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.     

Taxonomic review and key to European Ichneumon flies (Hymenoptera, Ichneumonidae), parasitoids of gall-forming sawflies of the genera Pontania Costa, Phyllocolpa Benson, and Euura Newman (Hymenoptera, Tenthredinidae) on willows: Part I

Extensive material on parasitoids of the family Ichneumonidae reared from gall-making sawflies on Salix is identified. The general host—parasite list includes 75 species of hosts—Pontania (40 spp.), Phyllocolpa (21 spp.), Euura (14 spp.), their food plants (33 species of Salix), and their parasitoids of the family Ichneumonidae; many data on hosts and parasitoids are recorded for the first time. The complex of ichneumonids on gallforming sawflies is composed of about 45 species belonging to 15 genera of 6 subfamilies. The annotated list of about 30 species of ichneumonids (except for species of Saotis) reared from gall-making sawflies is given with data on the literature, studied material, distribution, and hosts. Two new species Anoncus gallicola sp. n. and Rhinotorus brachycerus sp. n. are described. A key to the species of the Ichneumonidae of this complex is given.     

Phylogeny of the bee family Melittidae (Hymenoptera: Anthophila) based on combined molecular and morphological data

The bee family Melittidae comprises a small, but biologically fascinating, group of mostly oligolectic bees, some of which are oil collecting. Phylogenetic relationships within this family are poorly understood and some genera cannot be placed with confidence at the subfamily level. We analysed melittid phylogeny using a combined dataset of five nuclear genes [28S, elongation factor‐1α (EF‐1α, F2 copy), long‐wavelength rhodopsin, Na‐K ATPase and RNA polymerase II] spanning 4842 bp plus 68 adult morphological characters. Our study included 25% of the species‐level diversity and 81% of the generic‐level diversity and included all previously recognized tribes and subfamilies. We analysed the dataset using parsimony, maximum likelihood and Bayesian methods. All methods yielded congruent results. All topologies recovered the three previously recognized subfamilies (Dasypodainae, Melittinae, Meganomiinae), but two genera (Afrodasypoda and Promelitta) are transferred from Dasypodainae to Melittinae. On the basis of our tree topologies we identify four tribes (Dasypodaini comb.n. , Hesperapini stat.n. , Macropidini comb.n. and Melittini), only one of which (Melittini) matches a widely used classification. Lastly, we discuss the evolution of host‐plant association in the light of our new phylogenetic hypothesis. Our results strongly support multiple independent origins of oil‐collecting behaviour in the Melittinae.     

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 in the cricket tribe Xenogryllini (Orthoptera,Gryllidae, Eneopterinae) and description of the Indian genus Indigryllus gen. nov.

The subfamily Eneopterinae is known greatly for its diversified acoustic modalities and disjunct distribution. Within Eneopterinae, tribe Lebinthini is the most studied group, due to its highest species diversity (ca. 150 species in 12 genera), endemic distribution on the islands of Southeast Asia and of the South West Pacific, males’ ability to produce high‐frequency calling songs, and evolution of females’ vibrational response. To investigate the distribution pattern and diversification of acoustic and behavioral attributes in a larger frame, clear understanding of phylogenetic relationships within other tribes of Eneopterinae is vital. In this study, we focus on the tribe Xenogryllini, sister group of Lebinthini. Xenogryllini, as opposed to Lebinthini, is known by fewer species (11 species in two genera), distributed widely in continental Asia and Africa, and for producing low‐frequency calling songs. We describe a new genus Indigryllus with a new species of the tribe Xenogryllini, discovered from the southwest of India. We used eight molecular genetic markers to reconstruct the phylogenetic relationships. The resultant phylogenetic tree is used to compare and discuss distribution patterns and acoustic modalities between Lebinthini and Xenogryllini.     

