Generic revision and phylogenetic analysis of the Metriorrhynchinae ( Coleóptera : Lycidae )

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 Kasantsev, 1988 (= Xylobanellus Kleine, 1930), Samanga Pic, 1921 (= Broxylus C.O. Waterhouse, 1879), Strophicus C. O. Waterhouse, 1879 (= Enylus C.O. Waterhouse, 1879), and Tapromenoeus 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, Protaphes Kleine, 1926, and Parapyropterus Kleine, 1926 are proposed to be junior subjective synonyms of Lycoprogenthes Pic, 1915. Redescriptions of Metriorrhynchinae genera and a key to genera are provided.


INTRODUCTION
The Metriorrhynchinae is the largest subfamily of Lyci dae, both in the number of genera and number of species.Kleine (1933) listed in the last World Catalogue about 1200 species in 41 genera now classified in this subfamily.That is, in terms species this subfamily make up over 40 percent of the Lycidae.Most Metrior rhynchinae are from Southeast Asia, the Indonesian Archipelago, New Guinea and Northern Australia.Fur ther species occur in Africa, Madagascar, temperate Asia, Ceylon, and India.Only one species occurs in Eastern Europe and in Siberia.Within Lycidae, Metrior rhynchinae are defined by their circular phallobase , flat, conspicuous, unpaired vaginal gland in most representatives , and a pronotum with lanceolate median areola, usually with lateral and frontolateral pronotal carinae .Metrior rhynchinae appear to be close to the Erotinae as they have very similar pronotal carinae in some forms, structurally similar elytral costae, male and female genitalia similar in some features, and generally similar larvae (Bocak and Matsuda, in press).Fabricius (1775) described the first species that is now placed in Metriorrhynchinae (Pyrochroa serraticornis now classified in Trichalus) and later proposed several additional species (Fabricius, 1801).Numerous entomolo gists working on the material collected by scientific expe ditions in the first half of the XIXth century (Dalman, Fahraeus, Macleay, Hope, Klug, and Boisduval) fol lowed Fabricius.They described few species and classi fied them in then genera Lycus F., 1787, Calopteron Castelnau, 1838, or Dictyoptera Latreille, 1829. Guerin-Meneville (1830, 1838) and Castelnau (1838), were the first to introduce the genera of Metriorrhynchinae to lycid classification: Cladophorus Guerin-Meneville, 1830, Metriorhynchus Guerin-Meneville, 1838, and Porrostoma Castelnau, 1838.Various authors disagreed with the dating of their descriptions, and the validity of these taxa was clarified much later by Bocak (1998c).Numerous authors added further species to various genera (e. g.Erichson, Blanchard, Thomson, Walker, Motschulsky, Redtenbacher, Kiesenwetter, and Kirsch).C.O. Water house (1877, 1879) alone proposed thirteen new genera which still form the backbone of Metriorrhynchinae clas sification.He described also numerous new species from different parts of the Metriorrhynchinae range of distribu tion.Further species were proposed in the last decades of the XIXth century (Blackburn, Bourgeois, Fairmaire, Gorham, Harold, Kirsch, Kiesenwetter, Lea, W. M. Leay, and Schaufuss).Only two authors, R. Kleine and M. Pic, contributed substantially to the knowledge of Lycidae before World War II.They described hundreds of new species and several new genera.Kleine and Pic defined genera mostly on the basis of the presence or absence of secondary costae, flabellate antennae or the number of pronotal carinae.Congruence of these criteria with other characters needs to be established.Unlike Pic, Kleine dis sected genitalia, but preferred other characters for classi fication.Consequently, Kleine (1926a) classified Metriorrhynchus parallelus in Cladophorus on the basis of its flabellate antennae, although it is the type species of Metriorrhynchus, which he recognised.
Only a few species were described after World War II.Nakane (1968Nakane ( , 1969aNakane ( , b, 1971Nakane ( , 1980) ) studied the fauna of Japan and Taiwan, Kasantsev (1988) described the genus Rossioptera (= Xylobanellus Kleine) from Russia, but he did not classify it in the Metriorrhynchinae.Bocak & Bocakova (1987b, 1989, 1990b, 1991, 1999), Bocak (1998aBocak ( , b, d, e, 1999aBocak ( , b, 2000aBocak ( , b, c, d, 2001a, b), b), and Bocak & Matsuda (1998) published several geographi cally limited revisions and descriptions of new species from the Oriental Region and New Guinea.Bocak (1998c) solved the long lasting uncertainty about the status of the genera described by Guérin-Méneville.Calder (1998) compiled a catalogue of Australian Lycidae and proposed 85 new combinations within the Porrostoma.These new combinations were not based on the study of type specimens, and therefore the generic classi fication of Australian Lycidae remains obscure.
At present, the entire classification of the Metriorrhynchinae at the species and generic levels is chaotic.A large part of this confusion results from the inadequate nature of Pic's descriptions and unclear reasons for his proposals.He thought that as his descriptions were long they would be useful (W. Wittmer, pers. comm.).As Kleine usually ignored Pic's descriptions, revisions of most genera are urgently needed.
The species richness and high endemism in the transi tional Austro-Oriental zone, as well as the restriction of Lycidae to moist forests and usually to mountainous habi tats, suggest that Metriorrhynchinae could be a valuable model group for evolutionary and zoogeographical stud ies.This characteristic is combined with the fact that Lycidae are models in numerous mimicry complexes.Despite all this, the poor knowledge of Metriorrhynchinae has prevented them from being used for more general studies.Research on this group has suffered from the absence of comprehensive definitions of genera as well as the doubtful phylogenetic affinities between genera and tribes.This paper should establish the basis for further study of this extremely diverse group.

Natural history
Metriorrhynchinae are one of the commonest groups of Lycidae in tropical Asia and Australia.The adults live mostly on the leaves of shrubs and herbs in dense forest canopy.Although most specimens in museum collections were collected individually or by sweeping of the lower strata of tropical forests, Metriorrhynchinae are active also in the upper strata as shown by the material obtained by insecticidal fumigation in Sulawesi (Bocak, 2000c).They are generally slow moving and fly only occasionally and reluctantly.The highest dispersal activity was observed in early morning and shortly before sunset.Some species are attracted to light but the proportion of Lycidae collected at lights is generally low.Unlike Lyci nae, the Metriorrhynchinae are rarely attracted to flowers.Leptotrichalus are the only Metriorrhynchinae regularly collected on flowers in South East Asia.Flowers in Aus tralia attract at least some species of Porrostoma and Trichalus.Larvae were collected in decaying wood.Lycidae generally seek moist conditions and inhabit rotten trunks or branches in shaded places, often laying on the ground and in close contact with moist soil.Several larvae of Metriorrhynchinae have been described.Burakowski (1988) described the larva and pupa of Xylobanellus erythropterus; Hayashi (1954), Fukuda & Hayashi (1981), and Hayashi (1986) described several Japanese species of Cautires.Potozkaja (1981) described the larva of Xylobanus angusticollis (Motschulsky).Other larvae of Metriorrhynchinae were reported recently by Bocak & Matsuda (in press).Metriorrhynchinae are often involved in Mullerian mimicry complexes, Lawrence & Britton (1991) mentioned those from Australia.Moore & Brown (1981) studied the antifeedants in the body of Porrostoma rhippidium (W.M. Leay, 1827).

Taxa
Altogether 45 genera were classified in the Metriorrhynchinae and Bocakova (in press) transferred Broxylus to the Metrior rhynchinae: for a list of genera see Appendix 1.The phyloge netic analysis deals with 34 genera of Metriorrhynchinae.One of Metriorrhynchinae genera was found to be a member of the Erotinae (Pseudosynchonnus).Four genera were synonymised prior the analysis, because they did not differ from genera included in the analysis: Dilolycus (= Metriorrhynchus), Flabelloporrostoma (= Metriorrhynchus), Samanga (= Broxylus), and Strophicus (= Synchonnus).One genus was not found in Pic's collection and its identity is unknown (Falsolucidota).The genera Mimoxylobanus, Oriomum, Cladophorinus, Xylobanomorphus, Xylobanomimus, and Malacolycus are available only as single specimens of one sex, and therefore they were excluded from the analysis.Their redescriptions and discussion of their possible systematic position are given in the taxonomic part.The proportion of unknown character states is very high for these genera and this uncertainty collapsed the basal part of the tree of Metriorrhynchini to the unresolved bush.Wiens (1998) discussed the consequence of a high proportion of missing data.All the genera excluded from the analysis as well as genus of unknown identity are monotypic, therefore their exclusion did not restrict substantially the scope of the analysis.All type species of Metriorrhynchinae genera were studied except Falsolucidota Pic, 1921b, which should be deposited in the National Museum of Natural History in Paris, but was not found in Pic's collection.Bocak & Bocakova (1990a) reclassified the Metriorrhynchi nae.Evidence for monophyly was sought in the morphology of the ingroup and trees were rooted using the outgroup method (Watrous & Wheeler, 1981).A sister-group for the Metrior rhynchinae was not suggested by previous authors and the pos sibility of a close relationship between Erotinae and Metriorrhynchinae based on larval characters was briefly dis cussed by Bocak & Matsuda (in press).In addition, Erotinae and Metriorrhynchinae have the same arrangement of elytral costae, carinae on the pronotum and slender parameres (present only in Figs 1-2: Cladograms. 1 -the strict consensus tree derived from three trees provided by an analysis of the matrix in Table 2 (mh*, bb*); 2 -the strict consensus tree based on two trees obtained after application of successive weighting.The numbers above branches correspond to node numbers in Tab. 3.
Conderini within the Metriorrhynchinae).The phylogeny of Lycidae has not yet been analysed, and therefore multiple out groups were chosen and the impact of switching to different out groups tested.The following taxa were used as outgroups: Dictyoptera (represented by D. aurora (Herbst)) and Pyropterus (P.nigroruber (Degeer)) from Erotinae as presumably the closest sister-group, Lyponia (L.(Ponyalis) laticornis Fairmaire), which is considered to be the most basal member of Platerodinae (Bocakova, 2001), and Calochromus (Calochromus sp.from Yunnan) from the Calochrominae.Dictyoptera and Pyropterus represent genera that have and lack secondary elytral costae.This character was thought to be important for the generic classification of Lycidae.The selected outgroups represent all potential relatives of Metriorrhynchinae.The remaining subfamilies of Lycidae differ substantially in adult and larval characters (Lycinae: Lycini and Macrolycini) and in addition some of them have neotenous females (Duliticola, Platerodrilus, presumably Ateliinae, Leptolycinae: Dexorini).

