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Phylogeny of the Caenogastropoda (Mollusca), Based on Comparative Morphology

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Phylogeny of the Caenogastropoda (Mollusca), Based on Comparative Morphology TABLE OF CONTENTS Abstract .......................................................................................................................................................161 Introduction ................................................................................................................................................162 Materials and Methods ................................................................................................................................163 Discussion of Characters .............................................................................................................................175 Shell (Characters 1‑52) ..........................................................................................................................175 Head‑foot (Characters 73‑147) ..............................................................................................................177 Operculum (Characters 53‑72) ..............................................................................................................180 Pallial organs (Characters 148‑228) .......................................................................................................181 Circulatory system (Characters 240‑261) ...............................................................................................185 Excretory system (Characters 262‑290)..................................................................................................185 Digestive system (Characters 291‑539) ..................................................................................................187 Foregut (Characters 291‑302) ..........................................................................................................187 Buccal mass (Characters 303‑326) ...................................................................................................190 Odontophore (Characters 327‑406) .................................................................................................190 Radula (Characters 407‑444) ...........................................................................................................197 Salivary glands (Characters 445‑461) ...............................................................................................200 Esophagus (Characters 462‑489)......................................................................................................200 Stomach (Characters 490‑516) ........................................................................................................203 Intestine (Characters 517‑539) ........................................................................................................206 Reproductive system (Characters 540‑648) ............................................................................................207 Male (Characters 540‑582) ..............................................................................................................207 Female (Characters 583‑648) ...........................................................................................................208 Central nervous system (Characters 649‑673) ........................................................................................210 Cladistic Analysis ........................................................................................................................................212 Discussion of the Cladogram .......................................................................................................................212 Extra characters, not utilized directly in the present study ......................................................................217 Analysis of the obtained cladogram and the taxonomy ...........................................................................219 Paleontology ..........................................................................................................................................225 Comparison with molecular studies .......................................................................................................226 Conclusions ................................................................................................................................................227 Resumo .......................................................................................................................................................227 Acknowledgments .......................................................................................................................................228 References ...................................................................................................................................................228 Appendix 1. .................................................................................................................................................238 Appendix 2 ..................................................................................................................................................267 Appendix 3 ..................................................................................................................................................309 Appendix 4 ..................................................................................................................................................318 Volume 42(4):161‑323, 2011 PHYLOGENY OF THE CAENOGASTROPODA (MOLLUSCA), BASED ON COMPARATIVE MORPHOLOGY 1 LUIZ RICARDO L. SIMONE ABSTRACT The systematics, classification and phylogeny of the Caenogastropoda are revised based on an analysis of the morphology of representatives of all branches. The basis of this work is the de- tailed examination of the morphology of 305 species, most of which are reported on in detail elsewhere. Representatives of most caenogastropod families were included (comprising 270 spe- cies), and 35 outgroup taxa. A phylogenetic analysis based upon 676 morphological characters, with 2291 states (1915 of which are apomorphic states), is presented. The characters comprise every organ system and many are discussed in detail. The polarization is based on a pool of non-caenogastropods, comprising 27 representatives of Heterobranchia, Neritimorpha, Vetigas- tropoda, Cocculiniformia and Patellogastropoda. Additionally, eight representatives of other classes are also included. The root is based on the representative of Polyplacophora. A few char- acters were included in order to organize the outgroups, to find the position of Caenogastropoda among them, and to find the synapomorphies of Caenogastropoda. A strict consensus cladogram of the 48 most parsimonious trees (Fig. 20; length of 3036, CI = 51 and RI = 94) is presented, a synopsis of which is: ((((((Cyclophoroidea2 (Ampullarioidea5 (Viviparoidea15 (Cerithioidea19 (Rissooidea41 (Stromboidea47 (Calyptraeoidea67 (Naticoidea97 (Cypraeoidea118 (Tonnoidea149 (Conoidea179 (Cancellarioidea222 – Muricoidea212)))))))))))) HeterobranchiaV) Neritimor- phaU) VetigastropodaL) CocculiniformiaJ) Patellogastropoda) (superscripts indicating the nodes at Fig. 20). The monophyly of the Caenogastropoda is supported by 60 synapomorphies. Based on this cladogram, the systematics, evolution and paleontology of the group are discussed, as along with comparisons with other phylogenetic studies involving Caenogastropoda, includ- ing those obtained from molecular data. The list of characters is included in Appendix 1; the matrix of characters in Appendix 2; and the list of synapomorphies of each node are presented in Appendix 3 in form of symbols. As the phylogeny obtained in this analysis has a high degree of concordance with other recent phylogenetic studies, a number of new supra-superfamiliar taxa are introduced: Siphonogastropoda (Cypraeoidea + Peogastropoda); Adenogastropoda (Naticoidea + Siphonogastropoda); Rhynchogastropoda (Calyptraeoidea + Adenogastropoda); Strombogastropoda (Stromboidea + Rhynchogastropoda); Peogastropoda (Tonnoidea + Neo- gastropoda); Epiathroidea (Viviparoidea + Sorbeoconcha); Hydrogastropoda (Ampullarioidea + Epiathroidea); Adenogonogastropoda (Neritimorpha + Apogastropoda) (Appendix 4). Key‑Words: Gastropoda; Caenogastropoda; Phylogeny; Morphology; Anatomy; Systematics. 1. Museu de Zoologia, Universidade de São Paulo. Caixa Postal 42.494, 04218‑970, São Paulo, SP, Brasil. E‑mails: [email protected], [email protected] 162 Simone, L.R.L.: Morphological Phylogeny of Caenogastropoda INTRODUCTION (as pectinibranchés), and Latinized by Goldfuss (1820) (Cox, 1960b; Bouchet, personal communication). Pec‑ “Caenogastropoda – the field of success” (Haszprunar, tinibranchia has been also called Monotocardia in 1988a: 415) some classifications (e.g., in Encyclopaedia Britannica; Marcus, 1958; Golikov & Starobogatov, 1987, 1988). Despite some initial inconsistencies (e.g., the Probably because of above mentioned confu‑ inclusion of trochoids) when introduced by Cuvier sion and due to the informal tendency for erecting (1814), the name Pectinibranchia generally encom‑ supra‑familial taxa with the suffix – gastropoda (as the passed most of the Meso‑ and all Neogastropoda as case of the Stenoglossa replaced for Neogastropoda), later defined by Thiele (1929) (the latter taxon was Cox (1960a) introduced the epithet Caenogastropoda called Stenoglossa by that author). As the name sug‑ for replacing the previous Pectinibranchia. The ap‑ gests, the taxon is an assemblage of monopectinate parent objective of the author was
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