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Phylogenetic and Phenetic Systematics of The 195 PHYLOGENETICAND PHENETICSYSTEMATICS OF THE OPISTHOP0ROUSOLIGOCHAETA (ANNELIDA: CLITELLATA) B.G.M. Janieson Departnent of Zoology University of Queensland Brisbane, Australia 4067 Received September20, L977 ABSTMCT: The nethods of Hennig for deducing phylogeny have been adapted for computer and a phylogran has been constructed together with a stereo- phylogran utilizing principle coordinates, for alL farnilies of opisthopor- ous oligochaetes, that is, the Oligochaeta with the exception of the Lunbriculida and Tubificina. A phenogran based on the sane attributes conpares unfavourably with the phyLogralnsin establishing an acceptable classification., Hennigrs principle that sister-groups be given equal rank has not been followed for every group to avoid elevation of the more plesionorph, basal cLades to inacceptabl.y high ranks, the 0ligochaeta being retained as a Subclass of the class Clitellata. Three orders are recognized: the LumbricuLida and Tubificida, which were not conputed and the affinities of which require further investigation, and the Haplotaxida, computed. The Order Haplotaxida corresponds preciseLy with the Suborder Opisthopora of Michaelsen or the Sectio Diplotesticulata of Yanaguchi. Four suborders of the Haplotaxida are recognized, the Haplotaxina, Alluroidina, Monil.igastrina and Lunbricina. The Haplotaxina and Monili- gastrina retain each a single superfanily and fanily. The Alluroidina contains the superfamiJ.y All"uroidoidea with the fanilies Alluroididae and Syngenodrilidae. The Lurnbricina consists of five superfaniLies. Of these the Biwadril-oidea and Sparganophiloidea superfan. nov. contain each a single genus and fanily. The Almoidea superfam. nov. contains the fam- ilies Alnidae and Criodrilidae and, provisional.ly, the Lutodril"idae. The Lumbricoidea is constituted by the Lunbricidae (redefined to include the DiporodriLidae as a subfanily), Konarekionidae, Kynotidae, Glossos- colecidae s.strict., Microchaetidae and Hor:nogastridae(redefined to include the Ailoscolecidae as a subfaniLy, with the Hormogastrinae and Vignysinae fornerly included). The Megascolecoidea,as previousl.y, contains the Megascolecidae, Eudrilidae and Ocnerodrilidae, although the phylogram in- dicates that the Ocnerodrilidae should be placed in a separate and prin- itive superfamily of the Lurnbricina. *t( Introduction A higher, suprafanilial, classification of the 0ligochaeta has been attempted by Benham(1890), Beddard (1895, 1901), MichaeLsen (1921' 1928' L929, 1930), Meyer (1929) and Yarnaguchi (1953) and Brinkhurst and Janieson (1971). Of these, those of Michaelsen (1950), of Yamaguchi and of the latter two authors are the nost pertinent to a contenporary discussion of the phylogeny of the earthworms. A study of the phylogeny of earth- worns cannot be divorced fron consideration of that of the Haplotaxidae and the Alluroididae, the total conplex comprising the Opisthopora of Michaelsen, (1930), the Subsectio Diplotesticulata of Yanaguchi (1953), or the order Moniligastrida and the order Haplotaxida, ignoring the sub- order iubificina, in the systen of Brinkhurst and Janieson. Janieson (L976) has reduced the orders of the Oligochaeta to two, the Lumbriculida 1/r* Evolutionary Theory 32L95-233 (I"larch' 1978) The editors thank C. J. Goodnight and another referee for help in evaluating this paper. by The University of Chicago for the author O1978, r-96 B.G.M. JAMIESON and the Haplotaxida, by inclusion of the Moniligastridae as a suborder Moniligastrina in the Haplotaxida following reinterpretation of the nature of the moniligastrid testis-sac and construction of a phytogeny of the Moniligastridae according to the principLes of Hennig. Gates (e.g. L972) recognized no categories between the order Oligo- chaeta and the fanilial ranks, and therefore attenpted no groupings (other than an alphabetic listing) of the fanilies Acanthodrilidae, Allur- oididae, Criodrilidae, Eudrilidae, Glossoscolecidae (s.strict.) Haplotaxidae, Hornogastridae, Lunbricidae, Megascolecidae, Microchaetidre,, s. Gates, Moniligastridae, Ocnerodrilidae, Octochaetidae and Sparganophilidae. In L976, however, Gates recognized a suprafanilial grouping, the Lumbricoidea, more restricted in constitution than that of Brinkhurst, and Janieson, to include those fanilies known or supposed to have ovaries with a single egg string, the Lr-unbricidae, Biwadrilidae, Konarekionidae, Lutodrilidae, Spar- ganophilidae, Hornogastridae, Ailoscolecidae and Diporodrilidae, alL holarc. tic groups. IncLusion of the Biwadrilidae went contrary to a statenent of Nagaseand Nonura (L937:348) for the tfpe-species that the ovaryrrbecones divided into lobesrr and the author is unable to trace the evidence for regarding the ovaries of the Ail.oscolecidae and Diporodril.idae as single- stringed. The renaining families, with the possible exception of the Hornogastridae, are weLl knownto possess single-stringed ovaries, however. The only monothetic character of the Lurnbricoidea s. Gates is the ovariaf, one and the present study attenpts to elucidate the significance of this shared character. one of the rnajor differences between the systens proposed by Jarnieson (1971d, L976a) and the cLassification, al.beit soLely at a familial Level., of Gates lies in his treatnent of those fanilial or subfamiLial taxa tlpified by GlossoscoLen, Mienoeltaekn, and ALma. Janieson (1971d) has argued that Al,nn and other genera grouped with it should be renoved fron the Microchae- tidae, in which, on the basis largeLy of posttesticular spernathecae, they were placed by Michaelsen and Gates, and that the African genera Mtcroehae- tzn and IrLtogenia have their closest rel.atives in the S. ftnerican Glossos- colecidae such as Glossoseoles and PontoeeoLes. The S. American genera Dz.LLoerh,rcand GlyphidytLoerLue are regarded as belonging with ALma in the Alrninae (Janieson, 1971d) or ALnidae (Janieson, 1976a). CyLodtilus is grouped with ALmqthough as a separate tribe or subfanily but a special. affinity between Criodrilrc and the Lunbricidae is recognized (Janieson, 1971d). Omodeo(1956) anticipated the author in recognizing that ALma and Dtiloeyius were widely separate from MLeroeVwetrcand frorn the Glossos- colecid earthworms, although the group to which they belonged was not named. The family Microchaetidae was ?econstituted to contain CalLidy,i,Lus and GLyphidriLw" which the author considers closely reJ.ated to ALma and Dy,LLoe- ritso in addition to Mi.erocVnetus and TrttogerrLa. Kyrntus was also incLuded in the Microchaetidae in which Horrnogasterformed a subfanily, the Hornogas- trinae, Lwnbriew was considered by Onodeoto be descendedfron a form sirnilar to CrtodyiLtn, as supported by Jamieson (1971d). A further najor area of debate has been the classification of the Fanily Megascolecidaein sensu lato of Stephenson(1930). The alternative class- ifications of Stephenson(1930), Michaelsen (1933), Pickford (1937), Omodeo(1958), Lee (1959), Gates, (1959) and Sims (1966, L967) have been extensively reviewed by Sins (1966) and Janieson (1971b) and it will suf- fice here to note the chief areas of disagreenent, in the proposals of Gates (1959), Sims (1967) and Janieson (1971a). Gates (1959) recognized four distinct farnilies for the Megascolecidaes. Stephenson:the Megascolecidae (racemoseprostates), the AcanthodriLidae (with tubular prostates and neronephridia) and the OcnerodriLidae (prostates and nephridia as the L97 I ;i;b.c', rorgc.N sr|f.dOEH\, 'dqrd,rpoltrN (J rd c!.. O o o tr l') <t td o (! \o 6 (J uN -Co .d o F: O N \tor H | .l d O O '- .dr .d E 5 bO d I rA O <> d d N co € <t h tror aJ rd = . .d rd tr @ d o N I (d r F\ Od td F) d (S .. S I O \O fr !6u.l I <t (J I N E ti n dD u) t! \O o .. F- +).d I gl O O | <.. CA Cr.. 3! .drA c) ..Ln 5 S N OdaO H 6 r r4 t4 Aa r: i F- \Od d i- t 5 dor N . rl N .$ . dor o (!.' u) cr l'.., or rh < t <t :.: !A .r N ADa O.dot 'd frn <t d 6t.. d F- +J .. <'\t I F- | .6@ +, +rclt H dN O N .h.. N^ ri ,. ltr | ts F- \O E EH d DO ^ O 55 FF U)F N orr N $r4 d Z td i tr .Ot e C .. ,<F Ot (, _.Ol i F dd .N . =d ., Ot I C O C d O O . 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