56 



Y.I. KANTOR, A. MEDINSKAYA AND J.D. TAYLOR 



Table 1 List of crassispirine gastropods and the outgroup Gemmula 

 which have been sectioned, with details of their collection locations. 



Crassispira (Crassispira) incrassata (Sowerby, 1834). North side of 



Venado Island, Panama, (8°53'N, 79°36'W). ANSP A9695 T357. 

 Crassispira (Crassispira) maura (Sowerby, 1834). Between Tortota and 



Venado Island, Bay of Panama, Republic of Panama (8°5 1 '45N, 



79°35'40W), 9-10 m, ANSP A9613 T357. 

 Crassispira (Gibbaspira) dysoni (Reeve, 1846). Isla de Lobos, Gulf of 



Mexico, Mexico (21°28'N, 97°13'W). ANSP A9423T357. 

 Crassispira (Glossispira) harfordiana flucki (Brown & Pilsbry, 1913). 



Playa Benito, Campeche, Gulf of Mexico, Mexico (19°48'N, 90°36'W). 



ANSP 356707 A9734 T357. 

 Crassispira (?Crassispirella) Iatizonata (E.A. Smith, 1882). 1 km north of 



Holotown, St James, Barbados (13°1 l'N, 59°38'W). ANSP A9866G 



T357. 

 Crassispira (Monilispira) pluto Pilsbry & Lowe, 1932. West San Carlos 



Bay, Sonora, Mexico (27°57'N, 1 1 1°04'W). ANSP A9226 T357. 

 Crassispira (Striospira) kluthi Jordan, 1936. Venado Island, Panama 



(08°53'N, 79°36'W). ANSP 356323 A9698 T357. 

 Crassispira (Striospira) tepocana Dall, 1919. San Carlos, Guaymas, 



Sonora, Mexico, 35 m. ANSP A6670. 

 Crassispira (Striospira) xanti Hertlein & Strong, 195 1 . West side of 



Viradores sur, Bahia del Cocos, Costa Rica (10°34'45N, 85°43'35W). 



ANSP35791A10167T357. 

 Crassispira (Crassiclava) turricula (Sowerby, 1834). Off Nacascola, west 



side of Bahia Culebra, Bahia Culebra, Costa Rica (10°37'15N, 



85°41' 20W). ANSP A9753B T357. 

 Crassispira (Crassiclava) apicata (Reeve, 1845). Cactus Point, Prickly 



Pear Island, British Virgin Islands (18°30'55N, 64°22'30W). ANSP 



355532 A9461ET357. 

 Burchia spectabilis Sysoev & Taylor, 1997. Houtman Abrolhos Islands, 



Western Australia (see Sysoev & Taylor, 1997). BM(NH). 

 Miraclathurella bicanalifera (Sowerby, 1834). West side of Viradores Sur, 



Bahia del Cocos, Costa Rica (10°34.30'N, 85°43.40'W). ANSP 357806 



A9857AT357. 

 Hindsiclava andromeda (Dall, 1919). 3 miles southwest of Punta San 



Antonio, Sonora, Gulf of California, Mexico (27°54'N, 1 1 1°08'W). 



ANSP358149A10192GT357. 

 Hindsiclava militaris (Reeve, 1843) 2 miles east of Punta Doble, Sonora, 



Gulf of California, Mexico (27°55'N, 1 1 1°04'W). ANSP A10186 T357. 



West side of Bahia Culebra, off Nacascola, Bahia Culebra, Costa Rica 



(10°37'15N, 85°41'20W). ANSP 357487 A9753HT357. 

 Funa jeffreysi (Smith, 1875). South of Cape d'Aguilar, Hong Kong, 20 m. 



BM(NH). 

 Funa latisinuata (Smith, 1877). South of Cape d'Aguilar, Hong Kong, 



20 m. BM(NH). 

 Ptychobela suturalis (Gray, 1838). North Lantau Island, Hong Kong, 12 m. 



BM(NH). 

 Cheungbeia mindanensis (Smith, 1877). South of Cape d'Aguilar, Hong 



Kong, 20 m. BM(NH). 

 Cheungbeia robusta (Hinds, 1839). South of Cape d'Aguilar, Hong Kong, 



20 m. BM(NH). 

 Vexitomina garrardi (Laseron, 1954). 2-3 km E. of Malabar, Sydney, 



Australia (32°59.27'S, 150°16.48'E). AM. 

 Inquisitor cf adenicus Sysoev, 1996. Off Ras Madrakah, Oman 



(19°14.8'N, 56°26.8'E), 935 m. NMW. 