A comprehensive molecular phylogeny of tiger beetles (Coleoptera,Carabidae, Cicindelinae)

Tiger beetles are a remarkable group that captivates amateur entomologists, taxonomists and evolutionary biologists alike. This diverse clade of beetles comprises about 2300 currently described species found across the globe. Despite the charisma and scientific interest of this lineage, remarkably few studies have examined its phylogenetic relationships with large taxon sampling. Prior phylogenetic studies have focused on relationships within cicindeline tribes or genera, and none of the studies have included sufficient taxon sampling to conclusively examine broad species patterns across the entire subfamily. Studies that have attempted to reconstruct higher‐level relationships of Cicindelinae have yielded conflicting results. Here, we present the first taxonomically comprehensive molecular phylogeny of Cicindelinae to date, with the goal of creating a framework for future studies focusing on this important insect lineage. We utilized all available published molecular data, generating a final concatenated dataset including 328 cicindeline species, with molecular data sampled from six protein‐coding gene fragments and three ribosomal gene fragments. Our maximum‐likelihood phylogenetic inferences recover Cicindelinae as sister to the wrinkled bark beetles of the subfamily Rhysodinae. This new phylogenetic hypothesis for Cicindelinae contradicts our current understanding of tiger beetle phylogenetic relationships, with several tribes, subtribes and genera being inferred as paraphyletic. Most notably, the tribe Manticorini is recovered nested within Platychilini including the genera Amblycheila Say, Omus Eschscholtz, Picnochile Motschulsky and Platychile Macleay. The tribe Megacephalini is recovered as paraphyletic due to the placement of the monophyletic subtribe Oxycheilina as sister to Cicindelini, whereas the monophyletic Megacephalina is inferred as sister to Oxycheilina, Cicindelini and Collyridini. The tribe Collyridini is paraphyletic with the subtribes Collyridina and Tricondylina in one clade, and Ctenostomina in a second one. The tribe Cicindelini is recovered as monophyletic although several genera are inferred as para‐ or polyphyletic. Our results provide a novel phylogenetic framework to revise the classification of tiger beetles and to encourage the generation of focused molecular datasets that will permit investigation of the evolutionary history of this lineage through space and time.     

The cladistics and higher classification of the Pimpliformes (Hymenoptera: Ichneumonidae)

The monophyly of the ichneumonid clade Pimpliformes is established and the phylogenetic relationships of the eight component subfamilies are resolved. The clade (Acaenitinae + (Diacritinae + (Cylloceriinae + (Diplazontinae + Orthocentrinae)))) is the sister-lineage to the clade (Pimplinae + (Rhyssinae + Poemeniinae)). The Nearctic genus Cressonia Dasch is transferred to the Diacritinae from the Orthocentrinae. Tribes are not recognized in the Acaenitinae as the Coleocentrini (sensu Townes, 1971) is paraphyletic with respect to the Acaenitini. The Cylloceriinae is recognized as comprising three genera, Cylloceria Schiødte, Allomacrus Förster and Sweaterella gen.n. The Orthocentrinae, including the Helictinae of authors, is shown to be monophyletic, but the latter is clearly shown to be paraphyletic if the Orthocentrus genus-group is excluded. The Pimplinae comprises four monophyletic tribes: the Delomeristini, consisting of Delomerista Förster and Atractogaster Kriechbaumer; the Perithoini trib.n., which includes only Perithous Holmgren (= Hybomischos Baltazar syn.n.); the Pimplini, which includes the Theronia genus-group as well as the Pimpla genus-group; and the Ephialtini, which includes the Polysphinctini syn.n., a monophyletic group that previously rendered the restricted Ephialtini paraphyletic. The tribe Delomeristini is the sister-group to the clade (Ephialtini + (Perithoini + Pimplini)). The subfamily Poemeniinae is recognized as comprising three tribes: the Pseudorhyssini (trib.n.) which includes the single Holarctic genus Pseudorhyssa Merrill; the Rodrigamini (trib.n.) which includes only the Costa Rican genus Rodrigama Gauld; and the Poemeniini. The tribe Pseudorhyssini is the sister-group to the clade (Rodrigamini + Poemeniini). The phylogenetic inter-relationships of the genera of Poemeniini are resolved. A new genus from South Africa, Guptella (gen.n.) is described, and Achorocephalus Kriechbaumer is shown to be a synonym of Eugalta Cameron (syn.n.). The evolution of biological traits within the Pimpliformes is discussed with reference to the elucidated phylogeny, and zoogeographic patterns are outlined.     

