Superfamily ELATEROIDEA

Elateroidea

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Superfamily ELATEROIDEA

Click Beetles

Compiler and date details

31 December 1996 - Andrew A. Calder, CSIRO Entomology, Canberra, Australian Capital Territory, Australia

Introduction

The Elateroidea currently comprises about 1052 genera and 21 440 species (Lawrence 1982) worldwide. They form a mixed assemblage which includes some well known beetles that are commonly referred to as click beetles, fireflies or soldier beetles. The recognised Australian fauna, with 129 genera and 1105 species, represents 12% and 5% of the world's genera and species respectively; the families Rhinorhipidae, Brachyspectridae, Eucnemidae, Throscidae, Elateridae, Lycidae, Lampyridae and Cantharidae (Lawrence & Britton 1991) are represented. The family with the largest number of species is the Elateridae with 667 valid species recorded for Australia.

Members of the Elateroidea usually have a heterogeneous life cycle with long-lived larvae and short-lived adults that occupy different ecological niches (Lawrence & Newton 1982). The biology of most species is poorly known and the ecological information is to a large extent based on extrapolation from well known species in the relevant group, often from work done on overseas faunas. Adults are at least in part predacious or phytophagous, but some generally appear not to feed at all. Cantharid adults apparently feed on nectar and pollen or are carnivorous, feeding on other insects. Lampyrid adults generally do not feed at all and nothing is actually known about the feeding habits, if any, of the families Elateridae, Eucnemidae, Throscidae or Rhinorhipidae. Larvae are either phytophagous with many species being root feeders, saprophagous or predacious. Lycid larvae are thought to feed on slime moulds (Moore 1989) or soft, decaying organic matter.

Some species, particularly in the Elateridae, are of economic importance. A number of elaterids have larvae which are predominantly soil dwellers that feed on plant roots. Those that are commonly referred to as wireworms are destructive to pastures, cereals and root crops. Many species, particularly of Elateridae, Eucnemidae and Throscidae, are dependent on the existence of rotting wood in various stages of decay to complete their life cycles.

The Elateroidea treated in this work have been treated by earlier authors as belonging to three separate but related superfamilies: Artematopodoidea (incorrectly known as Artematopoidea), Elateroidea and Cantharoidea. The exact composition of the Elateroidea has been in a state of flux, undergoing partial reorganisation in the last few years due to the work of Lawrence and Newton. Crowson (1972) recognised two separate superfamilies, Elateroidea and Cantharoidea, subsequently adding a third superfamily, Artematopoidea (Crowson 1973), to accommodate both extant and fossil species from East Prussian Baltic amber. In doing so, Crowson noted that his new group was not distinguishable from the Elateroidea and Cantharoidea on the basis of any single derivative character and preferred to maintain three separate superfamilies due to uncertainty about the phylogenetic relationships. These same three superfamilies were maintained by Crowson (1981): Artematopodoidea (Artematopodidae, Callirhipidae, Brachyspectridae), Elateroidea (Cebrionidae, Elateridae, Throscidae, Eucnemidae) and Cantharoidea (Cneoglossidae, Plastoceridae, Omalisidae, Lycidae, Drilidae, Phengodidae, Telegeusidae, Lampyridae, Omethidae, Cantharidae).

Lawrence & Newton (1982) and Lawrence (1988) considered that the three superfamilies of Crowson, excluding the Callirhipidae, and more recently (Lawrence & Newton 1995) excluding the Cneoglossidae, form a monophyletic group. The recently recognised Australian family Rhinorhipidae (Lawrence 1988) was tentatively placed in the old Elateroidea, but later moved to incertae sedis within the series Elateriformia (Lawrence & Newton 1995). Initially, Lawrence's phylogenetic analyses showed the taxon to be basal to the Elateroidea (which included the old Artematopodoidea, Elateroidea and Cantharoidea), Psephenoidea or a clade comprising these two groups. Recently, Lawrence, Nikitsky & Kirejtshuk (1995) showed the taxon forming part of a clade containing Dascilloidea and Buprestoidea.

Within the Australian fauna, two distinct groups are recognisable in the Elateroidea sensu Lawrence. A group of families with heavily sclerotised integuments in which streamlining of the body has been developed to a high degree, with the transformation of the prothoracic interlocking mechanism to form a clicking mechanism that startles predators. The second group, loosely articulated beetles with a thin, flexible cuticle, have lost these mechanical defense systems and developed chemical defense mechanisms and aposematic colour patterns (Crowson 1981). Members of this second group were formerly included in the Cantharoidea.

