Superfamily SCARABAEOIDEA

<I>Onthophagus declivis</I> Harold, 1869

Onthophagus declivis Harold, 1869

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Superfamily SCARABAEOIDEA

Compiler and date details

2002 - Updated by Andrew A. Calder, CSIRO Entomology, Canberra, Australia

1992 - Gerasimos Cassis, Australian Museum, Sydney, New South Wales, Australia, W.W. Keith Houston, Australian Biological Resources Study, Canberra, Australian Capital Territory, Australia, Tom A. Weir and Barry P. Moore

Introduction

Members of the Scarabaeoidea (=Lamellicornia) are distinctive beetles. They generally have a stout, often large, body which may be brightly coloured. Many are of economic importance, some have an unusual biology and some are of cultural significance. It is a relatively uniform group, cosmopolitan in distribution, and currently composed of about 2,500 genera and 30,000 species. The recognised fauna of Australia comprises nearly 314 genera and over 2,620 species.

The family classification is unresolved; some groups are regarded as either distinct families or subfamilies. Iablokoff-Khnzorian (1977) recognised only six families, with the Scarabaeidae composed of 21 subfamilies. Crowson (1981) listed 10 families of Scarabaeoidea but did not consider the Diphyllostomatidae as a separate family. Lawrence (1982) considered the latter as a distinct family but included the Hybosoridae, Ochodaeidae and Glaphyridae as subfamilies of Scarabaeidae, recognising only eight families. Lawrence & Newton (1982) referred to these latter three groups as distinct families. The basic family classification used in this Catalogue is that of Lawrence & Britton (1991). However, the Bolboceratinae is now treated as a family following Browne & Scholtz (1995, 1999). The following families are not represented in the Australian fauna: the Nearctic Pleocomidae and Diphyllostomatidae, the widespread Ochodaeidae and the Glaresidae (Scholtz et al. 1987, 1988), and the Palaearctic, Nearctic and Neotropical Glaphyridae.

The relationship of the Scarabaeoidea to the remainder of the Coleoptera is still conjectural, although most authors consider that the Scarabaeoidea are derived from members of the Dascilloidea (Crowson 1981; Scholtz 1990). Crowson (1981) reviewed some of the knowledge of fossils of Scarabaeoidea, discussed a possible evolutionary pathway and suggested alternative interpretations for the changes in the structure and habits of the adults.

The phylogenetic relationships of the families and subfamilies is unclear. Iablokoff-Khnzorian (1977) analysed the phylogeny of these groups of Scarabaeoidea. Howden (1982) also reviewed the phylogeny of most of the higher taxa of the Scarabaeoidea and recognised the following sister-group relationships: Passalidae and Lucanidae; Pleocomidae and Geotrupidae (both as subfamilies of Scarabaeidae); Hybosoridae and Trogidae; Aphodiinae and Scarabaeinae; and the Cetoniinae, Melolonthinae, Dynastinae and Rutelinae. Lawrence & Newton (1982) state that the superfamily has two major sections based on the structure of the abdominal apex. The Lucanidae, Passalidae and Trogidae are considered to be related on the basis of the eighth segment being retracted. The remainder of the taxa have this segment exposed.

The Scarabaeoidea are distinguished from other beetles by a unique set of coxal and tibial adaptations for burrowing, a strong intrinsic spring mechanism for wing folding, a lamellate antennal club, and a distinctive grub-like larva (Lawrence & Newton 1982). Matthews (1984) discussed the highly developed antennae which can be closely apposed to protect the inner sensory surfaces when burrowing. Ritcher (1966) described the larvae of scarabaeoids in detail, with the main features being the C-shaped, stout body, three pairs of well-developed legs (except in Passalidae and in some Geotrupini), hypognathous head with strong mandibles, and a pair of three or four segmented antennae, and thorax and first eight abdominal segments usually with cribriform spiracles.

