22 



distinct from, the distal margin of 2BP; intervening membrane sclerotized. Anterior section 

 continuous with MP. Posterior section continuous with CuA. - BMP-BCuA Brace absent. 



Medial Basivenalia massive; strongly sclerotized. BMA broadly scaphoid: very large: 

 angled postero-proximad; flat but elevated above and separated from BMP by a deep 

 groove: lies anteriad, but not proximad, of BMP: fully fused to BMP. Anterior margin 

 concave. Proximal arch planate: straight: broad: enlarged antero-proximally. Distal arch 

 extremely long: allantoid; extends far distad. BMP planate: large: broadly fused to IBP 

 and BCuA: very weakly separated into anterior and posterior sections; discontinuous with 

 brace. Anterior margin concave; fused to IBP. Proximal margin deeply concave. Posterior 

 margin fused to BCuA and CuA; indistinct. Distal margin discontinuous with brace. 



Cubital Basivenalia large; ovoid or weakly deltoid: narrowly fused. Postero-distal margin 

 of BCuA fused to antero-proximal margin of BCuP. Suture line very weak but present. 

 BCuA about the same size as BCuP; oriented postero-distad. Distal margin with a very 

 deep embayment. Embayment strongly sclerotized. BCuP oriented antero-distad. - Cubitus 

 Anterior basally connected to BCuA along the postero-proximal margin. Anterior 

 interrupted by the BMP-CuA Brace. 



Basalare (Fig. 30) 



Head - HP lobe short; broad; weakly elevated. Dorsal surface truncate; weakly polished. 

 BScP lobe large: broad: oriented disto-ventrad: laterally ovoid. Dorsal surface broadly 

 ovoid: polished. Ventral surface polished. - Posterior Subcostal Basivenale ovoid; long; 

 polished; oriented proximo-ventrad. 



Discussion 



As with other morphological characters (see Scholtz 1990). passalids exhibit numerous 

 autapomorphic wing articulation and wing base character states. Monophyly of the 

 Passalidae is supported by the fact that all of the taxa in this family share the following 

 13 apomorphic character states of the wing articulation and wing base: 



1. lAx: the proximal arch is very strongly reduced along all margins, 



2. the distal arch strongly expanded anteriorly and distally: 



3. 2 Ax: the distal lobe is strikingly reduced posteriorly and apically. 



4. arises from the extreme anterior section of the ridge; 



5. 3 Ax: the tail is very weakly sclerotized. 



6. the demarcation between sclerites absent. 



7. the tail extends anteriad below the posterior margin of the neck, 



8. is separated from the head+neck by a tube-shaped stem; 



9. IBP: the proximal margin of BScA is interrupted by a deep and broad groove, giving 

 it a bi-lobed appearance which provides additional support for the lAx head teeth. 



10. the proximal and distal sections of BScA are indistinct from each other, 



11. the proximal arch of BR is extremely broad, very long and rectangular, 



12/13. BR. br and the br projection apices are fused to. and indistinguishable from BScA. 