The monophyly of this Elateroidea was alluded to by Lawrence & Newton (1982) and proposed formally by Lawrence (1988). The Elateroidea is distinguished by the presence of four Malpighian tubules, low grade type of wing folding (Hammond 1979), functional eighth abdominal spiracles and the absence of a mandibular mola and transverse metasternal suture. The propleurocoxal mechanism varies from an exposed trochantin and movable pleuron (e.g. Artematopodidae, Lycidae, Cantharidae, Lampyridae) to a highly reduced and concealed trochantin and rigid pleuron (e.g. Eucnemidae, Throscidae, Elateridae) with intermediate conditions found in Cerophytidae, Brachyspectridae or Cebrioninae and Plastoceridae (Lawrence & Newton 1982). The larva is distinctive in having a single large stemma on each side of the head, a bisetose tarsungulus and a feeding mechanism adapted for extraoral digestion (mola absent, ventral mouthparts connate and buccal cavity blocked by hairs). Connation of the basal four ventrites is typical of the taxa grouped in the former Elateroidea but this feature is absent in the soft-bodied forms comprising the Cantharoidea sensu Crowson. The freely articulated basal abdominal segments is attributed to neoteny (Lawrence & Newton 1995). The Cantharoidea were previously considered to be a monophyletic group, however it was pointed out by Lawrence & Newton (1995) that all distinguishing features that had been cited in evidence are either plesiomorphic or are associated with the malacoderm facies.

Fossils for the Elateroidea are mostly unknown from Australian sites. The only decribed forms being elytra of four species from the Upper Triassic Ipswich Series in Queensland, attributed to the Elateridae (Tillyard 1923), and an indeterminate cantharid from the early Cretaceous Koonwarra fossil bed in South Gippsland, Victoria (Jell & Duncan 1986). The latter bears striking similarities to recent ‘primitive’ Telephorus Schaeffer (= Cantharis Linnaeus) and an Heteromastix Boheman. However, representatives of the Elateroidea, particularly of Elateridae, are relatively well represented in overseas fossil beds. The Triassic beds (Mesozoic) of central Asia contain undisputed and undescribed Elateriformia (Kukolová-Peck 1991). The Jurassic and Cretaceous faunas of Russia and western Mongolia are known to contain members of extant Elateridae (Ponomarenko 1995). Fossils have been attributed plausibly to the family from the Lower Jurassic with the description by Dolin (1973, 1975) of several new genera and species that could be assigned to an extant subfamily, Agrypninae. The Elateridae are quite diverse in the Late Jurassic Kara Tau deposits (Dolin 1980) with some of the fossils being assigned by Dolin (1976) to recent subfamilies Cardiophorinae and Negastriinae, although Crowson (1981) considered some of these fossils to be allied to the Oestodinae (=Lissominae of Calder (1998)). Ponomarenko (1995) notes that the Elateridae was a more important group in the Eocene than at Present. Britton (1960) describes fossil species of Throscidae and Eucnemidae from the London Clay at Bognor Regis, Sussex, which is dated of Eocene age. The throscid could be assigned to an extant genus, Pactopus LeConte. Muona (1993) described several eucnemids and throscids from Baltic amber, some of which can also be attributed to extant genera. The amber from Baltic areas is thought to have originated in the Middle Eocene (Tertiary) (Larsson 1978) although Muona questions this dating for the amber. The Artematopodidae are known from the Middle Jurassic and Crowson (1973) described two new genera Electribius and Protartematopus from East Prussian Baltic Amber. Electribius Crowson has since been found to be extant in Mesoamerica (Lawrence 1995).

A generic overview of the Elateridae was published by Calder (1996). Ballantyne (1992) revised the Lampyridae for her doctoral dissertation. Muona (1991) revised the Indo-malesian tribe Galbitini and is currently monographing the Australian eucnemid fauna. The Throscidae, Lycidae and Cantharidae have received little attention since the publications of Lea and Cobos and are in need of modern revisionary work. Larvae of an undescribed species of Brachyspectra LeConte have been found in Winjana Gorge, WA (Lawrence & Britton 1991). This group is otherwise known only from North America, India and Malaysia. Keys to the families of larvae and adults found in Australia were provided by Lawrence & Britton (1991) and a pictorial key to the South Australian genera was given by Matthews (1985).

