Apoderus

Abstract: The phylogenetic relationships in the families Rhynchitidae and Attelabidae have been reconstructed. The main synapomorphies have been revealed. The morphologically advanced groups have been distinguished in the families studied. The family Attelabidae forms two large branches. The most advanced is the supertribe Rhynchititae, the representatives of which could adapt not only to development in various parts of a plant, but also to rolling leaf packages. This supertribe consists of eight well-defined tribes forming three groups: (1) Auletini and Minurini; (2) Cesauletini, Eugnamptini and Isotheini; (3) Pterocolini, Rhynchitini and Byctiscini. The families Rhynchitidae and Attelabidae (leafrolling weevils) belong to the most surprising beetles on our planet. Some of them have developed the ability to make leaf packages in which their larvae feed; others put eggs into fruits or vegetative parts of plants in which the larval development proceeds. The both families mostly inhabit forests and are associated with arboreal vegetation; those species that occur in open landscapes develop on herbs. These weevils are widely distributed over the planet, the most of species occur in the subtropical and tropical zones. These families are poorly investigated despite their wide distribution and a comparatively simple collecting both adults and larvae. The classification used until the present time was elaborated in the first half of the XX century by E. Voss. Unfortunately, when creating it, he used formal characters, therefore the classification is artificial: many closely allied species are placed in different genera, and close genera, in different tribes. Therefore Voss’s (1965) concept of the phylogeny of the Rhynchitidae and Attelabidae was largely erroneous. The problem of revealing the phylogenetic relationships in these groups remained unsolved. In last decades, the cladistic analysis has been widely used in the systematics and phylogeny of insects (Pavlinov, 1989, 1990; Rasnitsyn, 2002). Two authors (Sawada, 1993; Riedel, 2002) have undertaken attempts of the cladistic analysis of the Rhynchitidae and Attelabidae. Sawada (1993) proposed a phylogenetic hypothesis for species of the Rhynchitidae from Japan. He has managed to show that the tribes Eugnamptini and Isotheini, and also Rhynchitini and Byctiscini are sister-groups and form two lineages, both widely separated from the tribe Auletini. Sawada has made a number of mistakes. For example, Temnocerus japonicus (Morimoto) (tribe Rhynchitini) was united with species of the tribe Auletini; Teretriorhynchites amabilis (Roelofs) and Involvulus pilosus (Roelofs) (subtribe Rhynchitina) were grouped together with species of the subtribes Lasiorhynchitina, Temnocerina, and Perrhynchitina. The situation appeared worse with the final phylogenetic tree (Sawada, 1993) constructed on the basis of the method of minimisation of the number of characters (Sawada, 1988). This scheme reflected the traditional Voss’s classification where Isotheini are considered the most advanced tribe, and Eugnamptini are placed close to Rhynchitini. The erroneousness of the Sawada’s hypothesis may result primarily from including a small number of taxa in the analysis. Riedel (2002) has carried out a cladistic analysis of species of the tribe Euopsini of the New Guinea fauna (PAUP program). He has investigated representatives of various species-groups now promoted to genera (Legalov, 2003a). It should be noted that other Euopsini from the Oriental, Afrotropical, and Australian biogeographical regions have not been included in the analysis. The genera Epirhynchites (family Rhynchitidae), Attelabus, Lamprolabus, Euscelophilus (subfamRECONSTRUCTION OF THE PHYLOGENY OF THE RHYNCHITIDS ... 