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.     

A global phylogeny of apple snails: Gondwanan origin, generic relationships, and the influence of outgroup choice (Caenogastropoda: Ampullariidae)

Apple snails (Ampullariidae) are a diverse family of pantropical freshwater snails and an important evolutionary link to the common ancestor of the largest group of living gastropods, the Caenogastropoda. A clear understanding of relationships within the Ampullariidae, and identification of their sister taxon, is therefore important for interpreting gastropod evolution in general. Unfortunately, the overall pattern has been clouded by confused systematics within the family and equivocal results regarding the family's sister group relationships. To clarify the relationships among ampullariid genera and to evaluate the influence of including or excluding possible sister taxa, we used data from five genes, three nuclear and two mitochondrial, from representatives of all nine extant ampullariid genera, and species of Viviparidae, Cyclophoridae, and Campanilidae, to reconstruct the phylogeny of apple snails, and determine their affinities to these possible sister groups. The results obtained indicate that the Old and New World ampullariids are reciprocally monophyletic with probable Gondwanan origins. All four Old World genera, Afropomus, Saulea, Pila, and Lanistes, were recovered as monophyletic, but only Asolene, Felipponea, and Pomella were monophyletic among the five New World genera, with Marisa paraphyletic and Pomacea polyphyletic. Estimates of divergence times among New World taxa suggest that diversification began shortly after the separation of Africa and South America and has probably been influenced by hydrogeological events over the last 90 Myr. The sister group of the Ampullariidae remains unresolved, but analyses omitting certain outgroup taxa suggest the need for dense taxonomic sampling to increase phylogenetic accuracy within the ingroup. The results obtained also indicate that defining the sister group of the Ampullariidae and clarifying relationships among basal caenogastropods will require increased taxon sampling within these four families, and synthesis of both morphological and molecular data. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 98 , 61–76.     

Exploring myrmecophily based on the phylogenetic interrelationships of Falagonia Sharp, 1883 (Coleoptera: Staphylinidae: Aleocharinae) and allied genera

The taxonomy of Lomechusini Fleming has a complex history. Recent studies have shown that this group is polyphyletic; however, little is known about the evolutionary interrelationships among its constituent genera. The goals of the present study are to infer the phylogenetic relationships of Falagonia Sharp and closely related genera; to define the boundaries of those genera based on synapomorphic characters; and to explore the evolution of myrmecophily within the lineage. The phylogenetic analyses are based exclusively on morphological characters of adults. A total of 36 operational taxonomic units were used for the analysis. The best trees were selected based on maximum parsimony and Bayesian inference. During the parsimony reconstruction, different weighting strategies were used to recover the most robust phylogenetic hypothesis. Although minor differences were observed in the results of the different analyses, the topologies were consistent throughout. Several groups of genera proposed by Seevers (1965), such as the ‘Tetradonia’ and ‘Ecitopora’ groups, were not recovered. Thus, these may represent nonmonophyletic groups that were based on nonsynapomorphic diagnostic characters. Our analyses consistently recovered the genera Asheidium Santiago‐Jiménez, Delgadoidium Santiago‐Jiménez, Falagonia, Newtonidium Santiago‐Jiménez, Pseudofalagonia Santiago‐Jiménez, Sharpidium Santiago‐Jiménez, Tetradonia Wasmann and Thayeridium Santiago‐Jiménez, forming a monophyletic group that we have called the ‘Asheidium complex’. Falagonia mexicana Sharp shows seven autapomorphies, none of which were used to establish the genus. Based on the phylogenetic results, myrmecophily has evolved independently at least three times within the lineage. This study, based on morphological characters, is one of the first approaches towards gaining an understanding of the phylogenetic relationships within the polyphyletic tribe Lomechusini.     

