taxonID	type	description	language	source
E329D91AFFD4FFBD656BFA8CFCC8FD1A.taxon	discussion	The Aquamegadrili, named by Jamieson (1988) for aquatic crassiclitellates (see Introduction for geographic distribution), consist of the Sparganophilidae, Biwadrilidae, Almidae (mostly warm tropics, including Criodrilus, Mediterranean region, etc.) and Lutodrilidae (Southern Neartic). In the likelihood ratio tests the 28 S data-set is inconsistent with monophyly of the Aquamegadrili but this is entirely due to the inclusion of Komarekiona as sister-taxon of Sparganophilus. The 28 S data are inconsistent with partition of crassiclitellates into Aquamegadrili and Terrimegadrili (see families listed in Introduction); instead the representatives (Sparganophilus, Criodrilus and Lutodrilus) of the original aquamegadrile taxa, with Komarekiona, lie within a paraphyletic Terrimegadrili (Figs 2; 3). The Lumbricoidea is incompatible with the 28 S data but principally because the glossoscolecid sp. never groups with the remainder in the best trees but groups equivocally with Eudrilus. However the unity of the majority of the remaining Lumbricoidea will necessarily require more sampling.	en	Jamieson, Barrie G. M., Tillier, Simon, Tillier, Annie, Justine, Jean-Lou, Ling, Edmund, James, Sam, Mcdonald, Keith, Hugall, Andrew F. (2002): Phylogeny of the Megascolecidae and Crassiclitellata (Annelida, Oligochaeta): combined versus partitioned analysis using nuclear (28 S) and mitochondrial (12 S, 16 S) rDNA. Zoosystema 24 (4): 707-734, DOI: 10.5281/zenodo.4524860
E329D91AFFD4FFBD656BFC0CFE34FA9B.taxon	discussion	That the Crassiclitellata of Jamieson (1988), named for families in which the clitellum is multilayered, is a monophylum is confirmed in the molecular analyses. The MP majority rule consensus tree for 549 bp of 28 S only (Fig. 2) gives 100 % bootstrap support for the Crassiclitellata versus outgroup taxa. Maximum likelihood analysis differs little from a MP bootstrap tree. Eudrilus again associates with the glossoscolecid Gen. sp. but not with Microchaetus.	en	Jamieson, Barrie G. M., Tillier, Simon, Tillier, Annie, Justine, Jean-Lou, Ling, Edmund, James, Sam, Mcdonald, Keith, Hugall, Andrew F. (2002): Phylogeny of the Megascolecidae and Crassiclitellata (Annelida, Oligochaeta): combined versus partitioned analysis using nuclear (28 S) and mitochondrial (12 S, 16 S) rDNA. Zoosystema 24 (4): 707-734, DOI: 10.5281/zenodo.4524860
E329D91AFFD4FFBD67BEFD0CFBEAFABA.taxon	description	The family Eudrilidae has formerly been associated with the Megascolecidae and Ocnerodrilidae in a superfamily Megascolecoidea (Jamieson 1978) or in a separate superfamily Eudriloidea as the sistergroup of the Lumbricoidea + Megascolecoidea s. s., in the morphocladistic analysis (Jamieson 1988). In the present analysis, Eudrilus always has the glossoscolecid Gen. sp. (a lumbricoid sensu Jamieson 1978, 1988) as its sister-taxon, with moderate BS support of 63 - 84 % (MP, ML, Figs 2; 3). The other glossoscolecid, Pontoscolex, may or may not link with these. There is no molecular support here for regarding eudrilids as the unique sister-group of the Ocnerodrilidae + Megascolecidae assemblage. Omodeo (2000) derived eudrilids independently (from alluroidids), thus also noting their distinctness but that origin goes contrary to the present confirmation of the monophyletic nature of the Crassiclitellata.	en	Jamieson, Barrie G. M., Tillier, Simon, Tillier, Annie, Justine, Jean-Lou, Ling, Edmund, James, Sam, Mcdonald, Keith, Hugall, Andrew F. (2002): Phylogeny of the Megascolecidae and Crassiclitellata (Annelida, Oligochaeta): combined versus partitioned analysis using nuclear (28 S) and mitochondrial (12 S, 16 S) rDNA. Zoosystema 24 (4): 707-734, DOI: 10.5281/zenodo.4524860
