Advisory Committee on the Marine Environment ICES CM 1998/ACME:8 Ref.: E REPORT OF THE {)/O ICESIHELCOM BENTHOS TAXONOMIC WORKSHOP Roskilde, Denmark 4-7 November 1997 This re port 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
Transcript
Advisory Committee on the Marine Environment ICES CM 1998/ACME:8Ref.: E
REPORT OF THE
{)/O
ICESIHELCOM BENTHOS TAXONOMIC WORKSHOP
Roskilde, Denmark4-7 November 1997
This report is not to be quoted without prior consuItation with theGeneral Secretary. The document is areport of an expert groupunder the auspices of the International Council for the Exploration ofthe 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
bookeye
Thünen
TABLE OF CONTENTS
Section Page
-CIRRATULIDAE FROM THE KATIEGAT, 0RESUND, AND BALTIC SEA 22
A TAXONOMIC SCHEME 33
MORPHOLOGICAL FEATURES 37
THE LUMBRINERIDAE OF THE KATIEGAT AND THE BALTIC SEA 38
THE SPIONIDAE OF THE KATIEGAT AND THE BALTIC SEA 39
INTRODUCTION 1
2 TERMS OF REFERENCE 1
3 PROGRAMME FOR THE WORKSHOP 13.1 Phyllodocidae 13.2 Nephlyidae, Nereidae, and Polynoidae 13.3 Porifera 13.4 Sampling Techniques and Preservalion Practices 13.5 Pholoidae, Cirratulidae, and Ampharetidae 23.6 Gastropoda: Hydrobia. and Potamopyrgos 23.7 Gastropoda: Tectibranch Opistobranchia 23.8 Bivalvia: The Cardium-complex 23.9 Polychaeta: Spionidae and Flabelligeridae 23.10 Amphipoda 33.11 Feedback and General Discussion on Future Workshops 33.12 Closing ofthe Workshop 3
4 REFERENCES 3
LIST OF WORKSHOP PARTICIPANTS 4
AGENDA 6
PHYLLODOCIDAE 7
SELECTIVE LIST OF REFERENCES FOR POLYCHAETA FROM THE KATIEGATAND THE BALTIC SEA 9
IDENTIFICATION OF SPONGES 10
FINAL NOTES 21
ANNEX 1:
ANNEX 2:
• ANNEX 3:
ANNEX 4:
ANNEX 5:
ANNEX 6:
ANNEX 7:
ANNEX 8:
ANNEX 9:
ANNEX 10:
ANNEX 11:
1 INTRODUCTION
The ICESIHELCOM Benthos Taxonomie Workshop (WKBT) was the outcome of work initiated by theICESIHELCOM Steering Group on Quality Assurance of Biological Measurements (SGQAB). An intercalibrationexercise carried out by the Steering Group indicated that misidentification of species often occurs in the monitoring ofbenthie macrofauna. The aim of the Workshop was to minimize the possibility for species misidentification and to raisethe 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. DannyEibye-Jacobsen made a short presentation on practical issues of the Workshop. Taxonomie experts from the ZoologicalMuseum of Copenhagen acted as Iecturers.
Twenty people (the maximum number) participated in the Workshop. The names and addresses of Workshopparticipants 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 anddetermine 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 anoverview of the morphological features which are important for distinguishing the different species. A list of these arefound 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. Thecharacteristic morphological features were presented and Rainer (1991) was recommended for use as the key for thegenus 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 Porifera
ale Tendal gave a lecture on the phylum Porifera. An overview of the taxonomie problems with sponges was given asweil as a key to the speeies of sponges found in the Baltie Sea. The presentation by ale Tendal appears in Annex 5. Thelecture 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 zoologiealmuseums were on the programme. A range of sampling gear (Van Veen, HAPS, Smith-McIntyre) is apparently used inthe 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 adredge to supplement the traditional infauna sampling, although this gear is normally not quantitative. It was noted that aquantitative dredge has been developed in Holland (Bergman and Santbrink, 1994).
Different fixation teehniques were diseussed. Although fixation in formaldehyde is generaIly reeommended in theHELCOM 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 % ofthe sampies for quality assuranee purposes. This practice is recommended.
The eoIlection of referenee-speeimens was reeommended. It was further suggested that the Zoological Museum shouldmake 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 forhistological 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. Taxonomieproblems within these families were presented as weIl as keys for identifying species. These keys are attached as Annex6. 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 andecological aspeets of these genera was given. Copies of drawings of these speeies are attached as Annex 7. The leeturewas foIlowed by a practical session on preserved speeimens.
A lecture on Tectibranch Opistobranchia was given by Tom Schi~dte. The taxonomie problems in this group werepresented. A taxonomie scheme of the group is attached as Annex 8. The lecture was followed by a praetical session onprcserved specimens.
3.7 Gastropoda: Tectibranch Opistobranchia .,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 werepresented, 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 themorphologieal features which is important for the distinetion of the different speeies. A list of morphological featuresare 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 problemfor 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 futureworkshops.
