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WYPHA,T1

Tom LXV

OTTITCR)

MOCHBA • 1986

300J10r14 11ECK1411 WYPHAJI 1986, TOM LXV, imin. 2

11,11K 595.142.3:592/599:001.4

0 ,ITI1A1'HOCTIPIECKHX TIPH3HAKAX II0,73,CEMEHCTB CEMEOCTBA TUBIFICIDAE (OLIGOCHAETA)

H. Il. 011110TEHOBA

nau6o.riee BaNO1b1C npuanamr, rio xoTophim cocTannena oupe2..e-,rinTemman Ta6.nrma no,Rcemeilm Tubificiclae: nammue fl.iu oTcyreTnire Kpyri-rmix ue.nomornuron B 110.10CTI1 xapaNrcp npm:permennu, (1)opma 11 glIC:10

1.1BOCTaTIVICCKIIX /1:e.'ic3 na Kamc.itom my;KcKom rono;tylcre, no.no>uenne nop CC-

menpuemmucon, xapaRTep mymautux 110.110B1,IN, KACTOR. ,fl,.11S1 Anyx mopcum po-;Loo coaRarro 110B0C no.rIcemericTeo NOOtkadrilirlae. Flp11130,11,11TC51 COCTaB (na vporme poRon) ncex cemn uoncemericTn. flommneno ormcanne Svetlovia nzacii-?cita Cel:anovskaja B pe3y.nuraTe nepenceneRonaung romoTuna 3T0F0 BuRa.

FIonsTKit HaBecTH TIOpSIAOK B cemericTBe Tubificidae, rIpeCTaBHTeJITI KO-Toporo Hnipoko pacripocTpaneHbi B npecnbix 11 mopcHnx BoAax, npepnpinut-maoncb ,paBno. IlepBonana.nbno cpep,ii .ry6iulniunp, paaminamt ,H,Ba Hop,- cemeiicTsa (Eisen, 1879): Tubificinae H Telmatodrilinae. HecHo.nbRo nowke-(Stolc, 1885, 1888) 6bh.rui coaftaHm ente Ba nop,cemeficTBa: Ilyodrilinae-Stolc, 1885 H Bothrioneurinae Stolc, 1888. Ilyodrilinae ne,paBito 6bmo ne-penmenoBano Fpa6be (Hrabé, 1963) B Rhyacodrilinae Ha ocHonatinn TOr0,.

tiTO TIOTOM ero coyAwn Ilyodrilus coccineus Vejd., Tenepb cHHotnim Rhya-codrilus coccineus (Vejd.). artaro,RapH miTenclubmainin Hcc.flep,oBannik nomep,mte ,a,Ba p,ecnTii.neTHH onlica• M110r0 11013bIX (1)opm Ty6n#H,H,p. Oc-mucnimatine rIOCTO5IT-1110 naHaruntBaiouterocH mopci)oJIortmecHoro maTepttana riptitto,amo K pCB1l3IITM cTapbtx HOMITal, ,II,Har11030B, Hpirrepnes. Me1I s-1m 1n H ripe,pcTaB.IctutH 0 noitcemeiicTBax Ty611(1n1luip,.

Fpa6be (Hrabé, 1966) npe.p.Tio)kna Butte.rinTb poa Brancldura Bedd., o6- ..riaRatouttifi cBoeo6pa3timm cTpoennem n000noro annapaTa H HexoTopumn Heo6bi4nibtma comaTtmecimmii npitaitaxamii, B ontesibitoe nop,cemeficuto Branchittrinae, a aaTem (Hrabé, 1967) Aulodrilus Bretsch.—B no,pcemeil-cno AulocIrilinae. 1-IcKationcHan (1972) npnauma no,acemeilcua, BbiReneli-Hue Fpa6be, no He comacitaacb c Bbtpe.riennem nojtcemeiicTBa Telmatodri-linae, paccmaTpuBast naBecTubte B item pop,bi Telmaiodrilus Eisen H Ale-xandrovia Hrabé B cocTaBc Ty6H4ytiumt. • BpiitiKxepcT (Brinkhurst, 1971) opratutHoBaJi p.nst rpynnbi mopcHnx cpopm ente gallo noacemeficno Phallo-drilinae, a II3BeCTI-IbIll paHee mopcmii poji, C/itenio Sav. OTRO,TIILTI OT Tubi-ficinae B oco6oe TIO,II,COMeriCTBO Clitellinae. Bnponem, B 1979 r. (Brinkhurst, Baker, 1979) 3TOT po iu citoBa 6but Bonpauten B noRcemericTm Tubificinae. Kpome Toro, 3TOT }ice arrrop (Brinkiturst, 1971) nomecTihn Bothrioneurum Stolc B noftcemeiicTim Rhyacodrilinae, ynpaafflum TM ca/vibrm MOHOTHrIII-nenoe'noRcemeticTeo Bothrioneurinae, a B ,pa.gbiteruuem (Brinkhurst, 1981) nepeBe.n B 9TO >He noRceiviencTm pop, Brancldura Becld., ammit,ititpoBaB,

' BTopoe moHoTHruptecHoe nopcemeficuo Branchiurinae. H, HaHoHeu, Dpceyc -(Erseus, 1982) comaa nome noRcemeticno Limnodriloidinae Ha ocnoBe HeCKOJIbKIIX mopcmix po,poB.

06-bem nop,cemeileTB ne OCTaBaTICS1 110CTOSIIIHIDTM. npOlICXORHJ10 . IIOTIOJI-tienne cocTaBa muortix HORCOMerICTB, oco6eHrio mopcxux, BHoBb olTHcalllthI-

Mil it Hamelleune ero B peay.nbTaTe nepemetue•unl 11 peBH31.1i y›He.

194

113BeCTIIIDIX poacm. ,II,J151 060C110BaHH51 no,""cemericTH B xanecTise upliTeplies 11C110,T1b30BaJIIICb pasHme HpHsHaull, HO raamimm o6pa3om OCO6eHHOCTII

crrpoemin reHepaTHBHoro annapaTa. 3ti3eH (Eisen, 1885), Hanpnmep, npli-BOASI upaTullit ,rmarHos no,acemacTB Telmatodrilinae H Tubificinae, ynomli-Haer T1J151 nepBoro no,acemeticTBa Hecuozbiço npocTaTimecuux weaes Ha aTpint H 0c06y10 cTpyuTypy neHnambHoro o6pa3oBami5T, a 'tan sToporo Hoa-cemeiicTisa— oitHy npocraTimecuyio ›ueaesy Ha annli H inlyio neHHaabitylo cTpywrypy uau Haii6onee xapauTepnbie ripHsHaun 3TTIX noacemericTB. ILIToabu (Stolc, 1885) AJISI pas,aeaeHlin Ilyodrilinae (ceinac Rhyacodrili-nae) H Tubificinae npuaneimeT H Taime HpnsHaim, uax cpopma npocTaTittle-cmit aœ.nesm, Haannlle Hari oTcyTcmie cnepmaTocpopoB, cnoco6 pa3BHTH51 511111, no THny Naidomorpha (Rhyacodrilinae) uia rio Tnny BbICIIIIIX nepBert Tubificinae, Haatrinte HJIII OTCyTCT11311e TIOJIOBbIXILleTHHOK, a Tau>ue cpopma ne- miaabHoro annapaTa.

Fpa6be (Hrabê, 1963, 1966, 1967, 1979), 1%1Horo ca,eaaminlik JTJ151 cpop-mHpoisainin OCHOBHbIX npe,acTameimil B o6oacTH mopcpo.normi H cHcTema-THKH aintroxeT, TanKe onHpaeTcn H a npusnault, xapauTepnsymiune no.noBoil armapaT,.a upome Toro, ncnoabsyeT 1IeCKOJIbK0 comaninecurix OH ripHsHaeT caeasimune npnaimun Han6once Ba}I{Hb1M11 fi,J151 pasagaernin Ty6n4unnta Ha noacemeficTisa: 1) (Popma (ancpcpysHan Han 060C06J1e11H351)

H 1IHCJIO (o,rtHa HJIII HeCKOJILIKO) npocTaTimecunx xœ,ries; 2) (Popma mr}I{CKIIX ITOJIOBbIX meTox (cnepmmi Hall cnepmaTo3efirmm); 3) noomemte nop ce-menpnemmtuou (y nepeattero ,iiiiccermmeHTa Han B cepe,aime cermellTa); 4) crloco6 o6pa3oBaHlin iiiu (cospeBaime 511111, B tIaCTIFITILIX mininumx Han noo)niionue); 5) HaaHnHe 11.011 oTcyTcre.He HcTininoro nemica; 6) cpopma Hecppn,imeis (c mennwo6pa3Hbim nocTcenTaae 311XIITpelI)1II01.0 Tuna UJIM B BH)I,e HeCJIHTbIX neTeab); 7) Haanyme HJIII OTCyTCTBlie B TIOJIOCTH Tena upyri-H1,1X 1]eJ1OMOLITIT013; 8) Hamme HJ1H OTCyTCTB}Ie CHHHHbIX Hop, a upome TOPO, npH Bbmeaeium no,acemericTB Bothrioneurinae n Branchiurinae,— He-xoTopme npnsimun, npncyunie po,aam Bothrioneurunt n Branchtura.

BpimmepcT (Brinkhurst, 1971) eamicrseHno Bamimmil u y1o6i1mmn elniTaeT H 7 - fi npitsHault, KoTopme HmeroT, HO ero MHeHIII0, ,npelimy-Ills eCTII0 B TOM, IITO 11011TH Hensmermo ynommtaloTcH B ornicaminx H Ha HX OCHOBe cemelicTuo JierKo guarren Ha rpyruibi. ()Alla», mu. MbI orme-t-mail BbIIIIe, cam BTOP 6bra He concem yaosaersopeH cncTerviml Ty6Hdinu1ia H Bnoc.ne,acTBHH npoitomuan yaynmaTb ee, npousuo,an nepecTaHornm. KcTa-TH, B pa6oTe 1981 r. (Brinkhurst, 1981) OH IICTIOJIb30B3J1 B etiarHose Rhya-codrilinae eute ojuiii rpa6beiscunii ripimHax, a iimemio — rio.nonœnne - Hop cemenpnemimuois.