Morphology-based phylogeny of the treehopper family Membracidae (Hemiptera: Cicadomorpha: Membracoidea)

A parsimony‐based phylogenetic analysis of eighty‐three morphological characters of adults and immatures of seventy representatives of the tribes and subfamilies of Membracidae and two outgroup taxa was conducted to evaluate the status and relationships of these taxa. Centrotinae apparently gave rise to Nessorhinini and Oxyrhachini (both formerly treated as subfamilies, now syn.n. and syn.reinst., respectively, of Centrotinae). In contrast to previous analyses, a clade comprising Nicomiinae, Centronodinae, Centrodontinae, and the unplaced genera Holdgatiella Evans, Euwalkeria Goding and Antillotolania Ramos was recovered, but relationships within this clade were not well resolved. Nodonica bispinigera, gen.n. and sp.n., is described and placed in Centrodontini based on its sister‐group relationship to a clade comprising previously described genera of this tribe. Membracinae and Heteronotinae were consistently monophyletic. Neither Darninae nor Smiliinae, as previously defined, was monophyletic on the maximally parsimonious cladograms, but constraining both as monophyletic groups required only one additional step. The monophyly of Stegaspidinae, including Deiroderes Ramos (unplaced in Membracidae), was supported on some but not all equally parsimonious cladograms. More detailed analyses of individual subfamilies, as well as morphological data on the undescribed immatures of several membracid tribes and genera, will be needed to elucidate relationships among tribes and genera. A key to the subfamilies and tribes is provided.     

Molecular systematics of the Cactaceae

Bayesian, maximum‐likelihood, and maximum‐parsimony phylogenies, constructed using nucleotide sequences from the plastid gene region trnK‐matK, are employed to investigate relationships within the Cactaceae. These phylogenies sample 666 plants representing 532 of the 1438 species recognized in the family. All four subfamilies, all nine tribes, and 69% of currently recognized genera of Cactaceae are sampled. We found strong support for three of the four currently recognized subfamilies, although relationships between subfamilies were not well defined. Major clades recovered within the largest subfamilies, Opuntioideae and Cactoideae, are reviewed; only three of the nine currently accepted tribes delimited within these subfamilies, the Cacteae, Rhipsalideae, and Opuntieae, are monophyletic, although the Opuntieae were recovered in only the Bayesian and maximum‐likelihood analyses, not in the maximum‐parsimony analysis, and more data are needed to reveal the status of the Cylindropuntieae, which may yet be monophyletic. Of the 42 genera with more than one exemplar in our study, only 17 were monophyletic; 14 of these genera were from subfamily Cactoideae and three from subfamily Opuntioideae. We present a synopsis of the status of the currently recognized genera.
© The Willi Hennig Society 2011.     

Morphological phylogeny of army ants and other dorylomorphs (Hymenoptera: Formicidae)

Abstract. The dorylomorph group of ants comprises the three subfamilies of army ants (Aenictinae, Dorylinae, Ecitoninae) together with the subfamilies Aenictogitoninae, Cerapachyinae, and Leptanilloidinae. We describe new morphological characters and synthesize data from the literature in order to present the first hypothesis of phylogenetic relationships among all dorylomorph genera. These data include the first available character information from the newly discovered male caste of Leptanilloidinae. We used ant taxa from Leptanillinae, Myrmeciinae, and the poneromorph (Ponerinae sensu lato) subfamilies Amblyoponinae, Ectatomminae, and Paraponerinae as outgroups. We scored a total of 126 characters from twenty-two terminal taxa and used these data to conduct maximum parsimony and bootstrap analyses. The single most-parsimonious tree and bootstrap results support a single origin of army ants. The Old World army ant genus Dorylus forms a monophyletic group with the enigmatic genus Aenictogiton, which is currently known only from males; the second Old World army ant genus Aenictus is sister to this clade. This result generates the prediction that females of Aenictogiton, when discovered, will be observed to possess the army ant syndrome of behavioural and reproductive traits. The monophyly of the New World army ants (Ecitoninae) is supported very strongly, and within this group the genera Eciton, Nomamyrmex, and Labidus form a robust clade. The monophyly of Leptanilloidinae is also upheld. The subfamily Cerapachyinae appears paraphyletic, although this conclusion is not supported by strong bootstrap results. Relationships among genera of Cerapachyinae similarly are not resolved robustly, although parsimony results suggest clades consisting of (Acanthostichus + Cylindromyrmex) and (Cerapachys + Sphinctomyrmex). We tested for the effect of incompletely known taxa by conducting a secondary analysis in which the two genera containing ∼50% missing character data (Aenictogiton and Asphinctanilloides) were removed. The strict consensus of the seventeen most-parsimonious trees from this secondary analysis is poorly resolved outside the army ants and contains no clades conflicting with the primary analysis. The position of Leptanilla shifts from forming the sister group to Leptanilloidinae (without high bootstrap support) in the primary analysis, to falling within a polytomy at the base of the root of the dorylomorphs when incompletely known taxa are removed. This instability suggests that the placement of Leptanilla within the dorylomorphs in our primary analysis may be spurious.     