Mitochondrial and nuclear markers support the monophyly of Dolichopodidae and suggest a rapid origin of the subfamilies (Diptera: Empidoidea)

Abstract. The Dolichopodidae is a species‐rich dipteran group with almost 7000 described species. The monophyly of the subfamilies and their relationships remain largely unknown because the polarities of key morphological characters are unclear and molecular data are available only for 9 of the 19 proposed subfamilies. Here we test whether molecular data from two nuclear (18S, 28S) and four mitochondrial (12S, 16S, Cytb, COI) genes can resolve the higher‐level relationships within the family. Our study is based on 76 Oriental species from 12 dolichopodid subfamilies and uses eight species of Empididae and Hybotidae as outgroups. Parsimony and likelihood analyses confirm the monophyly of the Dolichopodidae, as well as the monophyly of five of the ten subfamilies represented by more than two species [Sympycninae, Sciapodinae, Dolichopodinae, Hydrophorinae (excluding tribe Aphrosylini), Neurigoninae]. There is strong support for restoring the tribe Aphrosylini as a separate subfamily Aphrosylinae. The monophyly of Medeterinae, Peloropeodinae and Diaphorinae is dependent on which tree reconstruction technique is used, how indels are coded, and whether the fast‐evolving sites are excluded. Overall, we find that our sample of Oriental species is largely compatible with the subfamily concepts that were developed for the northern temperate fauna. However, our data provide little support for relationships between the subfamilies. Branch lengths, saturation, and distance plots suggest that this is probably the result of the rapid origin of dolichopodid subfamilies over a relatively short time. We find that genera that are difficult to place into subfamilies based on morphological characters are generally also difficult to place using molecular data. We predict that a dense, balanced taxon sample and protein‐encoding nuclear genes will be needed to resolve the higher‐level relationships in the Dolichopodidae.     

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.     

Evolution of blindness in scolopendromorph centipedes (Chilopoda: Scolopendromorpha): insight from an expanded sampling of molecular data

Relative to its diversity (34 genera, 700 species), Scolopendromorpha has been undersampled in molecular phylogenetic analyses compared with the other chilopod orders. Previous analyses based on morphology have not resolved several key controversies in systematics and evolutionary morphology unambiguously. Here we apply new molecular and morphological data to scolopendromorph phylogenetics, with a focus on the evolution of blindness. The taxonomic sample includes 19 genera, many lacking previous molecular data, and diverse, cosmopolitan genera of Scolopendridae are sampled by multiple species. Phylogenetic analysis with Direct Optimization used 94 morphological characters and ca. 4.5 kb of sequence data from two nuclear (18S and 28S rRNA) and two mitochondrial (16S rRNA and COI) loci. A single most‐parsimonious cladogram selected after sensitivity analyses resolves Scolopendromorpha as monophyletic, and divides it into a blind clade of three families (Plutoniumidae, Cryptopidae, Scolopocryptopidae) and its ocellate sister group, Scolopendridae. Some species‐rich, cosmopolitan genera (Cormocephalus, Otostigmus, Scolopendra) in Scolopendridae are non‐monophyletic, and in several instances (e.g. New and Old World Scolopendra) relationships are more congruent with geographical distributions than with traditional classifications. The tribe Asanadini is particularly subject to parameter‐sensitivity, nesting in the combined analysis within Scolopendrini but as sister to all other Scolopendrinae for molecular data alone. The total‐evidence tree unambiguously optimizes trunk segmentation: a 23‐segmented trunk has a single origin in the blind clade. © The Willi Hennig Society 2011.     