 Inquisitor aemula (Angas, 1877). Outer Lagoon, 10 m, Noumea, New 



Caledonia. BM(NH). 

 Inquisitor latifasciata (Sowerby, 1870). Off Cape d'Aguilar, Hong Kong, 



25-30 m. BM(NH). 

 Epidirona gabensis (Hedley, 1922). 2 km East of Long Bay, Sydney, New 



South Wales, Australia, 66 m. AM. 

 Antiguraleus morganus (Barnard, 1958). Off Mendu Point 300 m, 



Transkei, South Africa (32°21.8'S, 29°0.0'E). NM. 

 Paraguraleus costatus (Hedley, 1922). 28 km east of south head of Little 



Bay, Sydney, NSW, Australia (33°58'54"S, 151 33'38"E), 183-192 m. 



AM. 

 Naudedrillia praetermissa (Smith, 1904). Off Sandy Point, Transkei, 



South Africa (32°37.4'S, 28°36.9'E), 90 m. NM. 

 Nquma scalpta Kilburn, 1988. Off Park Rynie, Natal, South Africa 



(30°23.2'S, 30°50.8'E), 140 m. NM. 

 Turridrupa bijubata (Reeve, 1843). 30 m, Beacon Island, Houtman 



Abrolhos Islands, Western Australia. BM(NH). 

 Haedropleura septangularis (Montagu, 1803). Torre San Giovanni, BR, 



Italy, 12 m. BM(NH). 

 Outgroup: subfamily Turrinae: 



Gemmula deshayesii (Doumet, 1839). Off Cape d'Aguilar, Hong Kong, 20 

 m. BM(NH). 



Abbreviations: AM, Australian Museum; ANSP, Academy of Natural Sciences, 

 Philadelphia; BM(NH). Natural History Museum, London; NM. Natal Museum; 

 NMW, National Museum of Wales. 



are known from around the continental margins of the Indo-W. 

 Pacific Ocean (Wells, 1994; Taylor & Wells, 1994; Sysoev, 1996). 

 The relationships of the Crassispirinae to other conoideans are 

 uncertain and those of the genera within the Crassispirinae are 

 totally unknown; thus it is highly uncertain how the common genera 

 of the East Pacific such as Crassispira, Hindsiclava etc. are related 

 to some of the common Indo-W. Pacific genera such as Inquisitor, 

 and Funa. 



In order to establish the anatomical range of the Crassispirinae 

 and evaluate characters which might be used to establish relation- 

 ships both within the group and with other conoideans, we studied 

 by serial sections the foreguts of 35 species. These were collected 

 within the East Pacific, Caribbean, Mediterranean and Indo-Pacific 

 provinces from a wide variety of habitats ranging from intertidal to 

 abyssal depths. Additionally, we used Gemmula deshayesi (sub- 

 family Turrinae) as an outgroup. This is the most extensive 

 comparative study yet attempted of the anatomy of any group of 

 Conoidea. 



MATERIAL AND METHODS 



Details of all the species used in the study are listed in Table 1 . For 

 all species longitudinal serial sections were made of the foregut, cut 

 at 8 urn and mostly stained in green Masson's trichrome. Radulae 

 were cleaned with a dilute sodium hypochlorite solution and exam- 

 ined by SEM. 



ANATOMICAL DESCRIPTIONS 



Abbreviations used in the anatomical figures 



asg acinous salivary gland 



asgm modified acinous salivary gland 



be buccal cavity 



blp buccal lips 



blpi invertible buccal lips 



bm buccal mass 



bs buccal sac 



bt buccal tube 



bts buccal tube sphincter, 



btsa anterior sphincter of the buccal tube 



btsi intermediate sphincter of the buccal tube 



cf circular fold, surrounding opening of buccal sac 



cfbt circular fold of the buccal tube 



con circumoesophageal nerve ring 



ct connective tissue of the buccal mass wall 



ctl connective tissue layer of the muscular bulb 



ebt sac-ike enlargement of the buccal tube 



ep epithelial pad 



gre glandular part of the rhynchodeum 



iep invaginated epithelium of the proboscis tip 



ipt inverted proboscis tip 



ire non-glandular invertible part of the rhynchodeum 



lmb lumen of the muscular bulb 



mebt thickened muscular wall of sac-like enlargement of buccal 



tube 



ngre non-glandular non-invertible part of the rhynchodeum 



oe oesophagus 



oel oesophageal loop 



p proboscis 





