on posterior margin of elytra becoming submarginal towards middorsum. Dorsal tentaculat cirri usually with 5 distinct . papillae on inner side ;nornata 28 1998 WKBTReport ." i'~ .....'" Q • 't.-." Remarks Small specimens of Pholoe can be very difficult to identify and good microscopes are absolutely necessary. \Vhile most species can be identified on characters ofthe anterior end, it is much easier to arrive at a certain identification if the specimen has at least some of the middle and posterior segTrients and elytra. As pignient or lack of such is usually an easy way to limit the choices, staining with Rose Bengal should be avoided ifat aII possible. Pltoloe assimilis usually lack interocular pigment, but this is present in some individuals or p'opulations. When such pigment is present the species may resemble smaIl P. baltica, but the latter has a distinct facial tubercle and neuropodia have numerous papillae (stylodes) distally, whereas these are lacking in P. assimilis. Because of its small size (most specimens are 5 mm or less), the species has often been considered a juvenile ofother species. The species is figured by Bick & Gosselck (1985: 208, fig. 2a-c, as P. minuta). P. baltiea is the species most often identified as P. minuta (Fabricius, 1780), e.g., Hartmann-Schröder (1971, 1996), but this is a species from SW Greenland and not known to occur in N Europe (M.E. Petersen, unpub!.). P. baltiea is closely related to P. longa (O.F. Müller, 1776), from S\V Greenland, and may be identical with it, but rather than introduce the name ofa Greenlandic species that may actually be distinct, I prefer to use the younger name .for a species described from the area (the 0resund) until the taxonomy of the P. longa group is settled. This way, the worst thing that can happen is that all P. balliea should be referred to P. longa. IfP. longa is used for N European material and the two species really are distinct, which s~ems to be the case, it will be necessary to specify which P. longa are in fact P. baltiea. Apart from the dark pigment between the eyes, the species is not pigmented, but the scales'. are ,often covered with a rusty accretion whieh can be brushed off. The species is figured by Hartmann-Sehröder (1971: 79, fig. 24a-d; and 1996: 74, fig. 24a-d [same as in 1971 edition], a11 as P. minuta), and Chambers (1985: 16. figs. 13a-b, 18a d, pls.A1-2, B1-2 (SEM), all as P. inornata). P. inornata always has at least some, usually 5, distinet papillae on the dorsal tentaeu lar cirri, a few on the ventral tc and one or more on the ventral cirri, hut the numher and development ofpapillae may vary. It sometimes has eolorless elytra and a smudge ofpigment hetween the eyes, and thus superficially resembles P. baltiea, hut can be distinguished by the ahsence ofstylodes (papiIlae) on the distal part of the neuropodia, the presenee ofpapillae on te and ve and the ineonspieuous faeial tubercle (eonspicuous in P. balliea). A left elytron ofthe speeies is figured in Gillandt (1979:23, fig. 3, as P. minuta). P. pdilida was described from Scotland and is also known from Danish waters, including the 0resund. It is figured in Chambers (1985: figs. l3e-d, 18e-h, p~ A~-4, B3-4). References Bick, A. & F. Gosselck. 