Molecular phylogeny of Nitidulidae: assessment of subfamilial and tribal classification and formalization of the family Cybocephalidae (Coleoptera: Cucujoidea)

We present a molecular phylogeny of Nitidulidae based on thirty ingroup taxa representing eight of the ten currently recognized subfamilies. Approximately 10 K base pairs from seven loci (12S, 16S, 18S, 28S, COI, COII and H3) were used for the phylogenetic reconstruction. The phylogeny supports the following main conclusions: (i) Cybocephalidae are formally recognized as a distinct family not closely related to Nitidulidae and its constituent taxa are defined; (ii) Kateretidae are sister to Nitidulidae; (iii) Cryptarchinae are monophyletic and sister to the remaining nitidulid subfamilies; (iv) subfamily Prometopinae stat. res. is reinstated and defined, to accommodate taxa allied to Axyra Erichson, Prometopia Erichson and Megauchenia MacLeay; (v) Amphicrossinae, Carpophilinae and Epuraeinae are shown to be closely related taxa within a well‐supported monophyletic clade; (vi) tribal affinities and respective monophyly within Nitidulinae are poorly resolved by our data and must be more rigorously tested as there was little or no support for prior morphologically based tribes or genus‐level complexes; (vii) Nitidulinae are found to be paraphyletic with respect to Cillaeinae and Meligethinae, suggesting that they should either be subsumed as tribes, or Nitidulinae should be divided into several subfamilies to preserve the status of Cillaeinae and Meligethinae; (viii) Teichostethus Sharp stat. res. is not a synonym of Hebascus Erichson and the former is reinstated as a valid genus. These conclusions and emendations are discussed in detail and presented within a morphological framework.     

Molecular phylogenetics of Erebidae (Lepidoptera,Noctuoidea)

As a step towards understanding the higher‐level phylogeny and evolutionary affinities of quadrifid noctuoid moths, we have undertaken the first large‐scale molecular phylogenetic analysis of the moth family Erebidae, including almost all subfamilies, as well as most tribes and subtribes. DNA sequence data for one mitochondrial gene (COI) and seven nuclear genes (EF‐1α, wingless, RpS5, IDH, MDH, GAPDH and CAD) were analysed for a total of 237 taxa, principally type genera of higher taxa. Data matrices (6407 bp in total) were analysed by parsimony with equal weighting and model‐based evolutionary methods (maximum likelihood), which revealed a well‐resolved skeleton phylogenetic hypothesis with 18 major lineages, which we treat here as subfamilies of Erebidae. We thus present a new phylogeny for Erebidae consisting of 18 moderate to strongly supported subfamilies: Scoliopteryginae, Rivulinae, Anobinae, Hypeninae, Lymantriinae, Pangraptinae, Herminiinae, Aganainae, Arctiinae, Calpinae, Hypocalinae, Eulepidotinae, Toxocampinae, Tinoliinae, Scolecocampinae, Hypenodinae, Boletobiinae and Erebinae. Where possible, each monophyletic lineage is diagnosed by autapomorphic morphological character states, and within each subfamily, monophyletic tribes and subtribes can be circumscribed, most of which can also be diagnosed by morphological apomorphies. All additional taxa sampled fell within one of the four previously recognized quadrifid families (mostly into Erebidae), which are now found to include two unusual monobasic taxa from New Guinea: Cocytiinae (now in Erebidae: Erebinae) and Eucocytiinae (now in Noctuidae: Pantheinae).     