Several partieipants expressed that the idea behind taxonomie workshops is good. Several suggestions to improve thequality 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 heldduring off years;
participation in taxonomie workshops should be optional for institutes collecting monitoring data in the OSPARarea.
3.12 Closing ofthe Workshop
The Chairman thanked the staff of the Zoological Museum for their engaged teaehing and for arranging a very fruitfulWorkshop. The partieipants were acknowledged for their active participation and positive attitude. The Workshop wasadjoumed.
4 REFERENCES
Bergman, M.J.N., and Santbrink, van, J.W. 1994. A new benthos dredge ('Triple-D') for quantitative sampling ofinfauna 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, includingthe deseription of N. pu Ich 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 MarineEnvironmental 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]. Boks 33FIN-00931 HelsinkiFinland
PoriferaGeneral discussion on sampling techniques, preservation practices andinteractions between monitoring units and zoological museumsLunchPolychaeta: Pholoidae, Cirratulidae, Ampharetidae
prostomium anterior to eyes not particularly dark with dark pigment
segment 1 ventrallynot 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 ofpresent 3 dark spots each parapodium ..
I Note that in both species the ventral cirri ofthe midbody are longer than the neuropodium. Itis important that a parapodium be removed to correctly judge this character, as all ventral cirriappear pointed in dorsal view.2 As on more posterior segments, where such middorsal spots are found in both species.3 .- This charader is only useful with live material, as the greenish-yellow pigment alwaysdisappears when the animals are transferred from formalin to alcohol... These two spots are sometimes connected across the dorsum as a thin transverse band, but thisis still easily distinguishable from the middorsal spot found in P. maculata.
sides of prostomium inconvex straight or slightly concave
dorsal view
ventral tentacular cirrislightly thicker than dorsal not thicker than dorsal
tentacular cirri tentacular cirri
dorsal cirri of midbody 2about 4 times as large as about twice as large as
neuropodium neuropodium
colour of living 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 differentfrom those originally described (both from Greenland). Material from Iceland and the FaroeIslands appears to show intermediate morphologies, so species clines may be involved. Untilfurther work is done, I would suggest that both names can routinely be used for specimens fromthe 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 inthe literature, dorsal cirri broader than long in E. flam and longer than broad in E. IOllga, is veryunreliable aS these differences are only apparent on very large specimens; otherwise the dorsalcirri are about as long as wide in both species. .3 Paradoxically, specimens ofE. tonga ?ften turn Y§Y dark following preservation.
conspicuously broader thanlong, with a drawn-out tip
about 4 times as large asneuropodium
asymmetrical and sharplypointed
often with dark pigment
with conspicuous transversesegmental bands
slightly longer than broad, tipnot drawn out
about twice as large asneuropodium
almost symmetrical andbluntly pointed
not particularly dark
without or with only weaklydeveloped transverse
segmental bands
8
1 These characters only reliable on animals longer than 10 mm.
/998 WKBT Report
- -----------------
ANNEX 4
SELECTIVE LIST OF REFERENCES FOR POLYCIIAETA FROM TIIE KATTEGAT AND TIIEBALTIC-
Holthe, T. 1991. Identification of Annelida Polychaeta from northern European and adjacentArctic waters. - GWlIleria 66: 1-30. [An excellent paper with a family by family listingof relevant literature. ]
Nephtyidae:Rainer, S. F. 1991. The genus Neph(l's (Polychaeta: Phyllodocida) of northern 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. [InNorwegian, but with excellent figures and based on a large amount of material. Possiblyavailable from: Zoologisk Institutt, Universitetet i Osol, Postboks 1050, Blindern, Oslo3, 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 of Oslo. [InNorwegian, 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 northwesternAtlantic surveys with descriptions of a new genus and two new specie?- j)roceedillgs0/ the Biological Socie(l' 0/ WashillgtOIl 105: 185-218. [With good information on thelatest swarm oflumbrinerid genera and the best description ofScole/oJ11ajragilis.]
Spionidae:Ramberg, 1. P. & T. A. Schram. 1982. A systematic review of the Oslofjord species of Polydora
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 reexamination ofP. ockelmalllli Pleijel, 1985 (Polychaeta, Spionidae). - Oplwlia 35: 209217.
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 of spongcs
a) Taxonomical problems with sponges
Sponges (Porifera) is one of the few large phylae (about 6000 living species) where many higher category taxa have not yet beenfinally outlined. Over the last 20 years orders have been erected,split and laid down again,and the same goes for families, not tospeak of genera·and species. There are several reasons for this:
1) Many original descriptions of sponge species are short and insufficient with todays eyes, and sometimes the original materialhas disappeared or is insufficiently labelIed, so there is doubtabout the identity of the single specimens in relation to the description ("The type specimen problem") .
2) Historically, terminology has been different in different languages ahd as used by various "schools", making it difficult tocompare descriptions especially of very similar species or specimens .where small differences are important (llThe terminology problem) .
3) Ideas on species variation has changed several times over time,in some periods leading to unjustified splitting into too many speeies, in others to unrealisting synonYmisation ("The splitterlumper problem)".