1985. Arheitsschlüssel zur Bestimmung der Polychaeten der OstseelIdentification key for the polychaetes ofthe Baltic Sea. - Mitteilungen aus dem Zoologischen Museum in Berlin 61 (2): 171 272. Chambers, S. 1985. Polychaetes from Scottish Waters. Part 1. Families Aphroditidae, Sigalionidae and Polyodontidae. - Royal Scottish Museum Studies, Royal Scottish Museum, Edinburgh, 38 pp. Gillan~t, L.. 1979. Zur Systematik, Autökologie und Biologie der Polychaeten des Helgoländer Felslitorals. Mitteilungen aus dem Hamburgischen Zoolgischen Museum und Institut 76: 19-73. Hanmann-Schräder, G. 1971. Annelida, Borstenwürmer, Polychaeta. - Die Tierwelt Deutschlands 58: 1-594. Hanmann-Schräder, G. 1996. Annelida, Borstenwürmer, Polychaeta. 2., neubearbeitete Auflage. - Die Tierwelt Deutschlands 58: 1-648. 29 1998 lVKBT Report Ampharetidae from the Kattegat, 0resund, and BaItic Keys and Dotes for identifying some of the species :' l\lary E. Petersen 2 November 1997 - Not to be cited without written pennission from the author. Zoological Museum, University ofCopenhagen, Universitetsparken 15, DK-2100 Copenhagen 0, Denmark Tel +45-35 32 10 17 - Fax +45-3532 10 10 - E-mail: [email protected] Species ofAmpharete Malmgren, 1866 from tbe Kattegat, 0resund, and Baltic Species included, type localities, principal synonyms: Alnpharete aClitifrons (Grube, 1860) (Greenland) :, Syn.: Amplzarete grubei MaIrngren, 1866 Aillpharete baltica Eliason, 1955 (Baltic bennen Bornbolm and Bleldnge) Altlp/raretefalcata Eliason, 1955 (Bohuslän, Swedish west coast) Amp/raretejinmarchica (Sars, 1864 [1866]) (Ramfjorden, Troms, Nonvay) Syn.: Ampharete aretlea Malmgren, 1866 'Amp/rarete lindstroemi Malmgren, ~867 sensu Hessle, 1917 (Bohuslän, Sweden) " Key to above species ofAmpharete 1. Pygidial cirri and papillae long, slender, and pointed 2 Pygidial cirri long, pygidial papiIIae short and blunt 3 2.' Abdominal neuropodia and last 2-5 thoracic neuropodia with dorsal cirrus that is ~ distinctly longer than neuropodium aeutifrons rJ Abdominal neuropodia without dorsal cirrus (smaller specirnens) or cirrus maxirnally half -l.-\\.a.- c\.'.J. as leng as neuro-podium (larger specimens) baltica ~r:J-{ -5 .''-,c . 3. Paleae long, conspicuous. Pygidial cirri each with small eye at base 4 Paleae minute, about 6-8 per side,' short and stout with pointed tips, may be hidden by .bases ofbranchiae : faleata 'I I 4: Paleae 10-12 per side re Sars (1866),12-16 per side re Holthe (1986), relatively thick and broad, markedly and abruptly curved at tip. Groups of branchiae nearly touching mid- dorsally jinmarchiea Paleae about15 per side (about 7 per side re Holthe 1986), slender, fmely tapering, not abruptly curved at tip. Groups ofbranchiae widely separated lindstroemi Remarks During the late 1970s, problems encountered while attempting to identify material from the 0resund prompted me to make a pictorial key to the species ofAmpharete. This was based on the key to· Ampharete in Holthe (1975) und supplemented with infonnation from original descriptions and my o\~ observat~o.ns on material from Scandinavian waters. Later the more detailed MIOS volume on Terebellomorpha (Holthe 1986) was published. and more recently Hartmann-Schräder (1996) has provided a more comprehensive key to Ampharete in her 30 1998 WKBT Report .' '; : :.... '" revised edition ofDie Tierewelt Deutschlands (the 1971 edition did not treat A. haltiea or A. lindstroemi as distinct species). I have only recently acquired the 1996 edition und have not yet had time to try out the keys. The key presented here is based on my earlier key and own observations plus the keys and information in the above sources, which should be consulted for additional infonnation and figures. Amplrarete aeutifrons - Up to 45 mrn lang (Holthe 1986); up to 80 mm, 13 mrn wide in Arctie waters (Malmgren 1866: 