5 



ABSTRACT 



Two character suites, the hind wing articulation, comprised of the first, second and third 

 axillaries and the median plate, and the hind wing base, comprised of the first and second 

 basal plates, are described for Glaresidae, Passalidae, Diphyllostomatidae, Lucanidae, 

 Glaphyridae, Trogidae, Bolboceratidae, Pleocomidae, Geotrupidae, Hybosoridae, 

 Ochodaeidae, Ceratocanthidae and Scarabaeidae (Aphodiinae, Aegialiinae, Aulonocne- 

 minae, Orphninae, Melolonthinae, Acoma, Oncerinae, Chasmatopterinae, Hopliinae, 

 Rutelinae, Dynastinae, Cetoniinae, Trichiinae and Valginae). Due to the magnitude of this 

 study, large number of characters and high degree of variability of the structures, it was 

 not possible, at this early stage, to adequately analyse the phylogenetic content of the 

 various character states. However, some notes concerning genealogical relationships 

 among the major taxa are given. 



INTRODUCTION 



The Scarabaeoidea are one of the largest and most variable superfamilies of Coleoptera. 

 Members of the superfamily vary tremendously in size, facies and habits but are united 

 by several unique characters (Lawrence & Britton 1991). Most adults are robust, short- 

 legged beetles with a typically lamellate antennal club, highly modified prothorax with 

 large coxae, usually dentate tibiae, strong intrinsic wing-folding mechanism, the second 

 abdominal sternite represented only by a lateral portion, the eighth tergite forming a true 

 pygidium and four Malpighian tubules. Hind coxal plates are absent. Larvae are grub-like 

 and usually C-shaped, with well-developed antennae and legs. They are without 

 urogomphi. Scarabaeoids feed on a wide range of plant and animal matter and dung. This 

 varies from detritus and most types of dung through lower plants to virtually all higher 

 plant tissues and carrion to predation on other insects. Their habits range from free-living 

 through fairly sophisticated forms of brood care to sub-social behaviour. 

 The Scarabaeoidea are morphologically one of the best-studied beetle groups. There have 

 been many broadly based comparative studies covering most major structures. These 

 include: antennae (Iablokoff-Khnzorian 1977), antennal sensilla (Meinecke 1975), eye 

 (Caveney 1986), mouthparts (Nel & Scholtz 1990), prothorax (Hlavac 1975), coxae 

 (Ritcher 1969c; Hlavac 1975), spiracles (Richer 1969a,b), wing venation (Crowson 1967; 

 Iablokoff-Khnzorian 1977), alimentary canal (unpublished), metendosternite (Crowson 

 1938; Iablokoff-Khnzorian 1977), male genitalia (d'Hotman & Scholtz 1990a,b), female 

 genitalia (Tanner 1927; Holloway 1972; Lawrence & Newton 1982; unpublished), 

 ovarioles (Ritcher & Baker 1974), karyotype (Smith & Virkki 1978; Yadav & Pillai 1979), 

 central nervous system (Iablokoff-Khnzorian 1977; unpublished), spermatozoan number 

 (Virkki 1969), malphigian tubules (Caveney 1986) and larvae (Areekull 1957; Ritcher 

 1966; Hinton 1967; Costa et al. 1988) to name a few. These character suites were recently 

 reviewed and phylogenetically assessed by Scholtz (1990). 



The current project is one of a series by members of our research team in which various 

 morphological structures of the major groups of Scarabaeoidea are being studied to 



