EVOLUTIONARY SIGNIFICANCE OF THE LOSS OF THE CHLOROPLAST-DNA INVERTED REPEAT IN THE LEGUMINOSAE SUBFAMILY PAPILIONOIDEAE

The distribution of a rare chloroplast-DNA structural mutation, the loss of a large inverted repeat, has been determined for 95 species representing 77 genera and 25 of the 31 tribes in the legume subfamily Papilionoideae. This mutation, which is regarded as a derived feature of singular origin within the subfamily, marks a group comprising six temperate tribes, the Galegeae, Hedysareae, Carmichaelieae, Vicieae, Cicereae, and Trifolieae, an assemblage traditionally considered to be monophyletic. This mutation also occurs in the chloroplast genome of Wisteria, a member of the tropical tribe Millettieae whose other members so far surveyed lack the mutation. These new DNA data, together with traditional evidence, support the hypothesis that Wisteria is an unspecialized member of a lineage that gave rise to the temperate tribes marked by the chloroplast-DNA mutation; the probable paraphylesis of Millettieae is revealed. Two other tribes, Loteae and Coronilleae (traditionally regarded as a derived element of the aforesaid temperate tribes) do not possess this chloroplast-DNA structural mutation and, therefore, presumably represent a distinct temperate lineage. This hypothesis is supported by additional evidence from pollen, inflorescence, and root-nodule morphology that suggests that the Loteae and Coronilleae share a more recent ancestry with tropical tribes such as Phaseoleae and Millettieae than with other temperate tribes.     

Genetic characterization of wild leguminous nodular bacteria living in the South Urals

Genetic diversity and phylogeny of rhizobia that nodulate 18 species of wild-growing bean plants of South Urals from 8 genera belonging to 4 tribes (Loteae, Genisteae, Galegeaev and Hedysareae) was studied. It was demonstrated that for the wild-growing plants of Galegeae and Hedysareae tribes symbiotic interaction with various strains of nodule bacteria that closely related to bacteria of Mesorhizobium sp. was typical of the plants of Genisteae tribe--to bacteria of Bradyrhizobium sp. In the nodules of Lortus ucrainicus from Loteae tribe we have found a rhizobium that is closely related to the bacteria of Mesorhizobium sp., and at Coronilla varia rhizobia strains obtained by us were close by sequence of a 16S pRNA gene to Rhizobium sp. In the nodules of some kinds of the investigated plants we found also minor species of a rhizobia, which structure is under the great influence of conditions of the host plant growth.     

THE SISTER GROUP OF THE PALEOTROPICAL TRIBE ARGOSTEMMATEAE: A REDEFINED NEOTROPICAL TRIBE HAMELIEAE (RUBIACEAE, RUBIOIDEAE)

Abstract— The sister group of the monophyletic tribe Argostemmateae, including Argostemma and Neurocalyx , is not the tribe Hedyotideae, or a part of it, as earlier assumed, but the monophyletic tribe Hamelieae, here redefined. It comprises the genera Hamelia, Hoffmannia (here including Xerococcus), Deppea (here including Schenckia), Pinarophyllon , and Omiltemia . The two tribes Argostemmateae and Hamelieae are a monophyletic group characterized by seven synapomorphies. A cladogram is presented showing the relationships of the seven genera based on 31 characters. One group of characters, the aestivation of the corolla, is shown to be much more complicated than hitherto assumed. In the Hamelieae the aestivation can be imbricate (to the right or to the left and quincuncial or not), contorted (to the right), alternative, or even valvate. The Argostemmateae occur in the Old World Tropics, and their distribution supports the hypothesis that at least parts of southeast Asia are a part of Gondwanaland. The vicariant pattern within the Argostemmateae indicates that the large area from west Africa through northern India to southeast Asia is vicariant to Ceylon and southern India. The distribution of the Old World tribe Argostemmateae is vicariant to the essentially Central American tribe Hamelieae.     

