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WORKING CLASSIFICATION OF THE GASTROPODA 273
in number of teeth. Oligomeha, Davisiana and Guttula
have sensory papillae on the cephalic tentacles (Waren,
unpubl), confirming their inclusion in the Vetigastropoda.
3' Hickman & McLean (1990) recognised the affinities of
Chilidontini and Calliotropini, by them recognised as tribes
in the Trochidae, and outlined their similarities, while they
considered the systematic position of Cataeginae uncer-
tain, due to the highly apomorphic radula of the type spe-
cies of Cataegis. Waren & Bouchet (1993) described a
less modified radula in Cataegis meroglypta McLean &
Quinn, 1987, but were still uncertain about the position.
Recent discovery of an undescribed species on sunken
wood in the Solomon Islands, with a chilodontine -
calliotropine type of radula, as understood by Hickman &
McLean (1990), makes it possible to, at least provision-
ally, conclude relations between these taxa.
38 We follow McLean & Hickman (1990) in regarding
Eucyclus as a vetigastropod related to Chilodontidae.
^ Content based on Tracey et al. (1 993). All fossil "archeo-
gastropods" without slit and selenizone were classified
" by Bändel & Fryda (1996) in a
Trochomorpha ". They did not allocate Palaeozoic taxa
morphogroup
to any particular superfamily. Classification based on
Hickman & McLean (1990) with modifications,
* Classification based on Marshall (1995)
*' Proconulidae ranked as a family of Trochoidea after
Gründel {2000a).
''^
It is uncertain whether Solariellidae should be classi-
fied in the Trochoidea or Seguenzioidea. The reduction
in number of both lateral and marginal teeth may indi-
cate seguenziid relations; in the absence of other infor-
mation we have maintained a placement in Trochoidea.
"3 Placement of Velainellidae in Trochoidea after Le Renard
(pers comm ).
** Classiftcation of Colloniinae adapted from Monari et al.
(1995). Helicocryptinae synonymized with Ataphrinae
by Monari et al., re-established as valid subfamily of
Colloniidae by Gründel (2003)
"5 Skeneinae treated as a subfamily of Turbinidae based
on radula and 16S sequences (Waren, unpublished).
* Tegulinae placed in Turbinidae based on radula and 16S
sequences (Waren, unpublished).
''''
Phasianellidae is treated as a separate family on the basis
of shell structure (Woodring, 1928; Robertson, 1985;
Marcus & Marcus, 1960; Hedegaard 1990), with
Tricoliidae also separated as distinct by some of these
authors. Also, Bändel & Geldmacher (1996) have produced
a phylogenetic scenario with Tricolia completely
independent from Turbo (Phasianella not included).
Sperm ultrastructure also differs between examined
turbinids and Tricolia {Phasianella sperm not known)
(Hodgson & Foster, 1992) Tricolia was supposed to be
unique in Trochoidea in having two shell muscles (Marcus
& Marcus, 1960), but this unusual condition may be re-
lated to the elongate shape of the operculum; it also oc-
curs in trochid limpets.
* Young specimens of Phasianella have the same
commarginal spiral line on the outside of the operculum
as Gabnelona and Eugabrielona, suggesting that the
latter genera are paedomorphic phasianellids. Large
species of Phasianella and Tricolia have the same tendency
to loss of the central tooth, oherwise known mainly
from patellogastropods.
*• The name Cocculiniformia originally encompassed all
cocculiniform taxa (Haszprunar, 1987). Molecular work
based on Coccopigya as representative of Cocculinidae
and Notocrater as representative of Pseudococculinidae
resulted in Cocculinoidea being polyphyletic (Colgan et
al , 2003), with Lepetelloidea now placed in Vetigastropoda
and Cocculinoidea placed outside them The
name Cocculiniformia is kept here in a restricted sense
to mark the distinctiveness of Cocculinoidea.
^ Cocculinidae usually appears as a very distinct clade
both in analyses based on morphology and on molecular
data Relations to Neritimorpha have been suggested
(e.g.. Ponder & Lindberg, 1997) and, more recently, to
Patello- and Vetigastropoda (Colgan et al., 2003).
5' Inclusion of Bathysciadiidae in Cocculinoidea after
Strong et al. (2003), based on admittedly poor informa-
tion on Bathysciadium. Bathysciadiidae share with
Lepetellidae and Addisoniidae the habit of discarding
the protoconch at the size of 0.3-0.6 mm, and this may
indicate a currently unrecognized relationship
^ Classification based on Moore (1960), with additions
from Bändel {1992a), Tracey et al. (1993), Bändel &
Fryda (1999), Fryda (1998c, 1999a) introduced two taxa,
Cyrtoneritimorpha and Cycloneritimorpha, within the
Neritimorpha. Cyrtoneritimorpha includes Ordovician-
Permian gastropods with fishhook-like protoconchs
Cycloneritimorpha unites all post-Palaeozoic Neritimorpha
and may possibly also include the Palaeozoic
Platyceratoidea and Nerrhenoidea.
^ Oriostomatidae included in Euomphaloidea by
Vostokova & Pchelintsev (in Pchelintsev & Korobkov,
1960).
^ Hypothesized by P J. Wagner (2002) to belong to the
"euomphaline subclade".
^ Based on molecular data, Kano et al. (2002) produced
a phylogeny of the Recent Nentimorpha recognizing four
clades: Hydrocenidae; Helicinidae + Neritiliidae;
Neritidae + Phenacolepadidae; Neritopsidae + Titiscani-
idae. Their groupings are followed here, with the result-
ing clades ranked as superfamilies.
* Classification based on Keen [in Moore, 1960].
^ The Carboniferous Dawsonellidae are regarded by Kano
et al. (2002) to be derived from an ancient Neritimorpha
before the first bifurcation of the Neritopsoidea, and to
be convergent in shell form with the Helicinidae.
^ Deianiridae placed in Neritoidea by Bändel & Fryda
(1999), regarded as the sister taxon of the Helicinidae
by Kano et al. (2002).
^ Neritiliidae ranked as family after Kano & Kase (2002).