4) Character deficiency, viz. low number of and high variation intaxonomically usable characters, is a wellknown problem in manysponge taxa. It has lead to a search for new charaeter complexeswithin fields as reproduction, genetics and bioehemistry, so farwith'some success in single cases, but difficult to carry throughfor a larger number of species ("The eharaeter problem").
5) While most sponge species show only little variatiorr within the 4tsame population or between different populations, other species mayvary literally from one side of an individual to another, certainlybetween 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 problem)".
b) The identification proeedure
Three categories of information should be considered:
1) The locality, substrate, depth, and exposure to current andlight.
2) The general outer morphological characteristics, at best infresh 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 moreexact terms, but nevertheless the appearence of a species can beexperienced as very characteristic, once you are familiar with thelocal fauna, because only one or a few species of the genus arerepresented.
3) Skeletal characteristics such as presence and geometric patternof spongin and spicule fibers, and the number, dimensions and arrangement of spicule types.
Generally spoken, the organization of the skeleton, theposition of the spicules in the skeleton, and the kinds of spicules 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 spicules present and their dimensions as a good starting point foridentification. Within some groups, for example Haliclona, wherevery few characters are at hand, it is, however, always necessaryto 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 spongetaken from different parts of the sponge body and squeezed in water(ar alcohol) between slide and cover glass. Always start with thelow power objective of the compound microscope to get an overview.This kind of preparation is fast to make, and is not permanent. Therisk-is that skeletal arrangement is dif-ficult to make out becauseof distorting, and that small or rare spicule types are notdiscovered.
A better idea of the skeleton can one get from handmade sections, which should"always be made from specimens which have beenhardened instrong alcohol for at least some days. Use a razor blade and cut as thin as possible. One section should be perpendicularto the surface. In case of cylindrical growthform both tranverseand longitudinal sections should be taken. A section made parallelto the surface, including dermal membrane mayaIso be useful. Suchsections can be studied in water or alcohol at low power, and thentrown away,~of they can be taken taken through adehydration andmounting procedure to give permanent slide preparations.
Clean spicule preparations are necessary when thefi~e structure of the spicules is to be studied and when numerous measurements are taken. A small piece of sponge tissue (or several smallpieces originating from different parts of the sponge (in all, thesize of a pea) should be boiled in a few"ml concentrated nitricacid). In the case of calcareous sponges chlorine or a similar basic bleaching agent can be used. After boiling (in the case of nitric acid, when the acid besomes clear again or when no more brown fumes are emanated), the remaining acid is pipetted off, and thespicules are cleaned in destilled water (be careful not to loosetoo many spicules during the procedure). Then the spicules can berinsed.in 96% alcohol. With a pipette a portion is spread on a slide, the alcohol burned off (or let to dry), and the dry slide withspicules"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 booksreferred to in the list of "local" literature), or try to find amatching spicule picture. When an idea of the species name has beenreached, always read the description very carefully in order tocontrol. The key does only give a limited degree of certainty. Itis always a possibility that there are more species in the areathan keyed out, either because they were not found at the time thekey ~as constructed, or the'key was been made for a different geographical area. Be aware that because of numerous synonymizationproblems the same species may be mentioned under different genericor 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, andannotated bibliography and key (working title) .
Hansson, H.G., 1994: Sydskandinaviska marina flercelliga evertebrater. - 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 DN1997-1. Directorate for Nature Management, TrondheJ:m, -Norway.409 pp.
Veethaak, A.D., R.J.A. Cronie & R.W.M. van Soest, 1982: Ecologyand distribution of two sympatric, closely related spongespecies, Halichondria,panicea (PalIas, 1766) and H. bowerbanki Burton, 1930 (Porifera, Demospongiae), with remarks ontheir speciacion. - Bijdragen tot de Dierkunde 52: 82-102.
Weerdt, .W. H. de, 1986: A systematic revision of the north-easternAtlantic shallow-water Haplosclerida (Porifera, Demospon-'giae) , part II: Chalinidae. - Beaufortia 36: 81 -165.
12/998 lVKBT Report
Baltic sponges
List of species recorded from the Danish belts to The Bottom Sea
Species in paranthesis have been recorded only once, and that inolder time. Species marked with an asterisk have not been found inthe area but they are expected to occur there.
1a Without spicules Ha~isarca dujardini1b With spicules 2
2a Spicules calcitic; at least some of them triradiates 32b Spicules siliceous; are rod-shaped 7
3a Form as a little bush of thinwalled tubes; at the open end thetubes may have a corona of spicules, but that is always shorterthan the tube diameter 4
3b (Form as a little thick-walled urn with an oscular corona ofspicules longer than diameter of the osculum ~. Sycon ci~iatum)
4a The spicules comprises a rod-shaped type S4b (No rod-shaped type present 6)
Sa Mostly triradiates; if there are tetraradiates the fourth rayis rudimentary Leucoso~enia complicata
Sb tetraradiates common and their fourth ray is weIl developed ..,~.................................... '.' Leucosolenia botryoides ,.,
If you believe that a specimen resembles a species but "something is wrong" there are severalways 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 becertain.