Phylogeny of Malpighiaceae: evidence from chloroplast ndhF and trnl-F nucleotide sequences

The Malpighiaceae are a family of ～1250 species of predominantly New World tropical flowering plants. Infrafamilial classification has long been based on fruit characters. Phylogenetic analyses of chloroplast DNA nucleotide sequences were analyzed to help resolve the phylogeny of Malpighiaceae. A total of 79 species, representing 58 of the 65 currently recognized genera, were studied. The 3' region of the gene ndhF was sequenced for 77 species and the noncoding intergenic spacer region trnL-F was sequenced for 65 species; both sequences were obtained for the outgroup, Humiria (Humiriaceae). Phylogenetic relationships inferred from these data sets are largely congruent with one another and with results from combined analyses. The family is divided into two major clades, recognized here as the subfamilies Byrsonimoideae (New World only) and Malpighioideae (New World and Old World). Niedenzu's tribes are all polyphyletic, suggesting extensive convergence on similar fruit types; only de Jussieu's tribe Gaudichaudieae and Anderson's tribes Acmanthereae and Galphimieae are monophyletic. Fleshy fruits evolved three times in the family and bristly fruits at least three times. Among the wing-fruited vines, which constitute more than half the diversity in the family, genera with dorsal-winged samaras are fairly well resolved, while the resolution of taxa with lateral-winged samaras is poor. The trees suggest a shift from radially symmetrical pollen arrangement to globally symmetrical pollen at the base of one of the clades within the Malpighioideae. The Old World taxa fall into at least six and as many as nine clades.     

Review of the systematics and phylogenetic relationships of the Sabethini (Diptera: Culicidae)

This paper reviews the historical treatment of the tribe Sabethini and genus-group taxa and examines the unusual life histories associated with the group. Although recognized by taxonomists as distinct, the taxonomic position of sabethines has been questioned and their rank within the family Culicidae unstable. In order to evaluate the current status of the classification of the tribe a cladistic analysis is performed. Thirty-seven taxa are selected from within the Sabethini and two outgroups were chosen from the tribe Aedini. Exemplars are selected from genus-group taxa world-wide and new and traditional character systems examined in larval, pupal and adult life stages. The results firmly establish the sabethines as a monophyletic group. However, the genera Runchomyia , Tripteroides and Wyeomyia are not demonstrably monophyletic. In addition, the data support the New World taxa as a monophyletic group to a paraphyletic assemblage of Old World taxa. The pattern displayed by the cladogram suggests the ability to vector arboviruses has arisen more than once in mosquitoes.     

Chloroplast DNA restriction site variation in theVernonieae (Asteraceae), an initial appraisal of the relationship of New and Old World taxa and the monophyly ofVernonia

Chloroplast DNA restriction site variation was examined for 35 taxa in theVernonieae and four outgroup tribes, using 17 restriction enzymes mapped for ca. 900 restriction sites per species; 139 mutations were found to be phylogenetically informative. Phylogenetic trees were constructed using Wagner and weighted parsimony, and evaluated by bootstrap and decay analyses. Relationships of Old and New World taxa indicate complex geographical relationships; there was no clear geographic separation by hemisphere. The relationships between Old and New World Vernonias found here support prior morphological analyses. The sister group to all New and most Old World taxa was composed of a small group of Old World species including yellow-flowered, trinervate-leaved species previously postulated to be basal in the tribe. The majority of both New and Old World taxa are derived from a lineage beginning with the monotypic genusStokesia, an endemic of the southeastern United States. The genusVernonia was also found to be paraphyletic within both the New and Old World. Available data do not support either the separation ofVernonia or the tribeVernonieae into geographically distinct lineages. The pattern of relationships within theVernonieae for taxa from North America, Asia, Africa, Central and South America is most similar to that of several other groups of both plants and animals with a boreotropical origin, rather than an origin in Gondwanaland. Such a pattern of distribution suggests more ancient vicariant events than are routinely postulated for theAsteraceae.     