Phylogeny, character evolution, and classification of Sapotaceae (Ericales)

We present the first cladistic study of the largely tropical family Sapotaceae based on both morphological and molecular data. The data were analyzed with standard parsimony and parsimony jackknife algorithms using equally and successive weighted characters. Sapotaceae are confirmed to constitute two main evolutionary lineages corresponding to the tribes Isonandreae‐Mimusopeae‐Sideroxyleae and Chrysophylleae‐Omphalocarpeae. The Sideroxyleae are monophyletic, Isonandreae are polyphyletic as presently circumscribed, and as suggested by the analyses, the subtribe Mimusopeae‐Mimusopinae has evolved within the Mimusopeae‐Manilkarinae, which hence is also paraphyletic. Generic limits must be altered within Sideroxyleae with the current members Argania, Nesoluma and Sideroxylon. Argania cannot be maintained at a generic level unless a narrower generic concept is adopted for Sideroxylon. Nesoluma cannot be upheld in a narrow or broad generic concept of Sideroxylon. The large tribe Chrysophylleae circumscribes genera such as Chrysophyllum, Pouteria, Synsepalum, and Xantolis, but the tribe is monophyletic only if the taxa from Omphalocarpeae are also included. Neither Chrysophyllum nor Pouteria are monophyletic in their current definitions. The results indicate that the African taxa of Pouteria are monophyletic and distinguishable from the South American taxa. Resurrection of Planchonella, corresponding to Pouteria section Oligotheca, is proposed. The African genera Synsepalum and Englerophytum form a monophyletic group, but their generic limits are uncertain. Classification of the Asian genus Xantolis is particularly interesting. Morphology alone is indecisive regarding Xantolis relationships, the combined unweighted data of molecules and morphology indicates a sister position to Isonandreae‐Mimusopeae‐Sideroxyleae, whereas molecular data alone, as well as successive weighted combined data point to a sister position to Chrysophylleae‐Omphalocarpeae. An amended subfamily classification is proposed corresponding to the monophyletic groups: Sarcospermatoideae (Sarcosperma), Sapotoideae (Isonandreae‐Mimusopeae‐Sideroxyleae) and Chrysophylloideae (Chrysophylleae‐Omphalocarpeae), where Sapotoideae circumscribes the tribes Sapoteae and Sideroxyleae as well as two or three as yet unnamed lineages. Morphological characters are often highly homoplasious and unambiguous synapomorphies cannot be identified for subfamilies or tribes, which we believe are the reason for the variations seen between different classifications of Sapotaceae. © The Willi Hennig Society 2005.     

What are the closest relatives of the hot-spring snakes (Colubridae, Thermophis), the relict species endemic to the Tibetan Plateau?

We conducted phylogenetic analyses to identify the closest related living relatives of the Xizang and Sichuan hot-spring snakes (T. baileyi and T. zhaoermii) endemic to the Tibetan Plateau, using mitochondrial DNA sequences (cyt b, ND4) from eight specimens, together with sequences from 95 additional caenophidian and five henophidian genera that were downloaded from GenBank. Phylogenetic trees were obtained using Bayesian Inference and Maximum likelihood methods. Results suggest that hot-spring snakes, which are adapted to high and cold environments, were clustered in the monophyletic Xenodontinae. Xenodontinae is one of the largest subfamilies of colubrid snakes, with about 90 genera and more than 500 species known, and are primarily tropical snakes previously thought to be restricted to the New World. Our data failed to provide any evidence that the New World xenodontines diverged from Thermophis and dispersed into the New World, also failed to suggest a colonization of Asia by New World xenodontines by dispersal from the New World. An alternative plausible scenario may be that Thermophis and the New World xenodontines evolved independently in Asia and America, respectively, after the divergence of their common ancestor. The divergence of the two species in Thermophis was caused by the barrier of the Hengduan Mountains, and the speciation had almost occurred when Tibetan Plateau attained present elevation.     

Molecular systematics and evolution of the Ptinidae (Coleoptera: Bostrichoidea) and related families

The Ptinidae (Coleoptera: Bostrichoidea) are a cosmopolitan, ecologically diverse, but poorly known group of Coleoptera and, excluding a few economic pests, species are rarely encountered. This first broad phylogenetic study of the Ptinidae s.l. (i.e. including both the spider beetles and anobiids) examines relationships based on DNA sequence data from two mitochondrial genes (16S and COI) and one nuclear gene (28S), using out‐group taxa from both the Bostrichidae and Dermestidae. Topologies varied depending on the genes used and whether data were analysed with either parsimony or Bayesian methods. Generally the two mitochondrial genes supported relationships near the tips of the phylogeny, whereas the nuclear gene supported the basal relationships. The monophyly of the Ptinidae was not inferred by all of the gene combinations and analysis methods, although the combined Ptinidae and Bostrichidae have a single origin in all cases. Alternative relationships include the Ptinidae s.s. (i.e. Ptininae and Gibbiinae) as sister to the anobiids (i.e. the nine remaining subfamilies of Ptinidae s.l.) + Bostrichidae, or the Bostrichidae as sister to the Ptinidae s.s.+ anobiids. Most of the larger subfamilies within the Ptinidae are not monophyletic. Further analysis with more taxa and more genes will be required to clarify and decide upon the best hypothesis of relationships found within the clades of the Bostrichidae and Ptinidae. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 165 , 88–108.     

