Subfamily Scarabaeinae

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Subfamily Scarabaeinae

Compiler and date details

2017 - ABRS

1 December 2002 - Gerasimos Cassis & Tom A. Weir; updated by Andrew A. Calder, CSIRO Entomology, Canberra, Australia

Introduction

The Scarabaeinae are a large cosmopolitan subfamily with a world fauna of some 220 genera and 4,600 species. The subfamily is considered to be monophyletic, with features in common with the Aphodiinae and the Geotrupidae (Halffter & Edmonds 1982). Gemminger & Harold (1869) provided the first world catalogue for the subfamily and the most modern updates are those of Gillet (1911) and Boucomont & Gillet (1927). The suprageneric classification used in this Catalogue is that of Halffter & Edmonds (1982).

The Australian endemic fauna consists of only three recognised tribes: Coprini, Onthophagini and Scarabaeini. The Coprini (62 genera) and Onthophagini (13 genera) are cosmopolitan tribes. The genus Onthophagus Latreille has been divided into 17 subgenera (Halffter & Edmonds 1982), but the Australian species have not been referred to any particular subgenus. The Scarabaeini has a worldwide distribution also, and comprises 101 genera.

Gunter & Weir (2017) in a study that describes 2 new genera and six new species, and creates a series of new generic combinations state, in discussing tribal classification, '[t]here is accumulating evidence from both molecular and morphological phylogenies that tribal classification within Scarabaeinae does not reflect natural groupings, with the most recent phylogeny proposing a revised classification system. On the basis of molecular evidence and morphological synapomorphies, Tarasov & Dimitrov (2016) revised the concepts for the tribes Deltochilini, Dichotomiini, and Coprini providing diagnostic characters to facilitate future classification. Significantly, all members of the Australasian clade, including 19 Australian genera, are removed from the tribes Deltochilini and Coprini, and are treated as incertae sedis.' Given that they signal further studies on the Australian fauna, no changes are made presently (October 2017) to the tribal classification employed here, and the two new genera are assigned to Scarabaeini.

The first Australian scarabaeines described were Scarabaeus quadripustulatus Fabricius, 1775 and Scarabaeus novaehollandiae Fabricius, 1775, from specimens collected in 1770 at Endeavour River on Cook's voyage of exploration. Macleay (1864, 1871, 1887, 1888) described 45 new species, while Blackburn (1890, 1892a, 1892b, 1892c, 1893, 1894, 1895, 1900a, 1900b, 1903, 1904, 1907) described 38 new species and provided a key to Onthophagus species in his 1903 paper. Lea (1923) described 30 new species and listed all the Australian species in the subfamily but did not provide keys. Paulian (1934, 1935) gave the first synoptic survey of the canthonine fauna of the Australian Region, provided keys to genera and species, and described one new genus and seven new species. Paulian (1933) also dealt with the coprine genera Coptodactyla Burmeister and Thyregis Blackburn, while Carter (1936) mentioned Demarziella Balthasar (as Pedaria Castelnau) and Aptenocanthon Matthews (as Panelus Lewis). Paulian (1937a, 1937b) proposed a further 10 new species of Onthophagus. Frey (1959, 1963, 1969, 1970) described nine new species of Onthophagus and Balthasar (1965) reviewed part of Coptodactyla. The most recent major works on the Australian Scarabaeinae are those by Matthews on the Onthophagini (Matthews 1972), Scarabaeini (Matthews 1974) and Coprini (Matthews 1976).

Matthews & Stebnicka (1986) reviewed Demarziella and transferred it from Aphodiinae to Scarabaeinae. Storey (1977) described six new species of Onthophagus and in later works described a new Aptenocanthon species (Storey 1984) and one new species in Aulacopris White (Storey 1986). Storey & Weir (1990) added a further 20 new species of Onthophagus and Storey & Monteith (2000) described five new species of Wet Tropics Aptenocanthon Matthews. Reid & Storey (2000) revised Temnoplectron Westwood adding another five new species from the Wet Tropics region of Queensland while Reid (2000) revised a group of cryptic species of Coptodactyla Burmeister. Matthews & Weir (2002) recorded two new species of the ball-rolling dung beetle genus Lepanus Balthasar from South Australia. Matthews (1984) provided an illustrated key to the scarabaeinine genera of South Australia.