Phylogeny and biogeography of Neotropical spittlebugs (Hemiptera: Cercopidae: Ischnorhininae): revised tribal classification based on morphological data

The spittlebug family Cercopidae is currently divided into two subfamilies: the paraphyletic Old World Cercopinae and the monophyletic New World Ischnorhininae. The most recent classification scheme proposed by Fennah in 1968 divided the New World Cercopidae into four tribes: Tomaspidini, Ischnorhinini, Hyboscartini and Neaenini. Herein we present a phylogenetic analysis of Ischnorhininae using 108 morphological characters and including 53 of the 59 recognized genera, to evaluate the tribal‐level classification and understand the processes underlying the current distributional patterns of these genera. We found significant support for the monophyly of many Neotropical genera, but Fennah's tribal classification is revised because tribes Neaenini, Ischnorhinini and Tomaspidini were recovered as polyphyletic. Hyboscartini was synonymized with Tomaspidini. A taxonomic key to tribes and genera of Neotropical spittlebugs is provided based mostly on recovered apomorphies. The biogeographical analysis suggests a Neotropical origin of ischnorhinines, more specifically in northwestern South America. This was possibly coincident spatially and temporally with the origin of grasses, with ancestral range expansions southward to the Amazonian and Paraná regions, and posterior vicariant events, possibly related to the expansion of forests in the Chacoan region, the South America diagonal of open formations. Dispersals to the Chacoan region and to the Nearctic region are hypothesized to have occurred only within genera. In the Chacoan region it is associated with more recent events, such as the diversification of C4 grasses and establishment of the savannas and seasonally dry forests.     

A Cladistic Analysis of the New World Species of Lotus L. (Fabaceae, Loteae)

The genus Lotus L. is a monophyletic group diagnosed by the possession of a standard claw with thickened infolded margin, stamens diadelphous, and the style hardened from the base. It comprises approximately 200 species distributed throughout the world. A cladistic analysis of the New World species was performed using 39 morphological and anatomical characters (29 from seed morphology and anatomy, 1 from plant habit, 1 from leaf morphology, 6 from flower morphology, and 2 from fruit morphology). Dorycnium, Edentolotus, Krokeria , and Pedrosia , of the Old World, and 28 species of the New World were considered terminal taxa. Tetragonolobus Scop. was chosen to root the cladograms and Dorycnium Mill. to reroot them. With Tetragonolobus the analysis yielded 15 equally parsimonious trees, each with a length of 74 steps, a consistency index of 0.62, and a retention index of 0.89. The 15 initial trees and the strict consensus tree defined 12 monophyletic groups. All terminal taxa form a monophyletic group diagnosed by the presence of a radicular lobe discernible to conspicuous (character 10); rim aril thick (character 13); stipules absent (character 31); and style simple and nondilated (character 36). The New World species form a monophyletic group on the basis of the seed relationship of length to width in hilar view 1.5:1 to 2:1 (character 5); micropyle linear-deltoid to bifurcate (character 19); and keel erostrate (character 33). Identical monophyletic groups were obtained when Dorycnium was used as root. These results are discussed in the context of data on cytology and morphology.     

A phylogeny of the "evil tribe" (Vernonieae: Compositae) reveals Old/New World long distance dispersal: support from separate and combined congruent datasets (trnL-F, ndhF, ITS)

The Vernonieae is one of the major tribes of the largest family of flowering plants, the sunflower family (Compositae or Asteraceae), with ca. 25,000 species. While the family's basal members (the Barnadesioideae) are found in South America, the tribe Vernonieae originated in the area of southern Africa/Madagascar. Its sister tribe, the Liabeae, is New World, however. This is the only such New/Old World sister tribe pairing anywhere in the family. The Vernonieae is now found on islands and continents worldwide and includes more than 1500 taxa. The Vernonieae has been called the "evil tribe" because overlapping character states make taxonomic delimitations difficult at all levels from the species to the subtribe for the majority of taxa. Juxtaposed with these difficult-to-separate entities are monotypic genera with highly distinctive morphologies and no obvious affinities to any other members of the tribe. The taxonomic frustration generated by these contrary circumstances has resulted in a lack of any phylogeny for the tribe until now. A combined approach using DNA sequence data from two chloroplast regions, the ndhF gene and the noncoding spacer trnL-F, and from the nuclear rDNA ITS region for 90 taxa from throughout the world was used to reconstruct the evolutionary history of the tribe. The data were analyzed separately and in combination using maximum parsimony (MP), minimum evolution neighbor-joining (NJ), and Bayesian analysis, the latter producing the best resolved and most strongly supported tree. In general, the phylogeny shows Old World taxa to be basal and New World taxa to be derived, but this is not always the case. Old and New World species are found together in two separate and only distantly related clades. This is best explained by long-distance dispersal with a minimum of two trans-oceanic exchanges. Meso/Central America has had an important role in ancient dispersals between the Old and New World and more recent movements from South to North America in the New World.     