ECJIII paccmaTpmsaTb Branchiura H Bathrioneurum B cocTaBe pua -

KO)i.pHJIHH, tu° MbI cluiTaem npaisHablimm, TO 4-a, 641, 7-ri H 841 nplisHauH B MX Tese (HmeioTcn nacTlitnime minium", meanwo6pa3Hme nocTcenTaae HedipHalies, upyritime uscaomou•Tbi, crunnibie Hopm) ripncyinn TOJILKO sTomy no,acemeilcm, HpHTom Heo6n3aTe.nbito ;tan .Bcero pua cpopm. Tau, nacrin-HbIX 1HtI11IIKOB HeT y Bothrionettrunt, Rhizodritus, Paranadrilus, ,H,J151 Heuo-TOpb1X ponoB nemBecTell xapauTep o6pa3orsami5I 001111TOB; meuncoo6pa3nmx HocTcenThae necppnatieB lier y Rhyacodriloides, Rhizocirilus, Branchiura, 1JIIHeKoTopux poaois neT Aa•ibix; CIIIII1HbIe nopm ormeneHm TOJIIDKO y

Epirodrilus, Rhyacôdrilus, Bothrioneurutn, y OCTaJIMIbIX p0,a0B nopm J11360

OTCyTCTBy10T, J11360 0 HM HeT cHeaeindu ITeJIOMOIIHTIDI H3BeCTITbI JI,J151 18 po- )1,0B, OTCyTeTByIOT Hnoraa y Quiloro po,aa, TIOCTOSIIIHO y Tpex poaos. Taimm 06pa3om, nepetniczeHHue JIHIlIb noRnepuinsaloT cBoeo6pasHe pua- KO,i(plIJIIIH.

ITTO uacaeTcn cpopmbi lieHHca riplisHax), TO OH MO}KeT runicrcno- BaTb B npeeezax ()Riions no,acemeticrisa B Bitae nceimo- Ham ucTiumoro ne-mica (13 riecuanburix no,acemeticTBax). Tau, cpeaa cpaamoapHaim 6 pOgOB HmeloT npocTme nopm, 4 poila —ncenaonermcm, 5— lleITIICH (113 flUX 2 po,aa C XIIT11110BIDIMII o6KzafiKamii, Kau y Ty4)I161111HH); cpeari TuibmaTo,ruipHami

3* 195

1 pon, nmeeT neimc, 1 — ncen,gonennc; cpmn pnaKoepHarm 3 po,na Hmemyr nefulc, 8— nopm, 10— ncen,noneunc.

AnaJmn orpomnoro n pa3noo6pa3noro mopcko.norrinecKoro maTepHa.na no Tyclm6mm,nam, ony(5AnKonannoro B noc,nen,nee spemH, y6e>m,uaeT B na>K- HOCTII CJle,UyIOILUIX npnnuaKon pange.neunn aToro cemeheTda na no,nce- mencTna: 1) na.nunne (KaK nan6o.nee yrunnepca.nnnuil npH3HaK pnaKo,n- plum') HAIL OTCyTCTBIre Kpyrinnix B HOJIOCTIT Tula; 2) xapaK- Tep npliKpen.nennn, cpopma 11 gam° npocTaTnnecKnx Amen na Ka>mom my>(-CKOM ronoywre; 3) no.no>KeHne nop cemenpuemmmon; 4) xapaKTep my>K-CICI1X HOJIOBbIX mneToK. Ha ocnone 3T1'IX npunnaKon cocTamena me,nyrannan onpe,ne.nnTe.nnuan Taônnu,a no,ncemeticTs Tubificidae.

1 (2) Kpynnue (nnor,.na cpannirreobno memme, HO mHorounc.nennue) ne-TIOMOHHTbI B HOJIOCTII Te.na, Kai( npanmno, eCTb (nun,' neT, TO upocTa- TnnecKan. >Ke.,Tena ,ancl)(pynnan) Rhyacodrilinae

2 (1) Kpyrmux HejlOMOIIIITOB 3(8) OTBepCTIIR cemenpnemunKon y nepmnero (n ange ncrannottennn

Hero) ,auccennmeirra (n oT,ne.nnunix peRKnx mynanx nocepe,nnue cermenTa).

4(5) Ha Ka)Kaom my>KcKom rono,nyKTe 60.11bLile Anyx npocTaTHHecKux >Ke-J1e3 Telmatodrilinae

5(4) Ha Ka.>raRom my>KcKom rono,nyKTe He 6o.nbine ,anyx npocTaTnnecKrix >icemen.

6(7) flpocTaTnnecKne >Ke.nenbi, emu ecTn, KorvmaKTffle, nacTo• cTe6mb-naTnie, pe.nœ c innpoicum ocnonannem Phallodrilinae

7 (6) 0,nua npocTarnnecKan )Ke.ne3a co cTe6e.nbKom, nTopan — ,amwn-na51 MIT oTcyTcTnyeT NootIcadrilinae subfam. n.

8(3) OTnepcTun cemenpuemunKon •npnmepno B cepeiinne cermenTa (pe,n-1:0 CaBIII137TbI xnepe,rui BAH KnaRn).

9(10) °Ana npocTaTnnecKan >manena, umpoKo npmxpermennaH K npocTa-TnnecKofi Hporana,aKe (prostatic pad) mum BAH ero AnneprnKy.na Lhnnodriloidinae

10(9) On,na npocTaTnnecKan >Ke.nena, npllKpen.nennan K mum c nomonwo CTe6e,TIbKa.

11 (12) HmeloTcH cnepmnn Aulodrilinae 12(11) HmetoTcH cnepmaTonetirmnr Tubificinae

Hapyr,ny c npnnnaKamn, yKanamibuinT B onpeRe.nnTeannoti Ta6,nnue, OT-menim pn,n Apyrnx Ba>Kiinix npHnnaKon, a TaK>Ke o6'nem nogcemericTn.

RHYACODRILINAE FIRABL:, 1963

ComaTnnecKne ineTnnKH pannoo6panHnie: Anyny6qaTme, Qrmo3y64aThie, Tpex3y6naTme, imor,na c znramenTom, coegnnmoui,nm inDKniul 3y6en co CTBOJIOM werrnmm; Beepnbre, BOJIOCHbIe. LIOJIOBbIe lleTHHR1I nacTo nmerayrcn. 1/Inorn,a nmeroTcH crumble nopm. Heckpn,nim nacTo c meunwo6pa3ubuvr nocT-cenTage. Aly>Kcime rono,nywrrar C ,U(Hclupy311o1l mneTKamn HJIH rpynnamn meToK) npocTaTnnecKort >Ke.neaori. HIIOUB HMelOTCSI Kony.nnun-onnue >Kmenni, nacTo rucTo.norrinecim cxo,unbie C npocTaTnnecmoil TKaIlb10 (nanpumep, y Macquaridrilus). Hnorea npocTaTntiecKnx K,neToK neT.My>K-exile HOJIOBble oTnepcTnn B nun npocTnix nop HJIH ncen,nonenncon (nuor,aa nammoneHumx B umomnnecKne meumn), nacrosanne neuncm pe,mm. Ceme-npnemnnKu napnbre, pen‘ Ko nenapnue, oTKpunatoTcH B X cermerrre y nepe21- Hero (oneub pe,nKo naRnero) finccermmenTa, infor,na n nocTepnopuori nacrin IX cermenTa, unorna OTCyTCTByI0T. Ileterodriloides nmeeT 2 napni cemenpn-emunKon B XI H XII cermenTax. Cnepmnn B Blue cno6o,anoii maccni, onenb pe,n,Ko B BMW opraunnonannbix nytwon. flpn oorenene y neKoTopux pogon o6pa3yroun MOpyJII)I 001I,IITOB.

B BTO no,acemeticTno nomemenni cae,nyloume pub': Epirodrilus Hr., Rhyacodrilus Bret., Svetlovia "ekanovskaj a, Paranadrilus Gavrilov, Pro-

196

tuberodrilus Giani et Martinez-Ansemil,. Branchiura Beddard, Bothrioneu-rufn *Étole, Peristodrilus Brinkhurst, Rhizoclrilus Smith, Monopylephorus Lev., Rhyacodriloides &kanovskaja, Jolydrilus Marcus, Ainucirilus Fino-genova (---.Vadicola Baker), Lymphacliaeta Snimschikova, Heterodrilus Pierant., Heterodriloides Erseus, Gieredrilus Erseus, Macquaridrilus Jami-eson, Heronidrilus Erseus et Jam.

Ue.nomourrrbt (cambiti xapaxTepribul nprisitaK 9Toro non scemeikTna, OT-cyTerrspoLunii y SM{ OCTaJ1b1111X no,ncemeticTH) He •arirreribr y Rhyacoclrilo-ides, Branchiura, Lytnphachaeta, ne Heurta oTmenaioTcri y Rhizodrilus pa-cificus. He IICKJ110 ,10I10, npnypotienbi K onpeng.nermum CTaRlIBM

IK•3liennoro turima II B Kame-To nepHonsu moryT OTCyTCTBOBaTb.

P0,11, Svetlovia 5m.n co3nan IleKanômKoil (1975) A.ng a5licca.nbiroro 5ail-Kanbcrwro muta S. maculata. 143yneinre Hamir romTnna npine.no K neKoTo-pomy immenemuo H ,nonomiemno ero nepHoomicamm.

