Dear Colleagues,
Dr. Read has alerted us to the monumental work A Higher Level Classificatio=
n of All Living Organisms (Ruggiero et al. 2015) and to the section on Anne=
lida, including an erratum. The authors are to be admired for undertaking t=
he great amount of work necessary to compile this work. It is described as =
a =93consensus classification=94 but for Clitellata one can find no referen=
ce in the paper to the major researchers on the taxonomy and phylogeny of t=
he group whereas for the Polychaeta major authorities are appropriately cit=
ed. As a result the =93classification=94 of the =93Superorder N.N.=94 is li=
mited to what appears to be an alphabetic listing of orders. One might ques=
tion the use of the term =93order=94 for taxa which in many cases have been=
shown from morphocladistic and molecular analyses to lie at different leve=
ls in cladistic phylograms. The classification of clitellates is certainly =
not in the parlous state suggested in this work. The elimination of the ent=
ire grouping of earthworms (Crassiclitellata) as an afterthought in an erra=
tum was perhaps due to the perceived necessity to recognize only the albeit=
artificial orders. With great respect for this valuable work I respectfull=
y include below some notes on the various taxa, with particular reference t=
o the Metagynophora, and a few references in support of my comments.
Kind regards,
Barrie Jamieson
Notes on A Higher Level Classification of All Living Organisms.
Subclass Hirudinea
Correctly included in the Clitellata but not here recognized as a subclass.=
Considered to lie in a lumbriculid clade (e.g. Jamieson 2006; Sidall et al=
. 2001, 2006; James and Davidson 2012) and.
Arynchobdellida. Included in the Hirudinida by Sidall et al. (2001); Sidall=
et al. (2006).
Rhynchobdellida. Not recognized as a formal taxon by Sidall et al. (2006).
Subclass Oligochaeta. This name, if retained, should also include the =91Hi=
rudinea=92 of A Higher Level Classification of All Living Organisms. Replac=
ed by the name Clitellata (Erseus and Kallersjo 2003; Jamieson 2006).
Lumbriculida. Considered (Jamieson 2006) to be basal in a lumbriculid, bran=
chiobdellid, acanthobdellid, hirudinid clade, as supported from 18S molecul=
ar analyses (Erseus and Kallersjo 2003; James and Davidson 2012).
Acanthobdellida. Near to but more basal than the Branchiobdellida in morpho=
logical and molecular studies. Not included in the true leeches, Hirudinida=
(e.g. Sidall et al 2001; Erseus and Kallersjo 2003; Sidall et al. 2006). S=
ister taxon of the more plesiomorphic Lumbriculidae in molecular analysis (=
James and Davidson 2012).
Branchiobdellida. Valid grouping. Not, contrary to A Higher Classification.=
. , in the Oligochaeta s. strict. Sister to the Hirudinea (Jamieson 2006) o=
r within the Hirudinida-Acanthobdellida clade (James and Davidson 2012) but=
excluded from, though close to, the Hirudinida by Erseus and Kallersjo (20=
03) and Sidall et al. (2006).
Capilloventrida. Capilloventer shown to be an isolated and very basal taxon=
by Erseus and Kallersjo (2003) and James and Davidson (2012). Considered t=
he most basal oligochaetes, under the provisional name Eoclitellata, by Jam=
ieson (2006). I do not understand the argument that such a distinctive leve=
l of organization as Capilloventer displays should not be considered at lea=
st ordinal when this is supported in phylogenetic analyses but I leave this=
to =93microdile=94 experts.
Haplotaxida .Obsolete. =93The ordinal taxon Haplotaxida, as presently under=
stood, could be polyphyletic and should be provisionally abandoned in favou=
r of the monophyletic Metagynophora=94, which is supported in molecular ana=
lysis (James and Davidson 2012).
Tubificida. Contains a large number of =91microdile=92 aquatic families (se=
e Jamieson 2006), see Erseus and Kallersjo (2003) and James and Davidson (2=
012), for 18S analyses.
Enchytraeida. Grouped with the branchiobdelid-Hirudinida clade by 18S analy=
sis (James and Davidson 2012). Also considered more basal than tubificids f=
rom general morphology (Jamieson 1988) and from sperm ultrastructure (Jamie=
son 1981, 1983; Ferraguti and Erseus 1999) but sister of the earthworms (Cr=
assiclitellata) in molecular analysis by Sidall et al. (2001) and Erseus an=
d Kallersjo (2003), moniligastrids and alluroidids not considered. Sister t=
o the Metagynophora, when these taxa considered, in Jamieson (2006).