E329D91AFFD4FFBE67BEFAECFED9FCBA.taxon	discussion	In the morphocladistic analysis (Jamieson 1988), the superfamily Megascolecoidea contained only the Megascolecidae, with the two subfamilies Acanthodrilinae and Megascolecinae, and excluded the Ocnerodrilidae and Eudrilidae. However, all three of these families had been tentatively included in the superfamily Megascolecoidea in Jamieson (1978, 1980). Recognition by Lee (1959), Jamieson (1971 a-c, 1978, 1980), and Sims (1966, 1967) of an Acanthodrilinae + Megascolecinae assemblage together with the Ocnerodril (- inae) (- idae) is endorsed in the present molecular analyses. Monophyly of the ocnerodrile exemplar Eukerria saltensis with the Megascolecidae (Acanthodrilinae + Megascolecinae) is supported in all cladistic analyses that have additional outgroups. This relationship of the Ocnerodrilidae with the Megascolecidae has> 90 % BS support in MP trees (see Figs 2; 4, also Jamieson 2000). In view of the sister-group relationship of the Eukerria saltensis with the Megascolecidae s. l., it appears that the family Ocnerodrilidae may be included in the Megascolecoidea, rather than having a suprafamilial rank of its own, but this needs wider sampling to test monophyly, and the relationships of the two ocnerodrile tribes Ocnerodrilini and Malabarini.	en	Jamieson, Barrie G. M., Tillier, Simon, Tillier, Annie, Justine, Jean-Lou, Ling, Edmund, James, Sam, Mcdonald, Keith, Hugall, Andrew F. (2002): Phylogeny of the Megascolecidae and Crassiclitellata (Annelida, Oligochaeta): combined versus partitioned analysis using nuclear (28 S) and mitochondrial (12 S, 16 S) rDNA. Zoosystema 24 (4): 707-734, DOI: 10.5281/zenodo.4524860
E329D91AFFD7FFBE67BEFCCCFB72F93A.taxon	discussion	The definition of the subfamily Acanthodrilinae sensu Jamieson (1971 a) differs fundamentally from that of Gates (1959, 1972). Prostates are not only tubular but may also (rarely) be racemose and nephridia are not only holonephridia but may also be meronephridia. Unlike many acanthodrilids of Gates, the prostates usually do not discharge on segment 18 (doing so in Rhododrilus), typically opening on segments 17 and 19, as in the type-genus. Megascolecinae with homeotic displacement of male pores may correspond with this definition but show their affinities with megascolecines in other respects. Posterior nephridia in Acanthodrilinae sensu Jamieson (1971 a) lacked the median funnel diagnostic of the Dichogastrini within the Megascolecinae sensu Jamieson. The alimentary and vascular systems differed from those of the Ocnerodrilinae in some of which, as in Eukerria Michaelsen, 1935, the male and prostate pores have the acanthodrilin arrangement. It has been shown in the molecular analysis that dichogastrins with acanthodrilin male pores must be transferred to the Acanthodrilinae and that the dichogastrin nephridial condition has arisen more than once (see below).	en	Jamieson, Barrie G. M., Tillier, Simon, Tillier, Annie, Justine, Jean-Lou, Ling, Edmund, James, Sam, Mcdonald, Keith, Hugall, Andrew F. (2002): Phylogeny of the Megascolecidae and Crassiclitellata (Annelida, Oligochaeta): combined versus partitioned analysis using nuclear (28 S) and mitochondrial (12 S, 16 S) rDNA. Zoosystema 24 (4): 707-734, DOI: 10.5281/zenodo.4524860
E329D91AFFD7FFBE656BFCECFDA5FA7A.taxon	discussion	With regard to previous classifications of the Megascolecidae the following conclusions can be drawn from the present molecular analyses. The widely used system for internal classification of the Megascolecidae of Gates (1959, 1972) cannot be sustained, as already argued by Lee (1959, 1970) and Jamieson (1971 a-c). Diagnosis of Megascolecidae in the restricted sense of Gates (1972) by the possession of racemose prostates (with holo- or meronephridia) is not supported in any MP trees, with multiple evolution of racemose and tubular prostates implied by the molecular phylogenies (Figs 3; 4). However, bootstrap values are low in the relevant section of the trees and, as no species with tubular prostates has a high bootstrap linkage with a species with racemose prostates [notwithstanding the Terrisswalkerius athertonensis-Didymogaster clade]. Likelihood ratio tests are equivocal on this but we suggest this is more a reflection on the SH test as overly conservative.	en	Jamieson, Barrie G. M., Tillier, Simon, Tillier, Annie, Justine, Jean-Lou, Ling, Edmund, James, Sam, Mcdonald, Keith, Hugall, Andrew F. (2002): Phylogeny of the Megascolecidae and Crassiclitellata (Annelida, Oligochaeta): combined versus partitioned analysis using nuclear (28 S) and mitochondrial (12 S, 16 S) rDNA. Zoosystema 24 (4): 707-734, DOI: 10.5281/zenodo.4524860