Svetlovia maculataelcanovskaja

FOTIOTHII. CememniKoH 2 napbr H IX cermerrre, KaK oTmetraeT LleKanon-cKan, II fronomurrenbrian napa meribumx no pa3mepy B VIII cermenTe. Cmi-pa.nbno 3aKpyliennbi1i cemympoHoir RJ111110ii OK0.110 250 MKM BriaaaeT anii-KJIbHO c6oKy B TIIf, KoTopuil COCTOTIT 113 OBaJlb110fi amnyabi, y3Koro cemsm3HepraTanbnoro 'canula ii maccHHHoro ncel3,noneunca, a He nemrca, KaK yKa3aHo H nepHoomicamm (pricyHoK, A). Amnyza aTprm c xce.ne3HcTbrm anwreariem (3a iicKmonermem swramHoro Korma) H pecHnliKamH, R.arma ee OK0.110 200, umprma maKcHma.nbrlo 115 MKM, Tommina mycKy.nbHoil o6mna,rr-KH amnyou ,/to 7,5 MKM. MycKyoncTbril cemsninepraTumbril KaHaz ,AJIHHOff

oKo.no 40 H umprinoil 17,5-30 MKM ruvieeT otreHb y3KHri npocHeT (2,5— 5 mKm). nceHrronemica 137,5 mKm, nmpHria H iterrrpaabnoil tracTit 75 MKM. Mnoroniicaelinue mbnuelmbre BOJIOKITa npliKpenonior nceHRonemic (oui' qacTnutio nmelorcn II na cemnunepraTezhnom Kana.ne) K cTenKe Tula. FIennaebnble TIO3a,aII ITCCB,11,011CIITICO13. D- TOT po,n, mopciro.no- runecKH 5,1m3oK K Epirodrilus, Ho OTTIIItlaeTCSI OT Hero Ancpcirepernmanirei1 annanbriori Tpy6Krl Ha amny.ny, cemsinnepraTemiimil Karlaz, nceHironennc,

Tame Haannirem pecHnneK BnyTpir aMr131.111)1 must, parmomepnoil T0,11111,11- Hoti cTeimn qacTeil annaabiloil Tpy6Kn.

PHALLODRILINAE BRINKHURST, 1971

ComaTnnecKne ineTinum TOJIbK0 ,U,By3y6 ,1aTbIe, nnoura c ourameirrom, COBA11115110111IIM 3y6eu, co cr3o.nom meTinum, KaK IICKJ110 11e14110 HITOr-

,na •meeTcri TpeTIM 3y5etr. flo.noBbie meTnrucli nacTo HMEIOTCH. KuruKa y HemoTopmx BITROB oTcyTcuyeT. Ha KarK,nom rono,RyKTe y 60,1IbITIIIIICTBa po-ROB 2, pence 1 ripocTaTrinecKan xceze3a (unor,rra orcyTcrHyeT). Y ireKoTopmx po,non 9Kraohriari npocTaTa (ncen,noripocTaTa?) npliKpermena K ocuoHaruno neunca il (mit) K cTenKam neHumbribtx memKon. Cnepmnit cHo6oeriori Niac-coi1, KaK ncKnioneune — Kpyr,nbic cnepmaTo3etirmbr.

K 3Tomy no,ncemeficTiry OTH005ITC2 pOJJIDI: PhallOdrihIS Pierant., Bathycl-rilus Cook,,Aktedrilus Ku., Adelodrilus Cook, Coralliodrilus Erseus, Ba-cescuella Hr., Inanidrilus Erseus, Olavius Erseus, Jamiesoniella Erseus, Spiridion Kn., Atlanlidrilus Erseus, Uniporodrilus Erseus, Bermudrilus Erseus, Peosidrilus Baker et Erscus, Duridrilus Erseus.

HecmoTpri na licK.rnoturrem)noe cHoeo6paane neKoTophix HpI1311aKOB OT- ReabliblX porroB (nenapubni cemenpnemmiKH H XII cermerrre, °Ter- CTBIle KIIIIIKII, cemsmpono,n, 6e3 pecungeK H irp.), BTO onerib Kom- naKTilan o,uriopo,nuari rpyrina, B KoToport aerico npoc.nexumatoTcri CBY131I

merK,n,y po,namn. D-pceyc (Erseus, 1979) cnifraeT npoToTnnom mnornx pon,oH ciropmy, 6,rin3Kylo K Phallodrilus.

197

21e-rami crpoeatta Soellovia mandata (A), NoolkacIrilus lunnalus (B) ii Discordiproslalus longiselosus (13): A— Tepmintamblian Itatm, my)Ketcoro rolmaywra (carrirrammt.eil pa3pe3); B-13 --no Baker, 1982; 3. n-3amina npoc-ra-rugeeKaa )1( e.ne3a, ti. C aonymittuounael eyhtaa, 3f— mbanithi, At. 2— memiantime tiae-ra my)eekoro roecifiywra, d — myxcaoe norto-ace oTeepc-rne, n—ricertitonetinc, n. n—nepeRnan npocTaTaxteeKaa Imene3a, n. aJ11311ble 111,C111111(11, C —CeNICIlpnemmuc, C. (3— cemelutaa noponka, cii cuisit-11)0E10R, ais:. 2—

Dirraindian tiaCTb MyiKCK0r0 roHoRma, 311. 2— TO )Ke, 311TaelbllaSI tlaCTIa

NOOTKADRILINAE FINOGENOVA, SUBFAM. N. •

Comanmecime merFuncH Any3y6t1aThre HJILL ozm3y6LiaTbie. Hmeloren ne-Humbuble meTHHKH. i'Vly}Kcime rono,aywrm COCTOSIT M3 4 nacTeti, pa3r1ma-101.11,I1XC51 no nmpime, nplicyrcumio Mill OTCyTCTBH10 pecrarmoro . 911HTeJIHA, Toomtme mbnuennoro UMM; oxamumaloTcH neeH,RonemicamH. Ha swra.nbHom KoHne Kaw,uoro rono,aywra nmeeTeg eTe6e,ribilaTasi, a Ha 3HT3AbFIOM — )1,11(1)- (1)y:31-m51 npocTaTugeeKa51 x<e.ne3a. ,E(HcpcpysHasi npocTaTa moweT OTCYTCTB0- Ban), B 3TOM c.nytiae 3nirrefunI 9HTMJIBI1011 qacTH ro,rimennbuI H >Ke.riesu-eTbifi. Flepe,Lumf qacTb myAcKoro 1.0110,UYKTM —B X cermeHTe. CriepmHH CB0- 60110ii m aceoti (pneyHoK, B — B).

CocTaH noRcemencTria: Nootkadrilus Baker, Discordiprostatus Baker. Bernœp (Baker, 1982), onneammul o6a po,n(a, nomecTna Hx H no,wemeii-

CTBO Phallodrilinae Ha ocHoHamm ripncyTcum2 cTe6e.nbgaToii 3aJHefl npo- cTaTimeeKori (1)0pmb1 cemenpnemuirKoH, my.›KcKoro rollo,uywra H ne- HHJIbHMX Il OTCyTCT13F151 xpyrmbrx HeJlOMOLIHTOB. OH cmwraeT, Trro Nootkadrilus uau6o,aee 6,rumoK n Coralliodrilus Erseus, aTpufl KoToporo Jammu npocTaTmecKux }Ke.nea, H n Inanidrilus Erseus, nmeiomemy TOM,» samuoio npoeTaTy, neunanbubie meTHHKH zayx THHOB H MYCRYJITICTble ceme-npHemmum. Ha Ham H3rJ1si,us, e,,unneTHeuna eymecTHefman (Kpome OTCYTCT-ULM MCJIOMOHIITOB, xapawrepuoro H ,V151 ,apyrux noAcemeficTH, xpome pila - icojpiiJiiiu) ii o6masi e (1)ao,no1mmuHamll /lem — Hamme aulleti cTe6eob- itaTort HPOCTMTbr. WeJle3liCTIA aKe 911HTe.THITI aTpHsi npH OTCYTCTIMM npocTa- TuLtecmix mce,Tic3 H3HecTeH H ipm ,apyriix Ty6wpnuu,q (Hanpnmep, epe,THI pna- 1CO/!j)IIJIHH Svellovia, Epirodrilus), Kax H myexy,nHeTaz ribmwisHaH qacTb ce-

198

j

menpueminixos (Suetlovia). Kpome TOPO, Tpy,tuto HCKaTb p0,1:1,CTBO meA,ay Nootkadrilus, Bepownio, wropHinto TPT11BI11i4M CeM5111p0B0e, H nepe,runoto npocTarrintecKyio ACJIE3y, H Coralliodrilus, TaKAe BTOpHILHO yTpaTHHumm 06e npocTaTbi Hcxo,mioft tpopmbt, Ho o6.riajtaxnumm RocTa-roanta ,II,JIHHHbIM ce-mimposonsolvt.

Tem 6o.nee Tpyfuto 061,FIC1111Tb nommeHne H oaHom polie imam-lux npnatia- ROB ()KeJle3HCTbIrt 31-111Teallti ronoitywra npit OTCyTCTBIII1 npocTaTbt 3HTal1bH0

cTe6e.nbKoHast ripocTaTa 3KTaTIbH0) TaKitx Rsyx RocrraTottuo nporummyrbix, a He npitmitTHHHbix poutos cita.nooRpnottn, KaK Coralliodrilus ii Inanidrilus. Nootkadrilus n Discordiprostatus, Hecomiteinto, mopcpo.norntiecKH 6.11113KH

mew,ay co6ort, 3FIT3J1131-Ible qacTit EX aTpnea, no-HH,rtnmomy, romo.norwillibt, 1103TONly etu,e Tpy,ance nepeilTH oT cfm.A.no,aptuntri K Discorcliprostatus — cpopme c 0,111111M 113 Hatt6o.nee fliiMIITIIBHbiX THHOB npocTaTtit cpeett ()miro-xeT (a,m1.4y3noi1). Bo BCRICOM c.nyttae, TaKoù nosspaT K 6onee RpertHemy COCT051111110 9TOrl Hawitoil c.rpyKTypbt B npeRe.nax o,atioro noRcemeficTsa ripti OTCYTCTB1411 itpyrtix crientultuttecmix gepT (ka.rhao,apnanit npeitc-ramaeTcH mano HeposyrnbiNI.

