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J.D. TAYLOR, Y.I. KANTOR AND A.V. SYSOEV 



paired or fused. Single accessory salivary gland in Aforia. 



Remarks. Since the type-genus of the subfamily Turriculi- 

 nae, Turricula Schumacher, 1817, is transferred to the Clav- 

 atulinae (see above), the next available name for this group is 

 Cochlespirinae Powell, 1942. 



Subfamily Zonulispirinae McLean, 1971 



Shells rather small (15-25 mm), claviform. Anterior canal 

 usually short, sometimes moderately long. Predominantly 

 spiral scuplture, well developed. Protoconch multispiral, ini- 

 tially with smooth whorls, then with oblique axial riblets. 

 Anal sinus on the shoulder, often sub-tubular, with well 

 developed parietal callus. Operculum with terminal nucleus. 



Radula. With strong membrane and marginal teeth in each 

 row. Teeth semi-enrolled, to rolled, hollow teeth with narrow 

 base. Tips may be barbed or unbarbed. 



Foregut. Proboscis long, with a single distal buccal tube 

 sphincter. Buccal mass distal. Odontophore small with two 

 unfused cartilages. Buccal lips present. Salivary glands fused. 

 Anterior of venom gland ciliated. Oesophagus elongated 

 between the buccal mass and nerve ring. 



Subfamily Turrinae H. & A. Adams, 1853 



Shell usually of medium to large size (up to 110 mm), 

 fusiform. Anterior canal elongated and narrow, rarely trun- 

 cated. Anal sinus on the whorl periphery. Axial sculpture 

 weak or absent. Protoconch smooth in its initial part, subse- 

 quent whorls axially costate; paucispiral protoconchs smooth. 

 Operculum with terminal nucleus. Egg capsules dome- 

 shaped, operculate. 



Radula. Strong radular membrane, 2-3 teeth in each row. 

 Central tooth either well-developed, small or absent, quad- 

 rate to rectangular with a strong central cusp. Lateral teeth 

 absent. Marginal teeth of robust wishbone type. 



Foregut. Proboscis moderately long, rhynchostomal 

 sphincter posterior, a single distal buccal tube sphincter, 

 protrusive lips of buccal tube present. Buccal mass basal. 

 Odontophore small with fused cartilages. Salivary glands 

 paired. No accessory salivary glands. Anterior part of venom 

 gland ciliated. 



Family Conidae Fleming, 1822 



Radula consisting of hollow marginal teeth only. Radular 

 membrane absent. Radular diverticulum divided into short 

 and long arms. Odontophore absent. Radula and venom 

 gland may be absent. Salivary glands acinous or tubular. 

 Accessory salivary gland either present or absent. Operculum 

 either present or absent. 



Subfamily Clathurellinae H. & A. Adams, 1858 



Shell small to rather large, fusiform to biconic. Anterior canal 

 short or indistinct to moderately elongate. Sculpture pre- 

 dominantly spiral in most genera. Anal sinus deep to very 



shallow, on the shoulder slope or on the periphery. Col- 

 umella with or without pleats. Protoconch usually paucispiral, 

 smooth, sometimes carinate or weakly spirally ribbed, rarely 

 axially costate on its last whorl. Operculum with terminal 

 nucleus present, vestigial or absent. 



Radula. Awl- or harpoon-shaped marginal teeth, without 

 (very rarely with) solid base, tooth cavity opens terminally at 

 the proximal end in vast majority of species. 



Foregut. Proboscis short to long, 1 or 2 anterior buccal 

 tube sphincters, buccal mass basal. Short buccal lips in 

 Tropidoturris. Odontophore absent, radular caecum present 

 — divided by septum in Bathytoma (Micantapex) . Salivary 

 glands tubular in Borsonia, acinous in others, paired, single 

 or absent. Single accessory salivary gland present in some 

 species. Venom gland with uniform histology. No elongation 

 of oesophagus. 



Remarks. This subfamily comprises species classified by 

 other workers in the subfamilies Borsoninae and Clathurelli- 

 nae. Being very variable in both anatomical and shell charac- 

 ters, the subfamily may be of polyphyletic origin. More 

 species need to be studied anatomically before any satisfac- 

 tory classification can be attempted. The subfamily is defined 

 mainly by the character of the radular teeth. Several groups 

 of genera can be isolated within Clathurellinae according to 

 shell characters. 



'Clathurellid' group is characterized by medium-sized 

 shells (usually 10-25, up to 40 mm), with a moderately 

 elongate siphonal canal, and a well developed, often cancel- 

 late sculpture. Columella without pleats, but both inner and 

 outer lips may be denticulated; anal sinus deep located on the 

 shoulder. Protoconch usually multispiral, last whorls with a 

 pronounced medial carination and, sometimes, weak axial 

 lamellae on the lower half. A distinctive feature of this group 

 is densely granulated shell surface of most genera (except of 

 one subgenus of Glyphostoma and, probably, Nannodiella). 

 Operculum absent. Radular teeth long and slender, slightly 

 curved, without a solid base. 



'Bathytomid' group. Shell of medium to rather large size 

 (usually 20-30, up to 70 mm), more or less biconic. Sculpture 

 usually well developed, entirely spiral, ribs often gemmulated 

 by growth lines; typically there is a peripheral tuberculated 

 flange. Anal sinus rather deep, located on the whorl periph- 

 ery. Columellar pleats strong to obsolete. Protoconch of 

 1.5-3 whorls, smooth or minutely papillated. Operculum with 

 terminal nucleus. Radular teeth either long, with more or less 

 terminal opening, or short, with large cylindrical solid base 

 and lateral opening. 



'Borsoniid' group. Shell of rather small to medium size 

 (usually 15-25, up to 62 mm), fusiform. Anterior canal 

 moderately elongate, sometimes long. Both spiral and axial 

 sculpture may be present. Columellar pleats weak or absent. 

 Anal sinus on the shoulder slope. Protoconch of 1-2 smooth 

 whorls. Operculum fully developed, small or absent. Radular 

 teeth long, without solid base, open terminally, or, rarely, 

 short, with large cylindrical base, open laterally. Egg capsules 

 dome-shaped, with an operculum. 



'Mitromorphid' group. Shell small (usually 4-8, up to 17 

 mm), biconic and 'mitriform'. Anterior canal very short or 

 indistinct. Aperture narrow, columella with or without teeth, 

 outer lip usually denticulated, anal sinus shallow and subsu- 

 tural. Sculpture predominantly or entirely spiral. Protoconch 

 of 1.5-2 smooth whorls. Operculum absent. Radular teeth 



