Bull. nat. Hist. Mus. Lond. (Zool.) 63(1): 55-92 



Issued 26 June 1997 



Foregut anatomy and relationships of the 

 Crassispirinae (Gastropoda, Conoidea) 



YURI I. KANTOR AND ALEXANDRA MEDINSKAYA 



A.N. Severtzov Institute of Animal Evolutionary Morphology and Ecology, Russian Academy of Sciences, Lenin 

 Avenue 33, Moscow 1 17071, Russia 



JOHN D. TAYLOR 



Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD, UK 



CONTENTS 



Introduction 55 



Material and methods 56 



Anatomical descriptions 56 



Character analysis and relationships within Crassispirinae 84 



Discussion 87 



Conclusions 91 



Acknowledgements 91 



References 91 



SYNOPSIS. The foregut anatomy of 31 species from the conoidean subfamily Crassispirinae is described. Great variation is 

 found between species in the configuration of the foregut including features such as: the structure of the rhynchodeum, the 

 morphology of the proboscis, the position and number of buccal tube sphincters, the position and structure of the buccal mass, 

 the histology of the salivary glands and venom gland, the length of the oesophagus, the structure of the muscular bulb and the 

 morphology of the radular teeth. Many species have marginal teeth of the wishbone type, but teeth are paddle-shaped in Funa and 

 Vexitomina, harpoon-shaped in Cheungbeia and a hollow awl shape in Ptychohela. Many crassispirinae have rather similar shells, 

 but different anatomies. Some species with similar radulae have different anatomies and others with similar anatomy have widely 

 differing radulae. An analysis of relationships, using Gemmula as outgroup, shows that the various subgenera of Crassispira are 

 not monophyletic and should be raised to generic status. Also, Epidirona is polyphyletic with some species belonging in the 

 Turrinae, as do some species of Turridrupa. Additionally, some species classified in the genera Guraleus, Antiguraleus and 

 Paraguraleus, until recently classified as Mangeliinae, belong in the Crassispirinae. 



INTRODUCTION 



Gastropods of the superfamily Conoidea are notable for the posses- 

 sion of a large, coiled venom gland, together with the highly 

 modified radular teeth which are used to inject the venom into the 

 prey. Although Conns is the most well-known taxon, it represents 

 only a small part of the total diversity of the group which conserva- 

 tive estimates suggest as more than 4000 living species and 340 

 genera (Taylor, Kantor & Sysoev, 1993). Although most classifica- 

 tions (e.g. Powell, 1966; McLean, 1971) have been based largely on 

 shell and radular characters, Taylor et al. (1993) have recently 

 provided anatomical criteria, mainly derived from characters of the 

 foregut, for the definition of suprageneric taxa of conoideans. Al- 

 though their study involved anatomical investigation by serial sections 

 of more than 72 species of conoideans, this nevertheless represented 

 only a small fraction of the living genera and species. Moreover, 

 amongst the species studied so far, a wide disparity in the configura- 

 tion of the various organs of the foregut in the Conoidea has been 

 revealed (Taylor et al., 1993), with new arrangements still being 

 discovered ( Kantor &Tay lor, 1994;Taylor, 1994; Kantor and Sysoev, 

 1996). These preliminary studies suggested that the subfamily 

 Crassispirinae, one of four subfamilies ofTurridae possessing 'wish- 



bone' radular teeth, showed a wide variation in radular morphology 

 and foregut anatomy, including the possibility of further evolution- 

 ary pathways to the hypodermic feeding system. Moreover, within 

 the Crassispirinae, some species in the genera Inquisitor, Funa and 

 Ptychobela which possess rather similar shells, were shown to have 

 very different foregut and radular morphologies (Kilburn, 1989; 

 Taylor, 1994; Taylor & Wells, 1994). The problem of using shell 

 characters alone to classify conoideans has recently been high- 

 lighted in the case of the southern African species Antiguraleus 

 morgani, previously classified in the Mangeliinae on the basis of 

 shell morphology, but shown to be a likely crassispirinan on radular 

 characters (Kilburn, 1994). For these reasons we decided to investi- 

 gate the anatomy of the Crassispirinae in more detail. 



Currently, some 48 genera and subgenera have been assigned to 

 the Crassispirinae (Taylor et al., 1993; Taylor & Wells, 1994; 

 Kilburn, 1994) mainly on the evidence of radular characters, but in 

 some cases on shell morphology alone. The subfamily is diverse in 

 the tropical West America (McLean, 1971; Keen, 1971) with over 52 

 species recorded, and also in the Caribbean (Maes, 1983) and West 

 Africa (Femandes, Rolan & Otero-Schmitt, 1995). The Indo-Pacific 

 fauna is less well known and there are many undescribed species, but 

 Kilburn (1988; 1994) reports 54 species from Southern Africa, 

 including numerous new genera and species, and many other species 



©The Natural History Museum, 1997 



