Eur. J. Entorno!. 99: 315-351, 2002 ISSN 1210-5759 Generic revision and phylogenetic analysis of the Metriorrhynchinae (Coleóptera: Lycidae) Ladislav BOCAK Faculty of Science, Palacký University, tř. Svobody 26, 771 46 Olomouc, Czech Republic; e-mail: [email protected] Key words.Coleóptera, Metriorrhynchmae, generic revision, phylogeny, taxonomy, key, World Abstract. The subfamily Metriorrhynchinae is the most species-rich clade of Lycidae (Coleoptera). A recent proposal suggests that the Erotinae is a sister group of the Metriorrhynchinae. Within the Metriorrhynchinae, evidence is presented for the monophyly of the Conderini and Metriorrhynchini and their sister group position. The Trichalina, Hemiconderina and Metriorrhynchina form the tribe Metriorrhynchini. The relationships between the basal lineages of this group are poorly understood. Several clades are distin­ guished within the Metriorrhynchina, but there is only weak evidence supporting a relationships between them. The distribution of individual clades is discussed. Carathrix Kleine, 1926 (=Pseudodontocerus Pic, 1921), Dilolycus Kleine, 1926 (= Metriorrhynchus Gemminger et Harold, 1869),Flabelloporrostoma Pic, 1923 (= Metriorrhynchus Gemminger et Harold, 1869),Rossioptera Kasant- sev, 1988 (=Xylobanellus Kleine, 1930), Samanga Pic, 1921 (= Broxylus C.O. Waterhouse, 1879),Strophicus C. O. Waterhouse, 1879 (=Enylus C.O. Waterhouse, 1879), andTapromenoeus Bocak et Bocakova, 1989 (=Prometanoeus Kleine, 1925) are proposed as junior synonyms. Pseudosynchonnus Pic, 1922 is transferred to the Erotinae (Taphini) and Pseudosynchonnus Pic, 1922, Prota- phes Kleine, 1926, andParapyropterus Kleine, 1926 are proposed to be junior subjective synonyms ofLycoprogenthes Pic, 1915. Redescriptions of Metriorrhynchinae genera and a key to genera are provided. INTRODUCTION first to introduce the genera of Metriorrhynchinae to lycid classification: Cladophorus Guerin-Meneville, 1830, The Metriorrhynchinae is the largest subfamily of Lyci­ Metriorhynchus Guerin-Meneville, 1838, andPorrostoma dae, both in the number of genera and number of species. Castelnau, 1838. Various authors disagreed with the Kleine (1933) listed in the last World Catalogue about dating of their descriptions, and the validity of these taxa 1200 species in 41 genera now classified in this was clarified much later by Bocak (1998c). Numerous subfamily. That is, in terms species this subfamily make authors added further species to various genera (e. g. up over 40 percent of the Lycidae. Most Metrior­ Erichson, Blanchard, Thomson, Walker, Motschulsky, rhynchinae are from Southeast Asia, the Indonesian Redtenbacher, Kiesenwetter, and Kirsch). C.O. Water­ Archipelago, New Guinea and Northern Australia. Fur­ house (1877, 1879) alone proposed thirteen new genera ther species occur in Africa, Madagascar, temperate Asia, which still form the backbone of Metriorrhynchinae clas­ Ceylon, and India. Only one species occurs in Eastern sification. He described also numerous new species from Europe and in Siberia. Within Lycidae, Metrior­ different parts of the Metriorrhynchinae range of distribu­ rhynchinae are defined by their circular phallobase (Figs tion. Further species were proposed in the last decades of 91-132), flat, conspicuous, unpaired vaginal gland in the XIXth century (Blackburn, Bourgeois, Fairmaire, most representatives (Figs 143-160), and a pronotum Gorham, Harold, Kirsch, Kiesenwetter, Lea, W. M. Leay, with lanceolate median areola, usually with lateral and and Schaufuss). Only two authors, R. Kleine and M. Pic, frontolateral pronotal carinae (Figs 57-86). Metrior­ contributed substantially to the knowledge of Lycidae rhynchinae appear to be close to the Erotinae as they have before World War II. They described hundreds of new very similar pronotal carinae in some forms, structurally species and several new genera. Kleine and Pic defined similar elytral costae, male and female genitalia similar in genera mostly on the basis of the presence or absence of some features, and generally similar larvae (Bocak and secondary costae, flabellate antennae or the number of Matsuda, in press). pronotal carinae. Congruence of these criteria with other Fabricius (1775) described the first species that is now characters needs to be established. Unlike Pic, Kleine dis­ placed in Metriorrhynchinae (Pyrochroa serraticornis sected genitalia, but preferred other characters for classi­ now classified in Trichalus) and later proposed several fication. Consequently, Kleine (1926a) classified additional species (Fabricius, 1801). Numerous entomolo­ Metriorrhynchus parallelus in Cladophorus on the basis gists working on the material collected by scientific expe­ of its flabellate antennae, although it is the type species of