E329D91AFFD7FFBE656BF9ACFC23FCDA.taxon	discussion	The Octochaetidae were defined by Gates (1972) and, as the Octochaetinae, by Gates (1959) and Sims (1967), as all species with tubular prostates and more than one pair of nephridia per segment (meronephridia). Gates (1959, 1972) diagnosed the Acanthodril (- inae), (- idae), as having tubular prostates and a pair of nephridia (holonephridia) per segment. The relationship of the meronephric Neodiplotrem a with the holonephric Diplotrema in all trees (either 28 S, mtDNA or combined) argues against recognition of the Octochaet (- idae), (inae), and against the Acanthodril (- inae), (- idae) of Gates and of Sims. The Neodiplotrema + Diplotrema clade fully endorses the conclusion by Lee (1959, 1970) that phylogenetic pairs of holonephric with meronephric species of Acanthodrilinae are recognizable. We will now further consider subdivision of the Megascolecidae into the subfamilies Acanthodrilinae and Megascolecinae and division of the Megascolecinae into the tribes Perionychini, Dichogastrini, and Megascolecini in the classification of Jamieson (1971 a-c).	en	Jamieson, Barrie G. M., Tillier, Simon, Tillier, Annie, Justine, Jean-Lou, Ling, Edmund, James, Sam, Mcdonald, Keith, Hugall, Andrew F. (2002): Phylogeny of the Megascolecidae and Crassiclitellata (Annelida, Oligochaeta): combined versus partitioned analysis using nuclear (28 S) and mitochondrial (12 S, 16 S) rDNA. Zoosystema 24 (4): 707-734, DOI: 10.5281/zenodo.4524860
E329D91AFFD6FFBF656BFB2CFC9AFB3A.taxon	discussion	The Dichogastrini were defined by presence of a single stomate meronephridium median to astomate micromeronephridia on each side in caudal segments, in the absence of posterior enteronephry (Jamieson 1971 a). It has, however, repeatedly been questioned (e. g., Jamieson 1978, 1981; Dyne 1984) that dichogastrins with acanthodrilin male pores (here represented by Dichogaster) are monophyletic with those with megascolecin pores (e. g., Digaster). The present analyses relegate “ acanthodrilin dichogastrins ” (Dichogaster and Neodiplotrema) to the Acanthodrilinae and “ megascolecin dichogastrins ” (albeit represented only by Digaster and Didymogaster) to the Megascolecinae s. l. in combined and separate analyses for both mtDNA and 28 S with overall good support (e. g., BS support of 77 % in the combined ML tree, Fig. 4). The Nearctic Diplocardia longiseta appears to lie within this “ acanthodrilin ” clade (83 % BS, Fig. 3) but the only available data (the 28 S) are not sufficient for further resolution and more taxa are required. Diplocardia Garman, 1888 has been shown by James (1990) to be closely similar in morphology to Diplotrema. The available 12 S data support the argument (Jamieson 1995) that Rhododrilus glandifera, in the Wet Tropics of Queensland, is locally derived from a precursor with the acanthodrilin arrangement of male pores (probably Diplotrema with which it has a 99 % BS value in Fig. 3) though this requires confirmation from analysis of larger numbers of sequences. R. glandifera thus appears to deserve a subgeneric rank in Diplotrema or generic rank separately from Rhododrilus, the type-locality of which is in New Zealand. All trees from combined data endorse recognition of the Acanthodrilinae for worms with acanthodrilin male pores, including acanthodrilin Dichogastrini (Dichogaster) (though there is lack of resolution for 28 S, Fig. 2) but not those ocnerodriles (Eukerria) with acanthodrilin or other male terminalia. The ocnerodriles (albeit represented only by Eukerria) are phylogenetically distinct in the present study and are well-defined morphologically.	en	Jamieson, Barrie G. M., Tillier, Simon, Tillier, Annie, Justine, Jean-Lou, Ling, Edmund, James, Sam, Mcdonald, Keith, Hugall, Andrew F. (2002): Phylogeny of the Megascolecidae and Crassiclitellata (Annelida, Oligochaeta): combined versus partitioned analysis using nuclear (28 S) and mitochondrial (12 S, 16 S) rDNA. Zoosystema 24 (4): 707-734, DOI: 10.5281/zenodo.4524860
E329D91AFFD6FFB067BEFB6CFE18FDDA.taxon	discussion	The third tribe of the Megascolecidae, the Megascolecini, was defined by having male and prostate pores coincident on segment 18 (rarely segment 17), and meronephry in which a median stomate nephridium, if present, differed from those of dichogastrins in opening into the intestine (enteronephry). Prostates were racemose, tubular or tubuloracemose (Jamieson 1971 a-c). In contrast, Gates (1959, 1972) attributed only worms with racemose prostates, irrespective of nephridial types, to his restricted Megascolecidae. Resolution of the Megascolecini was not an aim of the present work and as few representatives have been included (Amynthas Kinberg, 1867, Begemius Easton, 1982, Propheretima Jamieson, 1995, and Spenceriella Michaelsen, 1907) results must be regarded with caution. However, none of the analyses supports retention of the Megascolecini as defined by Jamieson (1971 a-c). It is to be expected that the criteria of meronephry with enteronephry may have evolved more than once. A core of megascolecin genera, including among others Begemius, and Amynthas, is suspected to be monophyletic however (Jamieson 1981), though it no longer appears that Spenceriella is as close to the pheretimoids as previously argued.	