PllO/O€! ?ha/liea
B) You be1ieve it is not Photo€! haltiea, you don't know which species it is, but ofthe species YOllknow, 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
e-j C) Too many characters are missing and you don't feel very certain about Ph%e ha/liea at all.~,/'
Ph%e sp.
0) Too many characters are missing and you have no idea whether it is P/l0/OI! ha/liea. Oon'tglless 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
I thank all of you for your active participation - it has been a pleasure to arrange thisworkshop.
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
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, notaltemating 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 segmentsusually conspicuous, forming complete or partial einetures, often. prntruding at rightangles to body Cltaetozone (p. 3)
RemarksThere are presently 10 recognized genera of cirratulids (Blake 1996), but 4 of these: Cirriformia, Monticellina, Protocirrineris, and Timarete are not known to occur in the area underconsideration. Caulleriella? serrata Eliason, 1962 will not key out with the above key, but asit is still only known from a few specimens this is not likely to cause any problems. -
Kcy to spccies of Apltelocltaeta 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.
RemarksAphelochaeta was erected by Blake (1991) for bitentaculate cirratulids with exclusivelycapillary 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 arepresent in Danish waters, and perhaps more in the Kattegat-Baltic area. One of these appearsto 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 of only capillary chaetae suggests that Aphelochaeta is the eorrect genus, but theidentity ofMcIntosh's material (by Southem 1914 referred to Cirratllllls) and also the identityof Cirratu11iS filiformis Kefcrstein, 1862 (= Aphelochaeta) is presently not clear. A. vz\,'iparawas transferred to Aphelochaeta from Tharyx by Hartmann-Schröder (1996). In Danish watersI'have only seen the species in material from the Limfjord.
Apheloclzaeta sp. from eryptofaunal habitats such as Laminaria holdfasts is anunnamed hennaphroditic species with brood protection in the burrow and will be described byme (M. E. Petersen, in prep.).
T/iaryx mliitibranchiis [sie] (Grube, 1863) was reported from Helgoland by Hartmann-Schröder (1971) and Gillandt (1979); Hartmann-Schröder (1996) later referred it toAphelochaeta mllltibranchis. The speeies is not the one described by Grube as HeterocirnlSmliltibranchis Grube, 1863. Grube's type matenal has been examined and will be describedseparately. The speeies has not been seen in northem European waters.
Key to species of Calil/eriel/a Chamberlin, 19191. Cryptofaunal in Laminaria holdfasts; pygiditim entire; live animals bright yellow ..
..................................................: C parva Gillandt, 1979Infaunal in soft sediments; pygidium with two lobes; live animals olive green with goldensheen C bioculata (Keferstein, 1862)
RemarksCalilleriella pan'a was originally described as C bioclliata pan'a Gillandt, 1979, and raisedto 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 NEAmerican east coast (M. E. Petersen in Hartmann-Schräder 1996). The species is hennaphroditic and has brood protection in the burrow (petersen 1994); it may occur together withAphelochaeta 'sp. (see above). The above are the only two species of Cau/lerie/la presentlyknown from Danish waters.
Caulleriella caputesoeis (Saint-Joseph, 1888) is not a Calilleriella; the report byGillandt (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..
Chaetozone setosa has been widely interpreted and the name. has been used for a largeassortment of species from all parts of the world. The true C. setosa is a northern species andprobably not present in the area under consideration. At least two species of Chaetozone areeasily recognizable, but more are undoubtedly present. A lectotype of C. setosa has beenselected from Malmgren's material from Spitsbergen, and a redescription of the species is inpreparation (11. E. Petersen & S. F. Garcia-Martin, in prep.). For the present, the followingdesignations are suggested.
Key to some Kattegat-Baltic speciesof CltaetozoneI. 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
RemarksIn both species the prostomium is pointed, but it is relatively longer und a bit more slender inClzaetozolle sp. F group, where the first branchiae (branchiae 1) arise either level with thebase of the,tentaculophores or slightly anterior to these; the "tops" of the tentaculophores areofteh level with chaetiger 2. Aciculars of posterior segments are relatively short, and themeinbrunous "webbing" between posterior chaetae is not pronounced.
.: In Clzaetozol,e cf. setosa, the fIrSt -branchiae are nearly in line with thetentaculophores, 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 leftsides crossing dorsally where aciculars are best developed. Webbing between aciculars ofposterior segments is often prominent.
Key to species of Cirratulus Lamarck, 18181. ,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
RemarksCirratulus caudatus is a fairly large infaunal species; it is not uncommon, but is rarely takenas complete specimens. Although the original description implies that there were two groupsof tentaeles, none of the syntypes have more than two tentacles, which is also the case withmore 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 Laminariaholdfasts. It reproduces asexually by fragmentation (petersen 1991); regenerating specimensare'~ommon in late summer and earlyfall, but can be found )rear round. Sexual reproductionocc'urs in late sUf!lI11er; male and fe"?-ale epitokes with numerous long capillaries swim up off
24 1998 lVKBT Report
• t" f, ~,' .t' \. ..
