34 



Basalare (Fig.5 1 ) 



Head - HP lobe convex; dorsally elevated from neck; continuous with neck. Apex not 

 curved. Dorsal and ventral surfaces rough. BScP lobe claviform; convex; weakly elevated 

 from neck. Dorsal surface lenticular; slender; polished; lies below the ventral margin of 

 the HP lobe. Ventral surface polished. - Posterior Subcostal Basivenale weakly square- 

 shaped; polished. 



Discussion 



Monophyly of the Glaphyridae 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 arch apex of the tail is strongly curved ventrad; 



2. 2Ax: the apex of the dorso-distal ridge is extremely broad along its entire length, extends 

 anteriorly, and curves very weakly proximally from the dorso-distal lobe base, 



3. the subalare tendon attachment point is very short, very broad, and apically shallowly 

 and narrowly concave; 



4. IBP: the BR embayment is of normal size, but is more strongly deltoid, with all margins 

 straight; 



5. 2BP: the medial section of BMP is distinct from the remainder of BMP as an elevated 

 tube-shaped structure. 



The phylogenetic position of Glaphyridae within the Scarabaeoidea has been an issue of 

 some debate. This family was once accorded superfamily status due to its unusual 

 distinctiveness, but Hinton (1967) more realistically accorded family status. Based on male 

 and female genitalia, Zunino (1988) claimed that glaphyrids occupy an intermediate 

 position between Melolonthinae and Scarabaeidae. Holloway (1972) noted that, according 

 to Crowson's (1967) key, glaphyrids occupy a position close to Diphyllostomatidae. 

 d'Hotman & Scholtz (1990a,b) and Scholtz (1990) presented convincing evidence, from 

 numerous adult and larval characters, which suggests that glaphyrids are archaic. More 

 importantly, they found that the basal piece of the male genitalia is very similar to some 

 lucanid genera. 



Analysis of wing articulation and wing base characters indicates that Glaphyridae are the 

 sister group of Trogidae + Bolboceratidae + Pleocomidae, with whom they share a single 

 derived state of the hind wing base (Browne & Scholtz 1995). 



Trogidae 



Introduction 



The Trogidae are a monophyletic, cosmopolitan family, consisting of three genera and 

 about 300 species. Trogids have wide distribution, but occur mainly on the southern 

 continents in sandy arid regions (Scholtz 1986). Trox Fabricius occurs naturally in Europe, 

 North America and Africa. Omorgus Erichson occurs in South and North America, 