1 ENTOMOLOGICAL REVIEW Vol. 84 No. 7 2004 765 ily Attelabinae) and Apoderus (subfamily Apoderinae) have been taken as outgroups. Riedel has studied the most advanced representatives of the family Attelabidae, which was inevitably reflected in the result. In the cladogram, only close species were united. Relationships between groups of species have remained unrevealed. Surprising is Riedel’s conclusion: admitting that he had failed to reconstruct the phylogeny of the New Guinea fauna of Euopsini; he writes, however, that Apoderinae do not deserve the subfamily rank and should be considered a tribe. This judgment, based on the study of a single species, is obviously incorrect. My attempt to revise separate groups of species (Legalov, 2001) has faced the problem of a poor supraspecific classification, which made impossible revealing the relationships between the taxa. Therefore a decision was taken to start not stem-up but top-down. The supraspecific classification of the Rhynchitidae and Attelabidae of the world fauna (Legalov, 2003а) has been reconsidered, and their phylogenetic relationships were reconstructed. Some preliminary results of this research on the phylogeny of these groups have been published (Legalov, 2003). I investigated the material from state museums and private collections. The type or authenticated material of most of the supraspecific taxa of Rhynchitidae and Attelabidae was examined, totaling more than 20000 specimens of more than 700 species (36%), representing 82% of supraspecific taxa of Rhynchitidae, and 86%, of Attelabidae.. The choice of a program for the construction of cladograms was important. I compared the results obtained with use of the programs Hennig 86, Phylip, PAUP 2.4.1., and SYNAP 420. The first two do not provide an explanation of calculations and therefore have been rejected. PAUP and SYNAP provided similar results especially if characters were weighed. Yet SYNAP 420 not only marks branches by phylogenetic events and allows checking up calculations, but also provides two additional parameters (index of advance and index of phylogenetic relationship). For this reason, the reconstruction of the phylogeny was carried out with the SYNAP 420 program (Baikov, 1999). The following abbreviations are adopted in the paper. 1. Index of advance (IP) is a sum of advanced characters 2. Index of phylogenetic relationship (IPHR) is IP minus unique advanced characters. For identical IPHR, a preference was given to the polytomy (a coalescence of identical nodes), an appearance of a unique new character, and the minimum of reversions. When weighing characters, those most important were assigned the values of 2 or 3 points (see table). The matrix was not optimized, the value of a reversion was identified as -1; 0 is a plesiomorphic, and 1, an apomorphic state of the character. The results of the phylogeny reconstruction have been presented in a preceding publication (Legalov, 2003а). The Characteristic of Rhynchitidae and Principles of Cladograms Construction Rhynchitidae are ancient beetles known from the late Jurassic (Oxford epoch) (Legalov, 2003a). It might be assumed that originally they developed in reproductive organs of gymnosperms (similarly to Nemonychidae), but subsequently adapted to development on angiosperms. Rhynchitidae have formed five ecological groups (three groups including species which do not roll leaves and two groups which are classified as “leaf-rollers”) (Legalov, 2003a) which have obtained a considerable species-richness (1087 known species). The family Rhynchitidae is represented in the recent fauna by 1087 species from 250 supraspecific taxa (2 supertribes, 13 tribes, 18 subtribes, 141 genera and 76 subgenera). 33 species are known as fossil records (Legalov, 2003a). For revealing the phylogenetic relationships between tribes in the family Rhynchitidae a cladogram was constructed (figure), for which 47 morphological characters were used (table). 1. Body with setae (0), without setae (1). 2. Metallic sheen present (0), absent (1). 3. Rostrum long (0), short (1). 4. Tooth on exterior side of mandible absent (0), present (3). 5. Labial palps of female 3-segmented (0), 1or 2-segmented (1). 6. Proand mesocoxa approximate (0), widely separated (1). 7. Frons wide (0), narrow (1). 8. Eyes convex (0), almost flat (1). 9. Antennae attached subbasally (0), submedially or subapically (1). 10. Antennal club symmetrical (0), asymmetrical (1). 11. Club short (0), usually long (1). 12. Keel on sides of pronotum absent (0), present (1). 13. Elytra almost rectangular (0), rounded (1). 14. Sculpture of elytra smooth (0), coarse (1). 15. Prescutellar striole present in the majority of genera (0), absent or present only in primitive genera (1). 16. Striae on elytra present (0), absent (1). 17. Wings developed (0), reduced (1). 18. Spines directed forwards on prothorax of males absent (0), present (1). 19. 1st–5th ventrites free