Acknowledgements

The project that culminated in the production of the database and the published volume (Calder 1998) was supported by a grant from the Australian Biological Resources Study (ABRS) which is gratefully acknowledged. I am also indebted to Drs Keith Houston and Alice Wells for editorial advice. This Catalogue was produced using the newly developed taxonomic–bibliographic database Platypus that was developed by the Australian Biological Resources Study. Steve Shattuck is particularly thanked for his assistance.
The following two individuals are gratefully acknowledged for their input into two families included in this Catalogue. Dr Jyrki Muona and Dr Lesley Ballantyne for their advice on the Eucnemidae and Lampyridae respectively, which are currently the subjects of their personal research. Dr Muona allowed me access to his data and checklist of Australian Eucnemidae, while Dr Ballantyne provided access to her unpublished doctoral thesis. Many new synonymies and combinations were discovered during their studies and in an attempt to make this Catalogue as up to date and complete as possible these new nomenclatural decisions have been included and credited to either of these two individuals.

The preparation and data entry for this Catalogue was conducted in CSIRO Entomology, Canberra, and use of the Organisation's resources and facilities, particularly computing resources, is gratefully acknowledged.

I particularly thank the staff of the CSIRO Black Mountain Library for their help in locating the many obscure references encountered in the compilation of this Catalogue and for their patience in processing innumerable requests for inter-library loans, many from overseas, so that the original bibliographic reference could be verified.

The illustrations by F. Nanninga and S.P. Kim were taken from The Insects of Australia and are reproduced with permission from CSIRO Entomology and Melbourne University Press.

Database Notes

The information on the Australian Faunal Directory site for the Elateroidea is derived from the Zoological Catalogue of Australia database compiled on the Platypus software program. The original work was published on 14 May 1998 as (Calder, A.A., 1998) Subsequent minor changes to the database include citations for nomenclatural decisions published by Calder (1998).

The original orthography of all species names is given in preference to the emended version of those names that have been altered to conform to the rules of classical grammar as required by Article 34(b) (ICZN 1985). The taxa are arranged in alphabetic order within the next highest category. Synonyms are arranged in chronological order. All publications containing the original descriptions of genera and species have been seen. If a publication was not available for personal perusal in Australia, a photocopy of the relevant pages as well as the title page of the volume was obtained from various overseas library sources to verify the bibliographic reference.

Previous workers on the Australian fauna usually did not designate a holotype, even when dealing with a single specimen. Nor did they note the number of specimens they had before them at the time of description except for occasionally noting when they only had a single specimen. The work of both Carter and Lea present problems in this regard. In some instances the authors mentioned a holotype in the original description and the museum it was located in. On investigation it was found that the specimen involved was only marked 'type'. In such cases where there is no ambiguity, I have cited the existence of a holotype. However, occasionally I was confronted by more than one specimen on a single card with no definite indication of which specimen is the holotype, other than they are types. I have interpreted this situation as indicating the existence of a syntype series. On other occasions both authors failed to even designate any specimen as the 'type' or holotype, thus I have also interpreted this situation as indicating the existence of a syntype series in the absence of a definite statement that the new species description was based on a single specimen.

I have checked the location of type specimens in the Australian National Insect Collection (ANIC), Australian Museum, Sydney (AM), The Natural History Museum, London (BMNH), Museum of Victoria, Melbourne (NMV) and the South Australian Museum, Adelaide (SAMA). Where a lectotype is listed without paralectotypes, the existence or whereabouts of any paralectotypes is unknown. I thank the following curators and collection managers for their assistance and access to collections under their care: Dr Max Moulds (AM), Dr E.G. Matthews (SAMA), Dr Ken Walker and Ms Catriona McPhee (NMV) and Mr Martin Brendall and Mr Malcolm Kerley (BMNH).

The date of publication of several items cited in this database varies from that which is normally given. The following references have been helpful in this regard:
Griffin (1932) for the dates of publications and contents of the parts of Westwood's Introduction to the Modern Classification of Insects;
Cowan (1970) for the dates of the Atlas plates for Duperrey's Voyage autour du Monde on S.M. Corvette Coquille in 1822–1825;
Hayek (1983) for the date of publication of the fourth volume of Silbermann's Revue Entomologique;
the date of publication of Captain Phillip King's Narrative of a Survey of the Intertropical and western Coasts of Australia was taken from Common & Moulds (1973); and
Madge (1988) was used as the source for the publication dates of Dejean's Catalogues.

Common names for a several elateroid beetles have been taken from Naumann (1993).

Loaded into Platypus 2.0, 25 October 1999, then repaired, compacted.

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 DEG 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.