I'I
the bottom and spaw-n in the free water mass, after which the epitokes die. The species is notvery common in pure mud but may be present if there are dead shells. The epitokes weredescribed 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 aIarge, black-eyed, infaunal species common in British waters but never observed in theKattegat 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. Itsbiology has been studied in detail by Olive (1970, 1971, 1973); eggs are laid in an irregularjelly mass and development is direct (Stephenson 1950a). A color photo of a female fromDevon, 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 of Pomatocerostubes off Tjämö, western Sweden. The species resembles C. incertus but differs from it inbeing yellow with red eyes. To date no yellow C. incertus have ever been seen, even thoughnumerous live specimens have been observed. Cirratlilus sp. is small and only known from afew specimens; its status is unclear and it may be undescribed (M.E. Petersen, unpubl.). Anyadditional material would be most welcome.
Genus Dodecaceria Orsted, 1843Syn.: Zeppelina Vaillant, 1890Dodecaceria conc/rarum 0rsted, 1843.?Syn.: Dodecaceriafimbriata (Verrill, 1879)Syn.: Dodecaceria calilleryi Dehorne, 1933Not Dodecaceria conchamm sensu Gibson (see below)
RemarksOnly Dodecaceria concharllm is likely to be encountered in the Kattegat-Baltic area. Thespecies is common in dead shells that have been bored by species ofthe boring sponge Clionaor in dead bamacle tests or bivalve shells in Laminaria holdfasts. It also occurs in encrustingcoralline algae ("Lithothamnion " spp., now referred to other genera), but this is not a commonhabitat in Danish waters. The species reproduces asexually by schizogamy or fragmentation inlate summer and fall, and single middle segments or anterior or posterior ends may be foundregenerating the missing portions. The original segmentes) are usually recognizable by theirdarker 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 muchuseful information on this and other species of Dodecaceria, but unfortunately Gibson choseto 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 of0rsted's type localities (the 0resund off Hellebrek and the Kattegat off Jutland betweenFrederikshavn and Skagen) or in any Danish waters.
D. jimbriata (Verrill, 1879) is an eastern North American species described in thegenus Heterocirrus; it is either a junior synonym of the true D. concharum or a distinct butvery similar North American species (see George & Petersen 1991 for a review of the mainsynonyms); its name has no place in th~ taxonomy ofEuropean Dodecaceria.
D. cQulleryi Dehorne, 1933 was erected for an asexually reproducing species ofDodecaceria 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 byschizogemisis (multiple fission).
Zeppelilla Vaillant, 1890, is a genus earlier referred to the Ctenodrilidae and believedto reproduce only asexually; George & Petersen (1991) pointed out that most species ofZeppelina were referable to asexual regenerates or juveniles ofDodecaceria spp..
D. concltarum has a life cycle similar to that of Cirratu/us incertus. with asexualreproduction by fragmentation and sexual epitokes with gametes shed into the free watermasse The life span of the species was by Gibson & Clark (1976) estimated to be about 3years 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 notpresent in Danish waters, where the salinity appears to be too low for it (Gibson 1996).
RemarlcsTlzaryx killariellsis is the most common - perhaps the only - true Tharyx in Danish waters. Itis common in the \Vadden Sea and is the T. marioni of Farke (1979). The species-is weIlillustrated.in Southern (1914). It was transferred to Tharyx by Blake (1991); in HartmannSchröder (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 Apheloclzaeta (see comments under this).
References
Blake, J. A. 1991. Revision of some genera and species of Cirratulidae (polychaeta) from the western NorthAtlantic. - 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 fromthe eastern North Pacific. - In J. A. Blake, B. Hilbig, & P. H. Scott (eds.): Taxonomie Atlas of the BenthicFauna 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 ofChaeto::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. - BulletinBiologique 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: 207293.
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 inBremerhaven 18: 69-99.
26 1998 WKnT Report
George, J. D. & M. E. Petersen. 1991. Validity ofthe genus Zeppelina VaiIlant (polychaeta: Ctenodrilidae). -lnM. 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 thefamily 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 ofthe 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 of reproduction. - 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 Associationofthe U. K. 76: 625-635.
Gibson, P. H. & R. B. Clark. 1976. Reproduction in Dodecaceria cau/leryi (polychaeta: Cirratulidae). - Journalofthe 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 of some cirratulid genera.- Bulletin of Marine 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 hermaphroditismamong the Cirratulidae. - In J.-C. Dauvin, L. Laubier & D. J. Reish (eds.): Actes de la 4eme ConferenceInternationale 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 IrishAcademy 31: 1-160, pIs 1-15.
Stephenson, W.,.l950a. The development of Cirratulus cirratus (O.F. Müller). Report of the Dove MarineLaboratory for 1943, sero III, No. 11: 7-20.