Characters
I identified 72 characters, 67 potentially informative for the relationships within an ingroup and five of them constant within an ingroup and used for its definition.Polymorphic characters in terminal taxa were coded as "-" in the matrix, and all characters are described in Table 1.Autapomorphic characters of indi vidual genera were not included and they are discussed in the taxonomic part.When possible, several species of each genus were scored for each character.The genera included in the analysis were always available as type(s) or identified speci mens of type species of both sexes.In several cases, data from disarticulated closely related species were compared with the type species.Because of the danger of inappropriate sampling, the examination of several representatives of each genus was preferred.This was easy in case for recently revised genera, but difficult in the case of the very large genera used by M. Pic (Metriorrhynchus, Cladophorus, Cautires), for genera typologi cally consisting of very diverse groups of unrelated species (Procautires) or genera formally monotypic but based on numerous undescribed species.I preferred to score the group of species closely related to the type-species.An alpha-taxonomic study of such genera was beyond the scope of this paper.The present study is based exclusively on adults.Both male and female genitalia of all available type specimens were dis sected and the membranous parts stained lightly with chlorazol black.Important characters were drawn using an ocular grid-screen or illustrations were derived from photographs taken with an Olympus DP-10 or Camedia 3000 digital camera attached to an Olympus SZX-12 stereoscopic microscope.(13)(14)(15)(16)(18)(19)(20)(21)(22).

Results
When Calochromus, Lyponia, Dictyoptera, and Pyropterus were used to root cladograms, the analysis (mh*, bb*) yielded three parsimonious trees, each with 261 steps, a Cl = 0.39 and a RI = 0.69.The strict consensus tree derived from them was identical with one of the original trees and included a polytomy in Trichalina (Fig. 1), and therefore the length, CI and RI of these trees did not differ.When only one of the original outgroup taxa was designated as an outgroup, the topology of the tree did not change.The mapping of characters is given in table 3, and the number of clades in Fig. 1.
Further, successive weighting was applied to the matrix after mh* and bb* (again with Calochromus, Lyponia, Dictyoptera and Pyropterus as outgroups).Two trees with length 651, a CI = 69 and a RI = 86 were obtained.These trees differed in the position of Xylobanus, either being a sister-group of Cautires or Broxylus.The con sensus tree was slightly longer (length 653) and had iden tical indexes.The strict consensus tree differs in that it has a polytomy in the Cautires clade.

Discussion
The present analysis is the first attempt to investigate the phylogeny of the Metriorrhynchinae using a cladistic approach.The only classification of Lycidae (Bocak & Bocakova, 1990a) is not based on phylogenetic principles but on the need to diagnose supergeneric taxa for a com parative morphology of the major groups of Lycidae, and their identification.Nevertheless, the monophyly of the Metrirrhynchinae as understood by Bocak & Bocakova (1990a), as well as their constitutive tribes and subtribes is supported by the present phylogenetic analysis.The   2).The monophyly of the Metriorrhynchinae is supported by several synapomorphic characters: transverse gula (Fig. 11), considerably compressed antennae, antennomere 2 much shorter than antennomere 3 (Figs 29-51), unconstricted base of median areola , median areola connected to frontal margin of the pronotum by a keel , symmetrical bulges at base of pronotum absent, circular phallobase .The shape of the phallobase is very characteristic and unlike that of other Lycidae.In addition, the shape of the pro notal carinae is very characteristic and only specimens of Enylus is these an indistinct frontal keel, but even in such cases the general appearance of the pronotum is different.The very short antennomere 2 is possibly a synapomor phic character state, although Ateliinae have a very short antennomere 2 as have several Leptolycinae.Both groups differ substantially in the shape of their cranium, male genitalia and Ateliinae have pobably neotenous females (Bocak & Matsuda, in press).These groups are unrelated to the Metriorrhynchinae, therefore an independent origin of these character states in the Metriorrhynchinae is sug gested.
The Conderini, according to the present phylogenetic hypothesis (Figs 1, 2), have a basal placement within the Metriorrhynchinae.That the phallus has a laterally com pressed apical part, and a hook-like base, and there are processes on the phallobase support their monophyly.All these characters are unique within the Lycidae.Addition ally, the shape of the pronotal carinae is very character istic in most species (Figs 57,58), but the shape of the carinae is not the same in all specimens of some species and therefore was not coded in the matrix.Conderini share several symplesiomorphic character states with the  Tab.3, and the characters not mentioned here have under gone multiple reversions.The absence of parameres are hypothesised to be a homoplasy, because this character state has evolved independently in Lyponia (Platerodinae, char.54) and is known also in the Platerodini, which were not included as an outgroup in the analysis.Metriorrhynchini differ in this character from all similar Lycidae.However, the loss of parameres has occurred independ ently in other groups.
The tribe Metriorrhynchini is formed by three subtribes: Trichalina, Hemiconderina and Metrirrhynchina.Their relationship is unresolved.When no successive weighting was applied, the Trichalina had a basal position as a sister-group of the remaining Metriorrhynchini, and the Hemiconderina were the next basal clade.After succes sive weighting the relative position of these subtribes changed (Figs 1,2).But in all analyses these clades remained stable.Unfortunately the larvae of these groups are unknown and I have not found a character to corrobo rate the hypotheses.Both hypotheses can be interpreted in the same way in the formal classification.Kleine (1928Kleine ( , 1933) ) defined the Trichalina as a group of Metriorrhynchinae genera with a short elytral primary costa 1. Bocak (1998b) showed that the shortened costa 1 evolved several times in the Metriorrhynchinae and there fore restricted Trichalina to the terminal clade Trichalus + Microtrichalus + Flabellotrichalus + Schizotrichalus + Eniclases and provisionally classified Enylus and Diatrichalus as incertae sedis in the Metriorrhynchini.The pre sent analysis placed Enylus and Diatrichalus basally in this clade because of two synapomorphic character states: tubercles on the lateral margin of the pronotum (Figs 61-66) and the shape of the basal part of phallus (Figs 104,106).The tubercles may or may not be distinct in individual specimens but are regularly present, and together with the characteristically elevated lateral mar gins give the uniform pronotal appearance of the Tri chalina.Considering the topology of the trees produced by the present analysis (Figs 1, 2), Enylus and Diatri chalus must be classified in the Trichalina.
The Hemiconderina as proposed by Bocak & Bocakova (1990a) was monotypic.Later, Bocak (1999b) described Wakarumbia from Sulawesi put it in the Hemiconderina.This was supported by the present analysis and addition ally the Australian Achras was added to the terminal clade which is characterised by straight slender costae (char.27).In both cladograms (Figs 1, 2) Synchonnus is a basal member of the Hemiconderina.Although genera classified in the Hemiconderina have many similar char acters, it can not be defined as a monophyletic group, as the Hemiconderina clade has no autapomorphic character state.
That the Metriorrhynchina is monophylic is not well founded, as I have not found any synapomorphic char acter state to support the cladistic definition.In addition, the relationships between the individual clades within the Metriorrhynchina remain obscure.
Most members of this clade have a complex pattern of pronotal carinae forming four areoles on the frontal margin of the pronotum (i.e., frontolateral costae are pre sent on the pronotum, character 18).This character is useful for identifying most Metriorrhynchina.On the cladogram in Fig. 1 the Metriorrhynchina without basal clade Bulenides and Caenioxylobanus share the state 18.1, after successive weighting (Fig. 2) the state 18.1 is not shared by the basal Metriorrhynchus+Lobatang+Leptotrichalus clade.Additionally, this character is a homoplasy as it is a similar arrangement of carinae in Schizotrichalus and Eniclases (Fig. 66).The arrangement of carinae in Eniclases is quite different from that of most Metriorrhynchina -only two diverging keels are present (Fig. 66).Schizotrichalus has similar longitudinal carinae but its median frontal carina is forked apically and con nected with longitudinal carinae.Consequently, three carinae are attached to the frontal pronotal margin.Having no evidence to the contrary, both situations were scored as an identical character state.Further evidence will be needed to support or refute the Metriorrhynchina.As few larvae of the Metriorrhynchina have been described (Bocak & Matsuda, in press) they will not help to solve this ambiguity.The larvae of most known Cautires and Xylobanus are very similar and they share some characters with Metriorrhynchus.Porrostoma differs in having characteristic processes on the hind margins of its terga.Most of them share with Xylobanellus terga that are divided in two parts by a membranous line.
Only two constitutive clades give robust support to both cladograms (Figs 1,2): the Ditua group formed by almost exclusively Papuan genera and the Porrostoma group with a restricted Austro-Papuan distribution.The clade Bulenides + Caenioxylobanus is either a basal group (Fig. 1) with the Cautires clade as the next clade, or they are a basal group within the Cautires clade (Fig. 2).
The position of Metanoeus is variable and it is attached as a basal member either to the Porrostoma or Ditua clade.In any case, Metanoeus, has an Oriental distribu tion, and is not related to the Afro-Oriental clade (Cau tires, Bulenides, Prometanoeus, etc.).Some characters present only in some members of these genera could not be coded in the matrix: several Metriorrhynchus species share with Metanoeus very obtuse, indistinct pronotal carinae and both have shorter (Metriorrhynchus) to ves tigial (Metanoeus) valvifers.Further evidence is needed to clarify the position of Metanoeus.

