76 



articulation and wing base with Aegialiinae and Aphodiinae, which together form a 

 monophyletic group (Browne 1993). It is likely that these three taxa together form 

 Aphodiinae, as has been implied by other workers (Koshantschikov 1913; Scholtz 1990). 



Aulonocneminae share 46 apomorphic character states of the wing articulation and wing 

 base with Aphodiinae, Aegialiinae and Scarabaeinae (Browne 1993). This relationship has 

 been suggested previously (Stebnicka 1985; Cambefort 1987; Nel & Scholtz 1990; 

 d'Hotman & Scholtz 1990a). Many morphological characters support this relationship, 

 including those of the mouthparts (Nel & Scholtz 1990), male genitalia (d'Hotman & 

 Scholtz 1990a), and others from the head, thorax and abdomen (Stebnicka 1985). Only a 

 single diagnostic character separates aulonocnemines from Aphodiinae (Stebnicka 1985). 



Scarabaeinae 



Introduction 



The Scarabaeinae are a large subfamily of approximately 4500 species and 200 genera 

 (Halffter & Edmonds 1982). Members of the subfamily are generally known as the true 

 dung beetles. The Scarabaeinae include six tribes. Onthophagini, Coprini, Eurysternini. 

 Oniticellini, Onitini and Scarabaeini. 



The adults and larvae of most Scarabaeinae utilize subliquid and liquid contents of dung 

 and decaying vegetation, described by Stebnicka (1985) as coprophagy or soft saprophagy. 

 For further information on biology, the reader is referred to the recent work by Halffter & 

 Matthews (1966) and Halffter & Edmonds (1982). 



Hind Wing Articulation Description 



First Axillary (Fig. 122) 



Head - Dorsal surface strikingly reduced posteriorly; convex; all margins are equally 

 extremely narrow and very long. Antero-dorsal margin oriented weakly postero-distad; 

 reduced in width; planate. Antero-proximal margin with ventral enlargement reduced. 

 Postero-proximal margin enlargement reduced and absent. FSc2 base normal width. Apex 

 oriented postero-distad; rounded. Anterior surface strikingly narrow; strikingly short; not 

 waisted medially. FScl distinct; long and broad; oriented antero-dorsad; very strongly 

 elevated over the base of the ventrad projection. Dorsal and ventral surfaces convex. Apex 

 acute. Ventral projection long and narrow; enlarged mesally; deeply concave; strongly 

 oriented postero-distad, weakly ventrad and not curved posteriad. Dorsal surface concave. 

 Ventral surface convex. Apex narrow; rounded; weakly recurved. Concavity located in the 

 preapical area; surrounded by three unequally strong ridges. Distal embayment moderately 

 shallow; oriented mesad. FSc2 fused to the distal margin of the ventral projection; oriented 

 distad and weakly dorsad; reduced; small; round; completely planate; broad. Dorsal surface 

 not enlarged; convex. Ventral surface convex. Apex aciculate. Head and neck dorsal 

 surface extended anteriad. 



Neck very narrow; strikingly long; strongly oriented distad and ventrad; articulation with 

 2Ax narrow, reduced anteriorly; medially convex; discontinuous with tail. Proximal margin 