Stephenson, W. 1950b. An epitokous cirratulid occurring in the Cullercoats tanks. Report of the Dove MarineLaboratory for 1948, sero III, No. 11: 21-30. - .-
1998 WKBT Report 27
28
Pholoidae from the Kattegat, 0resund, and BaUicKeys 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
I.. \Vith eyes. Scales with or without pigment. FaciaI tubereie variable 2Without 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 3Scales without pigment. Elytral papilläe moniliform. Facial tubereie prominent, often aslarge as median antenna. Prostomium with dark pigment between eyes. Neuropodia withstylodes on distal part :::: : baltica
3. Prostomium with conspicuous black eyes, usually without other pigment. Elytra evenlypigmented brownish or blackish"often pale in deep water, covering dorsum completely,usually lying flat against dorsum. Elytral papillae marginal, slender, short and slightlycapitate anteriorly, elongated and tapering posteriorly. Tentacular cirri smooth orirregular, 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, leavingnarrow middorsal strip bare, often ruffled and not lying flat against dorsum. Elytralpapillae evenly tapering, never capitate, marginal laterally, on posterior margin of elytrabecoming submarginal towards middorsum. Dorsal tentaculat cirri usually with 5 distinct .papillae on inner side ;nornata
1998 WKBT Report
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." i'~ ..... '" Q • 't.-."
RemarksSmall specimens of Pholoe can be very difficult to identify and good microscopes areabsolutely necessary. \Vhile most species can be identified on characters ofthe anterior end, itis much easier to arrive at a certain identification if the specimen has at least some of themiddle and posterior segTrients and elytra. As pignient or lack of such is usually an easy wayto limit the choices, staining with Rose Bengal should be avoided ifat aII possible.
Pltoloe assimilis usually lack interocular pigment, but this is present in someindividuals 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 (mostspecimens are 5 mm or less), the species has often been considered a juvenile of other 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 tooccur 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 thename of a 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 groupis settled. This way, the worst thing that can happen is that all P. balliea should be referred toP. longa. If P. 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 thescales 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, 18ad, pls.A1-2, B1-2 (SEM), all as P. inornata).
P. inornata always has at least some, usually 5, distinet papillae on the dorsal tentaeular cirri, a few on the ventral tc and one or more on the ventral cirri, hut the numher anddevelopment of papillae may vary. It sometimes has eolorless elytra and a smudge ofpigmenthetween the eyes, and thus superficially resembles P. baltiea, hut can be distinguished by theahsence of stylodes (papiIlae) on the distal part of the neuropodia, the presenee of papillae onte 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 keyfor the polychaetes of the Baltic Sea. - Mitteilungen aus dem Zoologischen Museum in Berlin 61 (2): 171272.
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.
1998 lVKBT Report29
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Ampharetidae from the Kattegat, 0resund, and BaIticKeys 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
1. Pygidial cirri and papillae long, slender, and pointed 2Pygidial 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 rJAbdominal neuropodia without dorsal cirrus (smaller specirnens) or cirrus maxirnally half -l.-\\.a.- c\.'.J.as leng as neuro-podium (larger specimens) baltica -5 .' ~r:J-{
'-, c .
3. Paleae long, conspicuous. Pygidial cirri each with small eye at base 4Paleae minute, about 6-8 per side,' short and stout with pointed tips, may be hidden by
.bases ofbranchiae : faleata'II
4: Paleae 10-12 per side re Sars (1866),12-16 per side re Holthe (1986), relatively thick andbroad, markedly and abruptly curved at tip. Groups of branchiae nearly touching mid-dorsally jinmarchieaPaleae about15 per side (about 7 per side re Holthe 1986), slender, fmely tapering, notabruptly curved at tip. Groups ofbranchiae widely separated lindstroemi
RemarksDuring the late 1970s, problems encountered while attempting to identify material from the0resund prompted me to make a pictorial key to the species ofAmpharete. This was based onthe key to· Ampharete in Holthe (1975) und supplemented with infonnation from originaldescriptions and my o\~ observat~o.ns on material from Scandinavian waters. Later the moredetailed MIOS volume on Terebellomorpha (Holthe 1986) was published. and more recentlyHartmann-Schräder (1996) has provided a more comprehensive key to Ampharete in her
1998 WKBT Report
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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 notyet had time to try out the keys. The key presented here is based on my earlier key and ownobservations plus the keys and information in the above sources, which should be consultedfor additional infonnation and figures.
Amplrarete aeutifrons - Up to 45 mrn lang (Holthe 1986); up to 80 mm, 13 mrn wide inArctie 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 brackishwater; 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 of branehiae are elearly separated middorsally by a distance about equal to thewidth of2 branchiae. Paleae are evenly eurved and slender.
Ampharete baltiea - Up to 18 nun long (Holthe 1986). Prostomium acutely pointed 'indorsal view, with 2 small eyes. Usually 12 uneinigerous abdominal segments, but sometimeswith 13. Pygidial cirri without eyespots.
Very conunon in brackish water, this is probably the only speeies occurring in manyfjords and is often misidentified as A. aeutifrons. Larger specimens may develop a shortdorsal cirrus that extends about half the length of the neuropodiallobe, but on smaller wonnsthis is absent. The two groups of branchiae are very elose middorsallY,with branchiae of theinnennos;.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 easilyidentified by the very tiny pateae and the widely separated groups of branchiae. Anteriorthoracic notochaetae are bent, with a wide base abruptly eurving to asiender tip.
