152 



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



ing foregut anatomy. Additionally, brief descriptions of the 

 shell characters used are given in Appendix 1. 



Outgroup 



The relationships of the Conoidea to other Neogastropoda 

 are very unclear and there is no obvious sister group. In our 

 various analyses we used two outgroups. The first is Bentho- 

 bia the most primitive non-coniodean neogastropod known 

 (Kantor, 1991). This gastropod has a buccal mass situated at 

 the base of the proboscis, a muscular connection between the 

 radular retractors and columellar muscles, and a full set of 

 glands connected with the oesophagous. Additionally, we 

 used as a second outgroup a hypothetical ancestral taxon 

 consisting of the underived states, where known, of all the 

 characters used in the analysis. 



Method 



The data were analysed using version 3.0 of the PAUP 

 program (Swofford, 1991). Characters 2, 7, 23, 25, 26, 27, 28, 

 29, 30, 31, 33, 34, 35, 36, 37, 38, 41, 42 were treated as 

 unordered. The matrix of taxa and character states is shown 

 in Table 3. 



Results of phylogenetic analysis 



Although we have used many new anatomical characters, the 

 cladistic analysis gave rather disappointing results. The taxa 

 of 'lower conoideans' especially, were rather poorly resolved 

 with major branches supported by rather few weak charac- 

 ters. Additionally, small adjustments to the data set produced 

 rather large changes in tree topography and the number of 

 alternative trees generated. 



Despite these limitations we thought it worthwhile to 

 present the results of our analysis, which is the first for the 

 Conoidea to use anatomical characters. Future work will 

 extend on the character set shown in Table 3 and hopefully 

 improve the resolution of the analysis. 



A heuristic search with the matrix shown in Table 3 and 

 with Ancestor as outgroup, produced over 900 equally parsi- 

 monious trees (189 steps; consistency index = 0.296; 

 homoplasy index 0.704). A 50% majority rule consensus 

 cladogram derived from these trees is shown in Fig. 27. 

 Despite the large number of trees generated, most of the 

 trees are very similar to each other and most of the branches 

 are supported in 75-100% of the trees. Autapomorphies of 

 the internal nodes are listed in Table 4. 



The least-derived group are the two species of Drilliidae, 



Table 3. Matrix of taxa and character states used in the analysis. See Table 2 for further details of characters and caption to Fig. 27 for taxon 

 abbreviations. 



Ancestor 















(1 



7 







(1 































II 



(1 



1) 



II 



II 



(1 



II 



(1 



9 



n 



(I 



7 



7 



7 



(l 















1) 



7 



1 



7 



1 



3 







7 



7 







Benthobi 







1 















1 







































1 















1 











7 



1 







1 



7 



7 



2 



7 



7 







1 



4 















3 



















PseudomP 























1 



























1 











1 















1 















1 







1 



7 



7 























4 



1 







1 



1 







1 



1 







StrictiP 































7 



7 











1 



1 











1 















1 











7 



1 







1 



7 



7 



2 



7 



7 











4 















1 







1 



1 







ClavusUn 



























2 



















































1 











1 















7 



7 



















1 



4 















1 







1 



1 







SplendrC 



























1 



1 















































1 











1 















7 



7 























4 















1 







1 











ClavatuD 







2 















1 



1 























1 











1 















1 











2 



1 



1 



7 



1 



7 



















1 















1 



1 







1 



1 



1 



ClavatuC 























1 



1 



















1 



1 











1 



1 











1 











2 



1 



1 



7 



1 



7 



1 















1 















1 



1 







2 



1 



1 



ClionelS 























1 



2 



















1 



1 











1 















1 











2 



1 



1 



7 



1 



7 



1 















1 



4 















1 







1 



1 



1 



ToxicliT 























1 























1 



1 



1 







1 















1 











7 



1 



2 



7 



7 



1 























4 











1 



1 







1 



1 



1 



LophiotL 







1 



















1 















1 







1 



























1 











7 



1 



1 



7 



1 



7 



1 































2 



2 







2 











PolystiA 







1 



















1 























? 



























1 











7 



1 



1 



7 



1 



7 



















7 











1 



2 



2 







2 











TurricuN 























1 



2 















1 



1 



1 



1 







1 















1 











2 



1 



1 



7 



1 



7 



1 































2 



1 







2 



1 



1 



AforiaAb 























7 



1 























1 



1 



































2 



1 



1 



7 



1 



7 







1 



















7 



7 



2 



1 







1 











FunaLati 







1 















1 



2 























1 



1 







1 















1 











7 



1 



1 



7 







7 



1 















1 



4 











1 



1 







2 



1 







VexitomG 























1 



1 



1 



















1 











1 















1 











9 



1 



1 



7 







7 



1 















1 



4 



1 







1 



1 







1 











PilsbryN 























1 



1 



















1 



1 



1 







1 















1 











7 



1 



2 



7 



7 



1 



1 















1 



4 



1 











1 







1 



1 







MicantaP 



























1 























1 





































7 



1 



2 



7 



? 





