TITO Kacae-rcH Tonorpaclum myAcKoro TOHORyKTB, TO cpe,a,H 6o..nbatoro maTepna,na no (pa.n.noztpthatinam, HaKonaenHoro K nacTosnu,emy Hpemetut, He HafiTH ana.norun c ynommlyThimH po,aamit. Cam B9iiKep ninueT, tITO TpHH Nootkadrilus cambre czoAnble, a Discordiprostatus pança.nbitbr cper,H Ty611-4matnit. Bce HmtuecHaaantioe no6yA,LtaeT mem CO3,a3Tb fl,J151 3THX pOROB Ho-Hoe no,acemeficTrto. OTMeTIIM ente oju-ty peRxyto OCO5eHHOCTI3 HX mop4toao-11114— ripoHin-notienne myAcKoro ronoityKTa tlaCTI1qH0 113 XI B X cermeHT. D-Ta OCO6eHHOCTb Ii3BeCTHB y Rhizodrilus H Bothrioneurum B noRcemeticrEte Rhyacodrilinae.

TELMATODRILINAE EISEN, '1879

ComaTtutectme uteTinnut pa3HbIX TIH1OB: ,r1Hy3y6ttaTb1e, oRHo3y6qaTb1e, uteTKoo6pa3ti1le, itmocubie. flo.aoHbie meTunKii (men nmetoTcH. flpocTant-ixecKne /KeJ1C31,1 KONItlawruble, cTe6e.nbtiaTbie, 11X 1\10)KeT 61)1T1) Ha KaA...a,om min' Ro 16. ATpIIU aaKatitntHatoTcH nefutcamm 11311-1 nceHRoneHticamn. toTcH cnepmun miiii tiepeeo6pa3nme cnepmaToaefirmu. K aTomy noAcemeficT-Hy oTHocwrcH pub' Alexandrovia Hr. H Telrnatodrilus Eisen. XozmiquicT (Holinquist, 1974) npeiaraeT pas,aunurb Telmatodrilus iia ,Raa poRa.

LIMNODRILOIDINAE ERSEUS, 1982

Comaritieckue uteTuinot TOJIbK0 ABy3y6liarrbie, Hape,ruca iimeeTcyi oura-mewr muk,ay im)kuum 3y61Lom H CTB0.1101VI tuerniku. nOJIOBbIe IHeTHHKH pe,a-

B IX • cermetrre mo,a14mumporiau—H 3 E4 B3RYTblfl H )KeTIC311-

CTbIii 11 3111 cHa6 -Aett napofi .naTepo-Hewrpagbtibtx RuHewrillty.noH, nanpamen-EUX anepeR K RuccenumenTy 8/9. flpocrranulecKaa x(ezeaa ripnicpermeHa K

aTptiambHoll amny.ne, Kat; npartn.no, unipoKum ocHoHaHnem, Ho B enlIHhltIEbix

CJIrlaSIX C HOM0111,b10 L0BOJ1bIl0 yaKoro ccpunwrepa H cHaaatta c mo,inulunut-poBaHribim HilyTpennum anture.nnem amnymm, o6pa3ytotunm xapaicTepHyto npoma,axy c aeptincToii upyicrypoii (prostatic pad), HJII1 C aTpHa.nbublivi juittepTHxy.nom, TaKAe nmetoullim rpatty.nripoHattuyto Flpoicaa,u,Ky. ATplut

at;.21HatinatoTcH mante Heero nceHRonellncamn 1131 11 nopame, omettb pe,aKo Ha-ETO5IIII,I1M nenticom. Cuepmun B Blue cH0601tH0ii maccbt, riyiKoH 11 3111 peako ripoc -rbtx cneptvtaToaefirm.

CocTaH noacemeficTHa: Litnnodriloides Pierant., Marcusaedrilus Righi et Kanner, Tecticirilus Erseus, Thalassocirilides Brink. et Baker, Smithso-nidrilus Brinkhurst, Kaketio Righi et Kanner, Parakaketio Erseus, Doliod-rilus Ersetts.

AULODRILINAE HRABÈ, 1967 •

ComaTimeckne meTmum euy3y6qaTme, necoonnenbre, Beepnme, B0110C-Hme. Hanonme ineTimmi peeKu. CememinKon 1-3 riapm, sunnniKon 1 napa. flpocTaTimecKan .)Ke.neaa c yaimm inupoKum cTe6e.nbKom. ATpun oKan- q11Ba1oTcH neuncaM11, xoTopme nacTo nmeioT XHTHHOByI0 o6oJny-wy. Ceme-npneminimon 1, pe,uKo 2 napbr. BI Ubl Aulodrilus pa3111 1-10WalOTCSI B OCHOBHOM apxuTomneii, pme —110TIOBIJIM nyTem. nmoboti annapaT o6E,n1io FIT na HeCKOJIbK0 cermenTon K nepeenemy Komw. Ha 3a,n,Hem Konu,e Te.na infor,aa nmeeTcn necermenTnponaiman 3 01-l a, cpynKunonnpyloman KaK )Ka6pm.

K 3Tomy noecemeticuy OTHOCSITCSI meeyionme poem: iltdodrihts Bret-scher, Peipsidrilus Timm, Neoaulodrilus Giani, Martinez-Ansernil, Mou-bayed et Dia, Sketocirilus Karaman. floc.neenne eba poea ogenb 6J1I131(13, F103M0)K110, Fix cJieeyeT 06'be,LtHHHTb B oeim po,u.

TUBIFICINAE EISEN, 1879 .

Bbicume Ty614nuulebi, ComaTnuiecKne tneTiniKll eny3y6traTme, 0en03y6- tiaTme, Beeplime, BOJIOCHMe. Ilo.nonme ineTunKu, KaK upannzo, nemnoronitc-.nenum, nacTo HX lieT. KomnaKTilan cTe6e.nbliaTa5I npocTaTnnecKaR IKezeaa ompubaeTcsi B aTplin me,ananno IIJIII B 110J10BHFIe, morea OTCyT-cmen .ATpun 3axal1t111BaIOTC5l nenucamn, nomeumnimmil B nennainpnbre Kapmairm. Henncm moryT nmeTb xirrinionyio 060JIOLIKy BAH Tpy6Ky. BeTpe-tiaeTcH pa3Mn01cemie flTM apxuTommi.

CocTan noecemeficTba: Tubifex Lamarck, Limnodrilus Clap. Isochae-tides Hr., Potamothrix Vejd. et Mrazek, Psammoryctides Vejd., Ilyodrilus Eisen, Clitellio Say., Clitelloides Finogenova, Tubificoides La st., Spiro-sperme Eisen, Peloscolex Leidy, Baikalodrilas Holmquist, Haber Holmqu-ist, Antipodrilus Brinkhurst, Lycodrilus Grube, Lycodrilides Hr., Tabipe-r-ifer Semernoi, Arciodrilus Brinkh., Spirospermoides Dumnicka, Krened-rilus Dumnicka, Varichaetadrilus Brinkh. (=--11yodriloides Solc.)..

• B cocTaBe Tubificklae npnanaeTcg cemb no,ucemeiicTri, o6rbeenlinio- rim 72 poea. Phallodrilinae, Liinnodriloidinae, Nootkadrilinae mmionaloT TOJIbK0 mopckne IIJIII conolionaTonoeume 1)04 1,1 (c eeliminnumu Bruamir,

nTopimo B npecuoii noec), Auloclrilinae ii Telinatodrilinac ume-lOT TOJIIDKO flpCCHOBO,UllbIX npeecTanirre.neri, a Rhyacodrilinae H Tubificinae coeepwaT NaK ripecnonoenbig, Talc 11 mopcKire chopmbi 14, Kpome Toro, ne-CKOJIbl<0 poeou co cmemannumn apea.Tramn. Bcero B nacToninee spew! na-.etrirrbinaeTcn 35 mopcmix 11 33 npecirono,vibix poea Ty6i4nune, a TaKx<e 4 poea, o6nuix uni mopn 11 npecnbrx nu.

o qumoreneTnnecluix OTHOLLICIIIIRX 110,RCCMelICTB noKa ronopnTb Tpyeno, no, necomilenno, o6qiiic pnaKoeplunin nan6o,nee npumuTlinen. HecmoTpsi ira H313eCTIIbIC 11CpTbI criennam13aumi, aeecb MbI cTunmnaemcn c riall6oabninm ITHCJIOM UPT opramiaamlli. 3T0 OTCyTCTBHe o6o- co6dienn01l npocTaTiniecKoll 1KCJIe3BI, H nammile cnepmnen, H caa6a5l Julep-(i) epennuanum nenumbnoro annapaTa, meTamepan ronae (rianpnmep, Svetlovia, Bothrioneurum), il Briaeenne cemnriponoea na 3I(TaJ1bH0M ICOHHe aTpIIFI, II, neponTuo, pacno.noAenne oTnepcTiiii cemenpnemimKon B nepe,uneri liacTii X cermeirra, naaninie Kpyrnibix u e.110M011,11TOB, 06pa3onanne mopyn 00H- IITOB. ,Llpyrag oToinurrefibiran nepTa priaKoepiumn — mopq)morrinecKan

nponn.rinromaSICH, B tlaCTHOCTII, B Kparmem muor006p331111 cpopm II cTpyKTyp reilepaTunnoro annapaTa.

Han6o.nee 6Jli13K1i K pnaKO,UplIJIIHIaM (1)a.Turoejmulinnor II nyTKaepimulibi, y KoTopmx, necmoTpn Ira paanirrue OT,ReJII,HbIX nporpeccumbrx npumaKon, KaK, nanpumcp, KOMIlaKT1IbIC npocTaTbr. cao)Kno yupoennbili neinic (Akled-rilus), coxpammoch mnoro npumirrunlibIX HepT: B o6onx noecemerieTnax nopm cemenpuemnincou y enccennmenTa 9/10, ecTb criephum, nepe,uunn eluh-

200

;

c1iy3na5! npocTaTa y nyTKa,apilann, y (kaJumApLunni —BnaReinie cemimposo- ,aa xonen, napg,ay c nemicom npocTue nopm, a6eppaHT- no — meTamepnn rouai:G Bo,nee y,aa.nenbi OT pnaKoRplunin Anmno,apihrionan-

337.110pliJIHIIbI II TeAbmaTo,apnannm, XOT51 y inix ToAe 0)KHO Harun o6tuce c pnaKoziplumnamil: y Linmodriloidinae— npocTaTugecKasi )Keme3a XOT51 pme KomnaxTnag, 110 ripmpermena K aumo na ,21,0BOJIIDHO 0611.1HpHOM npocTpaucTse, y Telmatodrilinae-- OTBCpCTI1 51 cemenpnemmum y nepe,anero

11ccenumenTa, rpytneo6pa3nag cpopma mils', y Aulodrilinae— meTamem ronag; y Beex nepeimc,riennux no,a,cemeiicTs IIMBIOTCH cnepionn (y Te.nbmaTo-,apiunm,' napsi,ay co cnepmunmil, TaK>Ke gepseo6pa3ume cnepmarro3efirmm B po,ue Alexatzclrovia). Fi , naKonen,, y npeacTasnTe.neii CH,111)110 npoRminy'roro Tubificinae c Tpy,aom mœKno naihn annsecTpa.nbume gepTm, cKopee B Blue arrasn3ma. Flanpnmep, KaK ucKmotienue y eiumnimmx B1ILLOB Ty6ncluumn OT-BepCT1151 cemenpnemmums paCrIOJIO/KeIlbl 136,1111311 ,anccennmenTos 9/10 11.T1 H 10/11, a ne me,ananuo.