Phylogenetic relationships of major clades of Catostomidae (Teleostei: Cypriniformes) as inferred from mitochondrial SSU and LSU rDNA sequences

Suckers (Family Catostomidae) are holarctic in distribution and include 76 recent species in 14 genera, with 13 genera and 75 species occurring in North and Central America and Siberia. Although this group constitutes a significant component of many aquatic ecosystems, most historic systematic effort has been either alpha- or limited beta-level studies focusing on the two largest tribes within the family, the Catostomini and the Moxostomatini. A recent phylogenetic study based on morphological, biochemical, and early life history characters has advanced current understanding of relationships among catostomid fishes. To further examine phylogenetic relationships among basal lineages of catostomids, we sequenced the entire mitochondrial (mt) SSU and LSU rRNA genes from genera representing all subfamilies and tribes within Catostomidae. Phylogenetic analysis of gene sequences yielded monophyletic Catostomidae, Ictiobinae, and Catostominae and para- or polyphyletic Cycleptinae, with Myxocyprinus as the basal-most taxon and Cycleptus as either the next most-basal taxon or the taxon basal to the Catostominae. Relationships within the Catostominae were generally consistent with those proposed in the above-noted recent phylogenetic study although Thoburnia and Hypentelium were either a clade sister to or a grade group relative to Moxostoma and Scartomyzon. In all trees, Scartomyzon was paraphyletic and embedded within Moxostoma. Phylogenetic affinities of Erimyzon and Minytrema varied depending on data set and character weighting scheme employed. To better reflect phylogenetic relationships resolved in this extensive analysis, we propose the following changes to the classification of catostomids: formation of the new subfamily Myxocyprininae, containing Myxocyprinus from China; restriction of the Cycleptinae to the two species of Cycleptus from North America; restriction of the tribe Moxostomatini to Moxostoma and Scartomyzon; Erimyzon and Minytrema are incertae sedis within Catostominae; and resurrection of the tribe Thoburniini, containing Thoburnia and expanded to include Hypentelium.     

Higher‐level phylogeny of diving beetles (Coleoptera: Dytiscidae) based on larval characters

A comprehensive higher‐level phylogeny of diving beetles (Dytiscidae) based on larval characters is presented. Larval morphology and chaetotaxy of a broad range of genera and species was studied, covering all currently recognized subfamilies and tribes except for the small and geographically restricted Hydrodytinae, where the larva is unknown. The results suggest several significant conclusions with respect to the systematics of Dytiscidae including the following: monophyly of all currently recognized subfamilies, although Dytiscinae when considered in a broad context is rendered paraphyletic by Cybistrinae; currently recognized tribes are monophyletic except for Agabini, Hydroporini and Laccornellini; inter‐subfamily and inter‐tribe relationships generally show weak support, except for a few well supported clades; three distinct clades are recognized within Dytiscinae [Dytiscini sensu lato (i.e. including the genera Dytiscus Linnaeus and Hyderodes Hope), Hydaticini sensu lato, and Cybistrini]; and recognition of Pachydrini as a distinct tribe. Other less robust results include: Methlini sister to the rest of Hydroporinae; relative basal position of Laccornini, Hydrovatini and Laccornellini within Hydroporinae; close relationship of Agabinae and Copelatinae; Matinae nested deep within Dytiscidae, as sister to a large clade including Colymbetinae, Coptotominae, Lancetinae and Dytiscinae sensu lato; the sister‐group relationship of Agabetini and Laccophilini is confirmed. The results presented here are discussed and compared with previous phylogenetic hypotheses based on different datasets, and the evolution of some significant morphological features is discussed in light of the proposed phylogeny. All suprageneric taxa are diagnosed, including illustrations of all relevant synapomorphies, and a key to separate subfamilies and tribes is presented, both in traditional (paper) format and as an online Lucid interactive identification key.     

A combined morphological and molecular phylogeny of the genus Chironius Fitzinger, 1826 (Serpentes: Colubridae)

Chironius is one of the most speciose genera of the South American colubrid snakes. Although the genus represents a well‐known radiation of diurnal racers, its monophyly, affinities with other Neotropical colubrid genera, and intrageneric relationships are open questions. Here, we present a phylogenetic analysis of Chironius based on a data matrix that combines one nuclear (c‐mos) and two mitochondrial (12S and 16S rRNA) genes with 37 morphological characters derived from scutellation, skull, and hemipenial features. Phylogenetic relationships were inferred using maximum parsimony (MP) and maximum likelihood (ML). Our combined morphological and molecular analyses strongly support the monophyly of the genus Chironius and its sister‐group relationship with a clade formed by the genera Dendrophidion and Drymobius. Phylogenetic relationships within the genus Chironius is still controversial, although five clades are retrieved with medium to strong support. © 2014 The Linnean Society of London     

