Classification & Phylogeny
No doubt because of their common occurrence in Europe and distinctiveness, ladybirds were known to the earliest entomologists. Linnaeus in 1758 established the genus Coccinella and described 36 species that currently are assigned to several genera and some to different beetle families. Obviously for C. Linnaeus the overall rounded shape, spotted elytra and short, clubbed antennae were the distinguishing characteristics of the genus. Since Linnaeus' time, many new species have been described by many authors. However, it was about 100 years before two major works by the French entomologist, Etienne Mulsant (1848, 1850), set the foundations for the modern classification of Coccinellidae. Mulsant's publications included first ever supra-generic categories (Tribes) in Coccinellidae, keys to genera and descriptions of all (then) known world species. Whilst recognising many obvious errors or misinterpretations by Mulsant, we can but appreciate the ingenuity of Mulsant in recognising several natural groups and discovering important morphological characters in spite of limited optical equipment at his disposal. Mulsant's monograph was revised and improved by George Crotch. Crotch acquired most of the collections originally studied by Mulsant. Crotch (1874) provided a critical catalogue of the world taxa and proposed a slightly altered classification that departed from the division of Coccinellidae into the hairy 'Trichoisomides' and glabrous 'Gymnosomides' of Mulsant, and classified setose Epilachnides as a subgroup of the subfamily Coccinellidae. Crotch also described numerous new genera and species and synonymised many of the Mulsant taxa.
The end of 19th and the beginning of 20th Centuries saw numerous publications of two major workers, the German Julius Weise and the Frenchman A. Sicard in addition to single but important publications about Central European species by Ludwig Ganglbauer and North American fauna by Thomas Casey. Casey proposed many generic and tribal divisions of Coccinellidae that were based on his extensive collection of the North American beetles but also on examination of many world groups. Both Weise and Sicard published papers, refining the existing classification by describing new genera and proposing tribal groups.
According to Gordon (1985), Weise was apparently the first coccinellid taxonomist to realise that male genitalia could be useful to distinguish species. In the early 20th Century, one of Weise's pupils, Richard Korschefsky (1931, 1932) published the Coccinellidae portion of the 'Coleopterorum Catalogus', to date the only complete world catalogue of Coccinellidae that has ever been published.
The second half of the 20th Century was a 'Golden Age of Coccinellidae Taxonomy'. This was a period of intensive research on Coccinellidae by a very productive group of people, namely Hiroyuki Sasaji, Helmut Fürsch, Ryszard Bielawski, Robert Gordon Theodosius Dobzhansky, A.P. Kapur, Leopold Mader, Jean Chazeau, Matsuo Miyatake and Robert Pope, with many contributions on a local scale by other workers.
Certainly the publication of 'Phylogeny of the family Coccinellidae (Coleoptera)' by H. Sasaji in 1968 was a landmark in coccinellid research, comparable in significance with the Mulsant's monograph. Sasaji studied and analysed critically both adult and larval characters of Coccinellidae. He constructed a phylogenetic tree of the major groups and proposed a classification based on that tree. Other workers, mostly Fürsch and Gordon refined Sasaji's classification, and modified and extended it to the regions and groups of their interests. The most recent major contribution to the systematics of Coccinellidae were papers by Ivo Kovář (1996) who presented his ideas on phylogeny of Coccinellidae and briefly discussed major recognised groups and their supporting features. Kovář proposed a classification of Coccinellidae worldwide, recognising seven subfamilies and 38 tribes.
Although the cladistic morphology-based analyses and molecular studies are as yet incomplete and preliminary, there is sufficient evidence to support the first split of the branches of the coccinellid phylogeny that are here regarded as subfamilies. Microweiseinae includes three tribes (Microweiseini, Serangiini and Sukunahikonini) that share the unique asymmetrical tegmen with reduced and fused parameres, multi-cameral and sclerotised spermatheca, antennal insertions positioned close together between the eyes or in front of the eyes, and the antenna short, clubbed and comprising a maximum of 10 segments. Their larvae have the integument granulate, the mandible simple without a molar lobe and tibiotarsal apex with 2 spatulate setae; the pupa does not have urogomphi (Phuoc & Stehr 1974).There is probably no clear apomorphy uniting the rest of the ladybirds, here called Coccinellinae, except perhaps the symmetrical tegminal phallobase and well-developed and articulated parameres (reduced several times independently). The larval mandible has a molar lobe (lost in Epilachnini) and the tibiotarsal setae are clubbed or simple and usually in larger numbers.
Vandenberg (2002) discussed major problems with the current classification of Coccinellidae and, following Pope (1989), identified Coccidulinae-Scymninae as the most problematic area in Coccinellidae classification. From an Australian perspective her statement is very important as most of the Australian taxa belong to these two subfamilies (Pope 1989). Traditional characteristics, e.g. antennae short and eyes finely facetted in Scymninae versus antennae longer and eyes coarsely facetted in Coccidulinae do not hold for the southern taxa, many of which are intermediate (e.g. Scymnodes). Research has not so far provided satisfactory resolution in this area and we follow suggestion of Slipinski (2007) that all taxa previously classified in Coccidulinae and Scymninae that remained after removing distinct entities as Noviini, Scymniliini and Diomini have been lumped together into one tribal level taxon, here called Coccidulini, pending further results from morphological and molecular research.
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