Zoogeography
Unfortunately, the very fragmentary knowledge of Lycidae at the alpha taxonomic level and consequently the doubtful information on generic distribution limit the following discussion.I have used unpublished informa tion gathered by studying collections in the above institu tions.The data in the lycid catalogue (Kleine, 1933) are inapplicable.The typological species concept applied to most genera by Kleine and the problems originating from Pic's confused work means that these data has to be con firmed by studying the original type material.Information on the distribution of genera is given in the systematic part.
On the other hand, Lycidae appear incapable of crossing natural barriers such as sea straits or deserts.Lycidae are often limited to humid mountain habitats and when these mountains are isolated, Lycidae easily speciate.Unlike many other groups, there are very few lycid species with a very wide range.The few widely distrib- uted species are usually from lowlands and in this case the size of range is not impressive.I know only of one species that occurs in two zoogeographical regions.
The substantial part of the Metriorrhynchinae is an east Gondwanan element (distribution of Conderini is dis cussed further) with some representatives distributed in East Asia and a single species widely distributed from Cladophorus sp.; 145-146 -Synchonnus appositus.Scales 0.5 Russian Far East through Siberia to Belarussia and Poland (Fig. 3).The highest species and generic diversity occurs in the humid tropics of the Old World, but the generic species diversity is low in the Afrotropical Region.There are also substantial differences in the con tribution of Metriorrhynchinae to the total representation of Lycidae in the zoogeographic regions.
The striking observation is that Conderini as a basal group within the Metriorrhynchinae have the distribution similar to Erotinae, although they are not recorded from the Nearctic Region.Most species are known from the south-eastern part of the Palaearctic Region and the northern part of the Oriental region, fewer species occur distributed in the north-eastern part of the Palaearctic.Unlike the Erotinae they are not known from the Afrotropical andNearctic Regions (Fig. 3).
The range of Metriorrhynchini is limited to the Afrotropical, Oriental, Australian and eastern part of Palaearctic Regions.It seems reasonable to assume that the Metriorrhynchini dispersed to the Palaearctic Region after they evolved somewhere on the Gondwana continent.There are no endemic genera in the Palaearctic, and generally both the species and generic diversity of Palaearctic fauna are extremely low and decreases with the distance from the Oriental Region.Additionally, the Metriorrhynchinae form usually a small fragment of the Lycidae collected in the Palaearctic Region, unlike in the Oriental Region.The distribution of the Metriorrhynchini is given in Fig. 3.
Another important observation is that within the Metri orrhynchinae the two Australian lineages, Hemiconderina and Trichalina, have a basal position and that the diver sity of Australian fauna is much higher than Afrotropical one.The Hemiconderina are endemic to the Australian Region (Fig. 5) and within the subtribe the basal lineage represented by Synchonnus is restricted to continental Australia as well as Achras, which also has several plesiomorphic characters.The remaining genera of Hemicon derina are recorded from New Guinea and the Indonesian Archipelago east of the Wallace line.Both, Hemiconderis, known from New Guinea and the Yapen Island, and Wakarumbia, from Sulawesi and the Buton Island, have undergone considerable speciation.At pre sent, 23 Wakarumbia species are recorded from Sulawesi.Although, they show three different colour patterns, the substantial differences are in the male genitalia.Consid ering the tectonic history of Sulawesi and the limited range of Wakarumbia, we can presume that a part of Sulawesi of Australian origin or a part having contact with some part forming today New Guinea brought the ancestor of Wakarumbia to Sulawesi.I have not found any group of species restricted to one part of an island (Bocak, 1999b(Bocak, , 2000c(Bocak, , 2001b)), therefore the simultane ous speciation on various mountain ranges of widespread lowland species is possible.
The Trichalina are the other basal group of presumably Australian origin.All genera constituting the Trichalina are known from the Australian Region, only some of them, Diatrichalus and Microtrichalus are present also in the Oriental Region (Fig. 5).The highest diversity is gen erally found in tropical parts of the Australian Region.After the revision of the Papuan fauna almost forty Diatrichalus species are recorded from the Australian Region (Bocak, 2001a), and further species can be expected from Australia.On the other hand, only five Diatrichalus species were recently reported from the Phil ippines (Bocak, 2000a) and only one species from the Asian continent and the Great Sundas.The distribution of Microtrichalus is similar.Although its range reaches fur ther to the north and recently one species was found in southern Yunnan and northern Laos, the diversity of the Oriental fauna is very low and the number of species decreases gradually from the Wallace line to the north west.Congruent with patterns found in other groups and the tectonic history, Microtrichalus is rich in species in the Philippines (Bocak, 1998d).
The range of Metriorrhynchina is almost coincident with that of the Metriorrhynchinae (Fig. 3).The Palaearctic fauna of Metriorrhynchina is poor, both in number of species and genera.Only Cautires and Xylobanus are reported from China, Japan and the Russian Far East.Metriorrhynchina are quite rare in the Palaearctic Region and form a small part of the Palaearctic lycid fauna considering the number of species.Dispersal from the south offers a possible explanation for the present dis tribution ofMetriorrhynchina in the Palaearctic Region.
The Afrotropical fauna of Metriorrhynchina has not yet been revised and according to Kleine (1933) 192 species are reported from the Afrotropical Region.These species are classified in genera Cladophorus, Cautires, Procautires, Caenioxylobanus, and Xylobanus.I have studied available material in Pic's and Kleine's collections and I found that the Afrotropical species placed in Cladophorus and Procautires are not congeneric with Australian repre sentatives of these genera and should be placed in Cau tires.Afrotropical species of Stadenus were transferred to the Erotinae (Bocak & Bocakova, 1992).Therefore, only genera Cautires (about 150 species), Xylobanus (about 40 species), and Caenioxylobanus (2 species) are now con firmed for the Afrotropical Region.The Afrotropical fauna seems species rich, but multiple synonyms can be expected as Pic often described common species several times.Caenioxylobanus reported from Madagascar is the only genus endemic to the region.
The Afrotropical and Indian faunas of Metrior rhynchina are similar regarding the representation of gen era.Cautires, Xylobanus, and Prometaneus are the only genera occuring west of the low Brahmaputra River, and Prometanoeus is the only genus endemic to the western part of the Oriental Region.Cautires is a widely distrib uted genus known from West Africa to the Philippines (Fig. 4) and Japan.Xylobanus has a similar distribution, but unlike Cautires, Xylobanus crosses the Wallace line to the east.This genus has not yet been revised and the real relationship of the Australian representatives is unknown.
Metriorrhynchina from the eastern part of the Oriental Region are more diverse.In addition to Cautires and Xylobanus, two endemic genera are common in South East Asia -Metanoeus and Bulenides.Both do not cross the Wallace line (Fig. 4).Metriorrhynchus is the only genus in the Metriorrhynchina which has its centre of dis tribution in the Australian Region and reaches continental Asia.Although this genus is reported from India (Kleine, 1933), it was not found in collections from localities west ofBurma.
There are several endemic lineages of Metriorrhynchina in the Australian Region: Pseudodontocerus, Stadenus, Ditua, Cladophorus, Cautiromimus, Broxylus, Kassemia, Porrostoma, and Metriorrhynchoides.The ranges of these genera extend to the west in various degrees (Fig. 6).Metriorrhynchus and Lobatang have most of their repre sentatives in the Australian Region but a very small pro portion of them cross the Wallace line onto the Asian continent (Metriorrhynchus) or to the Philippines (Lobatang).The distribution of Leptotrichalus is a bit dif ferent.Most species of Leptotrichalus are reported from the Philippines, several from Sulawesi, the Lesser Sundas, Java, Sumatra and Borneo, and several from the Asian continent, only one of which occurs beyond the Isthmus of Kra (Bocak, 2000b).Recently, several unde scribed species were collected in Halmahera.This sup ports the conclusion about the relationship between Lobatang and Leptotrichalus inferred from morpho logical data in phylogenetic analysis.Lobatang, is a closely related genus known from the Papuan Subregion, Wallacea, and the Philippines.Most species are known from the eastern part of the region.Dispersal followed by speciation of an ancestor inhabiting islands north of Aus tralia possibly lead to the present distribution and species richness of Leptotrichalus.
Two clades are restricted to the Australian Region.The clade Porrostoma, Stadenus and Metriorrhynchoides is the most common group in Australia with few species known from New Guinea.The Ditua clade (node 27, Fig. 1) is recorded mainly from New Guinea and adjacent islands.Few species occur in Australia.Metanoeus has an ambiguous position in the present analysis and is the only genus endemic to the Oriental Region, but related to clades whose centre of distribution is the Australian Region.
There are substantial differences in the relative repre sentation of Metriorrhynchinae within the Lycidae.Metriorrhynchinae are relatively poorly represented in Afrotropical and Palaearctic Regions, where they usually form 10-20 per cent of collected specimens.They are more common in humid areas of South East Asia where they make up slightly over 50 per cent of the specimens.Their relative abundance increases to the east.East of the Wal lace line the Lycidae are almost exclusively represented by the Metriorrhynchinae.The Metriorrhynchinae form 95 per cent of the specimens collected in New Guinea and a substantial part of the Australian lycid fauna.Metrior rhynchinae form 90 per cent of the lycid species recorded from Australia.Dilolycinae Kleine, 1926: 186;Bocak & Bocakova, 1990a: 641. Dilolycini: Kleine, 1933: 84.Typegenus.Dilolycus Kleine, 1926a: 186.Haplothoracinae Kleine, 1926a: 95 (nomen nudum); Bocak & Bocakova, 1990a: 641.Type genus.Haplothorax Kleine, 1926a: 95 (nomen nudum).