363). Prostomium broadly rounded, usually with 2 small eyes. (11-) 12 uncinigerous abdominal segments. Pygidial cirri without eyespots. This speeies is eommon in marine localities but does not appear to occur in brackish water; reports of A. aeutifrons from brackish localities are probabty based on A. balliea. A. aeutifrons can easily be identified by the prominent dorsal cirri on the posterior neuropodia. The 2 groups ofbranehiae are elearly separated middorsally by a distance about equal to the width of2 branchiae. Paleae are evenly eurved and slender. Ampharete baltiea - Up to 18 nun long (Holthe 1986). Prostomium acutely pointed 'in dorsal view, with 2 small eyes. Usually 12 uneinigerous abdominal segments, but sometimes with 13. Pygidial cirri without eyespots. Very conunon in brackish water, this is probably the only speeies occurring in many fjords and is often misidentified as A. aeutifrons. Larger specimens may develop a short dorsal cirrus that extends about half the length of the neuropodiallobe, but on smaller wonns this is absent. The two groups of branchiae are very elose middorsallY,with branchiae of the innennos;.pair very elose, separated by aspace equal to or less than the width ofone branchia. Ampharete faleata - Up to 18 nun long (Holthe 1986). Prostomium broadly rounded, witJ:12 eyespots. 12 uncinigerous abdominal segments. Pygidial eirri without eyespots. Rare in the material I have seen, the species does not appear to be eonunon. It is easily identified by the very tiny pateae and the widely separated groups of branchiae. Anterior thoracic notochaetae are bent, with a wide base abruptly eurving to asiender tip. Ampharete finmarcltiea - Up to 50 rnm long (Holthe 1986). Prostomium broadly rounded, trilobed, with 2 eyespots. Branchial groups nearly touching middorsally; this also applies to A. balliea, but the two species are easily separated by differenees in the pygidial papillae. 12-13 uncinigerous abdominal segments. Malmgren (1867: 105) pointed out that his A. aretica Malmgren, 1866 was identical with Sars' species but also notes that it was actually published in 1866, and not 1864. Ampharete lindstroemi - Up to 12 mm long (Holthe 1986). Prostomium trilobed; with 2 6 eyespots (Holthe 1986). 12 uncinigerous aodominal segments. Not common, but specimens identified by rne as this speeies were present in material " [rom offVärö-Ringhals, Swedish west coast. Anobotlrrus Levinsen, 1884 Anobotlrrus gracilis ~lalmgren, 1866) Anohothrus gracilis is very cOrnn1on in the area treated and while specimens identified as this . species are ne~ly always correctly identified, smaller specimens appearr to be easily mistaken for species of Ampharete. There are marked differences in the buccal tentac1es of the two ; genera, smooth in Anobothrus and distinctly papillose in Ampharete, but these are not always obvious or easy to see. Although Anobothrus gracilis has relatively longer branchiae than Ampharete spp., it is most easily recognized by the prominent dorsal band ofeilia connecting the 5th from last thoraeie notopodia. If an "Ampharete" does not seem to fit the descriptiorf. \ 1998 WKBT Report 31 check the dorsum to make sure there is no such band ofcilia. Another difference that does not require dissection is the presence of transverse bands ofcilia on the anterior (frontal) surface ofthe innennost two pairs ofbranchiae (= pairs 1 and 2) in species ofAmpharete and no such ciliation in Anobothros (M. E. Petersen, unpubl.). Eliason (in Holthe 1986) has shown this ciliation in