Ampharete finmarcltiea - Up to 50 rnm long (Holthe 1986). Prostomium broadlyrounded, trilobed, with 2 eyespots. Branchial groups nearly touching middorsally; this alsoapplies to A. balliea, but the two species are easily separated by differenees in the pygidialpapillae. 12-13 uncinigerous abdominal segments.
Malmgren (1867: 105) pointed out that his A. aretica Malmgren, 1866 was identical withSars' 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 26 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.
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 mistakenfor 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 alwaysobvious or easy to see. Although Anobothrus gracilis has relatively longer branchiae thanAmpharete spp., it is most easily recognized by the prominent dorsal band of eilia connectingthe 5th from last thoraeie notopodia. If an "Ampharete" does not seem to fit the descriptiorf.
\
1998 WKBT Report 31
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check the dorsum to make sure there is no such band of cilia. Another difference that does notrequire dissection is the presence of transverse bands of cilia on the anterior (frontal) surfaceofthe innennost two pairs ofbranchiae (= pairs 1 and 2) in species ofAmpharete and no suchciliation in Anobothros (M. E. Petersen, unpubl.). Eliason (in Holthe 1986) has shown thisciliation in Ampharete baltica; I had discovered this independently before I was aware ofEliason's observations and have found it to be present on all species of Ampharete seen todate (M. E. Petersen, unpubl.).
References
Eliason, A. 1955. Neue oder wenig bekannte schwedische Ampharetiden (Polychaeta). - Meddelanden franGöteborgs Musei Zoologiska Avdelning 126 (Göteborgs Kungliga Vetenskaps- och Vitterhets-SamhällesHandlingar, 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 TiereweltDeutschlands 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.
Fig. 2. Hydrobia jenkinsi fra forskellige lokalitere~ i Danmark. (lverster: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 rccognizahlc 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
c
B
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).
MaxiUae IV and V fused as a large plate (3-4 times as large as III) on each side. Aciculayellow. Transitional setae (between limbate capillaries and hooded hooks) present onanterior setigers. Postsetallobes on posterior setigers strongly elongated and thin ..
. . . . . . . . . . . . . . . . . . ',' Abyssoninoe seopaMaxillae 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 onposterior setigers somewhat elongate, but not particularly thin 2
Acicula dark brown or black (in posterior setigers golden-coloured in juveniles), notaccompanied by more dorsal group ofthin acicub. Hooded hooks from setiger 26-35 inadults (but from 1 in small juveniles!! I). M.axillae IV "riding hat" shaped and t\vice aslarge as III. rvraxillae III with 1 tooth each or 2 teeth on one side onlv .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. SeoletomajragilisAcicula yellow, accompanied by more dorsal group ofthin acicula. Hooded hooks fromsetiger 1-5, also in adults. Maxillae IV triangular and only slightly larger than In.Maxillae III with 2 teeth on both sides Seoletoma impatiens
Seoletomajragilis (Müller, 1776)Syn:: L11mbrinerisjragilis - Hartmann-Schräder 1971Note: 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, NationalMuseum ofWales, Cardiffat a British workshop in 1990, incorporating information from varioussources, especially Ramberg & Schram (1982) and personal observations. Please note that thesekeys are specifically tailored to the geographic area under consideration: species of the samegenera may not key out correctly outside this area.)
I. Neuropodia of setiger I include I or 2 large curved spines SpiophanesNeuropodia of setiger I lack spines 2
2. Setiger 5 with strongly modified notosetae 3Setiger 5 without strongly modified setae 5
3. Gills start on setiger'2 BoccardiellaGills start on setiger 6-9 4
4. Setiger 5 in dorsal view only slightly longer than neighbouring setigers. Setiger 5 withtwo types of spines, arranged in two rows that form a U- or J-shape. Setiger 5 withoutcompanion setae. Neuropodial hooded hooks from setiger 8 Hooded hooks withsecondary tooth bending elose to main fang ..... . . . . . . . . . . . . .. PseudopolydoraSetiger 5 in dorsal view c1early longer than neighbouring setigers. Setiger 5 with only onetype of spine, arranged in a curved, almost horizontal row (a "flattened JII). Setiger 5oRen with companion setae. Neuropodial hooded hooks from setiger 7. Hooded hookswith secondary tooth having larger angle to main fang (subtle) Polydora
5. Prostomium with frontal horns (T- or Y-shaped) JvfalacocerosProstomium without frontal horns 6
6. Prostomium distally pointed 7Prostomium distally rounded or truncate 8
7. Gills only on anterior part of body (up to about 30 pairs). Gills c01'!!1?let~ly free fromnotopodial lamellae AonidesGills present almost to end of body. Gills at least partly fused to notopodial lamellae