ditions in the first half of the XIXth century (Dalman, Metriorrhynchus, which he recognised. Fahraeus, Macleay, Hope, Klug, and Boisduval) fol­ Only a few species were described after World War II. lowed Fabricius. They described few species and classi­ Nakane (1968, 1969a, b, 1971, 1980) studied the fauna of fied them in then genera Lycus F., 1787, Calopteron Japan and Taiwan, Kasantsev (1988) described the genus Castelnau, 1838, orDictyoptera Latreille, 1829. Guerin- Rossioptera (= Xylobanellus Kleine) from Russia, but he Meneville (1830, 1838) and Castelnau (1838), were thedid not classify it in the Metriorrhynchinae. Bocak & 315 Bocakova (1987b, 1989, 1990b, 1991, 1999), Bocakobserved in early morning and shortly before sunset. (1998a, b, d, e, 1999a, b, 2000a, b, c, d, 2001a, b), andSome species are attracted to light but the proportion of Bocak & Matsuda (1998) published several geographi­ Lycidae collected at lights is generally low. Unlike Lyci­ cally limited revisions and descriptions of new species nae, the Metriorrhynchinae are rarely attracted to flowers. from the Oriental Region and New Guinea. Bocak Leptotrichalus are the only Metriorrhynchinae regularly (1998c) solved the long lasting uncertainty about the collected on flowers in South East Asia. Flowers in Aus­ status of the genera described by Guérin-Méneville. tralia attract at least some species of Porrostoma and Tri- Calder (1998) compiled a catalogue of Australian Lycidae chalus. Larvae were collected in decaying wood. Lycidae and proposed 85 new combinations within the Porros- generally seek moist conditions and inhabit rotten trunks toma. These new combinations were not based on the or branches in shaded places, often laying on the ground study of type specimens, and therefore the generic classi­ and in close contact with moist soil. Several larvae of fication of Australian Lycidae remains obscure. Metriorrhynchinae have been described. Burakowski At present, the entire classification of the Metrior- (1988) described the larva and pupa Xylobanellusof rhynchinae at the species and generic levels is chaotic. A erythropterus; Hayashi (1954), Fukuda & Hayashi large part of this confusion results from the inadequate (1981), and Hayashi (1986) described several Japanese nature of Pic’s descriptions and unclear reasons for his species of Cautires. Potozkaja (1981) described the larva proposals. He thought that as his descriptions were long of Xylobanus angusticollis (Motschulsky). Other larvae they would be useful (W. Wittmer, pers. comm.). As of Metriorrhynchinae were reported recently by Bocak & Kleine usually ignored Pic’s descriptions, revisions of Matsuda (in press). Metriorrhynchinae are often involved most genera are urgently needed. in Mullerian mimicry complexes, Lawrence & Britton The subfamily Metriorrhynchinae was proposed by (1991) mentioned those from Australia. Moore & Brown Kleine (1926a), along with the subfamily Dilolycinae (1981) studied the antifeedants in the bodyof Porrostoma (=Haplothoracinae, nomen nudum). Later he described rhippidium (W. M. Leay, 1827). the subfamilies Trichalinae and Cladophorinae (Kleine, MATERIAL AND METHODS 1928). Kleine (1933) designated all these taxa as tribes of Lycinae. Bocak & Bocakova (1990a) proposed the tribe Taxa Conderini and subtribe Hemiconderina, synonymised Altogether 45 genera were classified in the Metriorrhynchinae Cladophorinae and Dilolycinae with Metriorrhynchinae and Bocakova (in press) transferred Broxylus to the Metrior­ and downgraded Trichalinae to the subtribal level. Metri­ rhynchinae: for a list of genera see Appendix 1. The phyloge­ orrhynchinae was made up of two tribes, Conderini and netic analysis deals with 34 genera of Metriorrhynchinae. One Metriorrhynchini, the later being divided to the subtribes of Metriorrhynchinae genera was found to be a member of the Erotinae (Pseudosynchonnus). Four genera were synonymised Trichalina, Metriorrhynchina and Hemiconderina. Bocak prior the analysis, because they did not differ from genera (1998b) revised the generic classification of the Tri- included in the analysis: Dilolycus (= Metriorrhynchus), Flabel- chalina. loporrostoma (= Metriorrhynchus), Samanga (= Broxylus), and The species richness and high endemism in the transi­ Strophicus (= Synchonnus). One genus was not found in Pic’s tional Austro-Oriental zone, as well as the restriction of collection and its identity is unknown (Falsolucidota). The Lycidae to moist forests and usually to mountainous habi­ genera Mimoxylobanus, Oriomum, Cladophorinus, Xylobano- tats, suggest that Metriorrhynchinae could be a valuable morphus, Xylobanomimus, and Malacolycus are available only model group for evolutionary and zoogeographical stud­ as single specimens of one sex, and therefore