Ampharete baltica; I had discovered this independently before I was aware of Eliason's observations and have found it to be present on all species of Ampharete seen to date (M. E. Petersen, unpubl.). References Eliason, A. 1955. Neue oder wenig bekannte schwedische Ampharetiden (Polychaeta). - Meddelanden fran Göteborgs Musei Zoologiska Avdelning 126 (Göteborgs Kungliga Vetenskaps- och Vitterhets-Samhälles Handlingar, FäIjden 6, sero A, 6 (16): 1-17. NB: Cover says sero B but headings in paper say sero A. Hartmann-Schräder, G. 1996. Annelida, Borstenwürmer, Polychaeta. 2., neubearbeitete Auflage.- Die Tierewelt Deutschlands 58: 1-648. Holthe, T. 1975. A Simple Key to the Northem European Species of Terebellomorphe [sie] Polychaeta. _ Universitetsforlaget, Oslo, Norway. 32 pp. Holthe, T. 1986. Polychaeta Terebellomorpha. - Marine Invertebrates ofScandinavia 7: 1-192. Malmgren, A. J. 1866. Nordiska Hafs-Annulater. - Färhandlingar Öfvers. af Kungliga Vetenskaps-Akademiet . fär 1865, Nr 5: 355-410, pIs. 18-29. Malmgren, A. J. 1867. Annulata Polychaeta Spetsbergiae, Groenlandiae, Islandiae et Scandinaviae Hactenus . Cognita.. - Frenckelliana, Helsingfors, Sweden. 1-127, pIs. 1-14. 32 1998 WKBT Report ANNEX 8 A TAXINOMICALSCHEME Fig.66. Typical shell am.l habitus nf frolll krt: IIrr!lllbill lflllmlll (?\lolltagu 1, 11. IIcglccla ~luus, amI 1/. IIh'ae I I'L'lI11 1998 WKBT Report 33 111" I" 11 , 7n~nl. Bond..." d.l. Fig. 2. Hydrobia jenkinsi fra forskellige lokalitere~ i Danmark. (lverste r:ekke, fra venstre til hojre: Vondaa, Ringkobing Fjord og Kylleb:ekken. Nederste r:ekke, fra venstre til hojre:- Egaa, Kolstrup Mergelgrav S og Binderup Aa. • 34 1998 WKBT Report H.ncglecla ~\ \) i cv®~~~~~ • ~~~~ H. ulvae ----Imm---- Fig. Gfio renes of the three Ilydrobia spcciC"S drawn from livc or prcscrvcd ma.tcrial but loasily rc cognizahlc in spiic of the diffcrl'"nt dcgrces of contra.ction or condition of prcscrv:ltion. 1998 WKBT Report 35 Fig. G5. Shdl variation in the three H.Jdrobia specia. The two shclls on the lcft in ("ach ,nil":! an' particularly common main type, A B c D Fig.67. O:ntral tooth of the radula in: At H)'drobia ulrQt; B, H. t'C7/roja; Ct H. TlLglccta; D, Potarru> PJT.(W jmJ.irtJi (MUlU J963). 36 1998 WKBT Report ANNEX 9 l\10RPIIOLOGICAL FEATURES ACTAEONllJAl: I~CTAcON TOf 1998 WKBT Report 37 ANNEX 10 THE LUMBRINERIDAE OF THE KATTEGAT AND THE BALTIC 1. MaxiUae IV and V fused as a large plate (3-4 times as large as III) on each side. Acicula yellow. Transitional setae (between limbate capillaries and hooded hooks) present on anterior setigers. Postsetallobes on posterior setigers strongly elongated and thin ...... ',' Abyssoninoe seopa Maxillae IV and V not fused (although IV may be quite large and V difficult to detect! still, IV is only twice as large as III). Transitional setae absent. Postsetal lobes on posterior setigers somewhat elongate, but not particularly thin 2 \, 2. Acicula dark brown or black (in posterior setigers golden-coloured in juveniles), not accompanied by more dorsal group ofthin acicub. Hooded hooks from setiger 26-35 in adults (but from 1 in small juveniles!! I). M.axillae IV "riding hat" shaped and t\vice as large as III. rvraxillae III with 1 tooth each or 2 teeth on one side onlv ...... Seoletomajragilis Acicula yellow, accompanied by more dorsal group ofthin acicula. Hooded hooks from setiger 1-5, also in adults. Maxillae IV triangular and only slightly larger than In. Maxillae III with 2 teeth on both sides Seoletoma impatiens , , C" SC~')0) \ \l.