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272
and content of superfamilies based on Tracey et ai
(1993); however, see Vostokova & Pchelintsev (in
Pcheiintsev & Korobkov, 1960) and P. J Wagner (2002)
for aiternative ciassifications.
A weakness in the classification of Palaeo- and Meso-
zoic gastropods is the automatic exclusion of fossils with
a multispiral protoconch from "archaeogastropods" and/
or Vetigastropoda, From a methodological point of view,
the absence of planktotrophy in early gastropods should
not be taken as a fact but as an hypothesis to be tested.
The Cambro-Devonian Clisospiroidea had multispiral
protoconchs, and it cannot be ruled out that the non-
planktotrophy of modern vetigastropods is derived rather
than plesiomorphic. The occurrence of an unquestionably
multispiral protoconch in a species of Mourlonia
[Eotomariidae] from the Devonian of Poland (Kaim in
press, pers. comm.) highlights this issue
2° Ataphridae seems to be the valid name for what has
hitherto been called Trochaclididae (Waren, unpubl).
^^ Classification of Eotomariidae based on Gordon &
Yochelson(1987).
22 Classification based on Keen [in Moore] (1960),
Christiaens (1973), and McLean (1984). The name
Deridobranchinae Gray, 1847 is based on Deridobranchus
argus Ehrenberg, 1831, a Red Sea species,
described by Ehrenberg as having an Emarginula type
animal and no shell The species has not been recog-
nized subsequently, and Deridobranchus and Deridobranchinae
have been omitted from classifications
23 Placement of Temnotropidae in Haliotoidea based on
presence of nacre (Bändel, 1991d).
2^* The relations between the taxa here included in
Lepetelloidea are uncertain. Morphological information
(Ponder & Lindberg, 1997) as well as molecular data
(Colgan et al., 2000) indicate a position within Vetigastropoda.
Lepetellidae and Addisoniidae (as well as
Bathysciadiidae, see Note 51 ) have the habit of discard-
ing the protoconch at a size of 3-0.6 mm. The inclusion
of the other families in Lepetelloidea is more
problematic
25 Haszprunar (1992) considered C/?onsie//a to be second-
25
arily coiled, but that seems unlikely (Ponder & Lindberg,
1997). The latter view is supported by more elaborately
coiled and sculptured taxa like Bichohstes (Chori-
stellinae), sensory bursicles in Choristes, presence of
eyes in at least one choristellid species (Waren, unpubl.),
and the parallel occurrence of Helicopelta, a coiled
addisoniid.
It seems unnecessary to use two families or even two
subfamilies to classify the two genera Lepetodrilus and
Gorgoleptis
^ Great similarity in protoconch, radular and ontogenetic
characters suggest close affinity of Lepetodrilidae and
Clypeosectidae (originally in Fissurelloidea), and this is
confirmed by molecular data (Geiger & Thacker, pers.
comm).
2^ Great similarity in protoconch, radular and ontogenetic
characters suggest close affinity of Lepetodrilidae and
BOUCHET&ROCROI
Sutilizonidae (originally in Scissurelloidea), and this is
confirmed by molecular data (Geiger & Thacker, pers.
comm.) Temnocinclis and Sutilizona have a radula of
typical scissurellid appearance (although the enlarged
fourth lateral tooth is missing); they differ mainly in shell
shape (protoconch not known in Temnocinclinae), but
are kept together by having a pair of monopectinate
ctenidia and the radula which has no clear demarcation
between the central and marginal field.
23 Murchisonioidea included in Caenogastropoda by Pon-
der & Waren (1988) and Bändel (1993b, 1997); in
Archaeogastropoda by Tracey et al. (1993) and Fryda &
Manda (1 997). Archaeogastropod-type protoconchs have
been found in the Devonian members of the included
families (Fryda & Manda, 1997; Fryda, unpubl. observ ).
* The systematic position of the Neomphaloidea remains
uncertain although close relations to the rest of the
Vetigastropoda from molecular data (McArthur & Koop,
1999; Colgan et al., 2000; Colgan et al., 2003; Waren
et al., 2003) and from morphology seem trustworthy.
The previously not noticed occurrence of sensory ctenidial
bursicles in Peltospiridae and Melanodrymiidae (Waren
et al., 2003) gives further support to close relations.
'' Content based on Tracey et al. (1993). All fossil
archeogastropods with slit and selenizone were classified
by Bändel & Fryda (1996) in a " morphogroup
Selenimorpha ". They did not allocate Palaeozoic taxa to
any particular superfamily
^ Classification based on Bändel (1993a) However, P. J.
Wagner (2002) noted that the Porcelliidae belong to the
Gosseletininae clade of the family Gosseletinidae (su-
perfamily Eotomarioidea).
^ Molecular data (Geiger & Thacker, in Geiger & Jansen,
2004, and pers. comm.) suggest that Scissurellidae are
not monophyletic. Scissurella + Sukashitrochus is the
sister group to Lepetodriloidea in a crown clade with
Haliotidae, and Anatoma is amongst the most basal
Vetigastropoda including Pleurotomariidae. Anatomidae
was treated at family rank by Geiger & Jansen (2004),
and for lack of a better alternative, we have classified it
as a family of Scissurelloidea The position of Larocheinae
is unsettled.
** Depressizoninae was based on a species known from
shells only. Its general similarity to species of Scissurella
(except having a more depressed shell) suggests much
closer affinity to Scissurella that to any other scissurelloid
group.
^ Anatomical information (Sasaki, 1998) refuted all previ-
ous speculations on caenogastropod affinity of
Seguenzioidea and confirmed basic vetigastropod
anatomy with several apomorphies. Some genera of
Seguenziidae, e.g. Ancistrobasis, closely approach shell
and external soft part morphology of Chilodontidae, as
exemplified by Calliotropis. 16S data support close affinity
of Cataegis, Calliotropis and Seguenzia (Waren
et al, 2003).
^ Guttulinae, Davisianinae, Putillinae, and Oligomehinae
are featureless, poorly known taxa. The radula (when
known) is, like in Seguenzia, characterised by a reduction