Phylogeny, classification, and biogeography of the cycloteline Therevinae (Insecta: Diptera: Thereavidae)

Twenty-one members of the Laurasian group of Therevinae (Diptera: Therevidae) are compared using 65 adult morphological characters. Cladistic analysis using parsimony on the 17 ingroup and 4 outgroup taxa provides a well-supported hypothesis of relationships among taxa within the Gyclotelini, tribe nov. The Cyclotelini is a monophyletic assemblage of mostly New World genera, including Anolinga , gen. nov. , Breviperna Irwin, Coleiana , gen. nov. , Crebraseta , gen. nov. , Cyclotelus Walker, Mesonana , gen.nov. , and Ozodiceromyia Bigot. In addition, three Old World genera, Ammothereva Lyneborg, Bugulaverpa , gen. nov. , and Procyclotelus Nagatomi & Lyneborg, are included in the tribe. These ten genera are divided into two monophyletic genus-groups, the Brevipema-group and the Cyclotelus-group. Keys are provided for the genera of Cyclotelini. The tribe, the two informal genus-groups, and all genera are diagnosed; five new genera and six new species are proposed. The biogeographical histories of the genera are discussed in terms of their cladistic relationships using methods of cladistic biogeography. Two major vicariant events account for the current distribution of the tribe. The first relates to the Beringian land bridge connecting western North America and eastern Asia. Second, New World cyclotelines were profoundly affected by the Early Eocene breakup of the archipelagic bridge between North and South America, and the distributions support the hypotheses favouring the continental origin of the Greater Antilles.     

Salvia (Lamiaceae) is not monophyletic: implications for the systematics, radiation, and ecological specializations of Salvia and tribe Mentheae

Salvia, with over 900 species from both the Old and New World, is the largest genus in the Lamiaceae. Unlike most members of the subfamily Nepetoideae to which it belongs, only two stamens are expressed in Salvia. Although the structure of these stamens is remarkably variable across the genus, generally each stamen has an elongate connective and divergent anther thecae, which form a lever mechanism important in pollination. In a preliminary investigation of infrageneric relationships within Salvia, the monophyly of the genus and its relationship to other members of the tribe Mentheae were investigated using the chloroplast DNA regions rbcL and trnL-F. Significant conclusions drawn from the data include: Salvia is not monophyletic, Rosmarinus and Perovskia together are sister to an Old World clade of Salvia, the section Audibertia is sister to subgenus Calosphace or the monotypic Asian genus Dorystaechas, and the New World members of section Heterosphace are sister to section Salviastrum. Owing to the non-monophyly of Salvia, relationships at the next clearly monophyletic level, tribe Mentheae, were investigated.     

THE GRANULAR INTERSTITIUM IN THE POLLEN OF SUBFAMILY PAPILIONOIDEAE (LEGUMINOSAE)

A wide range of transitional forms of granular interstitia from simple to complex and from random to ordered occur in the pollen of the subfamily Papilionoideae. Three main types are described: 1) large, widely spaced irregular granules (Type A); 2) densely packed groups of columellae and granules (Type B); and 3) a mass of more or less disorganized granules (Type C). In the genus Calopogonium (tribe Phaseoleae) all three types have been found in different species. Two of the types have been found in different species of the genus Psoralea (tribe Psoraleeae). Granular structures so far occur in six tribes: Desmodieae, Indigofereae, Loteae, Phaseoleae, Psoraleeae, and Vicieae. All of the tribes are regarded as being evolutionarily advanced in both macro and micro characters and many, but not all, show specialized pollen characters. It is concluded that the granular interstitium is a derived structure in papilionoid legumes.     

