26 



convex; separated from the posterior section by a weak impression. Posterior section 

 present; weakly convex; depressed below the anterior section. 



Cubital Basivenalia large; ovoid to deltoid; broadly fused. Posterior margin of BCuA fused 

 with anterior margin of BCuP. Suture line weak. BCuA about the same size as BCuP; 

 deltoid; convex; oriented weakly postero-distad; strongly sclerotized; an extension enters 

 CuA along the anterior margin, just proximal of the brace-vein junction; curved anteriad. 

 BCuP large; oriented distad; weakly sclerotized. Proximal margin forms a convex ridge. 

 Embayment shallow. - Cubitus Anterior fused to BCuA; only partially separated by a 

 weak sclerotized groove. Anterior section interrupted by the BMP-CuA Brace. 



Basalare (Fig. 37) 



Head - HP lobe digitate; very thin; small. Apex clavate; weakly curved ventrad. Base 

 adjacent to, but separated from, BScP lobe. BScP lobe claviform; convex; bulbous; 

 obscures neck: much larger than HP lobe. Dorsal surface moderately polished. Ventral 

 surface rough. - Posterior Subcostal Basivenale deltoid; polished. 



Discussion 



Monophyly of the Diphyllostomatidae is supported by the fact that all of the taxa in this 

 family share the following five apomorphic character states of the wing articulation and 

 wing base: 



1. lAx: the distal margin of the neck is deeply, but narrowly concave; 



2. 2 Ax: the dorso-proximal lobe base is greatly enlarged anteriorly and posteriorly, 



3. but medially to apically reduced, 



4. the distal margin of the subalare tendon attachment point is strongly separated from the 

 dorsal surface of the ventro-distal lobe, the latter extends posteriad beneath the former; 



5. 2BP: BCuA curves anteriad. 



Analysis of wing articulation and wing base characters indicates that Diphyllostomatidae 

 are a distinctive family warranting familial status and the sister group of Lucanidae 

 (Browne & Scholtz 1995). Diphyllostomatidae share six apomorphic character states of 

 the wing articulation and wing base with Passalidae, Lucanidae, Glaphyridae, Trogidae, 

 Bolboceratidae and Pleocomidae (Browne & Scholtz 1995). 



Lucanidae 



Introduction 



The Lucanidae are a large family with five large subfamilies and several small ones which 

 include approximately 100 genera and 750 species with virtually world-wide distribution. 

 Lucanidae are a well defined family whose monophyly is supported by many derived 

 characters (Crowson 1967, 1981; Howden 1982; Scholtz 1990). Caveney (1986) suggested, 

 based on synapomorphic ommatidium structure, that Lucanidae are probably more closely 

 related to Diphyllostomatidae. This relationship was reiterated by Scholtz (1990) and is 

 supported by wing articulation and wing base characters (Browne & Scholtz 1995). 