~I' 1 , Abyssoninoe seopa (Fauchald, 1974) . _cK.QlAib~~~ (tv~"- ~~r";" ,,~.l \' .\ Syn:rLlImbrinerisseopaFauC~4- t :"l";'·' :'Y"- LLllmbrineris scopa sWPG-Fauchald, 1974 (erected by Winsnes 1981) Seoletomajragilis (Müller, 1776) Syn:: L11mbrinerisjragilis - Hartmann-Schräder 1971 Note: In addition to the normal, short bulbous hooded hooks, juveniles have 1-3 hooks with a longer, slender blade on anterior parapodia. Seoletoma impatiens (Claparede, 1868) .' Syn: Lumbrineris tetrcmra - Hartmann-Schräder 1971 [Seoletoma tetraura Schmarda, 1861 is a species described from South Africa] ~ ~ 38 /998 WKBT Report ANNEX 11 THE SPIONIDAE OF THE KATTEGAT AND THE BALTIe (These keys are modified from information provided by Dr. Andrew S. Y. Mackie, National Museum ofWales, Cardiffat a British workshop in 1990, incorporating information from various sources, especially Ramberg & Schram (1982) and personal observations. Please note that these keys are specifically tailored to the geographic area under consideration: species of the same genera may not key out correctly outside this area.) I. Neuropodia ofsetiger I include I or 2 large curved spines Spiophanes Neuropodia ofsetiger I lack spines 2 2. Setiger 5 with strongly modified notosetae 3 Setiger 5 without strongly modified setae 5 3. Gills start on setiger'2 Boccardiella Gills start on setiger 6-9 4 4. Setiger 5 in dorsal view only slightly longer than neighbouring setigers. Setiger 5 with two types of spines, arranged in two rows that form a U- or J-shape. Setiger 5 without companion setae. Neuropodial hooded hooks from setiger 8 Hooded hooks with secondary tooth bending elose to main fang ...... Pseudopolydora Setiger 5 in dorsal view c1early longer than neighbouring setigers. Setiger 5 with only one type of spine, arranged in a curved, almost horizontal row (a "flattened JII). Setiger 5 oRen with companion setae. Neuropodial hooded hooks from setiger 7. Hooded hooks with secondary tooth having larger angle to main fang (subtle) Polydora 5. Prostomium with frontal horns (T- or Y-shaped) Jvfalacoceros Prostomium without frontal horns 6 6. Prostomium distally pointed 7 Prostomium distally rounded or truncate 8 7. Gills only on anterior part of body (up to about 30 pairs). Gills c01'!!1?let~ly free from notopodial lamellae Aonides Gills present almost to end of body. Gills at least partly fused to notopodial lamellae ...... Scolelepis 8. Gills only on middle and posterior setigers (except for an additional pair on setiger 2 of males) Pygospio Gills beginning on setiger I, 2 or 3 9 9. Gills present on at least 30 setigers 10 Gills only present on anterior 14 setigers (or less) 13 10. Gills from setiger I 11 Gills from setiger 2 12 1998 WKBT Report 39 11. Hooded hooks in neuropodia only Spio Hooded hooks in neuropodia and notopodia ,'v/aren:elleria 12. Prostomium broadly rounded or truncate. Caruncle extending as dorsal sensory ridge to at least setiger 26 Laonice Prostomium narrO\v and anteriorly slightly bilobed. Caruncle very short and difficult to distinguish lvficrospio 13. Only 1 pair ofgills, on setiger 1. Setiger 2 with dorsal fold Strehlospio At least 3 pairs ofgills, from setiger 2 or 3 14 14 3 pairs ofgills, starting on setiger 3 Aurospio At least 4 pairs ofgills, starting on setiger 2 15 15 At least 6 pairs ofgills, all smooth Prionospio (lvfinuspio) 4 pairs ofgills, 1st and 4th pair pinnate Prionospio (Prionospio) Aonitles C1aparede, 1864 1. At least 20 pairs ofgills. Prostomium with occipital antenna. Hooded hooks with one small tooth above main