FOREGUT ANATOMY OF CRASSISPIRINE GASTROPODS 



85 



Table 3 Distribution of character states for the crassispirine gastropods 

 examined. Details of characters listed in Table 2. 



Character 



11111111112 

 2345678901 234567890 



Inquisitor 



Hindsiclava Haedropleura 



Funa Vexitomina Cheun 9beia p^ cnobeia 



Fig. 31 Summary of major radular types found amongst the 

 Crassispirinae. Not to scale. 



They may, as in the Columbellidae (Guralnick & de Maintenon, 

 1997), be produced by different secretory cells from 'true' teeth. 

 Character 20: several types of marginal teeth are found within the 

 taxa examined. We have divided these into 6 major types, which 

 are summarized in Fig. 3 1. The Epidi rona type (State ())( Fig. 18a) 

 is a wishbone tooth of the clothes-peg type with the tooth bifurcat- 

 ing into two more or less equal limbs. The Inquisitor type (State 

 1 ) is the most widely distributed and consists of a robust, pointed, 

 major limb with a smaller, slender, secondary limb attached to it. 

 The Funa /Vexitomina type (State 2) consists of a long tooth with 

 a broad, pointed blade and a narrow shaft, with a very thin 

 accessory limb attached to the edge of the shaft. In the 

 Haedropleura type (State 3), the major limb has a trowel-like 

 distal end and a narrow shaft, with a slender accessory limb. The 

 Hindisclava type (State 4) has a flat, pointed, major limb and a 

 long, slender, detached 'handle-like' accessory limb. The 

 Cheungbeia type (State 5) is harpoon-like with a pointed, barbed 

 tip and a straight concave shaft, with a thin splint-like secondary 

 limb attached to the edge of the shaft. Finally, the Ptychobela type 

 (State 6) is awl-shaped and hollow, composed of two pieces which 

 are fused along one edge and twisted together. 



Outgroup 



As the outgroup for the Crassispirinae, we chose Gemmula deshayesii 

 from Hong Kong, which has the relatively underived foregut anatomy 

 typical of members of the subfamily Turrinae. Serial sections of the 

 foregut of this species were already available, as well as a published 



Crassispira incrassata 20200100201 



Crassispira maura 10200000001 



Crassispira dysoni 10200001301 



Crass, harford flucki 00000100320 



Crassispira latizonata 2 1 10 10 12 



Crassispira pluto 10000010100 



Crassispira kluthi 10 12 112 11 



Crassispira tepocana 10 2 12 1 12 1 



Crassispira xanti 10 2 1 12 10 1 



Crassispira turricula 2 1 1 10 1 12 



Crassispira apicata 10 1 10 12 



Hindsiclava andromeda 20201201100 



Hindsiclava nulilaris 2021 1000100 



Miraclathurella bicanal. 10 10 2 12 10 



Haedropleura septangularis 10 1 1 10 10 



Naudedrillia prateriss. 10000201200 



Nquma scalpta 10000201310 



Epidirona gabensis 00200001300 



Funa latisinuata 10 1110 10 110 



Funa Jeffrey mi 10 110 1110 



Inquisitor latifasciata 1 10 12 10 1 10 



Inquisitor aff. adenicus 10100001300 



Inquisitor aeimda 00000201220 



Ptychobela suiuralis 21001001020 



Cheungbeia mindanensis 20000100100 



Cheungbeia robusta 10000100200 



Antiguraleus morganus 10 10 1 10 



Guraleus costattu 10 10 1 10 2 



Burchia speclabilis 2 1 10 12 10 3 



Vexitomina garrardi 10201 100300 



Turridrupa bijubata 10 2 10 13 1 



Gemmula deshayesi 001000013 20 



10 10 

 10 10 

 10 10 

 10 10 



13 10 

 10 10 

 10 10 



14 10 

 10 10 

 14 10 

 14 10 

 14 10 

 14 10 

 10 10 

 0300 



13 10 



14 10 

 10 11 

 1110 

 12 10 

 1110 

 10 10 

 10 10 

 1010 

 10 10 

 1110 

 0200 

 10 10 

 1110 

 10 10 

 00 1 1 

 0000 



10 10 1 

 1 000 1 

 00001 

 1 000 1 

 1000 1 

 ??001 

 1000 1 

 1 0001 

 1000 1 

 10011 

 10011 

 10??? 

 10 104 

 1000 1 

 1 0003 

 10003 

 10001 

 10000 

 1 0002 

 1 0002 

 0000 1 



0000 1 



1 000 1 

 1 0006 

 1 0005 

 10005 

 1 0003 

 1 000 1 



0000 1 



1 0002 

 00200 

 1 0200 



description of the anatomy (Taylor, 1994). The characters of the 

 outgroup are recorded with the Crassispirine species in Table 3. 



Character distribution 



The distribution of the character states amongst the species studied 

 along with those of the outgroup are shown in the data matrix (Table 

 3). For Crassispira latizonata and Hindsiclava andromeda we had 

 insufficient material to investigate the radula and there are no 

 published descriptions. In the case of Crassispira pluto, the muscu- 

 lar bulb was lost from the sectioned material. 



Phylogenetic analysis 



This was performed using PAUP version 3.1 and McClade 3.04 for 

 subsequent analysis. Using the data matrix of Table 3 and the 

 heuristic sort option we obtained 24 equally parsimonious trees of 

 104 steps, with a Consistency Index of 0.33 and Retention Index of 

 0.54. A consensus tree generated from the 24 trees is shown in Fig. 

 32a and an example shown in Fig. 32b. Internal nodes of the tree are 

 supported by relatively few characters and there is a high level of 

 homoplasy. Epidirona gabensis and Turridrupa bijubata are undif- 

 ferentiated from the outgroup Gemmula in all trees. Whilst, 

 Crassispira dysoni and Inquisitor aff. adenicus which both possess 

 Inquisitor type teeth, are undifferentiated in the consensus tree but 

 form a weakly supported branch in most trees. Species classified 

 into different subgenera of Crassispira appear in widely separated 

 parts of the tree (Fig. 32b), apart from Crassispira (Crassispira) s. s. 

 and Crassispira (Striospira) which form a monophyletic clade. 



Because we sampled only 24 out of the 46 genera and subgenera 

 and because of the large amount of homoplasy, it would be prema- 