Molecular phylogeny of Anthyllis spp

For the genus Anthyllis (Fam. Fabaceae, tribe Loteae), with few exceptions, little information is available on the genetic variation among and within species. This genus contains 20 species distributed throughout Europe, Africa, and the Mediterranean basin. The most widespread species is A. vulneraria, and over 30 intraspecies taxa have been identified based on plant morphology. To study the molecular phylogeny of the genus, the sequences of the internal transcribed spacers ITS1 and ITS2 of the nuclear ribosomal DNA of 10 Anthyllis species, including 11 subspecies of A. vulneraria and three subspecies of A. montana, were obtained and analysed together with sequences of five other species of the genus obtained from GenBank. Our results suggest that the genus Anthyllis is not monophyletic and is divided in two main clades: the Anthyllis sensu strictu and the "tetraphylla clade". The former includes most of the Anthyllis species, and the latter includes three annual species more closely related to Lotus. All the taxa were also analysed according to seven chloroplast microsatellites, and these data closely confirm the results obtained with the ITS phylogeny.     

Molecular phylogeny, diversification and character evolution in Lupinus (Fabaceae) with special attention to Mediterranean and African lupines

Phylogenetic relationships within the complex genus Lupinus are estimated from internal transcribed spacer (ITS) sequences of the nuclear ribosomal DNA repeat. The molecular data supports Lupinus as a distinct monophyletic group within the tribe Genisteae. Different geographical lineages are revealed within Lupinus, which are each restricted to either the Old or the New World. In the New World, the ITS data support an eastern-western geographic disjunction of the lupines and the recognition of some well-supported clades. In the Old World, almost all the previously recognized taxa are taxonomically well differentiated. The homogeneous African rough-seeded lupines, Scabrispermae, are strongly supported as a monophyletic group, which is distinct from the diverse and heterogeneous circum-Mediterranean smooth-seeded ones. The latter appear to have evolved as two lineages, in which are revealed some intersectional relationships. Also ITS data allow the assessment of the phylogenetic position of the newly discovered species, L. anatolicus (in the Old World) and L. jaimehintoniana (the Mexican tree lupin). The ITS phylogeny suggests a rapid initial radiation of the lupines subsequent to their divergence from a common ancestor. Moreover, the results indicate that the annual and perennial habits have evolved many times in Lupinus and suggest a role for generation time in affecting the evolutionary history of lupines. Data on adaptive processes and character evolution are re-examined and discussed in the light of the ITS phylogeny.These studies were supported by the research unit UMR-CNRS 6553 Ecobio – University of Rennes (France), and by the NSERC of Canada (grant to R.J. Bayer, at the University of Alberta). They are greatfully acknowledged. All the persons that have contributed in different ways to this work on Lupinus, summarized in this presentation, are greatly thanked. Particularly, we would like to mention Malika Aïnouche, Roland Greinwald, André Huon, W.K. Swiecicki, Billie L. Turner and Ludger Witte for their contributions.     

虎尾草亚科系统发育关系的初步研究

运用广义形态学性状对虎尾草亚科（Chloridoideae）进行系统发育分析。内类群包括虎尾草亚科52属的69种植物,代表虎尾草亚科的主要类群;芦竹亚科（Arundinoideae）扁芒草族（Danthonieae）的Centropodia和Danthonia被选作外类群。分支分析表明,虎尾草亚科是一个单系类群。其严格一致树包括A、B、C、D、E5个分支。两个大族画眉草族（Eragrostideae）和虎尾草族（Chlorideae）代表虎尾草亚科内部类群分化的两个方向,分开处理较合理。细穗草族（Leptureae）放到虎尾草族中较合理。冠芒草族（Pappophoreae）是虎尾草亚科的基部类群,与画眉草族近缘。我们的研究支持虎尾草亚科从旧世界向新世界扩散的地理分布假说,并提供了虎尾草亚科属上类群的系统发育关系的框架。     