en	Jamieson, Barrie G. M., Tillier, Simon, Tillier, Annie, Justine, Jean-Lou, Ling, Edmund, James, Sam, Mcdonald, Keith, Hugall, Andrew F. (2002): Phylogeny of the Megascolecidae and Crassiclitellata (Annelida, Oligochaeta): combined versus partitioned analysis using nuclear (28 S) and mitochondrial (12 S, 16 S) rDNA. Zoosystema 24 (4): 707-734, DOI: 10.5281/zenodo.4524860
E329D91AFFD6FFBF656BFF4DFEFAFBFA.taxon	discussion	The tribe Perionychini (defined by megascolecin male pores and holonephridia, irrespective of prostate type) is represented here by five genera. There is no convincing support in any analysis for a unified or monophyletic Perionychini although there is evidence that some of these holonephric megascolecines are closely related. There is, however, much instability in perionychin relationships, reflected in low BS values and lack of consensus (Figs 3 - 5). In all analyses the “ perionychin ” (acanthodrile sensu Gates) Pontodrilus diverges at or near the base of the Megascolecinae s. l., and while exclusion from the Acanthodrilinae is upheld, its uncertain placement reflects its enigmatic affinities on morphological and ecological grounds. Pontodrilus is highly unusual among crassiclitellates in being euryhaline. It may be suspected of having had a long independent evolution. The tribe Perionychini, although a convenient taxonomic grouping, is thus a para- or polyphyletic assemblage in the molecular analyses. This has previously been suspected (Jamieson 1988) as the Perionychini is recognized on the basis of the symplesiomorphic possession of holonephridia (a condition seen throughout the Oligochaeta, whereas meronephry is virtually limited to Megascolecidae). It is therefore a grade rather than a clade.	en	Jamieson, Barrie G. M., Tillier, Simon, Tillier, Annie, Justine, Jean-Lou, Ling, Edmund, James, Sam, Mcdonald, Keith, Hugall, Andrew F. (2002): Phylogeny of the Megascolecidae and Crassiclitellata (Annelida, Oligochaeta): combined versus partitioned analysis using nuclear (28 S) and mitochondrial (12 S, 16 S) rDNA. Zoosystema 24 (4): 707-734, DOI: 10.5281/zenodo.4524860
E329D91AFFD9FFB0656BFDCDFE1AF93A.taxon	discussion	Morphological and molecular support for paraphyly or polyphyly of the Oligochaeta and inclusion within this group of leeches and branchiobdellids has been outlined in the Introduction. Our 28 S data are consistent with a leech-branchiobdellid-lumbriculid clade, within the Oligochaeta, less so for a leech-branchiobdellid grouping, considering the variation among methods. Although long branch / rate acceleration / base content has posed problems for the nuclear ribosomal genes analyses, consistency with data of different characteristics (mtDNA and COI) underlines these relationships. The lumbriculid relationship was proposed on morphological grounds by Michaelsen (1928 - 1932), Brinkhurst & Nemec (1986), Brinkhurst & Gelder (1989), Brinkhurst (1999 a) and, less certainly, by Brinkhurst (1999 b) and from molecular data by Siddall & Burreson (1998), Martin (2001) and Siddall et al. (2001). If accepted, the present phylogeny would confirm the Oligochaeta as a paraphyletic group as previously mooted (Jamieson et al. 1987; Jamieson 1988; Martin 2001), being merely the non-leech, nonbranchiobdellidan clitellates (Purschke et al. 1993). Inclusion of leeches and branchiobdellids within the Oligochaeta would thus render the name Oligochaeta synonymous with Clitellata (or Euclitellata of Jamieson 1983), as proposed by Siddall et al. (2001) and in a study of molecular phylogeny of the Tubificidae (subsuming the Naididae) by Erséus et al. (2002).	en	Jamieson, Barrie G. M., Tillier, Simon, Tillier, Annie, Justine, Jean-Lou, Ling, Edmund, James, Sam, Mcdonald, Keith, Hugall, Andrew F. (2002): Phylogeny of the Megascolecidae and Crassiclitellata (Annelida, Oligochaeta): combined versus partitioned analysis using nuclear (28 S) and mitochondrial (12 S, 16 S) rDNA. Zoosystema 24 (4): 707-734, DOI: 10.5281/zenodo.4524860
