Phylogeny of Mesoamerican freshwater mussels and a revised tribe‐level classification of the Ambleminae

Evolutionary and ecological hypotheses of the freshwater mussel subfamily Ambleminae are intensely geographically biased—a consequence of the complete exclusion of Mesoamerican taxa in phylogenetic reconstructions of the clade. We set out to integrate a portion of the Mesoamerican freshwater mussel assemblage into existing hypotheses of amblemine classification and evolution by generating a molecular phylogeny that includes four previously unsampled Mesoamerican genera and nine species endemic to that region. Given the traditionally hypothesized affinity to Nearctic mussels and the understanding that classification should reflect common ancestry, we predicted that (a) Mesoamerican genera would be recovered as members of the recognized tribes of the Ambleminae, and (b) genera would be supported as monophyletic. The mutilocus phylogeny (COI + 28S + 16S) reported herein does not fully support either of those hypotheses. Neither Cyrtonaias nor Psorula were supported as monophyletic and we predict several other Mesoamerica genera are also non‐monophyletic. The reconstructed phylogeny recovered four independent lineages of Mesoamerican freshwater mussels and these clades are distributed across the phylogeny of the Ambleminae, including the tribe Quadrulini (Megalonaias), Lampsilini (two lineages: Cyrtonaias explicata/Sphenonaias microdon, and Pachynaias), and a previously unrecognized, exclusively Mesoamerican and Rio Grande clade consisting of the genera Psoronaias, Psorula and Popenaias. The latter clade possesses several morphological characteristics that distinguish it from its sister taxon, tribe Lampsilini, and we recognize this newly identified Mesoamerican clade as a fifth tribe of the Ambleminae attributable to the Popenaiadini Heard & Guckert, 1970. This revised classification more completely recognizes the suprageneric diversity of the Ambleminae.     

Molecular phylogeny of the plant bugs (Heteroptera: Miridae) and the evolution of feeding habits

The first comprehensive cladistic analysis of Miridae, the plant bugs, is presented based on analysis of 3935 base pairs of mitochondrial (16S, COI) and nuclear (18S, 28SD3) DNA for 91 taxa in seven subfamilies. Data were analysed using maximum likelihood (ML), parsimony and Bayesian inference (BI) phylogenetic frameworks. The phylogenetic results are compared with previous hypotheses of higher relationships in the family using alternative hypothesis tests. A Bayesian relaxed molecular clock is used to examine divergence times, and ancestral feeding habits are reconstructed using parsimony and a Bayesian approach. Clades recovered in all analyses are as follows: Cimicomorpha, Miroidea and Miridae; Bryocorinae: Bryocorini; Stenodemini; Mirinae; Deraeocorinae (Clevinemini + Deraeocorini); Cylapinae; Isometopinae; Bryocorinae: Dicyphini; Orthotylini; Phylinae (Phylini + Pilophorini), and Phylinae as sister group to all the remaining mirid taxa. These results are largely congruent with former hypotheses based on morphological data with respect to the monophyly of various subfamilies and tribes; however, our results indicate that the subfamily Bryocorinae is not monophyletic, as the two tribes, Dicypini and Bryocorini, were separated in the phylogenetic results. Divergence time estimates indicate that the radiation of the Miridae began in the Permian; most genus‐level radiations within subfamilies began in the late Cretaceous, probably in response to the angiosperm radiation. Ancestral feeding state reconstructions based on Bayesian and parsimony inference were largely congruent and both reconstructed phytophagy as the ancestral state of the Miridae. Furthermore, the feeding habits of the common ancestors of Mirinae + Deraeocorinae, Bryocorinae + Cylapinae + Isometopinae + Orthotylinae, and the remaining taxa excluding Phylinae, were inferred as phytophagous. Therefore, at least three shifts from phytophagy or polyphagy to predation occurred within the Miridae. Additionally, based on the mirid host‐plant records, we discovered several trends, such as a strong relationship between host‐plant ranges and a facultative feeding habit. © The Willi Hennig Society 2011.     