The burrowing behaviour and associated adaptations are the most characteristic feature of the superfamily. The larvae are mostly soil dwellers or live in decaying matter on the soil surface and feed on a wide variety of organic matter, which ranges from carrion and dung, to humus, roots, rotten wood and fungi. Adults feed on leaves, nectar, pollen, sap, carrion and dung. Matthews (1984) postulates a number of evolutionary lines which exploited soil-asssociated food sources such as rotting wood (Lucanidae, Passalidae, Rutelinae, Dynastinae and Cetoniinae), humus and roots (Rutelinae, Dynastinae, Cetoniinae and Melolonthinae), carrion (Trogidae and Hybosoridae) and dung (Aphodiinae and Scarabaeinae). The feeding behaviour of the Australian Bolboceratidae is poorly known, although they are thought to feed on fungi and humus and one species is recorded as feeding on cattle dung (Carne 1965).

The Australian Scarabaeoidea fauna is perhaps the best described of any other equivalent beetle group (Matthews 1984). Most groups have had comprehensive revisionary work, with only the Cetoniinae in need of serious work. The Lucanidae and Passalidae have had little modern attention; much of the fauna, however, is described and understood. Lawrence & Britton (1991) gave a general review of the families and subfamilies and included important references and keys to the adults and larvae. Matthews (1984) gave a pictorial key to the genera of the South Australian Scarabaeoidea. Ritcher (1966) gave family, subfamily and generic keys to the larvae of the North American fauna. Howden (1981) discussed the zoogeography of some of the families and subfamilies of Scarabaeidae.

Acknowledgements

Preparation of this volume of the Catalogue formed part of the research conducted by the Division of Entomology, CSIRO, Canberra and the Division's resources and facilities were made available to us. The Australian Biological Resources Study supported part of this project through grants to Dr J.F. Lawrence.

The illustrations by F. Nanninga and S.P. Kim used in the family/subfamily introductions are from Lawrence & Britton (1991). They were kindly provided by and are reproduced with permission from the Division of Entomology, CSIRO and Melbourne University Press. The illustrations for the Cetoniinae and the Rutelinae were prepared for ABRS by A. Carter.

The authors wish to thank the many people who assisted us in various ways, especially: Dr J.F. Lawrence who initiated the project; Dr E.B. Britton for advice on the Melolonthinae; Dr K.L.H. Key, Dr D.K. McAlpine and Dr W.D.L. Ride for advice on nomenclatural problems; Dr M.J. Dallwitz, Dr E. Zurcher and Ms K. Pickerd for data base management; and the librarians at the CSIRO Black Mountain Library and the Australian Museum for their patience and help in finding many references.

Database Notes

This database was updated in 2002 by Dr Andrew Calder. For sections that have remained unchanged since the original compilation, only the original author/compiler names are given although the closing date is altered appropriately.

New names and new distributions given in relevant papers in Daccordi & Giachino (2003) added by ABRS 2007.

Limital Area

Distribution data in the Directory is by political and geographic region descriptors and serves as a guide to the distribution of a taxon. For details of a taxon's distribution, the reader should consult the cited references (if any) at genus and species levels.

Australia is defined as including Lord Howe Is., Norfolk Is., Cocos (Keeling) Ils, Christmas Is., Ashmore and Cartier Ils, Macquarie Is., Australian Antarctic Territory, Heard and McDonald Ils, and the waters associated with these land areas of Australian political responsibility. Political areas include the adjacent waters.

Terrestrial geographical terms are based on the drainage systems of continental Australia, while marine terms are self explanatory except as follows: the boundary between the coastal and oceanic zones is the 200 m contour; the Arafura Sea extends from Cape York to 124°E; and the boundary between the Tasman and Coral Seas is considered to be the latitude of Fraser Island, also regarded as the southern terminus of the Great Barrier Reef.

Distribution records, if any, outside of these areas are listed as extralimital. The distribution descriptors for each species are collated to genus level. Users are advised that extralimital distribution for some taxa may not be complete.