fang. Pygidium with numerous anal cirri .... A. oxycephala 10-11 pairs of gills. Prostomium without occipital antenna. Hooded hooks with two small teeth above main fang. Pygidillm with only 4 anal cirri ... A. paucihranchiata AOllides oxycephala (Sars, 1862) Aonides paucihranchiata SOlIthern, 1914 Aurospio l\'laciolek, 1981 Aurospio ball)-7l1ensis (Laubier, 1968) - Syn: Prionospio ockelmanni Pleijel, 1985 - Boccartliella ß1al,e & Klldenov, 1978 Boccardiella ligerica (Ferroniere, 1898) Syn: Polydora redeki Horst, 1920 Polydora (Boccardia) redeki - Hartmann-Schröder 1971 1.(/(JIlice Ma1mgl'cn, 1867 l.aoJ1ice ha!lllsieJ1sis Söderström, 1920 Syn: Laonice cirrala - Hartmann-Schröder 1971 (pars) 40 1998 WKBT Report • iUalacoceros Quatrefages, 1843 1. Hooded hooks with 2 small teeth above main fang, 20-25 of them per neuropodium. Pygidium with 15-30 anal cirri /'vI. vlilgaris Hooded hooks with 1 small tooth above main fang, less than 15 ofthem per neuro podium. Pygidium with less than 10 anal cirri 2 2. Neuropodia with 4-5 1100ks each. Anterior margin of prostomillm with smalI, separate indentation M fliligillOSliS Neuropodia with 7-12 hooks each. Anterior margin of prostomium without separate indentation (but with angle created by "horns") Ivl. tetracerus !vlalacocerosjuligilloslIs (Claparede, 1868) A4alacoceros tetracerus (Schmarda, 1861) /'vfalacoceros l'ulgaris (Johnston, 1827) iUarenzelleria Mesnil, 1896 I. Gills absent from posterior half ofbody. Notosetae ofsetigers 1 and 2 incilide some very long, conspicuous capillaries Al viridis Gills absent from posterior third ofbody. Notosetae ofsetigers land 2 include som long, but inconspicuous capillaries A{ wireni Marenzelleria viridis (Verrill, 1873) Marenzelleria wirelli Augener, 1913 A--ficrospio lVI esnil, t 896 /'vficrospio mecznikowia/1us (Claparede, 1869) .' Syn: Spio meczllikoll'ia/1lfS - Hartmann-Schräder 1971 Polydora Bose, t 802 1. Gills beginning on setigel' 8-9 ...... 2 Gills beginning on setigel' 7 3 2. Spines of setigel' 5 unidentate with no adornment (at the most a weak, clear subdistal . lamella on the concave side) P. caeca Spines of setigel' 5 bidentate with adornment of fine hai'rs 01' lamella between teeth (if lamella is present, it is on the convex side) ...... 4 3. Setiger I with neuro- and notosetae. Pygidium with 4 lobes 4 Setigel' 1 with nellrosetae only. Pygidillm clIp-shaped with dorsal notch 5 1998 WKBT Report 41 4. Spines of setiger 5 with fine hairs between two distal teeth. Gills absent on at least 30 posterior setigers ...... P. quadrilobata Spines of setiger 5 with "lamella" of fused hairs between two distal, widely diverging . teeth. Gills absent on 25 posterior setigers at the most P. calilleryi 5. Occipital antenna present on prostomium. Black pigment absent (exception: very small juveniles may have remnants oflarval pigmentation) P. comllta Occipital antenna absent on prostomium. Black pigment always present, at least on peristomium and pygidium P. ciliata Polydora caeca (0rsted, 1843) Polydora caul/eryi tvfesnil, 1897 Polydora ciliata (Johnston, 1838) Polydora C0rJ111ta Bosc, 1802 Syn: Polydora (Polydora) ligni - Hartmann-Schröder 1971 Polydora qlladrilohata Jacobi, 1883 Primwsp;o (IH;nllsp;o) Foster, 1971 1. 