63 



projects posteriad from neck. Dorsal surface ovoid; strongly convex; polished; elevated 

 from neck, but lies below the HP lobe. Ventral surface not polished. - Posterior Subcostal 

 Basivenale deltoid; polished. 



Discussion 



Ochodaeidae do not display any autapomorphic character states of the wing articulation 

 or wing base (Browne & Scholtz 1995). They share 12 apomorphic character states of the 

 wing articulation and wing base with Geotrupidae, Hybosoridae and Ceratocanthidae 

 (Browne & Scholtz 1995). 



Carlson & Ritcher (1974) implied that close relationship exists between Ochodaeidae and 

 Hybosoridae, Iablokoff-Khnzorian (1977) suggested that Aclopidae are more closely 

 related to Ochodaeidae, Crowson (1981) implied relationship between Ochodaeidae, 

 Hybosoridae and Geotrupidae, and Lawrence & Newton (1982) concluded that 

 Ochodaeidae, Hybosoridae and Ceratocanthidae appear to be closely related. Scholtz et 

 al. (1988) corroborated Lawrence & Newton (1982) and proposed that ochodaeids are the 

 sister group of Hybosoridae-Ceratocanthidae, a view confirmed by Browne (1993) and 

 reiterated here. 



Scarabaeidae 



The Scarabaeidae are a large family consisting of the following taxa: Aphodiinae, 

 Aegialiinae, Aulonocneminae, Orphninae, Melolonthinae, Acoma, Oncerinae, 

 Chasmatopterinae, Hopliinae, Rutelinae, Dynastinae, Cetoniinae, Trichiinae and Valginae 

 (Browne 1993). The family is characterized by the loss of functional spiracles, 

 membranization of pregenital segments, the loss of one of the free anal veins in the wing, 

 and by larvae with 4-jointed antennae and the absence of stridulatory organs on the legs 

 (Lawrence & Newton 1982). 



All scarabaeids share the following 23 apomorphic character states of the wing articulation: 



1. lAx: head proximal margin reduced, 



2. dorsal surface posteriorly reduced, 



3. anterior surface narrow, 



4. ventral projection long and narrow, 



5. embayment mesad, 



6. neck long; 



7. 2Ax: dorso-proximal and dorso-distal margins of dorso-proximal enlarged proximally 

 and distally, 



879. antero-median to posterior sections of the dorso-proximal ridge enlarged laterad over 

 dorso-distal ridge, 



10. anterior section of dorso-distal ridge basally curved distad, 



1 1 . posterior wing process ridge-shaped, enlarged to occupy both the anterior margin of 

 the ventro-proximal lobe to the postero-proximal corner of subalare tendon attachment 

 point, 