B pa3HbIX JIiIHIIHX KaK mopcKnx, TaK Il npecnoso,cmmx Ty61liin1um cxo,anoe pannTue ana,norrignmx myKTyp. 3T 0 0T1IOCHTC51: K

(Depenunamm Tepminia.nbnoil qacTil my)KcKoro rono,E,ywra OT npocTmx nop CJ10)KHO yeTpoeimoro nenuca (Phallodrilinae, Rhyacodrilinae, Limnodri.

loidinae); K BT0p11 ,11I01"1 pe,ayKunu npocTaTmiecKoll xœ.1e3b1 (Rhyacodrili-nae, Phallodrilinae, Tubificinae); K xonnenTpamin cuepmneB 113 xaoTune-cmoil maccm B opran1130aanume oupe,aezenumm o6pa3om nylnui ii ga..rtee B cnepmaTosetirmm (Rhyacodrilinae, Tehnatodrilinae, Phallodrilinae, Limno-driloidinae); K pa3BIITHIO nenapnoro cemenpnemnnKa 11.11I1 no.nnomy II0BeHMO cemenpnemmiKoe (Rhyacodrilinae, Phallodrilinae, Limnodriloidi-r.ae); K o6pa3onamno noKpoBamn cneunaJmoro ceKpeTa, K KoTopomy npn-AnnaloT qacTimm cy6cTpaTa (Telmatodrilinae, Phallodrilinae, Limnodri-loidinae, Tubificinae); K CJIITHI1IIIO HO.T1OBIDIX oTeepcnni B 0 11,110 (Rhyacodri-

Phallodrilinae, Limnodriloidinae) H K neKoTopmm ,apyrum npn3na-Kam. KaKoaa. npnpo,na 3T1IX n3menennii (pe3y.rwraT KonseprenTnoro 11,T 111

napumemnoro pammTim), noKa lle 13COFJUI 51C110, necmoTpil na TO, UTO B noc,ne,anee Bpem51 uaiun npe,acTamennsi o cTpyKTypubix noTenangx Ty6qm-

upeasbigaiino pacumplumcb. COBCEM ne,aaano (Erseus, 1980, 1981), nanpumep, cTamo 113BecTuo o c‘nytianx passirrnst cemenpnemimmor3 y Ty6n-(puni1. 3a mywcKumn HOJIOBBIMH oTsepcTi•mir B XII cermenTe. Oco6enno mnoro (palma HOBBIX, Roce,rie Heli3BeCTHIDIX 11.J1 51 Ty6mdmun,a mopcp000rntie-cmix oco6ennocTeii, mœKno Harun B ornicalmsrx mopcKux p0,11,0B, Irro came-geno MII0r1 B ,anarnosax mopcK•x no,acemelicTB (tu,eTinnui c JuiramenTom, ,unBepTimy,Trm Fia !mime 11 arrpnu, oTcynTane Kinum Ir T. e,.) . ,Umbnerauee BrimmaTeohnoe ncure,u,oBanne npecnosoimoii 11 mopcKoil ckaynm,• necomnen- HO, sneceT FICHOCTb BO Mriorile 13011p0CbI cncTemaTiŒn H clmaorenim Tori rpyrnim }KIIBOTI1b1X.

JIIITEPATYPA

tletcatioecKan O. B., 1972. Conpemeimoe COCTOYIHIle cHcTemarluot poRtnex Œrutroxerr (cem. Tubificidae).— B Boitime maoomentinçoque meppit. M.: Haylça, 3-32.— 1975. Howile Ty6mbluntiet (Oligochaeta, Tubificidae) us a6uccamt o3epa BailKadt.— Tp. HOA. nu-Ta AH CCCP, 18, 38, 112-130.

Baker H. R., 1982. Two new Phallodrilinae genera of marine Oligochaeta (Annelida: Tu-bificidae) from the Pacific Northeast.—Canad. J. Zool., 60, 10, 2487-2500.

Brinkhurst R. 0., 1971. Tubificidae.— In: Aquatic Oligochaeta of the World. Edinburgh: ,Oliver and Boyd, 1-860.-1981. A revision of the genus Monopylephorus and rede-finition of the subfamilies Rhyacodrilinae and Branchiminae (Tubificidae: Oligo-chaeta).—Canad. J. Zool., 59, 939-965.

Brinkhurst R. 0., Baker H. R., 1979. A review of the marine Tubificidae (Oligochaeta) of North Arnerica.-- Canad. J. Zool.„57, 1553-1569.

Eisen G., 1879. Preliminary report on genera and species of Tubificidae.— Bih. Kongl. Svenska Vetensk. Alçad. IIandl., 5, 16, 3-26.-1885. Oligochaetological researches. Comm. for 1883.—U. S. Comm. Fish. and Fisheries, pt XI, 879-964.

Erseus Cit., 1979. Inanidrilus bulbosus gen. et sp. n., a marine tubificid (Oligochaetal from Florida, U. S. A.— Zool. Scr., 8, 209-210.— 1980. New species of Phallodrilus.

201.

(Oligochaeta, Tubificidae) from the Arctic-Deep Sea and Norwegian Fjords.— Sar-sia, 65, 1, 57-60.— 1981. Taxomomic revision of the marine genus Heterocfrilus Pierant. (Oligochaeta, Tubificidae).— Zool. Scr., 10, 111-132.— 1982. Taxonomie re-vision of the marine genus Lintnodriloides (Oligochaeta: Tubificidae).— Verh. na-turwiss. Ver. Hamburg, (NF), 25, 207-277.

Holmquist Ch., 1974. On Alexandrovia otzegensis Hrabé from Alaska, with a revision of the Telmatodrilinae (Oligochaeta, Tubificidae).— Zool. Jb. Syst., 101, 249-268.

Hrabé S., 1963. On Rhyacodrilus lindbergi n. sp., a new cavernicolus species of the family Tubificidae (Oligochaeta) from Portugal.— Bol. Soc. Portug. Cienc. Nat., 10, 52— 56.— 1966. New or insufficiently known species of the family Tubificidae.— Publ. Fac. Sci. Univ. J. E. Purkynê, Brno, 470, 57-77.— 1967. Two new species of the family Tubificidae from the Black Sea, with remarks about various species of the subfam. Tubificinae.— Ibidem, 485, 311-365.— 1979. Vodni màloMétinatci (Oligochaeta) Ceslcoslovenska.— Acta Univ. Carolinae, Biologica. 1-2, 1-167.

Stoic A. , 1885. Vorlaufiger Bericht fiber Ilyodrilus coccineus. Ein Beitrag zur Kenntnis der Tubificiden.— Zool. Anz., 8, 638-644, 656-662.— 1888. Monografie C'eskYch .Tu-bificidù.— Abh. Bain. Ges. Wiss. ser. 7, 2, 11, 1-43.

31111 AH CCCP (Jlenuttepaâ) HocTynuna e pearls:1mo 16 uo.sepa 1984 a.

ON DIAGNOSTIC CHARACTERS OF SUBFAMI LIES IN THE FAMILY TUBIFICIDAE (OLIGOCHAETA)

N. P. FINOGENOVA

Zoological Institute, USSR Academy of Sciences (Leningrad)

Summary

The most important characters which have been used in the key of subfamilies in the family Tubificidae are the following: presence or absence of large coelomocytes in the body cavity; method of attachment; shape and number of the prostatic glands on each male gonoduct; localization of the seminal receptacle pores; type of the male genital cells. A new subfamily, the Nootkadrilinae subfarn. n., has been created for two 'marine genera, Noothadrilus Baker and Discordiprostatus Baker. A list of genera in all seven subfamilies is presented. The description of Svetlovia maculata Cekanovskaja is corrected and supplemented with new details on the basis of reinvestigation of the holotype of this species.

ISSN 0704-3716

Canadian Translation of Fisheries and Aquatic Sciences

No. 5245

Diagnostic characteritics of subfamilies of the Tubificidae family (Oligochaeta)

N. P. Finogenova

Original title: 0 diagnosticheskikh priznakakh podsemeistv semeistva Tubificidae (Oligochaeta)

In: Zoologicheskiy zhurnal Volume 65(2): 194-202, 1986

Original language: Russian

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Available from: Canada Institute for Scientific and Technical Information

National Research Council Ottawa, Ontario, Canada 'CIA 0S2

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1986 C

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194-202

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18

Year Année

Volume

1986 65

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Diagnostic characteristics of subfamilies of the Tubificidae family (Oligochaeta)

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0 diagnosticheskikh priznakakh podsemeistv semeistva Tubificidae (Oligochaeta)

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Zoologicheskiy zhurnal (Zoological Journal), 1986, v. 65, No. 2, pp. 194-202

Diae. nostic characteristics of subfamilies of the Tubificidae

familx (Oliochaeta)

by N.P. Finogenova

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The following important characteristics have been singled out and a

key to the subfamilies of the Tubificidae has been compiled on the basis of

them: the presence or absence of large coelomocytes in the body cavity,

the mode of attachment, form and number of prostatic glands on each male gonoduct, the location of the spermathecal pores, the nature of the male sexual cells. A new subfamily, Nootkadrilinae, has been established for two

marine genera. The composition (at the genus level) is given for all seven

subfamilies. The description of Svetlovia maculate Cekanovskaja has been

expanded following a review of the holotype of this species.