Pollination systems in the Asclepiadaceae: a survey and preliminary analysis

Using published and unpublished records of pollination in the dicot family Asclepiadaceae (the 'milkweeds') we offer a preliminary analysis of present-day pollination systems in the family. Variation in principal pollinators is apparent at and below the tribal level. The tribes Marsdenieae and Stapelieae and Asclepiadeae subtribe Gonolobineae are primarily Diptera-pollinated, a tentative conclusion also for the tribe Periploceae, and we emphasize the ubiquity and importance of fly pollination in the family. The rest of the tribe Asclepiadeae is pollinated in the main by Hymenoptera and Lepidoptera. Lack of data makes it impossible to draw even initial conclusions for the remaining tribes, Secamoneae and Fockeeae. Within the Asclepiadeae, there has been a trend towards more diverse pollination systems (incorporating butterflies and wasps) in the New World compared to the Old World. In terms of the taxonomic breadth of pollinators of individual species, Stapelieae is the most specialized tribe. We emphasize that this is only a preliminary account of pollination in the Asclepiadaceae, and detail areas where further work is urgently required.     

基于28S rDNA D2基因片段与形态特征的矛茧蜂亚科系统发育研究（膜翅目：茧蜂科）

本研究选取矛茧蜂亚科Doryctinae（昆虫纲Insecta：膜翅目Hymenoptera：茧蜂科Braconidae）的6族15属18种做内群,茧蜂科其它7亚科11属11种做外群,首次结合同源核糖体28S rDNA D2基因序列片段和100个形态学和解剖学特征对该亚科进行了系统发育学研究。利用“非圆口类"的小腹茧蜂亚科Microgastrinae为根,以PAUP*4.0和MrBayes 3.0B4软件分别应用最大简约法（MP）和贝叶斯法对矛茧蜂亚科的分子数据和分子数据与非分子数据的结合体进行了运算分析;并以PAUP*4.0对矛茧蜂亚科的28S rDNA D2基因序列片段的碱基组成与碱基替代情况进行了分析。结果表明：矛茧蜂亚科的28S rDNA D2基因序列片段的GC含量在39.33%～48.28%之间变动,而对于碱基替代情况来讲,矛茧蜂亚科各成员间序列变异位点上颠换（transversion）大于转换（transition）。不同的分析算法所产生的系统发育树都表明矛茧蜂亚科是一个界限分明的单系群;在矛茧蜂亚科内,除了吉丁茧蜂族Siragrini为单系群外,其他族（矛茧蜂族Doryctini和方头茧蜂族Hecabolini）都是并系群。对于矛茧蜂亚科内各属之间的相互亲缘关系,不同算法所得的系统发育树的拓扑结构不完全一致,表明矛茧蜂亚科内（属及族）的系统发育关系还有待于进一步研究。     

A preliminary phylogenetic analysis of the New World Helopini (Coleoptera,Tenebrionidae, Tenebrioninae) indicates the need for profound rearrangements of the classification

Helopini is a diverse tribe in the subfamily Tenebrioninae with a worldwide distribution. The New World helopine species have not been reviewed recently and several doubts emerge regarding their generic assignment as well as the naturalness of the tribe and subordinate taxa. To assess these questions, a preliminary cladistic analysis was conducted with emphasis on sampling the genera distributed in the New World, but including representatives from other regions. The parsimony analysis includes 30 ingroup species from America, Europe and Asia of the subtribes Helopina and Cylindrinotina, plus three outgroups, and 67 morphological characters. Construction of the matrix resulted in the discovery of morphological character states not previously reported for the tribe, particularly from the genitalia of New World species. A consensus of the 12 most parsimonious trees supports the monophyly of the tribe based on a unique combination of characters, including one synapomorphy. None of the subtribes or the genera of the New World represented by more than one species (Helops Fabricius, Nautes Pascoe and Tarpela Bates) were recovered as monophyletic. Helopina was recovered as paraphyletic in relation to Cylindrinotina. One Nearctic species of Helops and one Palearctic species of Tarpela (subtribe Helopina) were more closely related to species of Cylindrinotina. A relatively derived clade, mainly composed by Neotropical species, was found; it includes seven species of Tarpela, seven species of Nautes, and three species of Helops, two Nearctic and one Neotropical. Our results reveal the need to deeply re-evaluate the current classification of the tribe and subordinated taxa, but a broader taxon sampling and further character exploration is needed in order to fully recognize monophyletic groups at different taxonomic levels (from subtribes to genera).     