5-6 pairs of gills. Neuropodial postsetal lamella of setiger 2 ventrally prolonged. Transverse dorsal fold on setiger 10 P. (M.) cirrifera 6-13 pairs of gills. Neuropodial postsetal lamella of setiger 2 not ventrally prolonged. Transverse dorsal fold on setiger 14 P. (?vI.) cf mullibranchiata Priollospio (Minuspio) cirrifera Wiren, 1883 Syn: Prionospio cirrifera - Hartmann-Schröder 1971 (pars) Prionospio (Minllspio) cf mliitibranchiala Berkeley, 1927 Syn: Priollospio cirrifera - Hartmann-Schröder 1971 (pars) Pr;ollosp;O (Pr;ollosp;o) Malmgren, 1867 I. First pair of pinnate gills ofsimilar length as last pair. Setiger 7 with high dorsal crest. Notopodial postsetal lamellae ofsetiger 1 do not form a fold P. fallax First pair ofpinnate gills much larger than last pair. Setiger 7 with low (inconspicuous) dorsal crest. Notopodial postsetal lammelae of setiger I form a fold behind the prostomium P. sleenslmpi Prionospio fal/ax Söderström, 1920 Syn: Priollo.\pio malmgreni - Hartmann-Schräder 1971 Prio/lo.\pio sll!l!l1slrupi Malmgren, 1867 42 1998 WKBT Report Pseuc!ol'0!yc!ora Czerniavsky, 1881 I. Strong brown or black pigment present on prostomium, posterior segmt:nts and as bands on palps. Prostomium anteriorly rounded or with weak indentation .... P. pl/lchra Dark pigment absent (exception: very small juveniles may have remnants of larval pigmentation). Prostomium anteriorly with a deep incision P. anlennala Pselldopolydora alllellnala (Claparede, 1870) Pseudopolydora plllc!7ra (Carazzi, 1895) Pygosl'io C1allarede, 1863 Pygospio elegalls Claparede, 1863 Sco!elepis B1ainville, 1828 I. Anterior gills distally free ofnotopodial postsetallamellae S. squamala Änterior gills completely fused to notopodial postsetallamella 2 2.. - Pröstomium with occipital antenna. Hooded hooks without small teeth above main fang. Prostomium anteriorly somewhat rounded or trilobed. Neuropodial postsetallamellae of posterior segments as two separate lobes ...... S.JolioSliS Prostomium without occipital antenna. Hooded hooks with two small teeth above main fang. Prostomium anteriorly strongly pointed. Neuropodial postsetal lamellae of posterior segments as single lobe S. tridentata Scolelepis (Scolelepi.~)JOliOSliS (Audouin & Milne-Edwards, 1834) Scolelepis (Scolelepü) sqllamala (Müller, 1789) Scolelepis (Parascolelepi.\) tridenlata (Southern, 1914) .. Spio Fabricius, 1785 1. Neuropodial hooded hooks from setiger 1I (10 in juveniles). Camncle weil developed. (Gills ofsetiger 1 more than half as long as those ofsetiger 2) S. cf.jilicornis Neuropodial hooded hooks from setiger 13-21 (I 1-12 in juveniles). Caruncle weakly developed, not distinguishable. (Gills ofsetiger 1 ofvariable length) 2 2. Small light brown patches ofpigmentation present on posterior part ofprostomium and middorsally on anterior setigers S. armata Brown pigmentation absent 3 1998 WKBT Report 43 ,,', 3. Neuropodial hooded hooks from setiger 13-14 in adllits. Most hooded hooks with 1 tooth above main fang, but a few may have 2. Prostomium anteriorly narrowly rounded. Gills ofsetiger 1 more than half as long as those of setiger 2 S. martinensis Neuropodial hooded hooks from setiger 16-21 in adults. All hooded hooks with only 1 . tooth above main fang. Prostomium anteriorly broadly rounded. Gills of setiger I no more than halfas lang as those ofsetiger 2 S. goniocephala Spio armata (Thulin, 1957) Spio cfjilicol'l1is (Müller, 1776) Spio goniocephala (Thulin, 1957) Spio martinensis Mesnil, 1896 . Spioplzanes Gnlbe, 1860 I. Prostomillm anteriorly broadly rounded, withollt lateral "horns". Occipital antenna present on prostomium S. kroeyeri Prostomium with lateral "horns". Occiptial antenna absent S. hombyx Spiophanes hombyx: (Claparede, 1870) Spiophanes kroeyeri Grube, 1860 Strehlospio \Vebster, 1879 Streblospio shruhsolii (Buchanan, 1890) 44 1998 WKBT Report