162 



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



Subfamily Daphnellinae Deshayes, 1863 



Small to moderately large shells (usually 5-15 mm, deep-sea 

 species larger, up to 95 mm). Anal sinus sutural, shaped as a 

 reversed-L, or on the upper shoulder and varying in depth. 

 Sculpture variable, usually cancellate or with predominant 

 spirals, and often with a smooth shoulder. Protoconch usually 

 multispiral, rarely paucispiral, typically diagonally cancel- 

 lated, although some genera have spiral or axial ribbing. 

 Operculum absent. Egg capsules dome-shaped operculate. 



Radula. Radular teeth with large solid base and barbed or 

 unbarbed tips, tooth cavity opens laterally at the base. 

 Radula absent in some species. 



Foregut. Rhynchodeal introvert present in many species. 

 Rhynchodeal septum present in some species. Proboscis 

 usually short, often absent. Buccal mass basal. Radula appa- 

 ratus absent in many species, vestigial in Gymnobela emer- 

 toni. Radial muscles present in the rhynchodeal wall in 

 radula- and proboscis-less species. Buccal lips well devel- 

 oped, can be intverted into the buccal cavity. Salivary glands 

 paired tubular or absent. Accessory salkivary glands absent. 

 Venom apparatus absent in many species. In Daphnella 

 reeveana the anterior part of venom gland is ciliated. Muscu- 

 lar bulb can be single layered. 



Remarks. Although Thatcheria is sometimes classified in a 

 separate subfamily Thatcherinae, we failed to find any ana- 

 tomical or shell characters which would justify separation 

 from the Daphnellinae. 



advised on problems of phylogenetic analysis. Yuri Kantor and John 

 Taylor are grateful to the Royal Society, London and the Russian 

 Academy of Sciences, Moscow for grants which enabled them work 

 in London and Moscow respectively. Yuri Kantor and Alexander 

 Sysoev gratefully acknowledge a grant from the Soros Foundation 

 and the Russian Academy of Natural Sciences. 



APPENDIX 1 



Features of the shell 



Shell characters are still important for the systematics of 

 Conoidea, and thus should be included in the analysis. 

 However, there is probably no shell character which is 

 diagnostic of any single group. Moreover, there has been no 

 analysis of the adaptive or evolutionary significance of these 

 shell features. Nevertheless, a few shell characters appear to 

 be useful for the separation of clades. 



Shell shape 



This character which is concerned with overall shell shape is 

 the most subjective. We recognise five basic shell shapes: 

 1, fusiform shell; 2, cone-shaped shell; 3, turreted shell; 

 4, terebriform shell; 5, a large group of 'intermediate' states, 

 'biconic-fusiform', 'ovate-biconical', 'ovate-fusiform', 'clavi- 

 form', etc. characterized by rounded outlines of the shell, 

 which is more or less oval in its general profile. 



Subfamily Taraninae Casey, 1904 



Shell very small (up to 6 mm), ovate-fusiform. Anterior canal 

 rather short. Sculpture well developed. Anal sinus very broad 

 and shallow, situated on the shoulder or immediately below 

 it. Protoconch paucispiral, finely spirally striated, or with 

 spirally aligned granules. Operculum and radula absent. 



Foregut. Rhynchostomal sphincter absent, no radial 

 muscles in rhynchodeal wall. Proboscis absent. Buccal mass 

 undefined. Salivary glands absent. Venom apparatus absent. 



Remarks. This monotypic radula-less subfamily was rein- 

 stated (Kantor & Sysoev, 1989) because it differs in shell 

 characters from any other turrids lacking a radula. However, 

 the very simplified morphology makes the evaluation of the 

 status of the subfamily difficult. For the present we conserve 

 the subfamily, but are unsure of its status. 



Acknowledgements. For gifts and loan of material without which 

 this study could not have been completed, we are extremely grateful 

 to J.H. McLean, the late V. Maes, R.N. Kilburn, F.E.Wells, I. 

 Loch, R. L. Shimek, P. Bouchet, S. Gofas, B. Morton, W. F. 

 Ponder, G. Rosenberg, and B. Marshall. Other material was 

 obtained from the Institute of Oceanology, Academy of Sciences, 

 Moscow; Zoological Institute, Academy of Sciences, St Petersburg, 

 Zoological Museum, Moscow State University and we are very 

 grateful to the curators L.I. Moscalev, A.N. Mironov, B.I Sirenko 

 and D.L. Ivanov for their kind permission to use this material. David 

 Cooper expertly prepared many of the serial sections of gastropod 

 foreguts. Some unpublished material was made available by John 

 Miller and A.I Medinskaya. The optical photomicrographs were 

 made by Peter York. Andrew Smith and David Reid patiently 



Number of protoconch whorls 



Two types of protoconch can be recognised; the paucispiral 

 and multispiral. These types of the protoconch were into- 

 duced into turrid systematics by Powell (1942, 1966) and they 

 are widely used in taxonomy. Generally, this subdivision 

 coincides with that between planktotrophic and non- 

 planktotrophic modes of larval development, although there 

 are many exceptions to the rule among turrids (Bouchet, 

 1990). The character is considered as being of little phyloge- 

 netic importance (Bouchet, 1990), but a predominance of a 

 single type of the protoconch can be noted in some taxa. For 

 instance, most Daphnellinae and Conidae have multispiral 

 protoconchs, whilst the paucispiral type is a typical of the 

 Oenopotinae (Bogdanov, 1990). Protoconchs with 1-2 whorls 

 are here considered as paucispiral, and these with two or 

 more whorls as multispiral (Bouchet, 1990). 



Sculpture of the protoconch 



The pattern of protoconch sculpture has been widely used in 

 conoidean taxonomy since Powell (1942, 1966). Turrids have 

 a very wide variety of protoconch sculpture and at present, 

 we are unable to classify them into clearly defined types. 

 Thus we recognize only two major states of the character; 

 firstly protoconchs lacking or with only weakly defined sculp- 

 ture and secondly, protoconchs with well developed sculp- 

 ture. Some higher taxa may be characterized by the presence 

 or absence of protoconch sculpture. For example, the closely- 

 related Turricula and Clavatula usually possess a smooth 

 protoconch, whilst in the Turrinae it is usually axially costate. 

 The only type of the protoconch sculpture characteristic of a 

 single subfamily is the 'diagonally cancellated' form found 