8. Gills only on middle and posterior setigers (except for an additional pair on setiger 2 ofmales) PygospioGills beginning on setiger I, 2 or 3 9
9. Gills present on at least 30 setigers 10Gills only present on anterior 14 setigers (or less) 13
10. Gills from setiger IGills from setiger 2
1112
1998 WKBT Report 39
40
11. Hooded hooks in neuropodia only SpioHooded hooks in neuropodia and notopodia ,'v/aren:elleria
12. Prostomium broadly rounded or truncate. Caruncle extending as dorsal sensory ridge toat least setiger 26 LaoniceProstomium narrO\v and anteriorly slightly bilobed. Caruncle very short and difficult todistinguish lvficrospio
13. Only 1 pair ofgills, on setiger 1. Setiger 2 with dorsal fold StrehlospioAt least 3 pairs of gills, from setiger 2 or 3 14
14 3 pairs ofgills, starting on setiger 3 AurospioAt least 4 pairs ofgills, starting on setiger 2 15
15 At least 6 pairs ofgills, all smooth Prionospio (lvfinuspio)4 pairs of gills, 1st and 4th pair pinnate Prionospio (Prionospio)
Aonitles C1aparede, 1864
1. At least 20 pairs of gills. Prostomium with occipital antenna. Hooded hooks with onesmall tooth above main fang. Pygidium with numerous anal cirri .... A. oxycephala10-11 pairs of gills. Prostomium without occipital antenna. Hooded hooks with twosmall teeth above main fang. Pygidillm with only 4 anal cirri ... A. paucihranchiata
1. Hooded hooks with 2 small teeth above main fang, 20-25 of them per neuropodium.Pygidium with 15-30 anal cirri /'vI. vlilgarisHooded hooks with 1 small tooth above main fang, less than 15 of them per neuropodium.Pygidium with less than 10 anal cirri 2
2. Neuropodia with 4-5 1100ks each. Anterior margin of prostomillm with smalI, separateindentation M fliligillOSliSNeuropodia with 7-12 hooks each. Anterior margin of prostomium without separateindentation (but with angle created by "horns") Ivl. tetracerus
I. Gills absent from posterior half ofbody. Notosetae of setigers 1 and 2 incilide some verylong, conspicuous capillaries Al viridisGills absent from posterior third ofbody. Notosetae of setigers land 2 include som long,but inconspicuous capillaries A{ wireni
1. Gills beginning on set igel' 8-9Gills beginning on setigel' 7
........................................ 23
2. Spines of setigel' 5 unidentate with no adornment (at the most a weak, clear subdistal .lamella on the concave side) P. caecaSpines of setigel' 5 bidentate with adornment of fine hai'rs 01' lamella between teeth (iflamella is present, it is on the convex side) . . . . . . . . . . . .. 4
3. Setiger I with neuro- and notosetae. Pygidium with 4 lobes 4Setigel' 1 with nellrosetae only. Pygidillm clIp-shaped with dorsal notch 5
1998 WKBT Report 41
42
4. Spines of setiger 5 with fine hairs between two distal teeth. Gills absent on at least 30posterior setigers . . . . . . . . . . . . . .. P. quadrilobataSpines 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 smalljuveniles may have remnants oflarval pigmentation) P. comlltaOccipital antenna absent on prostomium. Black pigment always present, at least onperistomium and pygidium P. ciliata
1. 5-6 pairs of gills. Neuropodial postsetal lamella of setiger 2 ventrally prolonged.Transverse dorsal fold on setiger 10 P. (M.) cirrifera6-13 pairs of gills. Neuropodial postsetal lamella of setiger 2 not ventrally prolonged.Transverse dorsal fold on setiger 14 P. (?vI.) cf mullibranchiata
I. First pair of pinnate gills of similar length as last pair. Setiger 7 with high dorsal crest.Notopodial postsetal lamellae of setiger 1 do not form a fold P. fallaxFirst 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 theprostomium P. sleenslmpi
I. Strong brown or black pigment present on prostomium, posterior segmt:nts and as bandson palps. Prostomium anteriorly rounded or with weak indentation .... P. pl/lchraDark pigment absent (exception: very small juveniles may have remnants of larvalpigmentation). Prostomium anteriorly with a deep incision P. anlennala
1. Neuropodial hooded hooks from setiger 1I (10 in juveniles). Camncle weil developed.(Gills of setiger 1 more than half as long as those of setiger 2) S. cf.jilicornisNeuropodial hooded hooks from setiger 13-21 (I 1-12 in juveniles). Caruncle weaklydeveloped, not distinguishable. (Gills of setiger 1 ofvariable length) 2
2. Small light brown patches of pigmentation present on posterior part of prostomium andmiddorsally on anterior setigers S. armataBrown pigmentation absent 3
1998 WKBT Report 43
..
,,,'
3. Neuropodial hooded hooks from setiger 13-14 in adllits. Most hooded hooks with 1tooth above main fang, but a few may have 2. Prostomium anteriorly narrowly rounded.Gills of setiger 1 more than half as long as those of setiger 2 S. martinensisNeuropodial 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 nomore than half as lang as those of setiger 2 S. goniocephala
I. Prostomillm anteriorly broadly rounded, withollt lateral "horns". Occipital antennapresent on prostomium S. kroeyeriProstomium with lateral "horns". Occiptial antenna absent S. hombyx
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