72 



Discussion 



Although Aegialiinae do not exhibit any autapomorphic wing articulation and wing base 

 characters, they do share seven derived character states of the wing articulation and wing 

 base with Aulonocneminae 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). 



Many workers have suggested that aegialiines are most closely related to Aphodiinae 

 (Stebnicka 1985; Cambefort 1987; Nel & Scholtz 1990; d"Hotman & Scholtz 1990a). 

 Many primitive morphological characters support this relationship, including those of the 

 mouthparts (Nel & Scholtz 1990), male genitalia (d'Hotman & Scholtz 1990a), spiracles 

 (Ritcher 1969a,b), and chromosomes (Virkki 1967). This view is reiterated here. 

 Aegialiinae share 46 apomorphic character states of the wing articulation and wing base 

 with Aphodiinae, Aulonocneminae and Scarabaeinae (Browne 1993). 



Aulonocneminae 



Introduction 



This is a small subfamily comprising four genera and about 50 species, which occur mainly 

 on Madagascar but also on other Indian Ocean Islands, and in southern Africa. An 

 Aulonocnemis larva was described by Paulian & Lumaret (1974). Adult characters include 

 those of mouthparts that are not covered by the labrum or clypeus, a covered pygidium. 

 and eyes that are visible in ventral view. They are thought to be sapro-xylophagous 

 (Cambefort 1987). 



Hind Wing Articulation Description 



First Axillary (Fig. 115) 



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. 

 Po stem -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; short but broad. Dorsal and ventral surfaces convex. Apex 

 acute. Ventral projection moderately short and narrow; enlarged mesally; deeply concave; 

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

 concave. Ventral surface convex. Apex narrow; truncate. Concavity located in the preapical 

 area; surrounded by three unequally strong ridges. Distal embayment oriented mesad. FSc2 

 fused to the distal margin of the ventral projection; oriented distad and weakly dorsad; 

 reduced; small; round; planate. Dorsal surface not enlarged; convex. Ventral surface 

 convex. Apex rounded. Head and neck dorsal surface extended anteriad. 



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

 with 2 Ax naiTow, reduced anteriorly; continuous with tail. Proximal margin not curved 

 ventrad. Distal margin straight. Distal embayment very weakly concave. 