Phylogeny of Adramini (Diptera,Tephritidae) based on integrative evidence

Species of the tribe Adramini (Tephritidae: Trypetinae) are usually slender, and some specific species have eyes borne at the ends of their long stalks. This tribe is mainly distributed in the tropics and subtropics of the Afrotropical, Oriental and Australasian Regions; relatively few species occur in the Palearctic and Nearctic Regions. The phylogeny of the tribe Adramini is presented here based on analysis of morphological and molecular information (DNA sequences of nuclear 28S rDNA, mitochondrial COI and COII, and 16S rDNA genes) for its representative species in most genera. Three monophyletic groups (Adrama‐com‐group, Pelmatops‐com‐group and Dimeringophrys‐com‐group) were discovered in the combined morphological and molecular tree. The results showed moderate support for the monophyly of Adramini and strong support for most of its genera. However, Euphranta appears to be polyphyletic. Sapadrama, Celidodacus and Euphranta are basal taxon, and Coelopacidia, Soita and Trypanophion are closely related to the stalk‐eyed fruit flies (Pelmatops + Pseudopelmatops). A hypothesis regarding the morphology–function relationships for two main groups of Adramini (Adrama‐group and Pelmatops‐group) with different evolving probabilities is inferred. Sapadrama is proposed be removed from Adramini; a new genus, Ichneumonomacula Chen gen. n. and a new species Ichneumonmacula wangyongi Chen sp. n., are recognized and described, and a key to recognize the genera of Adramini around the world is provided.     

PHYLOGENY OF THE RUBIACEAE AND THE LOGANIACEAE: CONGRUENCE OR CONFLICT BETWEEN MORPHOLOGICAL AND MOLECULAR DATA?

Phylogenetic analyses of 33 genera of Rubiaceae were performed using morphological and a few chemical characters. Parsimony analysis based on 29 characters resulted in eight equally parsimonious trees, with a consistency index of 0.40 and a retention index of 0.69. These results were compared to a phylogenetic analysis of the same genera based on chloroplast DNA restriction site data. There are discrepancies between the two analyses, but if we consider groupings reflected in the present classification there is much congruency. With the exception of four genera, all the genera are positioned in the same group of taxa in the two analyses. Clades of taxa representing three of the four subfamilies (~the Antirheoideae, ~the Rubioideae, and the ~Ixoroideae) are monophyletic, while the fourth subfamily Cinchonoideae is shown to be paraphyletic. Both analyses support a widened tribe Chiococceae, including the former subtribe Portlandiinae (Condamineeae). Furthermore, in both analyses the tribe Hamelieae is placed outside the subfamily Rubioideae where it is now housed. In search for the most plausible sister group to the Rubiaceae, the genus Cinchona (Rubiaceae) was analyzed together with 13 genera of the Loganiaceae, Nerium (Apocynaceae), and Exacum (Gentianaceae). Cornus (Comaceae), Olea (Oleaceae), and these two genera together were used as outgroups. The analysis, including 25 characters, 16 taxa, and with Cornus and Olea together as an outgroup, resulted in four equally parsimonious trees, with a consistency index of 0.53 and a retention index of 0.62. The non-Loganiaceae taxa Cinchona (Rubiaceae), Nerium (Apocynaceae), and Exacum (Gentianaceae) were all found to have their closest relatives within the Loganiaceae indicating that the Loganiaceae are paraphyletic and ought to be reclassified. As a result of the morphological data the most plausible sister group to the Rubiaceae is the tribe Gelsemieae of the Loganiaceae.     