Matthews (1972) referred to the 'holotype' of a number of species for which the original descriptions either indicated syntypes or gave no information on the number of specimens. Cassis & Weir (1992) interpreted these cases as an incorrect reference to a holotype and Matthews' actions were therefore considered as lectotype designations (Article 74, ICZN 1985). For a number of species described by Macleay, Matthews (1972) did not correctly interpret the type situation. In these cases, Cassis and Weir (1992) either reinterpreted the data or cited the original information, this also being accepted in the current Catalogue.

The distribution of the Australian fauna is detailed in Matthews (1972, 1974, 1976) who correlated it with climate and vegetation types. Distribution information is also summarised by Howden (1981) and Doube et al. (1991). Most of the species of Onthophagus Onthophagini) are found in the northern and eastern regions of Australia (Matthews 1972: fig. 12; Storey 1977; Storey & Weir 1990). The number of species decreases toward the south and only nine species are found in south-western Australia, and eight in Tasmania. In the Coprini, the species of Demarziella and Coptodactyla occur largely within several hundred kilometres of the northern and eastern seaboards, whereas Thyregis has three species in south-eastern Australia and one along the western coast (Matthews 1976: figs 1 & 22). The Scarabaeini are restricted mainly to the eastern, northern and western coasts. Only three species occur in Victoria, three in South Australia and none in Tasmania (Matthews 1974: fig. 1 & 1984; Matthews & Weir 2002).

Labroma Sharp, Coproecus Reiche, Mentophilus Castelnau and Sauvagesinella Paulian are restricted to Western Australia, and Tesserodon Hope and Monoplistes Lansberge are found in the summer rainfall areas of northern Australia. Diorygopyx Matthews, Cephalodesmius Westwood, Aulacopris and Aptenocanthon occur in coastal New South Wales and southern Queensland and one species of Aulacopris extends into Victoria. The latter two genera also have relict distributions of one species each in the high rainfall mountainous areas of north Queensland. Pseudignambia Paulian & Pluot-Sigwalt and Boletoscapter Matthews are restricted to coastal Queensland while Canthonosoma Macleay inhabits the same areas but also extends further inland. Temnoplectron Westwood occurs in coastal Queensland and Northern Territory, and Lepanus Balthasar and Amphistomus Lansberge are widespread along the coast of New South Wales and the Northern Territory with species of the former genus in South Australia and south-western Australia.

The indigenous Australian fauna shows a high degree of endemicity at both generic and specific levels. Matthews (1972, 1976) suggested that this, coupled with the presence of archaic groups in the Coprini and Scarabaeini, gives the fauna a strongly insular aspect, unlike that of other regions of comparable size, but reminiscent of that of some continental islands. The indigenous fauna consists of 20 genera and 335 described species. The Coprini is represented by three genera and 22 species. Thyregis and Demarziella are endemic whereas three species of Coptodactyla are shared with New Guinea (Matthews 1976). The Onthophagini is represented only by the cosmopolitan Onthophagus, perhaps the largest genus of beetles known, which contains almost 200 Australian species (Matthews 1972; Storey 1977; Storey & Weir 1990). Most of these species are endemic, and at least seven are shared with New Guinea only. The Scarabaeini consists of 18 genera and 132 species. Eleven of the genera are endemic and the other five (Tesserodon, Amphistomus, Lepanus, Temnoplectron and Monoplistes) are shared with New Guinea (Matthews 1974; T.A.Weir pers. com.). Matthews (1976) postulated various origins for the Australian scarabaeine fauna. The genus Onthophagus, which contains species that are highly vagile, has Oriental affinities, whereas the Scarabaeini are thought to have Gondwanan affinities. The Coprini present a different pattern in having some affinities with the South American fauna although they appear to be most closely related to the Afrotropical and Holarctic genus Copris Müller, and the Afrotropical genera Pedaria and Paraphytus Harold.

Gunter & Weir (2017) erected three new genera and described six new species. In their introduction, they discuss recent developments in classification of Scarabaeinae especially in relation to the Australian fauna. They comment that Australian genera, apparently previously in two tribes, Deltochilini and Coprini, are now to be considered to be incertae sedis. Given that they signal ongoing studies on Australian species, the tribal classification in Scarabaeini is retained here for the present.

