Contr. Tert. Quatern. Geol. 32(4) 97-132 43 figs Leiden, December 1995 An outline of cassoidean phylogeny (Mollusca, Gastropoda) Frank Riedel Berlin, Germany Riedel, Frank. An outline of cassoidean phylogeny (Mollusca, Gastropoda). — Contr. Tert. Quatern. Geo!., 32(4): 97-132, 43 figs. Leiden, December 1995. The phylogeny of cassoidean gastropods is reviewed, incorporating most of the biological and palaeontological data from the literature. Several characters have been checked personally and some new data are presented and included in the cladistic analysis. The Laubierinioidea, Calyptraeoidea and Capuloidea are used as outgroups. Twenty-three apomorphies are discussed and used to define cassoid relations at the subfamily level. A classification is presented in which only three families are recognised. The Ranellidae contains the subfamilies Bursinae, Cymatiinae and Ranellinae. The Pisanianurinae is removed from the Ranellidae and attributed to the Laubierinioidea.The Cassidae include the Cassinae, Oocorythinae, Phaliinae and Tonninae. The Ranellinae and Oocorythinae are and considered the of their families. The third the both paraphyletic taxa are to represent stem-groups family, Personidae, cannot be subdivided and for anatomical evolved from Cretaceous into subfamilies reasons probably the same Early gastropod ancestor as the Ranellidae. have from Ranellidae the Late Cretaceous. The Cassidae (Oocorythinae) appears to branched off the (Ranellinae) during The first significant radiation of the Ranellidae/Cassidaebranch took place in the Eocene. The Tonninae represents the youngest branch of the phylogenetic tree. Key words — Neomesogastropoda, Cassoidea, ecology, morphology, fossil evidence, systematics. Dr F. Riedei, Freie Universitat Berlin, Institut fiir Palaontologie, MalteserstraBe 74-100, Haus D, D-12249 Berlin, Germany. Contents superfamily, some of them presenting a complete classifi- cation. Modern prosobranch classification schemes (e.g. Ponder & Warén, 1988; Vaught, 1989) recognise five Introduction p. 97 cassoid families: Bursidae, Cassidae, Ficidae, Ranellidae Material and methods p. 97 and Tonnidae. Beu (1988a; see also Beu & Maxwell, Classification p. 98 1990) convincingly showed that Distorsio and its allies, habitat and diet 99 Distribution, p. which previously had been attributed to the Ranellidae, masses and 102 Egg intracapsular development .... p. distinct This is represent a family. strongly supported Larval development p. 105 here and was adopted in the latest comprehensive publi- Protoconch and teleoconch p. 107 cation on the Cassoidea by Warén & Bouchet (1990), Anatomical features p. Ill who also introduced the Laubierinidae as a new cassoid Fossil record p. 120 family. The classification has been subsequently revised Phylogenetic cladogram and discussion p. 124 by Riedel (1994), who removed the Ficidae from the Acknowledgements p. 125 Cassoidea and by Bandel & Riedel (1994), who excluded References 125 p. the Laubierinidae. The remaining five families are the subject of this paper, which is an attempt at a clearly defined phylogenetic analysis on which a classification Introduction can be based. The Cassoidea (= Doliacea = Tonnoidea), containing such MATERIAL METHODS spectacular gastropods as the tritons and helmet-shells, AND have been the subject of research for a long time. Many authors have contributed the of to knowledge the Living animals were observed and collected by the author 98 Fig. 1. A: Initial whorl of shell with indication how to measure the maximum diameter (MD) and method of counting whorls; N - of I - initial axis B: of the last nonspiral shell, defining zero point. height protoconch whorl has to be correlated to get a better understanding of height/width ratio. (by snorkeling, diving and dredging) at several places in specimens were photographed using a reflex camera. The New Zealand (including Leigh Laboratory, Portobello method of counting whorls and measuring dimensions of Marine Station), Australia (Long Reef/Sydney, Heron protoconchs is illustrated in Fig. 1. Island, Lizard Island) and Bali (Indonesia). The snails usually were relaxed using a magnesium chloride solution CLASSIFICATION isotonic with seawater (3.5 %) and then fixed in 70 % ethanol or 5 to 10 % formalin. Klaus Bandel shells of The of the is of based (Hamburg, Germany) provided arrangement taxa course on Recent cassoids from the Caribbean, the Mediterranean phylogenetic considerations. However, in contrast to the and the Red Sea. Alan Beu (Lower Hutt, New Zealand) phylogenetic cladogram, it appears preferable to introduce and Bob Penniket (deceased, formerly Warkworth, New the classification as understood by the author, before the donated characters of certain discussed. Zealand) generously several