Remarks
Distribution.Palaearctic and Oriental Regions.Over 30 spe cies described at present; mainly from China and continental South East Asia.
Remarks.Two genera, Conderis and Xylobanellus are classified as Conderini.Their close relationship is well supported by the unique shape of male (Fig. 91) and female genitalia (Figs 133,142).They are easily distin guished by the presence or absence of secondary elytral costae.The absence of secondary costae is the only advanced feature found in Xylobanellus, but this occurs independently in many lineages within the Lycidae (sev eral occurences in the Metriorrhynchinae).Unfortunately, Conderis does not have any synapomorphic character state and therefore Conderis can be paraphyletic with respect to Xylobanellus.Pseudoconderis Pic, 1921a: 8, hors texte;Bocak, 1998a: 18. Type species.Pseudoconderis gorhami Pic, 1921a: 8;Bocak, 1998a: 18 (by subsequent designation).
Diagnosis.Body small to medium sized, most species cinnabar red.Antennae filiform to slightly serrate (Fig. 29).Pronotum with five areoles (Fig. 58), sometimes median areola reduced in size (as in Fig. 57), narrow, connected with frontal and lateral margins by stout cari nae.Elytra parallel-sided to slightly widened posteriorly, each elytron with four stout, longitudinal primary costae and five much weaker, sometimes interrupted, secondary costae.Transverse costae dense, sometimes irregular, bottom of elytral areoles bald.Terminal male abdominal sternite with hole in basal part.Phallus almost straight in middle, parameres long (Fig. 91).Valvifers with attached sclerites, widened in middle part, basal margin of coxites strengthened(Fig.133).Vaginamembranous.
Distribution.Eastern part of the Palaearctic Region (China, Japan), Oriental Region (mainly continental South East Asia, less common in Himalayas and India, as well as in Sundas; east ernmost record from Lombok).Over 20 species known from the range ofthe genus.
Remarks.Many Conderis species share the same char acteristic elytral structure.The transverse costae are usu ally very dense and the bottoms of elytral areoles are bald, mat and the surface has a fungus-like structure.This is the only apomorphic character of Conderis but not all species.
Distribution.Five species are placed in Xylobanellus, one occurring in Eastern Europe and Siberia, the rest in China, Thai land, Malaysia, Borneo and Java.
Remarks.All Xylobanellus species share several char acters, which potentially could be present in a common ancestor: secondary costae absent, rectangular reticulate cells and male segment A7 with straight margin.The similarly structured elytral costae is also encountered in unrelated lineages within the Metriorrhynchinae (e. g.Broxylus, Caenioxylobanus, Xylobanus) and in other sub families of Lycidae (Lycinae: Calopterini, part; Erotinae, part).Rossioptera is based on Eros erythropterus, previ ously classified in Dictyoptera.Bocak & Bocakova (1987a) transferred E. erythropterus to Xylobanellus.Rossioptera is considered to be a junior subjective synonym of Xylobanellus.

Remarks.
Metriorrhynchini are easily recognisable by the absence of parameres, presence of characteristic pro notal carinae forming up to seven areoles, flat unpaired vaginal gland and narrowly attached lateral glands.Metri orrhynchus Guérin-Méneville, 1838 was considered to be the type genus by Kleine (1926a).Bocak (1998c) dis cussed the validity of the name.
Distribution.Australian Region.Five species reported from Australia, New Guinea, and Jobi Island.
Remarks.Synchonnus has a basal position in Hemiconderina.As in Achras the male genitalia has a pair of spines (95)(96), but unlike this genus it has more robust body and obtuse pronotal carinae.Diagnosis.Body medium-sized.Head without rostrum.Antennae strongly compressed, slender, only antennomeres 3 and 4 broad, subsequent antennomeres narrow, acutely pointed.Maxillary palpi quite robust.Pronotum with almost straight basal margin and posterior angles acutely projected.Disc with five areoles.Elytra with strong primary costae on humeral third of elytra, much weaker further back, primary costae 1 and 3 much weaker apically, secondary costae on apical third of costa 3 absent, on costa 1 irregular.Transverse costae on elytra weak, irregular, elytral cells mostly slightly transverse.Male genitalia robust, short, with two large spines on basal part of internal sac.Ovipositor with short coxites and long, slender valvifers, vagina large, membranous (Fig. 151).

