FOREGUT ANATOMY AND CLASSIFICATION OF CONOIDEA 



157 



features were also used. Powell believed that the hypodermic 

 toxoglossate dentition could develop independently in differ- 

 ent lineages and, more importantly, that the appearance of 

 toxoglossate radula was not a significant reason for separating 

 groups at the subfamilial level. As a result, he classified some 

 taxa having quite different radular types (including both solid 

 and hollow marginal teeth) within a single subfamily. 



Morrison (1966) followed Thiele in recognizing a funda- 

 mental difference between groups with solid (= nontoxoglos- 

 sate) and hollow (= toxoglossate) marginal teeth. He 

 suggested a separation at the family level using the families 

 Turridae (with subfamilies Drilliinae, Clavatulinae and 

 'Lophiotominae or Crassispirinae'), Mangeliidae and 

 Tseudomelatominae'. 



The subfamily classification of Powell was considerably 

 revised by McLean (1971), who adhered strictly to the 

 principle of grouping together genera with the same type of 

 radula. He also added six subfamilies to Powell's classifica- 

 tion; three of these being described as new (Clathurellinae 

 H. & A. Adams, erroneously). Several subfamilies were 

 recognised (or retained after Powell) on shell characters, 

 but which share the same radular type, and some of these 

 seem to be rather poorly documented. However, McLean's 

 classification which includes 15 subfamilies is at present the 

 most detailed and well developed. 



In a continuing series of papers concerning South African 

 Turridae, Kilburn (1983, 1985, 1986, 1988), adopted a prag- 

 matic approach (Kilburn, 1983 p. 550 ' . . . any practical 

 subdivision is better than none ...'), and revised to some 

 extent the composition of subfamilies which he studied. He 

 also synonymized the Diptychomitrinae (= Mitrolumninae = 

 Mitromorphinae) with the Borsoniinae. 



Bogdanov (1986, 1987, 1990) described a new subfamily 

 Oenopotinae separating the operculate Oenopota and its 

 relatives from the Mangeliinae. Additionally, the subfam- 

 ily Taraninae was recently re-instated (Kantor & Sysoev, 

 1989). 



Some nomenclatural changes in the names and authorships 

 of several subfamilies were made by Cernohorsky (1972, 

 1985, 1987), and Ponder and Waren (1988). 



A different viewpoint was taken by Bouchet and Waren 

 (1980) in their study of North Atlantic deep-sea Turridae. 

 They avoided the use of any subfamilial divisions, considering 

 the present classification of Turridae to be artificial and based 

 mainly on (p. 5) ' . . . more or less randomly selected shell 

 characters'. 



At present there is no completely agreed classification of 

 Turridae, nor is there any agreement on which are the 

 taxonomically important characters. The existing variants of 

 turrid classification are based almost exclusively on shell, 

 radular and opercular features. 



The Terebridae have similarly been classified mainly on 

 shell characters. H. & A. Adams (1853) and Cossmann 

 (1896) divided the Terebridae into two subfamilies, includ- 

 ing the Pusionellinae as the second subfamily. Pusionella is 

 now known to belong to the turrid subfamily Clavatulinae. 

 A separate family, the Pervicaciidae, was proposed by 

 Rudman (1969) for Pervicacia tristis. However, Bratcher & 

 Cernohorsky (1987) included Pervicacia and similar forms 

 in the Terebridae. Taylor (1990) confirmed the distinctive- 

 ness of Pervicacia, and showed that many other terebrids 

 should be included in the family Pervicaciidae. 



The Conidae have long been considered as a fairly homo- 

 geneous group, the main problems have concerned the limits 



of the family and whether taxa such as Cryptoconus, Conor- 

 bis and Genota should be included. Cossmann (1896) for 

 example, included them in the subfamily Conorbinae within 

 the Conidae, whilst Powell (1966) includes this subfamily in 

 the Turridae. 



New classification proposed 



As a result of our analysis of foregut characters throughout all 

 the conoidean higher taxa we propose a new classification of 

 the superfamily. This classification represents a rather con- 

 servative compromise position. Although in principle the 

 classification should be based upon the results of the phyloge- 

 netic analysis, we were constrained by the rather poor 

 resolution obtained with our data set. Moreover, only a 

 rather small subset of conoidean species have been examined 

 in any detail. Information from taxa not included in the 

 cladistic analysis (mainly radular characters) has also been 

 used in constructing the classification. An example of the 

 problem is the family Turridae, which comprises the four 

 subfamilies with wishbone marginal teeth, plus the 

 Zonulispirinae. The cladistic analysis suggests two different 

 clades for these subfamilies. This is certainly possible, but the 

 branches are supported by rather few, and perhaps weak 

 apomorphies. Despite the deficiencies this is the first compre- 

 hensive classification of the Conoidea which includes ana- 

 tomical characters. Below we give descriptions of shell, 

 radula and foregut characters for each of the higher taxa that 

 we recognise. Some of the taxa have only provisional status. 

 For example, the subfamily Clathurellinae has been divided 

 up into five informal groups; it may well be polyphyletic, but 

 we have insufficient evidence to resolve the situation. Simi- 

 larly, we are uncertain of the status of the Conorbinae and 

 Taraninae. 



Summary of proposed classification 



Superfamily Conoidea 



Family Drilliidae (ICZN pending) 

 Family Terebridae 



Subfamily Pervicaciinae 

 Terebrinae 

 Family Pseudomelatomidae 



Family Strictispiridae 



Family Turridae 



Subfamily Clavatulinae 

 Crassispirinae 

 Zonulispirinae 

 Cochlespirinae 

 Turrinae 



Family Conidae 

 Subfamily Clathurellinae 

 Coninae 

 Conorbinae ? 

 Oenopotinae 

 Mangeliinae 

 Daphnellinae 

 Taraninae ? 