7 



4 











1 



2 







1 











BorsoniO 







1 











1 







1 























1 























1 













? 



1 



2 



7 



7 



1 

















7 















1 



1 







1 



1 







TomopleV 







2 



















2 























1 















1 



7 



7 



1 









7 



1 



2 



7 



7 



1 

















7 



4 



1 



1 







1 







2 











TropidoF 







1 



















1 























1 



1 















1 







1 









7 



1 



2 



7 



7 





















7 



2 











1 



1 







1 



1 







Ophiodel 







1 



















1 























1 



1 

































7 



1 



2 



7 



9 



1 











1 





? 















1 



1 







1 







1 ? 



AnarithM 







1 















1 



? 



7 



















1 



















7 







1 









7 



1 



2 



7 



7 





















7 



4 



1 











3 



1 











1 ? 



GlyphosC 







1 



















7 



7 



















1 



















1 







1 









7 



1 



2 



7 



7 



1 

















7 



4 



1 







1 



1 



1 



1 







1 ? 



EucithaS 































1 







1 



1 







1 























1 



1 









7 



1 



2 



7 



7 





















7 



4 



1 



1 







1 



1 



1 



1 



1 ? 



MangeliN 























1 



1 







1 



1 



1 







1 



1 



















1 



1 









? 



1 



2 



? 



7 















1 





7 



4 



1 



1 



1 



1 







1 







1 ? 



MangeliP 























1 



1 











1 



1 







1 



1 



















1 



1 









7 



1 



2 



7 



7 















1 





7 



4 



1 



1 



1 



1 







1 







1 ? 



OenopotL 







1 



















7 























1 



1 























1 









? 



1 



2 



7 



7 















1 





7 



4 







1 



1 



1 







1 











PhilberP 

















1 























1 







1 



1 



1 















1 



1 









7 



1 



2 



? 



7 











7 



7 





7 



4 



1 



1 



1 



3 



1 



1 







1 ? 



PhilberL 





















? 



7 



7 



7 















1 



1 











1 



7 



? 



1 



1 





7 



1 



7 



7 



7 



? 



2 



7 



? 





7 



4 



1 



1 



1 



3 



1 



1 







1 ? 



DaphnelR 













1 







1 



























1 



1 



1 











7 



1 



1 



1 





7 



1 



2 



7 



? 











1 



2 





7 



4 



1 



1 











1 



1 







1 ? 



GymnobeE 













1 



1 







7 



7 



7 















1 



1 



















1 



1 









7 



1 



2 



7 



7 







2 



7 



? 





7 



4 



1 



1 



1 











1 



1 



1 ? 



TeretioL 













1 



1 







7 



? 



7 















1 















1 



7 



7 



1 



1 





7 



1 



7 



7 



7 



? 



2 



7 



? 





7 



4 



1 



1 



1 















1 



1 ? 



AbyssobA 















1 



1 







? 



7 



7 















1 



1 











1 



? 



? 



1 



1 





7 



1 



7 



7 



? 



7 



2 



7 



? 





? 







? 



7 



2 















1 



1 ? 



Benthofa 







1 























1 



















1 





































7 



1 



2 



7 



7 



1 

















7 







7 







1 











1 











GenotaNi 































1 



















1 



7 























1 









7 



1 



2 



? 



7 



1 

















? 







1 







1 



1 







1 



1 



1 ? 



ThatcheM 







1 











































1 























1 



1 









7 



1 



2 



? 



7 











1 



2 





7 



2 



1 



1 



1 











1 







1 ? 



TaranisM 







2 







1 











? 



7 



? 















1 



1 











1 



? 



? 



1 



1 





7 



1 



7 



7 



7 



? 



2 



? 



7 





? 



4 







1 



1 



1 















1 ? 



ConusVen 



















1 











1 



















1 



1 















1 

















7 



1 



2 



7 



? 



1 

















7 



1 



1 











3 







1 











PervicaT 



1 











1 











7 



? 



7 















1 



1 



1 



1 















1 







7 



? 



1 







1 



? 



? 



2 



? 



? 







1 



3 















3 







1 











HastulaB 



1 







1 







1 







1 























1 







































7 



1 







7 



7 



7 























3 



1 











3 







2 











DuplicaC 



1 



2 



1 



1 











7 



7 



? 















1 



1 



1 



















1 











7 



1 







1 



7 



? 



1 



7 



? 











3 















3 







2 