Androconial hairbrushes of the Syntomis (Amata) phegea (L.) group (Lepidoptera, Ctenuchinae): a synapomorphic character supported by sequence data of the mitochondrial 16S rRNA gene

Males of several palaearctic Syntomis/Amata species (Lepidoptera: Arctiidae) possess androconial hairbrushes in connection with the foreleg coxa. The cuticular structure of these potentially behaviour-related and pheromone dissipating brushes is described. Such male-specific organs and signals play a crucial role in the female choice procedure. The presence of hairbrushes was found in 17 out of 28 inspected species of the tribe Syntomini. All members of the Syntomis phegea group (Europe to Central Asia, as well as Caspian, Caucasian and near-middle East species) have these structures, and only three oriental and south Asian, but none of three African species, carry this trait. The common genetic base of this morphological character is supported by an analysis of mitochondrial 16S rRNA from 19 representative taxa; species with hairbrushes form a monophyletic clade and the brushes are a synapomorphic character. This genetic finding corroborates the ethological significance of these organs. Phylogenetic data show a substantial genetic divergence between the tribe Ctenuchini (New World species) and the Old World Syntomini. Furthermore, DNA sequence data suggest a split of the genus Amata (sensu Obraztsov, 1966) in two distinct genera, Amata (without hairbrushes) and Syntomis (with hairbrushes).     

Phylogeny and historical biogeography of leafhopper subfamily Iassinae (Hemiptera: Cicadellidae) with a revised tribal classification based on morphological and molecular data

Phylogenetic relationships among major lineages of the leafhopper subfamily Iassinae were explored by analysing a dataset of 91 discrete morphological characters and DNA sequence data from nuclear 28S rDNA and histone H3 genes and mitochondrial 12S rDNA. Bayesian, maximum‐likelihood and maximum parsimony analyses yielded similar tree topologies that were well resolved with strong branch support except at the base of the tree, resulting in equivocal support for inclusion of Bythoniini as a tribe of Iassinae but strong support for the monophyly of Iassinae (excluding Bythoniini) and most previously recognized iassine tribes. Divergence times for recovered nodes were estimated using a Bayesian relaxed clock method with two fossil calibration points. The results suggest that the deepest divergences coincided with Gondwanan vicariant events but that more recent divergences resulted from long‐range dispersal and colonization. Biogeographical analyses suggest that the group most likely has a Neotropical origin. The following changes to the taxonomic classification are proposed: establishment of three new tribes, Batracomorphini trib.n. (based on type genus Batracomorphus Lewis), Hoplojassini trib.n. (based on type genus Hoplojassus Dietrich and including one other South American genus), Lipokrisnini trib.n. (based on type genus Lipokrisna Freytag and including two other endemic Caribbean genera); Krisnini is redefined to include only the Old World genera Krisna and Gessius; Iassini is redefined to include only the type genus and four endemic Afrotropical genera; Bascarrhinus Fowler and Platyhynna Berg, recently treated as genera incertae sedis, are placed in Hyalojassini; Thalattoscopus Kirkaldy is added to the previously monobasic tribe Trocnadini. Iassinae now includes 12 tribes, all of which appear to be monophyletic. Revised morphological diagnoses of the subfamily and each of the included tribes are provided and a key to tribes is also given. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:41295B68‐2DAB‐4C4F‐B260‐F7C054922173 .