30 



Discussion 



Monophyly of the Lucanidae is supported by the fact that all of the taxa in this family 

 share the following six apomorphic character states of the wing articulation, all confined 

 to 2Ax: 



1. the apex of the dorso-distal ridge is very strongly curved distad, 



2. reduced distally and anteriorly, 



3. revealing the distal and apical sections of the dorso-proximal ridge, the anterior section 

 of the dorso-distal ridge is recurved with the apex assuming a ventral orientation (in 

 addition to the distal curvature discussed above), 



4. the subapical to antero-median section of the dorso-distal ridge is strongly oriented 

 postero-distad, 



5. the median to posterior section of the dorso-distal ridge and lobe is strongly depressed 

 ventrad, curving beneath the antero-median section of the dorso-proximal ridge, giving 

 the appearance that the dorso-distal ridge and lobe twists around the dorso-proximal ridge 

 and lobe, 



6. the dorso-proximal ridge apex is lengthened anteriorly, beyond the ventro-distal ridge 

 apex. 



Lucanidae share six apomorphic character states of the wing articulation and wing base 

 with Passalidae, Diphyllostomatidae, Glaphyridae, Trogidae, Bolboceratidae and Pleoco- 

 midae (Browne & Scholtz 1995). 



Howden (1982) considered lucanids to be most closely related to Passalidae. Caveney 

 (1986) suggested, based on synapomorphic acone ommatidium structure between all 

 diphyllostomatids and many lucanids, that Lucanidae are probably more closely related to 

 Diphyllostomatidae, a view which is supported here (Browne & Scholtz 1995). 



The Lucanidae do not exhibit any wing articulation and wing base character states which 

 support Howden's (1982) proposal that it branched off early from other members of the 

 Scarabaeoidea and subsequently followed a separate evolutionary pathway, as do 

 Passalidae. 



Glaphyridae 



Introduction 



Glaphyridae are represented in the Holarctic Region by Glaphyrinae (with Amphicoma 

 and Lichnanthe the largest genera) and in South America by Lichniinae (Lichnia and 

 Cratoscelis). Adults are long-legged, hairy and brightly coloured. 



The males of Amphicoma spp. are pollen feeders and visit flowers (Crowson 1967), 

 whereas, according to Ritcher (1958), adults of Lichnanthe never feed. Ritcher (1958) 

 noted that larvae of Lichnanthe vulpina (Hertz) feed on cranberry roots and those of L. 

 rathvoni Le Conte on decaying leaves and other plant debris. The length of the life cycle 

 is 3-4 years (Ritcher 1966). 