Morphological phylogeny of gall‐forming aphids of the tribe Eriosomatini (Aphididae: Eriosomatinae)

Aphids of the tribe Eriosomatini are typically associated with the tree genera Ulmus and Zelkova as the primary host, on which they induce several types of leaf gall. To elucidate evolutionary changes in the aphid–host associations and gall morphology, phylogenetic relationships were inferred using 36 species (28 in the ingroup) and based on 52 morphological characters. Phylogenetic analysis with equal character weighting led to hundreds of most‐parsimonious trees, and the strict consensus of these was poorly resolved. However, the successive weighting of characters yielded three most‐parsimonious trees. The strict consensus of these supported the monophyly of the Eriosomatini and the monophyly of most genera. Reconstruction of the aphid–host associations on the consensus tree indicated that the ancestral Eriosomatini were associated with Zelkova and that the association with Ulmus evolved twice independently. Ancestral reconstruction suggests that galls of the leaf‐roll type are ancestral to those of the completely and incompletely closed pouch type, and that each type of pouch galls evolved independently. It is suggested that despite early diversification of the Eriosomatini on Zelkova species, Zelkova‐associated eriosomatines had become extinct or survived as relict parthenogens on the secondary host due to the elimination of Zelkova in most regions since the late Tertiary. In contrast, two large genera in the Eriosomatini, Eriosoma and Tetraneura, are associated with the largest elm section Ulmus whose elements are distributed widely in Eurasia, including boreal regions. Therefore, the available evidence suggests that the present species diversity and distribution patterns of the eriosomatines have been largely affected by the diversification and extinction of their host plants during the late Tertiary and Quaternary.     

Using molecules and morphology to infer the phylogenetic relationships and evolutionary history of the Dirini (Nymphalidae: Satyrinae), a tribe of butterflies endemic to Southern Africa

The first empirically supported phylogenetic hypothesis of relationships for the southern African endemic butterfly tribe Dirini is presented. Data derived from the morphology and ecology of the adults and immature stages (33 characters), and portions of the mitochondrial gene cytochrome oxidase I (COI) and the nuclear genes elongation factor 1α (EF1α) and wingless (WG) (totalling 1734 bp) were used to infer the relationships of the in‐group genera. An expanded molecular dataset using four genera from the Nymphalini and Satyrini to root the tree, and three genera from the Melanitini to test the monophyly of the tribe, was analysed using parsimony and Bayesian methods. Estimates of divergence times were calculated using two fossil calibrations under a relaxed molecular clock model. The monophyly of the tribe and each in‐group genus were strongly supported. Key findings are the sister‐taxon relationship of Aeropetes and Tarsocera, the apparent simultaneous or nearly simultaneous radiation of four lineages, the polyphyly of the species within Torynesis, and the apparent trans‐Atlantic dispersal of the ancestors of Manataria about 40 Ma. Estimates of divergence times indicate that the tribe has undergone two major radiations since its origin: the first when they left forest habitats in the mid–late Oligocene, shortly after the radiation of the grasses (Poaceae), and the second in the early‐middle Pliocene, coinciding with the aridification of southern Africa and the spread of conditions that favoured C₄ grasses over the C₃ grasses that dirine larvae prefer to eat. The high species diversity within the tribe appears to be partly a taxonomic artefact that may have resulted from the misinterpretation of climate‐related phenotypic variation within extant species. Relocation and breeding experiments should test this hypothesis.     

Molecular phylogenetics of Phyllanthaceae: evidence from plastid MATK and nuclear PHYC sequences

Plastid matK and a fragment of the low-copy nuclear gene PHYC were sequenced for 30 genera of Phyllanthaceae to evaluate tribal and generic delimitation. Resolution and bootstrap percentages obtained with matK are higher than that of PHYC, but both regions show nearly identical phylogenetic patterns. Phylogenetic relationships inferred from the independent and combined data are congruent and differ from previous, morphology-based classifications but are highly concordant with those of the plastid gene rbcL previously published. Phyllanthaceae is monophyletic and gives rise to two well-resolved clades (T and F) that could be recognized as subfamilies. DNA sequence data for Keayodendron and Zimmermanniopsis are presented for the first time. Keayodendron is misplaced in tribe Phyllantheae and belongs to the Bridelia alliance. Zimmermanniopsis is sister to Zimmermannia. Phyllanthus and Cleistanthus are paraphyletic. Savia and Phyllanthus subgenus Kirganelia are not monophyletic.