The majority of the introduced species in the Australian fauna were released intentionally as part of the CSIRO Dung Beetle Program. The basic aim of this project was to establish a dung beetle fauna in Australian pastures, which would rapidly bury the dung of domestic stock and thereby benefit pasture production and reduce the number of pest flies (Doube et al. 1991). Tropical southern Africa was the chief source of species for northern Australia. The Cape Province of South Africa and southern Europe provided species that were introduced into southern Australia. Bornemissza (1976, 1979) recorded the early successful establishment of beetles while their subsequent establishment and spread have been detailed in CSIRO Division of Entomology Annual Reports. Fifty-five species of dung beetle were received at CSIRO between 1968 and 1982. Of these, 41 species were released; to date 22 species are considered established and a further five have been recaptured but their status is uncertain (Doube et al. 1991). Only the latter 27 species were recorded in the Catalogue. It is estimated that at least one exotic species occurs in almost every Australian pasture and up to seven species occur together in a few localities (Doube et al. 1991). Some species, such as Onthophagus gazella (Fabricius), Onitis alexis Klug and Euoniticellus intermedius Reiche now occur over much of the continent while others appear to be restricted to northern or southern regions. The introduced species are distributed among the following tribes: Coprini—Copris Müller (4 species); Onitini—Bubas Mulsant (1 species) and Onitis Fabricius (8 species); Oniticellini—Euoniticellus Janssens (4 species) and Liatongus Reitter (1 species); Onthophagini—Onthophagus (6 species); and Scarabaeini—Sisyphus Latreille (3 species).

BIOLOGY
The Scarabaeinae are the most highly evolved of all the scarabaeid groups and feed on bacteria rich, semi-liquid foods, mainly dung. They use this material to make brood balls for the larvae which live in chambers or burrows in the ground (Matthews 1984; Lawrence & Britton 1991). Details of feeding behaviour and nest construction are known for only a few Australian species. Food preferences, as shown by numbers attracted to various types of bait, indicate that most of the species in the three indigenous tribes are coprophagous, while a small number of Onthophagus species are copro-necrophagous. Fourteen species of Onthophagus are known to feed on fungi and one species feeds on fallen fruits (Doube et al. 1991). Bornemissza (1971a, 1971b) described the nesting behaviour of Onthophagus compositus Lea and recorded mycetophagous breeding for Onthophagus dunningi Harold. Tyndale-Biscoe et al. (1981) published an ecological study of Onthophagus granulatus Boheman. Ridsdill Smith (1981) described aspects of the biology of Onthophagus ferox Harold. A few species of Onthophagus have unusual feeding behaviour. The adults of Onthophagus muticus Macleay and O. parvus Blanchard, for example, have prehensile claws that allow them to cling to the hair around the anus of macropods from which they seize the dung pellets as they are extruded (Matthews 1972). The species of Cephalodesmius differ in that the brood balls are formed from dead leaves and other plant material rather than dung (Monteith & Storey 1981). Halffter & Matthews (1966) described and summarised the various nidification behaviours for the subfamily, and Halffter (1977) and Halffter & Edmonds (1982) expanded on this and suggested evolutionary trends.

Diagnosis

The Australian Scarabaeinae adults are mainly black or brown in colour. They range in size from 1.5 mm for some Lepanus species up to 25 mm for Aulacopris species. The adult stage can be distinguished from other scarabaeids by a combination of the following character states: tarsal claws equal; elytra without a narrow membranous margin and leaving the pygidium exposed; abdominal spiracles located in pleural membrane and covered by elytra; midcoxae longitudinal and separated by more than their width; labrum concealed; scutellum absent (except in Thyregis); and hind tibia each with a single apical spur. They share with Aphodiinae the modified liquid-feeding mouthparts but are more robust and rounded in form, and usually exhibit conspicuous sexual dimorphism in the form of cephalic and/or pronotal armature in the male (Lawrence & Britton 1991). The larvae are distinguished by the possession of four-segmented antennae with a reduced apical segment (at most half as wide as penultimate), distinctly separated galea and lacinia, anal opening usually surrounded by fleshy lobes, two-segmented legs with claws small or absent, and a strongly hump-backed body (Ritcher 1966). Edmonds & Halffter (1987) have produced the most recent taxonomic review of the immature stages of the subfamily.

Published	As part of group	Action Date	Action Type	Compiler(s)
02-Nov-2017	Scarabaeinae	08-Oct-2017	MODIFIED
		12-Feb-2010	(import)

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