There have been many attempts to establish order in the family

Tubificidae, the members of which are widely distributed in fresh

and marine waters. Initially, two subfamilies, Tubificinae and

Telmatodrilinae, were distinguished among the tubificids (Eisen,

1879). Somewhat later (Stolc, 1885, 1888), another two subfami-

lies, Ilyodrilinae Stolc, 1885 and Bothrioneurinae Stolc, 1888,

were established. Ilyodrilinae was recently renamed Rhyacodrilinae

by Hrabe (1963) on the basis that Ilyodrilus coccineus Vejd., now

a synonym of Rhyacodrilus coccineus (Vejd.), served as its type

species. Many new forms of tubificids have been described over

the past two decades as a result of more intensive research.

SEC 5-25 (Rev. 82/11

*The numbers in the right-hand margin are the pages of the caurlime Russian text - translator

Interpretation of the constantly accumulating morphological

material resulted in revisions of old concepts, diagnoses and

criteria. Our concepts of the subfamilies of tubificids also

changed.

Hrabe (1966) proposed that the genus Branchiura Bedd., with

its distinctive structure of the genital organs and a number of

unusual somatic characters, be established as a separate subfamily,

Branchiurinae, and later (Hrabe, 1967) that Aulodrilus Bretsch. be

established as the subfamily Aulodrilinae. Chekanovskaya accepted

the subfamilies established by Hrabe, but did not agree with the

establishment of the subfamily Telmatodrilinae, regarding its

genera Telmatodrilus Eisen and Alezandrovia Hrabe as part of the

Tubificinae. Brinkhurst (1971) established another subfamily,

Phallodrilinae, for the group of marine forms, and removed the

previously known marine genus Clitellio Say. from the Tubificinae

and established it as an independent subfamily, Clitellinae. By

the way, this genus was again returned to the subfamily Tubifici-

nae in 1979 (Brinkhurst, Baker, 1979). Furthermore, the same

author (Brinkhurst, 1971) placed Bothrioneurum Stolc in the sub-

family Rhyacodrilinae, thus abolishing the monotypic subfamily

Bothrioneurinae, and later (Brinkhurst, 1981) assigned the genus

Branchiura Bedd. to the same subfamily, eliminating the second

monotypic subfamily Branchiurinae. Finally, Erseus (1982) estab-

lished the new subfamily Limnodriloidinae on the basis of several

marine genera.

The size of the subfamilies did not remain fixed. The compo-

sition of many of the subfamilies (especially the marine ones) was

expanded with redescribed forms, and altered as a result of the

(195) shifting and revision of already known genera. Various characters,

but mainly the structural characteristics of the generative organs

were used as criteria for substantiating the subfamilies. For

example, in his brief diagnosis of the subfamilies Telmatodrilinae

and Tubificinae, Eisen (1885) indicates several prostatic glands

on the atrium and a distinctive penial structure for the first

subfamily, and one prostatic gland on the atrium and a different

penial structure for the second subfamily, noting these as the

most characteristic features of these subfamilies. To differen-

tiate Ilyodrilinae (now Rhyacodrilinae) and Tubificinae, Stolc

(1885) also includes such characteristics as the form of the pro-

static gland, the presence or absence of spermatophores, Naidomorpha

(Rhyacodrilinae)-type or Tùbificinag-type development of the

eggs , the presence or absence of genital setae, as well as the

form of the penial apparatus.

Hrabe (1963, 1966, 1967, 1979), who has made a great contribu-

tion to the basic concepts of oligochaete morphology and systematics,

also proceeds from the features characterizing the genital organs,

and in addition uses several somatic characters. He believes that

the following characters are the most important for the differen-

tiation of tubificids into subfamilies: 1) the form (diffuse or

separate) and number (one or several) of prostatic glands; 2) the

form of the male sexual cells (spermatozoa or spermatozeugmata);

3) the location of the spermathecal pores (at the anterior dissepi-

ment or in the middle of the segment); 4) egg formation (maturation

of the eggs in partial ovaries or singly); 5) the presence or

absence of a true penis; 6) the form of the nephridia (with a

-4-

sacciform postseptale of the enchytraeid type or in the form of

unfused loops); 7) the presence or absence of large coelomocytes

in the body cavity; 8) the presence or absence of dorsal pores,

and in addition to this a number of characters typical of the genera

Bothrioneurum and Branchiura when distinguishing the subfamilies

Bothrioneurinae and Branchiurinae.

Brinkhurst (1971) believes that the only important and con-

venient characters are Nos. 1, 2 and 7 which, in his opinion, have

the advantage that they are almost always mentioned in the descrip-

tions of tubificids, and so the family is easily classified into

groups on the basis of these characters. However, as we have

already mentioned, the author himself was not quite satisfied with

the tubificid system and continued to improve it by means of re-

arrangements. By the way, in his 1981 paper (Brinkhurst, 1981),

he used yet another Hrabe character in the diagnosis of Rhyacod-

rilinae, namely the location of the spermathecal pores.

If we take the genera Branchiura and Bathrioneurum as part of

the Rhyacodrilinae (which we believe to be correct), then the 4th,

6th, 7th and 8th characters in their description (partial ovaries

sacciform postseptale of nephridia, large coelomocytes and dorsal

pores present) are typical only of this subfamily, and not neces-

sarily for the whole series of forms. For example, there are no

partial ovaries in Bothrioneurum, Rhizodrilus and Paranadrilus,

and for some genera the nature of oocyte formation is unknown;

there are no sacciform postseptale of the nephridia in Rhyacodri-

loides, Rhizodrilus and Branchiura, and for some genera there are

no data on this; dorsal pores are noted only in Eeirodrilus, Rhxa-

codrilus and Bothrioneurum, while the rest of the genera either

have none or nothing is known about them; coelomocytes are known

for 18 genera, are sometimes missing in one genus i and are never

found in three genera. Thus, the listed characteristics merely

emphasize the distinctiveness of the Rhyacodrilinae.

As to the form of the penis (5th characteristic), it may be

found, within the same subfamily, in the form of a pseudopenis

or a true penis (in several subfamilies). Among the Phallodrili-

nae, for example, 6 genera have simple pores, 4 genera pseudo-

penises, and 5 genera penises (2 of the latter with chitinous

cover as in the Tubificinae); among the Telmatodrilinae, one

genus has a penis, and one genus a pseudopenis; among the Rhya-

codrilinae, 3 genera have a penis, 8 genera pores, and 10 genera

a pseudopenis.

A study of the vast and diverse morphological material on

tubificids published recently has confirmed the importance of the

following characters to the classification of this family into

subfamilies: 1) the presence (as the most universal character for

Rhyacodrilinae) or absence of large coelomocytes in the body cavity;

2) the mode of attachment, the form and the number of prostatic

glands on each male gonoduct; 3) the location of the spermathecal

pores; 4) the nature of the male sexual cells. On the basis of

these characters, the following determination table has been drawn

up for the subfamilies of the Tubificidae.

1 (2) Large (sometimes comparatively small, but numerous)

coelomocytes usually present in body cavity (if not,

then prostatic gland diffuse) Rhyacodrilinae

2 (1) No large coelomocytes.

3 (8) Spermathecal apertures near anterior (as an exception,

posterior) dissepiment (in individual rare cases, in

middle of segment).

(19 (196)

4 (5) Each male gonoduct with more than two prostatic

glands Telmatodrilinae

5 (4) Each male gonoduct with not more than two prostatic

glands.

6 (7) If present, prostatic glands compact, often stalked,

less commonly with wide base Phallodrilinae

7 (6) One prostatic gland stalked, second one diffuse or

absent Nootkadrilinae subfam. n.

8 (3) Spermathecal apertures approximately in middle of segment

(rarely offset anteriorly or posteriorly)

9(10) One prostatic gland broadly attached to prostatic pad of

atrium or its diverticulum Limnodriloidinae

10(9) One prostatic gland attached to atrium by means of a

stalk.

11(12) Spermatozoa present Aulodrilinae

12(11) Spermatozeugmata present Tubificinae

Along with the characters indicated in the table, we shall

note a number of other important features, as well as the size

of the subfamily.

RHYACODRILINAE HRABE, 1963

The somatic setae differ; they may be forked, single—toothed,

three—toothed, sometimes with a ligament connecting the lower

tooth to the shaft of the seta, fan—shaped or hairlike. Genital

setae are often present. Dorsal setae are sometimes present.

The nephridia often have a sacciform postseptale. The male gono-

ducts have a diffuse (individual cells or groups of cells) prosta-

tic gland. Copulatory glands are sometimes present; they are often

histologically similar to the prostatic tissue (e.g. in Macluari-

drilus). Sometimes there are no prostatic cells. The male genital simple_

apparatus consists of pores or pseudopenises (sometimes in coelomic

sacs); true penises are rare. The spermathecae are paired, rarely

they unpaired', \--- open in segment X near the anterior (very rarely poster-

ior) dissepiment, sometimes in the posterior part of segment IX,

and sometimes they are absent. Heterodriloides has 2 pairs of

spermathecae in segments XI and XII. The spermatozo constitute

a free mass; they are very rarely found in organized clusters.

Morulae of oocytes form during oogenesis in some genera.

This subfamily includes the following genera: Epirodrilus Hr.,

Rhyacodrilus Bret., Svetlovia Cekanovskaja, Paranadrilus Gavrilov,

Protuberodrilus Giani et Martinez-Ansemil, Branchiura Beddard,

Bothrioneurum Stolc, Peristodrilus Brinkhurst, Rhizodrilus Smith,

MonouleEhorus Lev., Rhyacodriloides Cakanovskaja, Jolydrilus

Marcus, Ainudrilus Finogenova (=Vadicola Baker), LymEhachaeta

Snimschikova, Heterodrilus Pierant., Heterodriloides Erseus,

(197

Gieredrilus Erseus, Macauaridrilus Jamieson and Heronidrilus

Erseus et Jam.