Nomenclatural changes in <Emphasis Type="Italic">Lithospermum</Emphasis> (Boraginaceae) and related taxa following a reassessment of phylogenetic relationships

Lithospermum (Boraginaceae) comprises approximately 40 species in both the Old and New Worlds, with a center of diversity in the southwestern United States and Mexico. Using ten cpDNA regions, a phylogeny of Lithospermum and related taxa was reconstructed. Lithospermum (including New World and Old World species) and related New World members of Lithospermeae form a monophyletic group, with Macromeria, Onosmodium, Nomosa, Lasiarrhenum, and Psilolaemus nested among species of Lithospermum. New World Lithospermeae also is a monophyletic group, with Eurasian species of Lithospermum sister to this group. Because Lithospermum is not monophyletic without the inclusion of the other New World genera, species from these genera are transferred to Lithospermum, and appropriate nomenclatural changes are made. New combinations are Lithospermum album, Lithospermum barbigerum, Lithospermum dodrantale, Lithospermum exsertum, Lithospermum helleri, Lithospemum leonotis, Lithospermum notatum, Lithospermum oaxacanum, Lithospermum pinetorum, Lithospermum rosei, Lithospermum trinverium, and Lithospermum unicum; new names are Lithospermum chiapense, Lithospermum johnstonii, Lithospermum macromeria, Lithospermum onosmodium, Lithospermum rzedowskii, and Lithospermum turneri.     

PHYLOGENY OF THE EUGLENOID LORICATE GENERA TRACHELOMONAS AND STROMBOMONAS (EUGLENOPHYTA) INFERRED FROM NUCLEAR SSU AND LSU rDNA1

Previous studies using the nuclear SSU rDNA and partial LSU rDNA have demonstrated that the euglenoid loricate taxa form a monophyletic clade within the photosynthetic euglenoid lineage. It was unclear, however, whether the loricate genera Trachelomonas and Strombomonas were monophyletic. In order to determine the relationships among the loricate taxa, SSU and LSU nuclear rDNA sequences were obtained for eight Strombomonas and 25 Trachelomonas strains and combined in a multigene phylogenetic analysis. Conserved regions of the aligned data set were used to generate maximum‐likelihood (ML) and Bayesian phylogenies. Both methods recovered a strongly supported monophyletic loricate clade with Strombomonas and Trachelomonas species separated into two sister clades. Taxa in the genus Strombomonas sorted into three subclades. Within the genus Trachelomonas, five strongly supported subclades were recovered in all analyses. Key morphological features could be attributed to each of the subclades, with the major separation being that all of the spine‐bearing taxa were located in two sister subclades, while the more rounded, spineless taxa formed the remaining three subclades. The separation of genera and subclades was supported by 42 distinct molecular signatures (33 in Trachelomonas and nine in Strombomonas). The morphological and molecular data supported the retention of Trachelomonas and Strombomonas as separate loricate genera.     

Molecular phylogeny of Pamphagidae (Acridoidea,Orthoptera) from China based on mitochondrial cytochrome oxidase II sequences