Remarks. Achras was classified with Metriorrhynchus
and Metriorrhynchoides in the Metriorrhynchini (Kleine, 1933) and later was not studied but kept in the Metriorrhynchina (Bocak & Bocakova, 1990a).Based on the pre sent analysis it is transferred to the subtribe Hemiconderina.Achras is characterised by sclerotized, widened basal part of spermaduct and lateral sacks at base of vagina.The second character can very between species analogous to the situation in other genera.Achras seems to be very closely related to Wakarumbia but retains sev eral plesiomorphic character states, i. e. vestiges of secon dary costae, unrotated phallus, pair of basal spines on internal sac.
Wakarumbia and Achras both have characteristically shaped valvifers (Fig. 151), vagina with unpaired gland located on independent frontal sac, both genera have much weaker costae 1 and 3, but which extent the whole length of elytra.They have similar, characteristic labrum and slender lateral processes on mesoscutum.
Remarks.Hemiconderis is the only member of the Hemiconderina having entire secondary costae, shortened primary costa 3 and two spines on internal sac linearly arranged; the first located at base of phallus, the second on apical half.Basal part of valvifers of all females wid ened to some degree (Bocak & Bocakova, 1990b).H. bipustulatus Bocak & Bocakova, 1988 has additionally a shortened costa 1, but to a considerably smaller degree than encountered in Trichalina.
Diagnosis.Body small, very slender, delicate, with slender legs and antennae.Head without rostrum.Antennae slightly serrate to parallel-sided.Pronotum with five areoles, pronotal carinae straight and slender.Elytra almost parallel-sided, with four primary costae, primary costae 1 and 3 weak, secondary costae absent, only ves tiges present at base of elytra.Transverse costae well developed, regular, elytral cells mostly square-shaped.Phallus with one, complex basal spine on internal sac, in most species apical part of phallus rotated to various degrees.
Remarks.Wakarumbia is the only genus of Hemicon derina that lacks secondary costae.Apical part of phallus in most species is characteristically rotated and there is one complex spine at base of the phallus.
Distribution.Australian Region and eastern part of the Ori ental Region: Indochina, Thailand, southernmost Yunnan, Pen insular Malaysia, the Philippines, and the Great Sundas (Fig. 5).Highest diversity in New Guinea and Australia, the number of species decreases to the west of the range and only a few species are known from continental Asia.
Remarks.The genera with shortened elytral costa 1 previously classified in the tribe Trichalina were revised recently by Bocak (1998b) Diagnosis.Body small to medium-sized.Head small, without rostrum.Antennae strongly compressed, similar in both sexes, serrate or antennomeres almost parallel sided, never flabellate.Maxillary palpi robust, apical palpomere with an oblique apex, broad.Pronotum only with median areola, lateral carinae absent, weak lateral tuber cles on lateral margins of pronotum.Elytra with four pri mary costae, primary costa 1 strong over whole length, costae 2 and 4 much stouter than costae 1 and 3, costa 3 much weaker apically.Legs slender, strongly compressed.Male genitalia always with slender, long phallus, with pair of basal spines, partly sclerotized api cally, phallobasal membrane inconspicuous.Ovipositor with short, rather stout valvifers, fused with coxites (Fig. 137), vagina relatively short, bases of glandular ducts robust, accessory glands inserted laterally.Spermatheca slender, very long.Remarks.Enylus is the only genus classified in Trichalina not having a shortened elytral primary costa 1.Only the holotype of Strophicus nigellus was available, and therefore Strophicus was not included in the phyloge netic analysis.Strophicus is externally very similar to Enylus and the only differs in the size of its eyes: very small in males of Strophicus and very large in Enylus.The presence and/or absence of sexual dimorphism in eye size ise regularly observed in genera of Lycidae.This character is correlated with when in the day they fly.Males of species active at night have large eyes.There fore, I propose that Strophicus is a junior subjective synonym of Enylus.
Diagnosis.Body medium-sized, coloration variable but mostly black or black with metallic blue shine, only a few species have a brightly coloured pronotum and basal part of elytra.Never light concolour.Head small, without ros trum.Antennae strongly compressed, antennomeres parallel-sided to acutely serrate in both sexes, seldom very shortly flabellate in male.Maxillary palpi with broad apical palpomere bearing papillae at apex.Pronotum with one more or less broad longitudinal areola in the middle, lateral margins elevated.Elytral primary costa 1 consid erably shortened, reaching at most one third of elytral length, secondary costae irregular to completely absent in some species.Legs relatively strong and short, strongly compressed.Male genitalia with relatively short and robust phallus, internal sac lacks basal spines, regularly strongly sclerotized apically (Figs 100-101).Ovipositor with slender, long valvifers, sometimes incompletely fused at base, vagina relatively short, vaginal glands inserted laterally.Spermaduct robust at base, spermatheca slender, very long.
Distribution.Australian and eastern part of the Oriental Region.Only one species found in continental Malaysia, Java, Borneo and Sumatra.All other Oriental representatives occur in the Philippines.The highest diversity is found in New Guinea.Kleine (1933) placed six species in Diatrichalus.Later, it was found that many Oriental and Papuan species previously classified as Trichalus belonged to Diatrichalus (Bocak, 2000a(Bocak, , 2001a)).At present over 40 species are classified in this genus.
Remarks.In all Diatrichalus the basal part of spermaduct is widened and this unique character is considered to be apomorphic state.This genus originally included only species that lacked secondary costae.A study of most species showed that in Diatrichalus there are transitional states between strong secondary costae, weak irregular secondary costae and complete absence of costae.The internal sac and basal part of female spermaduct in Diatrichalus has a very characteristic and unique shape of and both species that have and lack secondary costae are included in this genus.Diagnosis.Body medium-sized, slightly widened posteriorly, antennae serrate in both sexes.Pronotum with one longitudinal median areola, first elytral primary costa shortened, secondary costae always well developed, regular.Male genitalia with slender phallus, internal sac membranous, with pair of spines at base, internal sac exposed, free.Ovipositor with extremely robust valvifers, fused in basal third, flat, closely attached to coxital processes.Vagina robust, lateral glands attached dorsally, their bases widened, unpaired vaginal gland short, spermathecal duct short, spermatheca large, bulbous.
Distribution.Australian Region: only continental Australia.
Diagnosis.Body small to medium-sized, very often with dorsal part of body at least partly yellow to lightly brown.Antennae serrate in both sexes, labial palpi slen der, apical palpomere slightly narrowed at apex, corn-pressed.Pronotum with one longitudinal median areola.First elytral primary costa shortened, secondary costae regularly present.Male genitalia very uniform within genus.Phallus slender, only basally completely sclerotized, ventrally open apically, internal sac exposed, with pair of sickle-like thorns at base, sometimes with V-shaped rod in membrane.Whole of male genitalia often very slightly pigmented and sclerotized, rendering structure of internal sac inconspicuous.Ovipositor with slender valvifers.Vagina slender to moderately robust, with dorsally attached lateral accessory glands whose bases are more sclerotized, proper glands very fine.Two lateral pockets of unknown function are present in lateral part of vagina.Very slender unpaired gland attached to base of vagina.
Distribution.Australian and Oriental Regions: Sunda Islands, the Philippines, Peninsular Malaysia, Thailand, Indo china, and the southernmost part ofYunnan.
Remarks.Many species formerly placed in Trichalus were transferred to Microtrichalus (Bocak, 1998d(Bocak, , 2000 d) d).At present, 40 species are classified in Microtrichalus, but the faunas of New Guinea and Australia, where the highest diversity is found, have not yet been revised.Lat eral pockets and unpaired basal vaginal gland are the unique apomorphic character states of Microtrichalus.
Stereotrichalus Kleine, 1926a: 183;Kleine, 1930a:  Diagnosis.Body medium sized to large, slightly wid ened posteriorly.Usually dark brown to black with humeral part of elytra lighter.Head without rostrum, antennae flabellate in male (Fig. 27).Pronotum with one longitudinal median areola, frontal and lateral margins of pronotum very often with dense short to very long pubes cence.Elytral primary costa 1 shortened, secondary costae always present.Male genitalia very slender, phallobase usually covered with extensive membrane.Phallus very long, narrow, sclerotized completely only basally, apically sclerotized dorsally, internal sac exposed, pig mented at apex, sometimes with strengthened rod in membrane, never with pair of spines at base.Vagina with two dorsally attached accessory glands, glands very fine, spermaduct long, curved, spermatheca lemon-shaped.
Distribution.Australian Region.Flabellotrichalus is recorded only from New Guinea and Moluccas.At present 13 species are classified in this genus.
Remarks.Long antennal lamellae in male, pubescence on pronotum, and shape of male genitalia are unique characters of Flabellotrichalus within the Trichalina.
Diagnosis.Body medium-sized.Head without rostrum, antennae with parallel-sided to serrate antennomeres 3-10, only a few species antennomeres flabellate in male.Labial palpi slender, apical palpomere parallel-sided.Pro notum divided by two divergent carinae into three fields (Fig. 66).Elytra with first elytral primary costa shortened, with complete secondary costae.Male genitalia with apparent keel on dorsal surface of phallus, with free, membranous internal sac, without sclerotised structures or spines (Fig. 106).Ovipositor with basal processes of cox ites attached to valvifers, lateral glands of vagina attached into the middle part of vagina, glands very fine.
Distribution.Australian Region.At present, 26 species are known from New Guinea and one from the Moluccas.Although I have seen very extensive lycid material from the Solomon Islands it contained no species ofthis genus.
Distribution.Eastern part of Oriental Region: Burma, Thai land, Indochina, Taiwan, Peninsular Malaysia, Sumatra, Java, Borneo, and the Philippines.
Remarks.Bulenides is characteristic in having only a small, narrow areola in middle of pronotum, the carinae forming the areola can be obtuse to absent and conse quently only a strong frontal keel is present on pronotum.The shape of pronotum is variable, but often distinctly narrowed frontally.The shape of pronotum is similar to that of Caenioxylobanus, which differs in the complete absence of secondary elytral costae and tibial spurs.Addi tionally, the form of the frontal margin of the pronotum in Bulenides and Caenioxylobanus is unique .This character state was confirmed in B. obsoletus (type species) and some other species but is absent from some species included in Bulenides.Only a complete revision of Bulenides will clarify the limit of the genus.
Diagnosis.Body medium-sized, completely dark brown to black.Head without rostrum.Antennae robust, strongly compressed, flabellate in male, serrate in female (Fig. 39).Maxillary palpi robust.Pronotum trapezoidal, with a distinct protruding frontal margin, which is formed by a robust vertical surface (Fig. 67).One longitudinal keel present on disc, anterior half of keel sharp and nar row, higher than wide at base, in basal part almost absent, transformed into a very wide, flat mound.Elytra flat, with four longitudinal costae and slender transverse costae.All costae very regular, slender and sharp.Reticulate cells quadrate to slightly transverse (Fig. 88), all costae shiny, black, bottom of cells velvet, dark brown.Male genitalia slender, lanceolate, internal sac with two spines on apical third.Ovipositor robust (Fig. 135), vagina membranous, spermatheca long, bulbous apically (Fig. 140).
Distribution.Afrotropical Region: Madagascar.Two species are endemic to Madagascar.
Distribution.Afrotropical, Australian, and Oriental Regions, Palaearctic Region: China, Japan, Korea, and Russian Far East.Almost 300 species are described forthis genus.

