so 



curved antero-ventrad beneath the BR proximal arch. Distal arch reduced; indistinct: 

 continuous with the anterior margins of both BMP and the BMP-BCuA brace. BMP 

 junction with BMA continuous and narrowly tubular; fused to the BMP-BCuA brace to 

 form a narrow, looping tube; separated from both IBP and BCu by membrane; anterior 

 margin strongly convex; extremely narrow; strongly elevated dorsad. Postero-distal section 

 reduced. 



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

 margin of BCuP. Suture line present. BCuA very small and ovoid; convex; oriented antero- 

 distad; moderately shifted anteriad to occupy the postero-distal section of 2BP: weakly 

 sclerotized. Proximal margin proximad of CuA antero-proximal margin. BCuP large; 

 deltoid; moderately short; very weakly convex; oriented postero-distad; very weakly 

 sclerotized. Distal embayment absent. - Cubitus Anterior continuous with BCuA. 



Basalare (Fig. 128) 



Head - HP lobe slender; continuous with neck. Apex narrowly rounded. Dorsal surface 

 strongly elevated from neck; polished. BScP lobe moderately broad; claviform; projects 

 dorsad and posteriad from neck. Dorsal surface ovoid; convex; polished; elevated from 

 neck but not as strongly as the HP lobe. Ventral surface not polished. - Posterior Subcostal 

 Basivenale deltoid; polished. 



Discussion 



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

 subfamily share the following 12 apomorphic character states of the wing articulation and 

 wing base: 



1. lAx: FScl is very strongly elevated, long and oriented antero-dorsad over the base of 

 the ventral projection, 



2. FSc2 is completely planate. 



3. the neck is medially convex, 



4. the distal arch of the tail is strongly reduced anteriorly, strongly elongated distally, 



5. strongly reduced posteriorly; 



6. 2Ax: the posterior section of the dorso-proximal ridge is strikingly enlarged laterad; 



7. FM1+FM2 - together with the proximal extension of the 3 Ax head are highly modified 

 to form a ball-and-socket joint; 



8. 3Ax: the antero-proximal section of the head is narrowly enlarged proximally to form 

 a long extension; 



9. IBP: the proximal arch of BR is strikingly enlarged, 



10. the distal extension of the proximal arch is strikingly enlarged antero- to postero- 

 distally, 



11. the BR embayment is strikingly narrow and falcate: 



12. 2BP: extremely short and narrow. 



Many workers have suggested that scarabaeines occupy an intermediate position between 

 Aphodiinae and Melolonthinae, most closely related to the the former (Howden 1982; 

 Scholtz 1990). Many larval and adult characters support this relationship (Howden 1982) 



