19 



Dorsal surface ovoid; weakly convex; polished: not depressed from neck. Ventral surface 

 polished. - Posterior Subcostal Basivenale deltoid, polished. 



Discussion 



Glaresidae exhibit the generalized scarabaeoid state of all characters, including those of 

 the wing articulation and wing base which leaves no doubt that Glaresidae are the most 

 archaic family in the Scarabaeoidea, quite likely very close in structure to the ancestral 

 scarabaeoid (Scholtz et al. 1994), and the sister group of all other scarabaeoids (Browne & 

 Scholtz 1995). This family does not display any autapomorphic characters of the hind 

 wing articulation or wing base (Browne 1991a. 1993; Scholtz et al. 1994; Browne & 

 Scholtz 1995). 



Passalidae 



Introduction 



The Passalidae are a virtually cosmopolitan family with approximately 40 genera and 500 

 species which are most abundant in tropical regions (Reyes-Castillo 1970). 

 The Passalidae are a well-defined family whose monophyly is supported by numerous 

 derived characters and there is little doubt that it is one of the more archaic scarabaeoid 

 families (Reyes-Castillo 1970). The numerous and unusual derived features exhibited by 

 Passalidae have led some workers to suggest that they be excluded completely from the 

 Scarabaeoidea (Reyes-Castillo pers. comm. 1990). This family is comprised of two 

 subfamilies, Aulacocyclinae and Passalinae (Reyes-Castillo 1970). 



Hind Wing Articulation Description 



First Axillary (Fig. 24) 



Head - Dorsal surface normal size. Antero-dorsal margin normal width; weakly deplanate: 

 oriented weakly postero-distad. Antero -proximal margin very strongly enlarged ventrally. 

 Po stew -proximal margin weakly enlarged proximally. FSc2 base normal width. Apex 

 rounded: oriented postero-distad. Anterior surface broad; very long; not waisted medially. 

 FScl very weak. Ventral projection tapers from the base to apex: short but of average 

 width: deeply concave: oriented disto-ventrad and curved posteriad. Apex narrow. 

 Concavity located in the preapical area; surrounded by three unequally strong ridges. Distal 

 embayment oriented dorsad. FSc2 deltoid; broad; very convex; oriented distad and weakly 

 dorsad. Dorsal margin not enlarged. Head and neck dorsal surface weakly curved 

 proximad. 



Neck normal width and length: weakly oriented antero-distad; broadly articulated with 

 2Ax. Proximal margin straight. Distal margin weakly concave. Distal embayment concave; 

 moderately narrow. 



Tail - Dorsal view: Proximal arch strongly reduced along all margins. Antero-proximal 

 margin concave. Dorsal surface weakly concave. Postero-proximal margin weakly convex; 

 weakly recurved. Articulation with PRR strong. Posterior margin concave. Distal arch 