Remarks.
Xylobanus is an extremely species rich and widespread genus.It is characterised by a pronotum with seven areoles (although the carinae can be obtuse to ves tigial on some parts of the pronotum, frontolateral and lat eral pronotal carinae are never completely absent) and elytra lack secondary costae (they are present at most ves tigially on narrow humeral part).Xylobanus and Cautires have a similar shaped pronotum and male and female genitalia.Their close relationship is supported by their very similar larvae (Potozkaja, 1981, Bocak & Matsuda, in press).The affinities with Broxylus are discussed fur ther.Several characters of Xylobanus are variable.The phallus is widely rounded apically and primary gonoporus is sclerotized in many species, including the type species Xylobanus costifer, but the apex is pointed and the gono porus membranous in X. corporaali Pic, 1922 from Sumatra.On the other hand these species both have phallus with a wide base.Xylobanus costifer has valvifers connected at base, as does one species from Bali and some other species, but this sclerite is vestigial or absent in other species.Also presence of spiculum gastrale is variable and some species have flabellate antennae (e. g.X. nigrimembris Pic, 1926d), or serrate antennae (X.humifer Walker, 1858).Only an extensive, detailed alphataxonomical study can clarify the concept of Xylobanus.The absence of secondary costae is an advanced feature, which has evolved several times in Lycidae.Diagnosis.Body medium-sized.Head small, antennae reach beyond the middle of the elytra, compressed.Palpi strongly compressed, maxillary palpi with long, spoon like apical palpomere, palpomere 4 as long as combined length of palpomeres 1-3, labial palpi short, apical pal pomere large (Fig. 15).Pronotum with partly reduced carinae, frontolateral carinae present on areola only, lat eral carinae weak (Fig. 70).Elytra flat, widest in apical tenth or strongly hemispherically widened, with four lon gitudinal, sharp primary costae and thin, sharp transverse costae of similar strength.Elytral cells quadrate to appar ently longitudinal (Fig. 90).Legs slender, strongly com pressed.Trochanters very slender, long.Male genitalia as figured (Fig. 111).
Remarks.Broxylus was classified in the Calopterini (Lycinae) and transferred to the Metriorrhynchinae recently by Bocakova (in press).Samanga and Broxylus are very closely related.They have similar shaped geni talia and large, flat apical palpomeres.These taxa also have quadrate elytral cells with distinct concave bottoms (these are unique in Metriorrhynchinae) and reduced lat eral carinae on the pronotum.Thanks to the considerably dilated elytra Broxylus has a very characteristic general appearance.The last mentioned character is the only dif ference between Broxylus and Samanga and therefore I propose Samanga to be a junior synonym of Broxylus.Elytra of Broxylus fenestratus are less dilated and similar to those of Xylobanus.
Broxylus is very close to Xylobanus from which it dif fers in the shape of its terminal palpomeres.We do not have reliable synapomorphic character state for Xylo banus, and therefore Broxylus may be a modified member of Xylobanus.
Distribution.Afrotropical and Oriental Regions; Palaearctic Region: Russian Far East, China, Japan, Taiwan.Although Cau tires was reported by Kleine (1926aKleine ( , 1933) ) from New Guinea, I have not found any Cautires from this region and Kleine's illus trations of male genitalia are similar to those of Cladophorus or Cautiromimus.In the south-east, the furthermost records of Cautires are from the Philippines, Borneo and Lesser Sundas (Sumbawa).I have not found any representative in Sulawesi or Moluccas and it is not present in New Guinean material.The highest diversity occurs in continental South East Asia, Sumatra, Java and Borneo.Cautires is the commonest lycid in Madagascar, but it is rare in continental Africa, although many species are reported from there.
Remarks.All characters used for the definition of Cau tires (seven pronotal areoles, entire elytral costae and shape of genitalia) are symplesiomorphic character states and at present no apomorphic character state is known.An alpha taxonomic study of the Cautires clade will lead to a better understanding of the relationships.
Distribution.Oriental Region: Ceylon.Two species classi fied in this genus.

Remarks. The shape of genitalia and pronotal carinae indicate a close relationship between Prometanoeus and
Cautires.They differ in absence and/or presence of flabellate antennae in male.The flabellate antennae evolved several times in Lycidae.The monotypic genera Prome tanoeus and Tapromenoeus are very closely related and although they are different in general appearance they have a similar shaped pronotum, pubescence on elytra, and a similar shaped aedeagus.Bocak & Bocakova (1989) based Tapromenoeus on the absence of secondary costae but the analysis showed that the reduction of sec ondary costae happened several times in the Metriorrhynchinae and is also often encountered in other sub families of Lycidae.Prometanoeus does not have any reliable apomorphic character state that separates it from Tapromenoeus.Therefore I consider Tapromenoeus to be ajunior subjective synonym of Prometanoeus.
Diagnosis.Body small to medium-sized, elytra at most slightly widened posteriorly.Head small, without rostrum, with large eyes even in females.Male antennal lamellae long, narrowly attached to basal parts of antennomeres (Fig. 40).Maxillary palpi relatively slender, apical antennomere parallel-sided.Pronotum with seven areoles, carinae obtuse (Fig. 73).Phallus wide basally (Fig. 113).Phallobasal membrane extensive, at least partly sclerotized.Internal sac with three pairs of spines.Vagina often considerably sclerotized, form of sclerotisation highly species specific, narrowed to apex (if vagina rounded, then basal part of spermaduct is robust), unpaired gland attached basally to apically, basal position correlated with sclerotisation of vagina, lateral glands large to medium sized, attached ventrally, when vagina sclerotized, laterally if membranous, basal part of glan dular ducts more or less sclerotized.Spermaduct usually extremely long, sometimes of medium length (in species with membranous vagina).Female valvifers considerably shortened, modified in plates (Fig. 150).
Distribution.Oriental Region.The highest diversity found in Borneo, Sumatra, and Peninsular Malaysia.Metanoeus is very rare in western part of Oriental Region.One species is common in Mindanao.Altogether over 20 species are classified in Metanoeus.
Remarks.The very obtuse pronotal carinae and ves tigial valvifers are characteristic of Metanoeus.Similar obtuse carinae are found in some Metriorrhynchus, which differ in the much shorter lamellae of male antennae, long slender phallus and only shortened valvifers.The scleroti sation of vagina, long processes on scutellum and long, basally attached male antennal lamellae reminiscent of some Papuan representatives of the Metriorrhynchinae.
No genera bearing similar characters were found within the range of Metanoeus.Further characters supporting the relationship of Metanoeus within Metriorrhynchina are need.Diagnosis.Body large, parallel-sided.Head small, with short, stout rostrum or without rostrum, galeae long, much longer than mandibles.Antennae serrate in both sexes or flabellate in male and serrate in female (Fig. 36).Maxillary palpi slender, long, apical palpomere slender, compressed, pointed at apex (as in Fig. 14).Pronotum with seven, in middle sometimes reduced, areoles on disc.Four strong elytral primary costae developed, costa 3 sometimes slightly shortened (by one eighth at most), slightly weaker in apical half of elytra.Secondary costae much weaker.Transverse costae dense, regular, a bit stronger than secondary costae, elytral cells transverse, regular.Phallus stout, internal sac without spines but with a sclerotised structure (Fig. 114) or similar structure accompanied by several pairs of short minute rods (Bocak, 1998c, Figs 1, 2).Ovipositor small, only partly sclerotized, with short valvifers (Fig. 134), vagina large, often at least partly sclerotized (Fig. 148;Bocak 1998c, Figs 6, 7).

Distribution. Australian Region and eastern part of Oriental
Region: Peninsular Malaysia, Thailand, Laos, Sundas, and the Philippines.Metriorrhynchus sericeus C. O. Waterhouse, 1879 is reported from India (Kleine, 1933), but Metriorrhynchus was not found in the recently collected material from India.Highest diversity is found in the Papuan Subregion and Australia, but fauna of these regions is badly in need of revision at the species level.Calder (1998) combined all Australian species of Metrior rhynchus with Porrostoma, because of the doubtful status of the name Metriorhynchus Guerin-Meneville, 1838.
Remarks.There are two types of vagina in Metrior rhynchus: M. parallelus (type-species) has a completely sclerotized vagina but species in the M. thoracicus Fabricius, 1801 group have a completely membranous vagina.These groups have a similar shaped paraproctus, very slender, reduced valvifers and a very strong membrane in which the ovipositor is enclosed.
Dilolycus was proposed for a species with considerably reduced carinae in the middle of the pronotum (Fig. 85).It has similar shaped male genitalia to the Papuan group of species (as in Bocak, 1998c, Figs 1, 2).Therefore I consider Dilolycus to be a junior subjective synonym of Metriorrhynchus.
The holotype of Metriorrhynchus mirabilis has dam aged antennae, only the three basal antennomeres of each antenna remain preserved, antennomere 3 is serrate.Two fragments of antennae were glued on the separate label: the first fragment was serrate, corresponding to the remaining antennomeres on the holotype, the second frag ment flabellate.It must be part of a different specimen, because it is made up by antennomeres 3-11.The name Flabelloporrostoma and its generic status are based on this incorrectly identified body part.Therefore, I propose Flabelloporrostoma to be a junior subjective synonym of Metriorrhynchus.
Diagnosis.Body slender, externally similar to Trichalus (general body form, shortened first elytral primary costa, only one longitudinal areola in middle of pronotum, Fig. 9), medium-sized to moderately large.Head without rostrum.Antennae long, strongly com pressed, antennomeres 3-10 almost parallel-sided to ser rate in both sexes.Maxillary palpi with slender apical palpomere.Pronotum with one longitudinal, median areola, lateral margins parallel-sided or pronotum much wider at base.Elytra flat, with shortened primary costa 1, usually with apparent secondary costae, in one species secondary costae distinctly weaker than transverse ones, irregular, interrupted.Legs long, slender, hind trochanters much wider apically.Phallus robust only at base, slender and apical two thirds very gradually narrowed, internal sac completely sclerotized.Phallobase robust, membrane partly sclerotized (Fig. 116).Ovipositor short and wide, valvifers robust, vagina wide, accessory glands inserted laterally, glandular ducts very short, unpaired apical gland as long as vagina.

