Towards a phylogeny of gastropod molluscs: an analysis using morphological characters

Morphological (including ultrastructural) and developmental characters utilized in recent literature are critically reviewed as the basis to reassess the phylogenetic relationships of gastropods. The purpose of this paper is to provide a framework of characters for future studies and a testable phylogenetic hypothesis. This is one of the fvst attempts to use such characters to assess the relationships of all major clades using parsimony methods. The analysis uses 1 17 characters and includes 40 taxa, predominantly ‘prosobranchs’. Five outgroup taxa are included, representing four conchiferan groups and Polyplacophora. Of the 1 17 characters reviewed and included in the analyses, nine are shell characters (four of these are shell structure), two opercular, two muscular, four ctenidial, 12 renopericardial and 24 reproductive (including 17 based on sperm and spermatogenesis), 27 of the digestive system, 32 of the nervous system and sense organs; the remainder are developmental (3) and of the foot and hypobranchial gland. In the initial analysis the data set included a mixture of binary and multistate characters with all characters unordered. These data were also analysed after scaling so that each character had equal weight. A third data set was constructed in which all characters were coded as binary characters. These analyses resulted in some implausible character transformations, mainly involving the regaining of lost pallial structures. Additional analyses were run on all three sets of data after removing five characters showing the most unlikely transformations. These analyses resulted in generally similar topologies. The robustness of the clades was tested using clade decay. The adaptive radiation of gastropods and their life history traits are briefly described and discussed and the terminology for simultaneous heririaphroditism refined. A scenario for the evolution of torsion equated with the fossil record is proposed and the effects of torsion and coiling on gastropods are discussed along with asymmetry imposed by limpet-shaped body forms. It is suggested that the first gastropods were ultradextral. The idea that heterochrony has played a major part in gastropod evolution is developed and discussed, particularly the paedomorphic stamp imposed on thc apogastropods. The veliger larvae of caenogastropods and heterobranchs are contrasted and found to differ in many respects. The evolution of planktotrophy within gastropods is discussed. Recent phylogenetic hypotheses for gastropods based on molecular data are generally in broad agreement with the present results. On the basis of our analyses we discuss the major monophyletic groups within gastropods. Gastropods appear to be a nionophyletic clade, and divide into two primary groups, the Eogastropoda (incorporating the patellogastropods and their (probably sinistrally coiled) ancestors and the Orthogastropoda - the remainder of the gastropods. Orthogastropoda comprises several well defined clades. The vetigastropod clade encompasses most of the groups previously included in the paraphyletic Archaeogastropoda (fissurellids, trochoideans, scissureiloideans, halioroideans pleurotomarioideans) as well as lepetodriloidean and lepetelloidean limpets and seguenzids. The location of the hot vent taxa Peltospiridae and Neomphalidae varies with each analysis, probably because there is a lack of ultrastructural data for these taxa and parallelism in many characters. They either form a paraphyletic or monophyletic group at or near the base of the \retigastropods or a clade with the neritopsines and cocculinoideans. The neritopsines [Neritoidea etc.) consistently form a clade with the cocculinoidean limpets, but their position on the tree also differs depending on the data set used and (in the case of the scaled data) whether or not the full suite of characters is used. They are either the sister to the rest of the orthogastropods or to the apogastropods. Caenogastropods piesogastropoda ( + architaenioglossan groups) + Neogastropoda] are consistently monophyletic as are the heterobranchs (‘Heterostropha’ + Opisthobranchia + Pulmonata). The caenogastropods and heterobranchs also form a clade in all the analyses and the name Apogastropoda is redefined to encompass this group. New taxa are proposed, Sorbeoconcha for the caenogastropods exclusive of the architaenioglossan taxa, and Hypsogastropoda for the ‘higher caenogastropods‘ ~ -- the Sorbeoconcha exclusive of the Cerithioidea and Campaniloidea.