Abstract Phylogenetic relationships of Pamphagidae were examined using cytochrome oxidase subunit II (COII) mtDNA sequences (684 bp). Twenty‐seven species of Acridoidea from 20 genera were sequenced to obtain mtDNA data, along with four species from the GenBank nucleotide database. The purpose of this study was analyzing the phylogenetic relationships among subfamilies within Pamphagidae and interpreting the phylogenetic position of this family within the Acridoidea superfamily. Phylogenetic trees were reconstructed using neighbor‐joining (NJ), maximum parsimony (MP) and Bayesian inference (BI) methods. The 684 bp analyzed fragment included 126 parsimony informative sites. Sequences diverged 1.0%–11.1% between genera within subfamilies, and 8.8%–12.3% between subfamilies. Amino acid sequence diverged 0–6.1% between genera within subfamilies, and 0.4%–7.5% between subfamilies. Our phylogenetic trees revealed the monophyly of Pamphagidae and three distinct major groups within this family. Moreover, several well supported and stable clades were found in Pamphagidae. The global clustering results were similar to that obtained through classical morphological classification: Prionotropisinae, Thrinchinae and Pamphaginae were monophyletic groups. However, the current genus Filchnerella (Prionotropisinae) was not a monophyletic group and the genus Asiotmethis (Prionotropisinae) was a sister group of the genus Thrinchus (Thrinchinae). Further molecular and morphological studies are required to clarify the phylogenetic relationships of the genera Filchnerella and Asiotmethis.     

Phylogenetic relationships of subfamilies and circumscription of tribes in the family Hesperiidae (Lepidoptera: Hesperioidea)

A comprehensive tribal‐level classification for the world’s subfamilies of Hesperiidae, the skipper butterflies, is proposed for the first time. Phylogenetic relationships between tribes and subfamilies are inferred using DNA sequence data from three gene regions (cytochrome oxidase subunit I‐subunit II, elongation factor‐1α and wingless). Monophyly of the family is strongly supported, as are some of the traditionally recognized subfamilies, with the following relationships: (Coeliadinae + (“Pyrginae” + (Heteropterinae + (Trapezitinae + Hesperiinae)))). The subfamily Pyrginae of contemporary authors was recovered as a paraphyletic grade of taxa. The formerly recognized subfamily Pyrrhopyginae, although monophyletic, is downgraded to a tribe of the “Pyrginae”. The former subfamily Megathyminae is an infra‐tribal group of the Hesperiinae. The Australian endemic Euschemon rafflesia is a hesperiid, possibly related to “Pyrginae” (Eudamini). Most of the traditionally recognized groups and subgroups of genera currently employed to partition the subfamilies of the Hesperiidae are not monophyletic. We recognize eight pyrgine and six hesperiine tribes, including the new tribe Moncini. © The Willi Hennig Society 2008.     

When molecules and morphology concur: the ‘Gondwanan’ midges (Diptera: Chironomidae)

A phylogeny of the Chironomidae subfamily Podonominae, significant in the history of phylogenetic biogeography, is estimated from an analysis of four genes. Fragments of two ribosomal genes (18S and 28S), one nuclear protein‐coding gene (CAD), and one mitochondrial protein‐coding gene (COI) were sequenced from specimens representing 13 of 15 genera, and analysed using mixed model Bayesian and maximum likelihood inference methods. Podonominae is monophyletic and sister to Tanypodinae – the shared development of the larval ligula is synapomorphic and diagnostic. Tribe Podonomini is monophyletic with the inclusion of Trichotanypus; tribe Boreochlini is a grade. Monophyly is confirmed for the genera Podonomus Philippi, Podonomopsis Brundin, Podochlus Brundin, Archaeochlus Brundin and Austrochlus Cranston, Edward & Cook: Parochlus Enderlein becomes monophyletic through the inclusion of Zelandochlus Brundin ( n.syn. ) with its type species, P. latipalpis (Brundin) n.comb. The ‘mandibulate’Archaeochlus plus Austrochlus is monophyletic with nonmandibulate Afrochlus weakly supported as a member of, or sister to, the African Archaeochlus. Subtending this group is Lasiodiamesa, although it associates in some analyses with the sister group Tanypodinae. Generic relationships coincide with those proposed based on morphology, particularly as understood via all life history stages of some problematic (autapomorphic, adult‐based) taxa. Divergence time analysis (beast ) allows inference of Mesozoic diversification of higher taxa in Podonominae, of appropriate timing for fragmentation of Gondwana, post‐African divergence, to have caused vicariance. Shallower nodes (within genera) imply both younger vicariance involving Antarctica and some recent dispersal, including southern to northern hemisphere movement in the New World. New Zealand taxa test controversial biogeographical relationships and show proximity to southern South America without direct Australian sister taxon pairs: dating implies persistence of midges through the ‘Oligocene’ bottleneck.     