Remarks.
Leptotrichalus and Lobatang has same shaped female genitalia, especially the shortened, wid ened valvifers.Lobatang is characterised by the unique modification of the internal sac, which has not been found in other Lycidae.
Diagnosis.Body medium-sized to large, slender, slightly widened posteriorly.Head small, without or with very short rostrum.Antennae compressed, antennomeres 3-10 parallel-sided to slightly serrate, never flabellate.Pronotum slender, long, apparently narrowed frontally (Fig. 75), only very slender longitudinal areola in middle of pronotum, sometimes less apparent lateral carinae pre sent on middle of lateral pronotal margin.Elytra slender, with considerably shortened primary costa 1 and with entire secondary costae.Legs very long, slender, strongly compressed.Male genitalia very diverse in shape, often slender, tube-like, internal sac without any basal spines, rarely with strengthened sclerotized parts.Phallobase often with large, well sclerotized membrane.Ovipositor with short valvifers, vagina short, accessory glands attached laterally, unpaired vaginal gland large, spermatheca bulb-like.
Distribution.Australian Region: Lesser Sundas, Moluccas, and Sulawesi; Oriental Region: Sunda Islands, the Philippines, Malaysia, Thailand and Indochina.Altogether almost 60 species are classified in Leptotrichalus with highest diversity in the Philippines.

Remarks. The relationship between Leptotrichalus and
Lobatang is supported by prolonged galees, very wide base of male sternum A8, strong, shortened valvifers and generally short and wide ovipositor and the slender and asymmetrical shape of basal part of phallus.They differ in the prolonged pronotum in Leptotrichalus and the sclerotized internal sac in Lobatang.Additionally, the apical maxillary palpomere is more slender in Leptotrichalus.Diagnosis.Body medium-sized.Head small, without rostrum.Antennae stout, serrate.Maxillary palpi with almost parallel-sided apical palpomere, without papillae, apex obliquely cut.Pronotum wide, almost straight at basal margin, widest before base, mostly only median areola present (see remarks), but at least small vestiges of frontolateral carinae are regularly present (Fig. 76).Pri mary costae strong, almost completely developed, costa 2 and 4 apparently stronger in apical part, other primary costae very slightly shortened, weaker.Phallus simple, slender, internal sac without spines or sclerotisation (Figs 117,118).Female genitalia with very robust and wide valvifers, which are firmly fused with coxites, vagina membranous, large (Figs 156,159).
Remarks.The very similar shape of male and female genitalia supports the affinity of Stadenus and Porros toma.Porrostoma differs in presence of rostrum, Sta denus has completely fused coxites and valvifers.Stadenus was originally defined by three areoles on the pronotal disc, but this character is unreliable.I have studied male and female Stadenus triareolatus in copula (BMNH).They are externally very similar and without any doubt they are conspecific.The female specimen has three areoles, the male seven areoles.But even the female specimen has some vestiges of frontolateral carinae.
Distribution.Australian Region.Porrostoma is the com monest lycid in Australia, where the highest diversity is found.Several species are known from New Guinea.
Remarks.Porrostoma was for long time considered a junior synonym of Metriorrhynchus.The name was used by C. O. Waterhouse (1877) and later sometimes incon sistently by Pic. Kleine (1933) listed Porrostoma as a junior synonym of Metriorrhynchus.Calder (1998) used Porrostoma as ajunior replacement name for Metriorhynchus Guerin-Meneville and proposed 85 new combina tions with Porrostoma (only Australian fauna).Bocak (1998c) replaced Metriorhynchus Guerin-Meneville by Metriorrhynchus Gemminger et Harold, used Porrostoma as a valid taxon and differentiated between Metriorrhyn chus and Porrostoma.The identity of most species classi fied in Porrostoma is unknown and this genus is badly in a need of revision.
Distribution.Australian Region: New Guinea, only four spe cies classified in Metriorrhynchoides.

Remarks. Metriorrhynchoides is closely related to
Porrostoma.Its pronotal carinae are very similar to those of Porrostoma and additionally they both have a long ros trum.Metriorrhynchoides is characterised by a widened apical part of elytra and costae 2 and 3 fused before apex of elytra.Additionally, Metriorrhynchoides species are a bright metallic blue with bright yellow body parts unlike Australian Porrostoma.All males of Metriorrhynchoides also have serrate antennae.I have not found an apomorphic feature defining Porrostoma, therefore it is very probable that Metriorrhynchoides is a subgroup of Por rostoma and will be synonymised with it in the future.

Remarks.
Cladophorus is defined by the form of its phallus and unique shape of phallobasal membrane (Fig. 121) , which has not yet been found in other genera of Metriorrhynchinae.At least several dozens of species with this type of aedeagus are known from New Guinea (Kleine, 1926a).On the other hand, most of almost 200 species previously placed in Cladophorus are not conge neric with the New Guinean species (all species from the Oriental, Palaearctic and Afrotropical Regions).Kleine classified as Cladophorus all species that have slender antennal lamellae (mostly Cautires).Pic often used the name Odontocerus Guerin-Meneville nec Stephens and these species were transferred to Cladophorus by Kleine (1933).Pic's concept is unclear and his descriptions do not allow identification bellow subfamily level.
Distribution.Australia Region: Australia, New Guinea and Moluccas.There are several Afrotropical and Oriental species now classified in Procautires but they are not congeneric with Procautires toxopei and should be classified with Cautires or another closely related genus.These species have less con spicuous secondary costae, but do not share other characters with the type species.
Remarks.Procautires is characterised by a unique modification of elytral secondary costae.With Kassemia and Cautiromimus it shares the same shape of phallus and sclerotisation of internal sac and with Cladophorus pig mented phallobasal membrane, although it is in this case less extensive and does not have the characteristic struc ture as in Fig. 121.Additionally, f have found a relatively large sclerite between coxites.Very small structure in similar position is present also in some Cautiromimus (Fig. 160).The sclerite between bases of coxites is reminicent of sclerite found in some species of Xylobanus (Fig. 154), but unlike in Xylobanus, the sclerite is not connected with valvifers and is located between bases of coxites.Therefore f do not consider these structures to be homologous.
Diagnosis.Body medium sized, relatively slender.Head small, without rostrum.Male antennae flabellate, female antennae serrate.Maxillary palpi with almost par allel sided apical palpomere, without papillae.Middle pronotal areola very narrow, deep, attached to the basal margin by very short keel, reaching almost 3/4 of pro notal length, frontolateral carinae very strong, but weak in frontal punctured area, absent in frontal quarter, lateral carinae are not connected with median areola (Fig. 81).Posterior half of pronotum shining, pubescent.Elytra par allel sided, with four primary and five secondary costae extending the whole length, primary costae stronger at humeri, weak in apical fifth.Transverse costae forming quadrate, rounded cells to slightly longitudinal rounded cells.Phallus stout, two plates at base of internal sac, pri mary gonoporus circular, sclerotized (Figs 122,123).Vagina very slender, its wall partly stiffened (Fig. 160).
Distribution.Australia Region: New Guinea, Moluccas.At present only C. reticulatus is classified here, but several unde scribed species exist in collections.
Distribution.Australian Region: New Guinea.Two species are classified here but several undescribed species exist in col lections.
Remarks.Ditua is reminiscent of Cautiromimus but differs in that male antennae are serrate and in the shape of pronotum.Ditua deplanata is a member of a very common lycid Mullerian mimicry complex and f have found in collections several not very closely related spe cies identified as Ditua dichroma.The name D. deplanata was up to now used as the junior synonym of Ditua dichroma along with two other names.The status of the synonyms of Ditua dichroma was not checked.

