CYLINDROBULLA AND ASCOBULLA IN THE WESTERN ATLANTIC (GASTROPODA, OPISTHOBRANCHIA, SACOGLOSSA) - SYSTEMATIC REVIEW, DESCRIPTION OF A NEW SPECIES, AND PHYLOGENETIC REANALYSIS
Pm. Mikkelsen, CYLINDROBULLA AND ASCOBULLA IN THE WESTERN ATLANTIC (GASTROPODA, OPISTHOBRANCHIA, SACOGLOSSA) - SYSTEMATIC REVIEW, DESCRIPTION OF A NEW SPECIES, AND PHYLOGENETIC REANALYSIS, Zoologica scripta, 27(1), 1998, pp. 49-71
Two often-confused western Atlantic shelled opisthobranchs - Cylindrob
ulla beauii P. Fischer, 1857, and Ascobulla ulla (Marcus & Marcus, 197
0) - are redescribed and compared, via a full literature review; scann
ing electron microscopy of shells and radulae, and serial histological
sectioning of soft anatomy. A third sympatric species, C. gigas, from
Florida, the Caribbean and Bermuda, is described as new, and also pro
vides the first larval development data for the genus. The overlapping
classificatory histories of the genera Cylindrobulla and Ascobulla ar
e summarized and discussed with reference to the Cephalaspidea, Sacogl
ossa (= Ascoglossa), Anaspidea, and Diaphanoidea. Two of the most-rece
nt studies (Jensen 1996a, Jensen 1996b; Mikkelsen 1996) have resulted
in conflicting conclusions about the relationships and classification
of Cylindrobulla. In the author's earlier phylogenetic analysis (Mikke
lsen 1996), Cylindrobulla formed a monophyletic group with the shelled
Sacoglossa with synapomorphies in digestive, reproductive, pallial, a
nd nervous system characters. Jensen's two-part study (Jensen 1996a, J
ensen 1996b) resulted in removal of Cylindrobulla to a new opisthobran
ch order, Cylindrobullacea. These two divergent cladistically produced
classifications were tested by critically re-evaluating Jensen's data
set, especially with regard to Cylindrobulla, and re-running the analy
sis. Thirteen of Jensen's 52 characters were recoded for Cylindrobulla
according to new morphological data and the new species described her
e. Other coding changes were required to update anatomical data for ot
her taxa, add supplementary characters and taxa, more clearly define t
he outgroup, and to critically re-evaluate the cladistic criteria behi
nd several characters. The resultant reanalysis produced a shorter tre
e using Fewer a priori assumptions, but nevertheless preserved the bas
ic tree topology, including three monophyletic sacoglossan clades, pre
sented by Jensen (1996b). Cylindrobulla, however, rejoined the shelled
sacoglossans (Oxynoacea), supported by two synapomorphies. This analy
sis therefore reconfirmed Cylindrobulla's proper placement in the Saco
glossa, rather than in the traditional Cephalaspidea or segregated in
its own monogeneric opisthobranch order. (C) 1998 The Norwegian Academ
y of Science and Letters.