Coelomocytes (the most characteristic feature of this sub-

family, which is not noted in any of the other subfamilies) are

not found in Rhyacodriloides, Branchiura or LymEhachaeta, and are

not always noted in Rhizodrilus Eacificus. It may be that they

are confined to certain stages of the life cycle, and so they may

be absent during certain periods.

The genus Svetlovia was established by Chekanovskaya (1975)

for the abyssal Baikal species S. maculata. Our study of the

holotype resulted in a change and expansion of its initial de-

scription.

Svetlovia maculata Cekanovskaja

Holotype. There are two pairs of testes in segment IX, as

noted by Chekanovskaya, and another pair of smaller ones in segment

VIII. The spiral vas deferens measuring about 250 pm in length

falls apically from the side into the atrium which consists of an

oval ampulla, a narrow ejaculatory duct and a massive pseudopenis,

and not penis as indicated in the initial description (Fig, A).

The atrial ampulla has a glandular epithelium (with the exception

of the ectal end) and cilia, measures about 200 pm in length and

at most 115 pm in width, and has a muscular lining up to 7.5 pm

thick. The muscular ejaculatory duct is about 40 pm long and

17.5-30 pm wide, and has a very narrow lumen (2.5-5 pm). The

pseudopenis measures 137.5 pm in length, and is 75 pm wide in its

central part. Numerous muscle fibres attach the pseudopenis (they

are partially present on the ejaculatory duct as well) to the body

wall. The penial setae are found behind the pseudopenises. This

genus is morphologically similar to ERirodrilus, but is distin-

guished from it by the differentiation of the atrial tube into

an ampulla, an ejaculatory duct and pseudopenis, as well as by the

presence of cilia inside the atrial ampulla and the uniform thick-

ness of the wall of the parts of the atrial tube.

PHALLODRILINAE BRINKHURST, 1971

The somatic setae are solely forked, sometimes with a ligament

connecting the lower tooth with the shaft of the seta; in exception-

al cases, there is sometimes a third tooth. Genital setae are

often present. Some species have no intestine. The gonoduct in

the majority of genera has 2, less commonly one prostatic gland

(sometimes absent). In some genera, the ectal prostate (pseudo-

prostate?) is attached to the base of the penis and (or) to the

walls of the penial bursae. The spermatozoa constitute a free

mass, and in exceptional cases round spermatozeugmata.

This subfamily includes the genera Phallodrilus Pierant.,

Bathydrilus Cook, Aktedrilus Kn., Adelodrilus Cook, Coralliodrilus

Erseus, Bacescuella Hr., Inanidrilus Erseus, Olavius Erseus, Jamie-

soniella Erseus, SEiridion Kn., Atlantidrilus Erseus, Uniporodrilus

Erseus, Bermudrilus Erseus, Peosidrilus Baker et Erseus and Duri-

drilus Erseus.

Despite the exceptional peculiarity of some of the features

of certain genera (unpaired atrium, spermathecae in segment XII,

the absence of an intestine, a wide vas deferens without cilia,

etc.), this is a highly compact homogeneous group in which we can

easily trace the relations between the genera. Erseus (1979)

regards the form similar to Phallodrilus as the prototype of many

of the genera.

NOOTKADRILINAE FINOGENOVA, SUBFAM. N.

The somatic setae are forked or single—toothed. Penial setae

are present. The male gonoducts consist of 4 parts which differ

in width, the presence or absence of a ciliated epithelium and the

thickness of the muscular layer; they end in.pseudopenises. There

is a stalked prostatic gland at the ectal end of each gonoduct,

and a diffuse one at the ental end. The diffuse prostate may be

absent, in which case the epithelium of the ental part is thickened

and glandular. The anterior part of the male gonoduct is found in

segment X. The spermatozoa constitute a free mass (Fig, B—C).

The subfamily is composed of Nootkadrilus Baker and Discordi-

Erostatus Baker.

Baker (1982), who described both genera, assigned them to the

subfamily Phallodrilinae on the basis of the presence of a stalked

-10—

posterior prostatic gland, the form of the spermathecae, male

gonoduct and penial setae, and the absence of large coelomocytes.

He believes that Nootkadrilus bears the greatest similarity to

Coralliodrilus Erseus, the atrium of which has no prostatic glands,

and to Inanidrilus Erseus which has only a posterior prostate,

two types of penial setae and muscular spermathecae. In our opinion,

the presence of a posterior stalked prostate is the only significant

feature (apart from the absence of coelomocytes, which is also

characteristic of other subfamilies with the exception of Rhyacod-

rilinae) common to the Phallodrilinae. A glandular atrial epithe-

lium in the absence of prostatic glands is also known for other

tubificids (e.g. Svetlovia and EEirodrilus of the Rhyacodrilinae),

as is the muscular efferent part of the spermathecae (Svetlovia). (19 (.

Furthermore, it is difficult to look for some relationship between

Nootkadrilus, which probably lost the vas deferens and anterior

prostatic gland a second time, and Coralliodrilus, which also lost

both prostates of the original form but does have a fairly long

vas deferens. It is especially difficult to explain the appearance

in the same genus of significant features (glandular epithelium

of the gonoduct with no prostate entally and a stalked prostate

ectally) of two fairly advanced, and not primitive phallodriline

genera such as Coralliodrilus and Inanidrilus. There is no doubt

that Nootkadrilus and DiscordiErostatus are morphologically similar,

the entai parts of their atria are apparently homologous, and so

it is even more difficult to pass from the Phallodrilinae to

Discordiprostatus, a form with one of the most primitive types of

prostates (diffuse) among the Oligochaeta. In any case, it is

-11-

very unlikely that such a regression of this important structure

to a more primitive state could have occurred within a single sub-

family in the absence of other specific phallodriline features.

Structural details of Svetlovia maculata (A), Nootkadrilus hamatus (B) and Discordiprostatus longisetosus (C): A - terminal part of male gonoduct (saggital section); B-C - after Baker, 1982; p.p. - posterior prostatic gland, c.p. - copulatory pouch, m - muscles, m.g. - median parts of male gonoduct, o - male genital aperture, p - pseudopenis, a.p. - anterior prostatic gland, p.s. - penial setae, s - spermatheca, s.f. - sperm funnel, v.d. - vas deferens, ec.g. - ectal part of male gonoduct, en.g. - entai part of male gonoduct

As to the topography of the male gonoduct, we can find no

analogy with the mentioned genera amidst the extensive material

that has so far been accumulated on the Phallodrilinae. Baker

himself writes that the atria of Nootkadrilus are the most complex,

and that DiscordiRrostatus are unique among the Tubificidae. All

this has prompted me to introduce a new subfamily for these genera.

I would like to note another of their rare morphological characters,

-12-

i.e. the partial penetration of the male gonoduct from segment XI

into segment X. This characteristic is known in Rhizodrilus and

Bothrioneurum in the subfamily Rhyacodrilinae.

TELMATODRILINAE EISEN, 1879

The somàtic setae are of different types, i.e. forked, single-

toothed, scopiform, hairlike. Genital setae are often present.

The prostatic glands are compact, stalked, and may number up to 16

on each atrium. The atria end in a penis or pseudopenis. Spermato-

zoa or vermiform spermatozeugmata are present. This subfamily

consists of the genera Alexandrovia Hr. and Telmatodrilus Eisen.

Holmquist (1974) proposes that Telmatodrilus be broken down into

two genera.

LIMNODRILOIDINAE ERSEUS, 1982

The somatic setae are exclusively forked, occasionally with

a ligament between the lower tooth and the shaft of the seta.

Genital setae are rarely encountered. The oesophagus in segment

IX is modified, either inflated and glandular, or with a pair of

lateroventral diverticula directed anteriorly toward dissepiment

8/9. The prostatic gland is attached to the atrial ampulla,

usually by its wide base, but in some cases with the help of a

fairly narrow sphincter, and is connected with the modified inter-

nal epithelium of the ampulla, which forms a characteristic lining

with a granular structure (prostatic pad), or with the atrial

diverticulum which also has a granular lining. The atria end

mostly in a pseudopenis or pores, very rarely in a true penis.

The spermatozoa constitute a free mass, clusters, or in rare cases

simple spermatozeugmata.

-13-

The subfamily includes Limnodriloides Pierant., Marcusaedri-

lus Righi et Kanner, Tectidrilus Erseus, Thalassodrilides Brink.

et Baker, Smithsonidrilus Brinkhurst, Kaketio Righi et Kanner,

Parakaketio Erseus and Doliodrilus Erseus.

AULODRILINAE HRABE, 1967

The somatic setae are forked, oar-shaped, fan-shaped, hairlike.

Genital setae are rare. There are 1-3 pairs of testes and one pair

of ovaries. The prostatic gland has a narrow or wide stalk. The

atria end in penises which often have a chitinous cover. One,

rarely two pairs of spermathecae are present. The species of Aulo-

drilus reproduce mainly architomously, less commonly sexually.

Their genital organs are usually displaced anteriorly by several

segments. At the posterior end of the body we sometimes find an

unsegmented zone which functions as gills.

This subfamily includes the genera Aulodrilus Bretscher,

PeiEsidrilus Timm, Neoaulodrilus Giani, Martinez-Ansemil, Moubayed

et Dia and Sketodrilus Karaman. The last two genera are very simi-

lar, so perhaps they should be combined in a single genus.

TUBIFICINAE EISEN, 1879

These are the higher tubificids. The somatic setae are forked,

single-toothed, fan-shaped, hairlike. Genital setae are usually

scarce, or often absent. The compact stalked prostatic gland

opens into the atrium medially or in the entai half, and is some-

times absent. The atria end in penises enclosed in penial bursae.

The penises may have a chitinous cover or tube. Architomous re-

production is encountered.

The subfamily includes the genera Tubifex Lamarck, Limnodrilus

Clap., Isochaetides Hr., Potamothrix Vejd. et Mrazek, Psammorxctides

-14-

Vejd., Ilxodrilus Eisen, Clitellio Say., Clitelloides Finogenova,

Tubificoides Last., SEirosEerma Eisen, Peloscolex Leidy, Baikalo-

drilus Holmquist, Haber Holmquist, Antipodrilus Brinkhurst, Lxco-

drilus Grube, Lxcodrilides Hr., TubiEenifer Semernoi, Arctodrilus

Brinkh., SEirosEermoides Dumnicka, Krenedrilus Dumnicka and Vari- -

chaetadrilus Brinkh. (=Ilxodriloides Sok.).