Kassemia Bocak, 1998
Kassemia Bocak, 1998b: 195.Type species.Kassemia oculata Bocak, 1998.Diagnosis.Body small to medium sized, parallel-sided, all representatives dark brown to brown with humeral quarter to half of elytra light yellow to brown, always dis tinctly much lighter than the rest of elytra.Head small, without rostrum, mandibles slender, long, three times longer than labrum, apical maxillary palpomere robust, securiform, broad.Antennae strongly compressed, flabel late in males and serrate in females.Pronotum with seven areoles.Elytral secondary costae completely absent, pri mary costa 1 shortened, costa 2 and 3 sometimes fused before apex.Legs relatively strong and short, strongly compressed.Male genitalia with relatively short and robust phallus, apical part widely open, internal sac strengthened at base with spine-like structure.Phallobasal membrane never sclerotized.Ovipositor with slender, long valvifers, vagina as long as valvifers, widest at apex, accessory glands inserted laterally.
Remarks.The combination of seven areoles on the pronotum, complete absence of elytral secondary costae, shape of male genitalia, and shortened primary costa 1 confirm the sure identification of Kassemia.The shape of male antennae and shape of male and female genitalia show little assotiation with shortened primary costa 1 in other genera.The shape of phallus indicates a close rela tionship with genera of the Cladophorus -Ditua -Cautiromimus clade.
Distribution.Australian Region: New Guinea.
Remarks.The flabellate female antennae are the autapomorphy of Pseudodontocerus and this character is not present in other Lycidae.Pseudodontocerus is character ised by the extremely long antennal lamellae in males, weaker lateral and frontolateral pronotal costae, and a slender body shape.I examined type species of Carathix and Pseudodontocerus as well as several closely related undescribed species, and have not found any difference supporting their separate generic status.Therefore, Cara thrix is considered to be a junior subjective synonym of Pseudodontocerus.

Oriomum Bocak, 1999
Oriomum Bocak, 1999a: 111.Type species.Oriomumfemoralis Bocak, 1999a: 111.Diagnosis.Body medium-sized, robust.Head rostrate, rostrum reaching to middle coxae.Antennomeres 1 to 5 slender, antennomere 6 of similar shape but slightly ser rate, antennomeres 7-9 short, triangular.Bottom side of antennomeres 1 and 3-4 with very long, dense hairs.Pro notum with 7 areoles.Elytra with four primary costae extending whole length, costa 1 weaker in posterior half.Secondary costae distinct in humeral area, weaker on rest of elytra, cells irregular.Legs significantly modified for clasping of female, with long dense pubesce.Femora of fore legs and mid legs with spines basally.Hind legs with bigger, triangular trochanters.Position of hind process of trochanter corresponds with the position of femoral spines on fore and mid legs.Phallus slender, parallel sided, with exposed, dense pubescence on apical part of internal sac, basal part of internal sac without sclerotisation.Female unknown.
Remarks.Oriomum differs from the remaining Metriorrhynchinae genera in the unique shape of its elytra with entire but apically distinctly weaker first primary costa, the presence of spines on hind margins of fore and mid femora, hind legs with acutely projected triangular tro chanters and opistognathous head with extremely long rostrum reaching the coxae of the mid legs, and with antennomeres of a very unusual shape.Oriomum seems to be closely related to the Porrostoma clade, but for the final decision on this a female is necessary.

Remarks.
Only the damaged holotype of Cladophor inus cyanescens was available for this study.Therefore the genus was not coded into the matrix and is now classi fied in Metriorrhynchina on the basis of the shape of male genitalia and general appearance.The species is easily recognisable by its metallic shine, seven pronotal areoles and flabellate antennae.

Remarks.
Only the holotype of Mimoxylobanus angus tatus was available.In general appearance and shape of phallus this species resembles Xylobanus.The exact posi tion of this genus within the Metriorrhynchina is because it differs substantially in the structure of internal sac.
Remarks.Xylobanomimus has very similar male geni talia, general appearance and structure of elytral cells to Malacolycus, but differs in the flabellate male antennae and strong fTontolateral carinae on the pronotum.Malaco lycus can be hypothesised as a sister group of Xyloba nomimus.Xylobanomimus is classified with Metriorrhynchina.
Remarks.Only the holotype of Malacolycus paululus is known.The pronotal carinae are considerably reduced and simple male genitalia are very similar to those of Xylobanomimus.Therefore, Malacolycus is placed in the Metriorrhynchina.The body form and structure of elytra resembles that of Xylobanomorphus, but they differ in male genitalia (Figs 129,130).
Distribution.Australian Region: New Guinea.
Remarks.At present, only the holotype of Xylobano morphus transformis is known.This species resembles Xylobanomimus and Malacolycus in general appearance, but differs in the sclerotisation of the internal sac.Consid ering its similarity to Xylobanomimus, the absent fronto lateral pronotal carinae can be hypothesised as second arily reduced.Several common characters and similarity in general appearance support the affinity of Malacolycus, Xylobanomimus and Xylobanomorphus.All these taxa are known from single specimens of males.Further material is needed for study and provisionally Xylobanomorphus is classified incertae sedis in Metriorrhynchina.

Taxon dubium Falsolucidota Pic, 1921
Falsolucidota Pic, 1921b: 9, hors texte.Type species.Falsolucidota testaceicollis Pic, 1921b: 9 (by monotypy).R em ark s.The identity o f the monotypic genus Falsolucidota is unknown.The type specimen was not found in P ic's collection in Paris and the original description does not help in the identification o f this taxon.
Key to M etrio rrh y n c h in a e genera 1. Elytral primary costa 1 entire and strong in whole length, weaker in apical half or very slightly shortened; always reachingatleastbeyondhalfwayalongelytra ............ 10 -Elytral primary costa 1 considerably shortened, never reaching beyond half way along elytra, typically not longer than one third of elytral length (Fig. 9  Pseudosynchonnus Pic, 1922-syn. n. Pseudosynchonnus Pic, 1922: 13, hors  Pseudosynchonnus is here transferred from the M etrior rhynchinae to Erotinae and Lycoprogenthes from Calochrominae to Erotinae.I have compared Pseudosyn chonnus, Lycoprogenthes, Parapyropterus and Protaphes (the last two already sooner classified in Erotinae) and I have not found any substantial difference between them.Therefore, I consider genera Pseudosynchonnus, Parapyropterus, and Protaphes to be junior synonyms o f Lycoprogenthes.
A ppendix 1: L ist of type m aterial studied.

Figs 3
Figs 3-6: Distribution.3 -distribution of subfamily and tribes; 4 -eastern limit of range of Asian continental elements; 5 -distri bution of Hemiconderina and Trichalina; 6 -western limit of range of Australian elements.
. Only genera with the charac teristic slender phallus and dorsally attached lateral vaginal glands belonged in the Trichalina in the restricted sense, and the sister-group position of Diatrichalus and Enylus was confirmed.The present analysis of the whole subfamily Metriorrhynchinae confirmed the monophyly of the restricted Trichalina and also supports the close relationship of genera Enylus and Diatrichalus to the Tri chalina clade.Although the relationship of these genera needs confirming using larval characters and molecular data, the genera Enylus and Diatrichalus are here classi fied in the Trichalina.Enylus C. O. Waterhouse, 1879 Enylus C. O. Waterhouse, 1879: 72.Type species.E. segregatus C. O. Waterhouse, 1879 (by monotypy).Strophicus C. O. Waterhouse, 1879: 73, syn.n.Type species.Strophicus nigellus C. O. Waterhouse, 1879: 73 (by monotypy).

Table 2 .
Data matrix

Table 3 .
Synapomorphies and homoplastic changes listed by nodes for the cladogram in Fig. 1.
Figs 29-51).Further apomorphic char acter states ofMetriorrhynchinae are given in Table 3. B. The present analysis supports the tribes Conderini and Metriorrhynchini.Three subtribes, Hemiconderina and Trichalina with a basal placement, form the Metrior rhynchini.The Hemiconderina concept is redefined.The genera Diatrichalus and Enylus are returned to the Trichalina.The resulting cladograms varied but some clusters within the Metriorrhynchina were relatively robust (Ditua clade, Porrostoma clade and Cautires clade).C.The distribution of clades was discussed.Conderini are restricted to East and South East Asia, Metriorrhynchini are considered to be an East Gondwanan element with only a few species reaching northern East Asia.All these species are members of widespread genera known from the Afrotropical and Oriental regions.All major clades classified as subtribes of Metriorrhynchina are either exclusively Australian or at least have most of their endemic taxa in the Austro-Papuan Region.D. The cladistic analysis showed that characters used for a long time to define genera (presence and/or absence of elytral secondary costae, shortened primary elytral costae, and serrate of flabellate antennae) evolved sev eral times in the Metriorrhynchinae.This study aimed to establish phylogenetic relationship between the genera of Metriorrhynchinae in order to study further individual clades within the subfamily.More extensive sampling and new sources of data, espe cially on immature stages and molecular data, will cor roborate or refuse the proposed phylogeny.Additionally, further studies are need to establish new combinations and to enlarge the data base available for cladistic analy ses.
Distribution.Australian Region: New South Wales, New Guinea, and Mysool Island.Only four species are at present classified in Enylus, two of them were originally classified in Strophicus.
cited 112 Trichalus species from the Oriental and Australian Regions.Recent studies indicate that most Oriental and Papuan species previously classified as Trichalus belong to Microtrichalus Schizotrichalus differs in five pronotal are oles from other genera classified in the in Trichalina. Remarks.
Afrotropical, Oriental, Australian Regions and eastern part of the Palaearctic Region.Coincident with distribu tion ofMetriorrhynchini (Fig.3). Distribution.
Caenioxylobanus have a unique shape and this genus differs from the closely related Bulenides in the absence of secondary costae and in the mat, velvet bottoms to the elytral cells.