38 



margin forms the anterior half of the BMP-BCuA brace; elevated from surrounding 

 membrane. Antero-distal margin aquiline; with many overlapping plates. Po stem -distal 

 section forms the posterior half of the BMP-BCuA brace; as equally elevated as the antero- 

 distal section. 



Cubital Basivenalia broadly fused. Posterior margin of BCuA fused with anterior margin 

 of BCuP. Suture line present. BCuA slightly smaller than BCuP; deltoid; strongly convex; 

 oriented distad; strongly sclerotized; lies proximal to the base of CuA. BCuP weakly 

 rectangular; less convex than BCuA; oriented distad; moderately sclerotized. Distal 

 embayment weak. - Cubitus Anterior arises from within a toroid concavity in BCuA. 



Basalare (Fig. 58) 



Head - HP lobe weakly claviform; convex; elevated from neck; continuous with neck. 

 Dorsal surface polished. BScP lobe moderately claviform; concave; weakly elevated from 

 neck. Dorsal surface lenticular; polished. Ventral surface rough. - Posterior Subcostal 

 Basivenale deltoid; polished. 



Discussion 



Monophyly of the Trogidae is supported by the fact that all of the taxa in this family share 

 the following four apomorphic character states of the wing articulation and wing base 

 (Browne et al. 1993 only recorded three, autapomorphy no. 1 is new): 



1. 2Ax: the subalare tendon attachment point is short, narrow and apically rounded; 



2. 2BP: with transverse crimps on the medial vein, mesal of the medial bridge and distal 

 of 2BP. 



3. the BMP-CuA brace is either reduced or modified (both are autapomorphic) depending 

 on the genus; 



4. RP3+4 is lost. 



Trogidae share six apomorphic character states of the wing articulation and wing base 

 with Passalidae, Diphyllostomatidae, Lucanidae, Glaphyridae, Bolboceratidae and 

 Pleocomidae (Browne & Scholtz 1995). Analysis of wing articulation and wing base 

 characters indicates that Trogidae occupy an intermediate position between Glaphyridae 

 and Bolboceratidae + Pleocomidae, more closely related to the latter (Browne & Scholtz 

 1995). 



Bolboceratidae 



Introduction 



The Bolboceratidae are a large, well-defined family whose monophyly is supported by- 

 numerous derived characters (Scholtz 1990; Browne 1991a,b; Scholtz & Browne, in press). 



Bolboceratidae, previously classified as one of four subfamilies of Geotrupidae. have 

 recently been elevated to family status since there is little evidence for the traditional view 

 that Geotrupidae constitute a monophyletic assemblage (Ritcher 1969a.b: Scholtz 1990: 