juvenile specimens taxa are This procedure with preserved protoconchs for further comparison. will hopefully avoid possible nomenclatural confusion. Additional and material borrowed from the The main other dry wet was changes or rearrangements, compared to Australian Museum (Sydney), the Institut fiir Hydro- classifications (e.g. Thiele, 1929; D.W. Taylor & Sohl, biologie und Fischereiwissenschaft (Hamburg), the Musee 1962; Golikov & Starobogatov, 1975; Beu, 1981, 1985, royal de l'Afrique Centrale (Tervuren), the National 1988a; Ponder & Waren, 1988; Beu & Maxwell, 1990; Museum of & New Zealand (Wellington) and the Zoo- Waren Bouchet, 1990) are briefly introduced, but will logisches Institut und Museum (Hamburg). be discussed in more detail elsewhere in the present Fossil material originating from the Campanian of paper. Mississippi (Coon Creek) and the Paleocene of Alabama The Ficidae Meek, 1864 have already been removed from the (Matthews Landing) was collected by Klaus Bandel in Cassoidea and been recognised as a superfamily co-operation with David T. Dockery (Jackson). Klaus (Riedel, 1994). The inclusion of the Laubierinidae Waren Bandel and colleagues provided Eocene gastropods from & Bouchet, 1990 and Pisanianurinae Waren & Bouchet, 1990 in the the Brazos River (Texas) and Oligocene gastropods from Cassoidea presents difficulties on account of Glimmerode which cassoids the absence of larval which unites (northern Germany) among a pallial appendage, with the other families Bandel & well-preserved protoconchs were examined. Several (compare Riedel, 1994). No excellently preserved gastropod faunas from various secondary periostracum on the larval shells is documented European Paleocene, Eocene and Miocene localities were in the literature (Waren & Bouchet, 1990). Moreover, the borrowed from the Institut royal des Sciences naturelles Laubierinidae and Pisanianurinae do not show the de Belgique (Brussels), the Nationaal Natuurhistorisch episodic growth of the teleoconch which is characteristic Museum (Leiden) and the Naturhistorisches Museum for cassoids. It is difficult to imagine that this character lost. other (Vienna). was Many gaps in knowledge have to be and Protoconchs, shells of juveniles and radulae were bridged other anatomical features are of uncertain mounted coated with the in the on stubs, gold and examined with interpretation, e.g. monopectinate osphradium the Laubierinidae aid of an electron microscope (CamScan). Larger or the asymmetrical osphradium in the 99 Pisanianurinae have not been recorded in the Cassoidea. 3a - Subfamily Cassinae Latreille, 1825 The Laubierinidae had already been removed from the Cassis Scopoli, 1777 Cassoidea (Bandel & Riedel, 1994) and the Pi- Cypraecassis Stutchbury, 1837 sanianurinae, which, in addition to conchological charac- 3b - Oocorythinae Fischer, 1885 also show anatomical features which Subfamily ters are very similar Dalium Dall, 1889 to those of laubierinids, should be provisionally placed Galeodea Link, 1807 within the superfamily Laubierinioidea.Alan Beu (pers. Oocorys Fischer, 1883 comm.) concurs with this view. Sconsia Gray, 1847 The tun-shells and its (Tonna relatives) are recognised as a subfamily of the Cassidae and the frog-shells (Bursa 3c - Subfamily Phaliinae Beu, 1981 and its the sister of the relatives) possibly represent group Casmaria H. & A. Adams, 1853 Cymatiinae. Sassia and Charonia have been removed Phalium Link, 1807 from the Cymatiinae and are attributed to the Ranellinae Semicassis Morch, 1852 but perhaps represent an independent ranellid branch 3d - Tonninae Suter, 1913 (subfamily). Dalium, Galeodea and Sconsia have usually Subfamily Eudolium Dali, 1889 been assigned to the Cassinae, but are here considered to Malea Valenciennes, 1883 be better placed in the Oocorythinae. Subgenera and Tonna Briinnich, 1772 extinct taxa not included in the classification are pre- sented below. Personopsis currently contains but a single Recent species, which, however, will be reassigned by DISTRIBUTION, HABITAT AND DIET Beu (pers. comm.) to a new personid genus. This means that Personopsis will have to be removed from the list of Personidae live in most warm oceans and are mainly when that Recent genera new name is coined. represented by the genus Distorsio (compare Clench & Turner, 1957; Lewis, 1972; Beu, 1985). Personopsis (?) is known exclusively from the West Pacific (Beu, 1985) Subclass Caenogastropoda Cox, 1959 and Distorsionella appears to be restricted to the southern Order 1991 Neomesogastropoda Bandel, West Pacific (Beu, 1978b, 1985). Suborder Troschelina Bandel & Riedel, 1994 Common species such as Distorsio anus (Linne, 1758) Superfamily Cassoidea Latreille, 1825 and Distorsio reticulata Roding, 1798, can be found in shallow-water habitats