Thus, the Tubificidae family is known to consist of seven sub-

families which together include 72 genera. Phallodrilinae, Limno-

driloidinae and Nootkadrilinae include only marine or brackish-

water genera (with individual species that inhabit fresh waters

as well); Aulodrilinae and Telmatodrilinae include only freshwater

species, while Rhyacodrilinae and Tubificinae contain both fresh-

water and marine forms, as well as several genera with mixed ranges.

At the present time, we know of 35 marine and 33 freshwater genera

of tubificids, as well as 4 genera common to both sea and fresh

waters.

It is still difficult to speak of the phylogenetic relations

of the subfamilies, but there is no doubt that the habit of the

Rhyacodrilinae is the most primitive. Despite the known features

of specialization, here we come up against the greatest number of

primitive features of organization, including the absence of a

separate prostatic gland, the presence of spermatozoa, weak dif-

ferentiation of the penial appartus, metamerism of the gonads

(e.g. Svetlovia, Bothrioneurum), the ectal entry of the vas deferens

into the atrium, probably the location of the spermathecal aper-

tures in the anterior part of segment X, the presence of large

coelomocytes and the formation of morulae of oocytes. Another

distinctive feature of the Rhyacodrilinae is morphological

-15-

tar

flexibility which is seen particularly in the extreme diversity

of the forms and structures of the generative apparatus.

The closest to the Rhyacodrilinae are the Phallodrilinae and

Nootkadrilinae which, despite the development of certain progres-

sive characters such as compact prostates and a complex penis

(Aktedrilus), have retained quite a number of primitive features;

both subfamilies have spermathecal pores at dissepiment 9/10, both

have spermatozoa, Nootkadrilinae have an anterior diffuse prostate, (

and Phallodrilinae have an ectal entry of the vas deferens into

the atrium, both a penis and simple pores and aberrant metamerism

of the gonads. The Limnodriloidinae, Aulodrilinae and Telmato-

drilinae are farther away from the Rhyacodrilinae, though they also

have much in common with them. Though it is narrower and compact,

the prostatic gland in Limnodriloidinae is attached quite exten-

sively to the atrium; in Telmatodrilinae, the spermathecal aper-

tures are located at the anterior dissepiment and the atrium is

pyriform; gonadal metamerism is observed in the Aulodrilinae; all

of these subfamilies have spermatozoa (in addition to spermatozoa,

vermiform spermatozeugmata are also found in the genus Alexandrovia

of the subfamily Telmatodrilinae). Finally, in the most advanced

subfamily Tubificinae, ancestral features can be found with diffi-

culty, sooner in the form of an atavism. For example, as an ex-

ception, the spermathecal apertures in individual species of the

Tubificinae are located near dissepiments 9/10 or 10/11, instead

of medially.

Analogous structures developed in a similar manner in dif-

ferent lines of both marine, and freshwater tubificids. This

applies to the differentiation of the terminal part of the male

gonoduct from simple pores to a complex penis (Phallodrilinae,

Rhyacodrilinae, Limnodriloidinae), the secondary reduction of the

prostatic gland (Rhyacodrilinae, Phallodrilinae, Tubificinae),

the concentration of spermatozoa from a disorganized mass into

organized clusters and then into spermatozeugmata (Rhyacodrilinae,

Telmatodrilinae, Phallodrilinae, Limnodriloidinae), the develop-

ment of an unpaired spermatheca or the complete disappearance of

spermathecae (Rhyacodrilinae, Phallodrilinae, Limnodriloidinae),

the formation by the integuments of a special secretion to which

particles of the substrate adhere (Telmatodrilinae, Phallodrilinae,

Limnodriloidinae, Tubificinae), the fusion of the genital apertures

into one (Rhyacodrilinae, Phallodrilinae, Limnodi:iloidinae), and

to a number of other characteristics. The nature of these changes

(the result of convergent or parallel development) is still not

always clear, despite the fact that our ideas of the structural

potentials of the Tubificidae have greatly expanded over the past

few years. For example, we learned just recently of the develop-

ment of spermathecae behind the male genital apertures in segment

XII in tubificids. A particularly large number of facts related

to new morphological characters previously unknown in tubificids

can be found in the descriptions of marine genera, which I have

noted in the diagnoses of the marine subfamilies (setae with a

ligament, diverticula on the intestine and atrium, the absence of

an intestine, etc.). Further detailed research of the freshwater

and marine fauna will definitely clarify many of the questions

pertaining to the systematics and phylogeny of this group of

animals.

— 1 7 —

REFERENCES

1. Chekanovskaya 0.V., 1972. Present state of the systematics of aquatic oligochaetes (fam. Tubificidae). IN: Aquatic Oligo- chaeta. Moscow, "Nauka" Publishers, pp. 3-32; —1975. New tubifi-cids (Oligochaeta, Tubificidae) from the abyssal zone of Lake Baikal. Tr. Limnol. in—ta AN SSSR, 18, 38, 112-130.

8d. Hrabe S., 1979. Aquatic Oligochaeta of Czechoslovakia. Acta Univ. Carolinae, Biologica, 1-2, 1-167.

9. Stolc A., 1885. Preliminary report on Ilyodrilus coccineus. Contribution to the knowledge of the Tubificidae. Zool. Anz., 8, 638-644, 656-662; — 1888. Monograph on Czech Tubificidae. Abh. Böhm. Ces. Wiss., ser. 7, 2, 11, 1-43.

Zoological Institute of the USSR Academy of Sciences (Leningrad)

Received on 16 Nov 1984

gem-wen:an O. B., 1972. CoapeNteunoe commute cucTemaTuRn 1301111131X oonroxeT (cem. Tubificidae).— B K11.: Bomme nia.lomenincosbie em'. M.: Hayka, 3-32.-1973. Hoame Ty6iiduittit1b1 (Oligochaeta, Tubificidae) 113 a6ucca.rut oaepa Bailka.n.— H0.1. 11H-Ta AH CCCP, 18, 38, 112-130.

2.. Baker H. R., 1982. Two new Phallodrilinae genera of marine Oligochaeta (Annelida: Tu- bilicidae) from the Pacific Northeast.— Caned. J. Zool., 60, 10, 2487-2500.

3. Brinkhurst R. 0., 1971. Tubificidae.— In: Aquatic Oligochaeta of the World. Edinburgh: ,Oliver and Boyd, 1-860.— 1981. A revision of the genus Afonopylephorus and rede-finition of the subfamilies Rhyacodrilinae and Branchiurinae (Taificidae: Oligo-chaeta).— Canad. J. Zool., 59, 939-965.

L, Brinkhurst R. 0., Baker II. R., 1979. A review of the marine Tubificidae (Oligochaeta) of North America. — Canad. J. Zool., 57, 1553- 1569.

5, Eisen G., 1879, Preliminary report on genera and species of Tubificidae.— Bih. Kongl. Svenska Vetensk. Akad. llandl., 5, 16, 3-26.-1885. Oligochaetological researches. Comm. for 1883.—U. S. Comm. Fish. and Fisheries, pt XI, 879-964.

b. Erseus Ch., 1979. Inonidrilus bulbosus gen. et sp. n., a marine tubificid (Oligochaetal from Florida, U. S. A.— Zool. Scr., 8, 209-210.— 1980. New species of Phallodrilus.

(011gochaeta, Tubificidae) from the Arctic-Deep Sea and Norwegian Fjords.— Sar-sia, 65, 1, 57-60.— 1981. Taxomomic revision of the marine genus Heterodrilus Pierant. (Oligochaeta, Tubificidae).— Zool. Scr., 10, 111-132.— 1982. Taxonomie re-vision of the marine genus Limnodriloides (Oligochaeta: Tubificidae).— Verh. na-turwiss. Ver. Hamburg, (NF), 25, 207-277.

'7. Holmquist Ch., 1974. On Alexandrovin onegensis Hrabé from Alaska, with a revision of the Telmatodrilinae (Oligochaeta, Tubificidae).— Zool. Jb. Syst., 101, 249-268.

$, Hrabé S., 1963. On Rhyacodrilus lindbergi n. sp., a new cavernicolus species of the family Tubificidae (Oligochaeta) from Portugal. — Bol. Soc. Portug. Cienc. Nat., 10, 52—

56.— 1966. New or insufficiently known species of the family Tubificidae.— Publ. Fac. Se Univ. J. E. Purkynè, Brno, 470, 57-77.— 1967. Two new species of the family Tubificidae from the Black Sea, with remarks about various species of the subfam. Tubificinae.— Ibidem, 485, 311-365.— 9. Vodni màloMétinatci (Oligochaeta) Ceskoslovenska.— Acta Univ. Carolinae, jo ogica. 1-2, 1-167.

9, stoic A., 1885. Vorlaufiger Bericht über Ilyodrilus coccineus. Ein Beitrag zur Kenntnis der Tubificiden.— Zool. Anz., 8, 638-644, 656-662.- 1888. Monografie éeskYch .Tu-bificidù.— Abh. Bôhm. Ges. Wiss. ser. 7, 2, 11, 1-43.

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ON DIAGNOSTIC CHARACTERS OF SUBFAMILIES IN THE FAMILY TUBIFICIDAE (OLIGOCHAETA)

N. P. F1NOGENOVA

Zoological Institute, USSR Academy of Sciences (Leningrad)

Summary

The most important characters v.rhich have been used in the key of subfamilies in the family Tubificidae are the following: presence or absence of large coelomocytes in the body cavity; method of attachment; shape and number of the prostatic glands on each male gonoduct; localization of the seminal receptacle pores; type of the male genital cells. A new subfamily, the Nootkadrilinae subfam. n., has been created for two 'marine genera, Nootkadrilus Baker and Discordiprostatus Baker. A list of genera in all seven

.subiamilies is presented. The description of Svetlovia maculata Cekanovskaja is corrected and supplemented with new details on the basis of reinvestigation of the holotype of this species.