Metagynophora Jamieson 1988. Defined by presence of a segment lacking gonad=
s between the last or only testes and the ovaries. Characterizes all =91meg=
adrile=92 oligochaetes (moniligastrids through Megascolecidae). Retained as=
a superorder in A Higher Level Classification of All Living Organisms.
Moniligastrida Jamieson 1988. Plesiomorph sister of the Opisthopora. Not ye=
t subject to molecular analysis. Retained as an order in A Higher Level Cla=
ssification of All Living Organisms.
Opisthopora Michaelsen 1932, Emend Jamieson 1988). Defined by movement of m=
ale pores from the plesiomorphic location in the segment immediately behind=
the testes to a more posterior location. Included as an order in A Higher =
Level Classification of All Living Organisms but removed in an erratum. Str=
ongly supported in morphocladistic (Jamieson 1988) and molecular analyses (=
Jamieson et al. 2002; James and Davidson 2012). Note that if Monigastridae =
are arbitrarily given ordinal rank this would apply also to their sister gr=
oup Opisthopora.
Alluroidina Jamieson 1988. With plesiomorphic single layered clitellum. Mal=
e pores in segment 13. No molecular studies. Sister to the Crassiclitellata=
.
Crassiclitellata. The true earthworms. Distinguished by a multilayered clit=
ellum. Clearly defined in morphocladistic analysis (Jamieson 1988). Strongl=
y supported in molecular analysis (Jamieson et al. 2002; Jamieson 2006; Jam=
es and Davidson 2012). Recognized as an order in A Higher Level Classificat=
ion of All Living Organisms but deleted in an erratum. For detailed classif=
ication of this large group see these works.
Literature cited
Erseus, C. and Kallersjo, M. 2003. 18S rDNA phylogeny of basal groups of Cl=
itellata (Annelida). Zoologica Scripta 33 (2): 187-196.
Jamieson, B. G. M. 1988. On the phylogeny and higher classification of the =
Oligochaeta. Cladsitics 4 (4): 367-402.
Jamieson, B. G. M., Tillier, S., Tillier, A., Justine, J.-L., Ling, E., Jam=
es, S., McDonald, K. and Hugall, A.F. 2002. Phylogeny of the Megascolecidae=
and Crassiclitellata (Annelida, Oligochaeta): combined versus portioned an=
alysis using nuclear (28S) and mitochondrial (12S, 16S) rDNA. Zoosystema 24=
(4): 707-734.
Jamieson, B. G. M. and Ferraguti, M. 2006. Non-leech Clitellata. Pp. 235-39=
2. In Reproductive Biology and Phylogeny of Annelida. Series Editor B.G.M. =
Jamieson. Volume 4. Editors Greg Rouse and Fredrik Pleijel. Science Publish=
ers, Enfield, New Hampshire, U.S.A. Jersey, Plymouth, U.K.
James, S. W. and Davidson, S. K. 2012. Molecular phylogeny of earthworms (A=
nnelida: Crassiclitellata) based on 28S, 18S and 16S gene sequences. Invert=
ebrate Systematics 26: 213-229.
Michaelsen, W. 1928. Oligochaeta. Pp. 1-118. In W. Kukenthal andT. Krubach =
(eds), Handbuch der Zoologie. 2. Plus appendices dates 1930, 1932). Walter =
de Gruyter and Co., Berlin.
Siddall, M. E., Apakupakul, K., Burreson, E. M., Coates, K. A., Ers=E9us, C=
., Gelder, S. R., Kallersjo, M. and Trapido, R. H. 2001. Validating Livanow=
: Molecular data agree that leeches, branchiobdellidans, and Acanthobdella =
peledina form a monophyletic group of oligochaetes. Molecular Phylogenetics=
and Evolution 21(3): 346-351.
Sidall, M. E., Bely, A. E. and Borda, E. 2006. Hirudinida. Pp. 393-429. In =
Reproductive Biology and Phylogeny of Annelida. Series Editor B.G.M. Jamies=
on. Volume 4. Editors Greg Rouse and Fredrik Pleijel. Science Publishers, E=
nfield, New Hampshire, U.S.A. Jersey, Plymouth, U.K.
Emeritus Professor BGM Jamieson
School of Biological Sciences
University of Queensland
St. Lucia
Brisbane
Queensland 4072
Australia
Mailing address:
1 Maxwell Court
Coolum Beach
Queensland 4573
Australia
Phone 61 (0)7 54461716
http://barriegmjamieson.com