Phylogeny and classification of Corylophidae (Coleoptera: Cucujoidea) with descriptions of new genera and larvae

Abstract Phylogenetic relationships within the family Corylophidae were investigated. Twenty ingroup taxa and six outgroups were included in a cladistic analysis, based on 48 characters derived from adult and larval morphology. Phylogenetic analysis confirms that Corylophidae are monophyletic within the superfamily Cucujoidea and may be subdivided into two subfamilies: the Australian Periptycinae and the cosmopolitan Corylophinae containing 10 tribes: Foadiini trib.n. , Cleidostethini, Aenigmaticini, Parmulini, Sericoderini, Peltinodini, Orthoperini, Corylophini, Teplinini and Rypobiini. All currently recognized family‐group taxa are thoroughly diagnosed, and keys to their identification based on adults and larvae are provided. Two new genera and three species are described: Weirus gen.n ., containing only W. tozer sp . n . (Australia: Queensland), and Stanus gen.n. , with the two species S. bowesteadi sp.n . (New Zealand) and S. tasmanicus sp.n. (Tasmania). The larvae of Pakalukodes bimaculatus?lipiński et al. from Queensland and of Stanus bowesteadi sp.n. from New Zealand are described and illustrated for the first time.     

Family‐level relationships of the spittlebugs and froghoppers (Hemiptera: Cicadomorpha: Cercopoidea)

The spittlebug superfamily Cercopoidea (Hemiptera: Cicadomorpha) comprises approximately 3000 phytophagous species (including some economically important pests of grass crops) classified among the families Cercopidae, Aphrophoridae, Epipygidae, Clastopteridae and Machaerotidae. However, the monophyly of these taxa has never been tested and the evolutionary relationships among these major lineages are unknown. Presented here are the results of the first ever phylogenetic investigation of the higher‐level relationships within Cercopoidea, based on DNA nucleotide sequence data from six loci (18S rDNA, 28S rDNA, histone 3, wingless, cytochrome oxidase I and cytochrome oxidase II) generated from exemplars of 109 spittlebug species representing all five described families, seven of eight subfamilies and 61 genera (eight additional exemplars, representing a selection of other Auchenorrhyncha taxa, were included as outgroups). The resulting topologies are used to evaluate the monophyly of each cercopoid family, and further to calculate divergence date estimates to examine the chronological origins and historical diversification of Cercopoidea. The results of this investigation suggest that: (i) four of the five described families are monophyletic; Epipygidae was recovered consistently as originating within Aphrophoridae; (ii) the exclusively Old World Machaerotidae is the most anciently diversified family of extant spittlebugs; (iii) New World Cercopidae (i.e. Ischnorhininae) constitute a derived monophyletic lineage; (iv) the genus Microsargane Fowler, classified currently within Aphrophoridae, actually belongs within Cercopidae; and (v) the origins of the major spittlebug lineages probably coincided with the breakup of Pangaea and, subsequently, Gondwana, as well as major floristic diversification such as the rise of angiosperms.     

Phylogeny of the tribe Empoascini (Hemiptera: Cicadellidae: Typhlocybinae) based on morphological characteristics,with reclassification of the Empoasca generic group

A morphology‐based phylogenetic analysis of the tribe Empoascini (Hemiptera: Cicadellidae: Typhlocybinae) is presented for 58 of 83 formerly recognized genera based on 99 morphological characters of adults. The results support excluding the New World Beamerana generic group from Empoascini. The remaining genera of Empoascini were recovered as a monophyletic sister group of Dikraneurini. Previously recognized tribes Jorumini and Helionini are derived from within Empoascini and are considered synonyms of the latter tribe. Three previously recognized informal generic groups, the Empoasca group, Alebroides group and Usharia group were paraphyletic but the Ficiana group was recovered as monophyletic based on five synapomorphies. Genera previously placed in the Alebroides group represent at least six independent lineages, indicating that the hind wing character separating this group from the Empoasca group (CuA and MP veins free) is highly homoplasious. Empoasca (sensu lato) is also paraphyletic. Thus, twelve previously recognized subgenera of Empoasca are elevated to genus status and five species groups of Empoasca from the New World are recognized as separate new genera. Sikkimasca Dworakowska, 1993 is treated as synonym of Marolda Dworakowska, 1977 based on the phylogeny. Biogeographic analysis suggests that Empoascini most likely first evolved in the Oriental region and spread to other biogeographic realms more recently by multiple independent invasions.