Phylogeny of the bees of the family Apidae based on larval characters with focus on the origin of cleptoparasitism (Hymenoptera: Apiformes)

Abstract.  Fifty-four genera of the bee family Apidae comprising almost all tribes were analysed based on 77 traditional and one new character of the mature larvae. Nine, especially cleptoparasitic species, were newly added. Analyses were performed by maximum parsimony and Bayesian inference. Trees inferred from the analysis of the complete dataset were rooted by taxa from the families Melittidae and Megachilidae. Unrooted trees inferred from the analysis of the partial dataset (excluding outgroup taxa) are also presented to preclude possible negative effects of the outgroup on the topology of the ingroup. Only the subfamily Nomadinae was statistically well supported. The monophyly of the subfamilies Xylocopinae and Apinae was not topologically recovered. The monophyly of the tribe Tetrapediini was supported, and this tribe was found to be related to xylocopine taxa. At the very least, larval morphology suggests that Tetrapedia is not a member of the subfamily Apinae. Our analyses support the monophyly of the Eucerine line (Emphorini, Eucerini, Exomalopsini, Tapinotaspidini) and of the Apine line (Anthophorini, Apini, Bombini, Centridini, Euglossini, Meliponini). All analyses support the monophyly of totally cleptoparasitic tribes of the subfamily Apinae. We named this group the Melectine line (Ericrocidini, Isepeolini, Melectini, Osirini, Protepeolini, Rhathymini). In previous studies all these cleptoparasitic tribes were considered independent evolutionary lineages. Our results suggest that their similarities with hosts in morphology and pattern are probably the result of convergence and host–parasite co-evolution than phylogenetic affinity. According to the present analysis, the cleptoparasitism has evolved independently only six times within the family Apidae.     

Molecular phylogeny of the fungus gnat family Mycetophilidae (Diptera,Mycetophiliformia)

Abstract A molecular phylogeny of the fungus gnat family Mycetophilidae based on the nuclear 18S, 28S, and the mitochondrial 16S rRNA genes is presented. The total alignment included 58 taxa and 1704 bp. The family was recovered as monophyletic in parsimony and Bayesian analyses. In the Bayesian analysis, Mycetophilinae and its two tribes, Mycetophilini and Exechiini, were monophyletic with good statistical support. The subfamily Mycomyinae was found consistently in a sister‐group relationship to Mycetophilinae. Gnoristinae was rendered paraphyletic, subtending Mycomyinae and Mycetophilinae. Within Gnoristinae, the genera Coelosia Winnertz, Boletina Staeger, Gnoriste Meigen group with Docosia Winnertz, usually considered to be a member of Leiinae. No support was found for the monophyly of the subfamilies Sciophilinae and Leiinae.     

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.     

A molecular phylogeny of Planorboidea (Gastropoda, Pulmonata): insights from enhanced taxon sampling

Planorbid gastropods are the most diverse group of limnic pulmonates, with both discoidal and highspired taxa. Phylogenetic relationships among these genera are confused and controversial. In particular, the monophyly of the limpet‐like taxa (traditionally Ancylidae) is disputed. Even recent molecular studies have concluded that substantially more work is necessary to solve the remaining issues concerning intergeneric phylogenetic relationships and higher taxa systematics. Planorbid snails are of great significance for humans as several members of this group are intermediate hosts of blood flukes (schistosomes) causing a chronic disease, schistosomiasis. We used the two independent molecular markers COI and 18S (concatenated dataset of 2837 nucleotide bp) to infer phylogenetic relationships of 26 genera (27 species) of Planorboidea, represented mostly by type species from mainly topotypical populations. With the majority of the taxa discussed not having been studied previously, this study attempted to test several hypotheses on planorbid phylogenetic relationships using Bayesian inference techniques. The monophyly of Planorboidea (= ‘Ancyloplanorbidae’) is strongly suggested on the basis of our extensive molecular analysis. Besides a distinct Burnupia clade, two major clades were recovered that correspond to family level taxa (traditional Bulinidae and Planorbidae). Considerable rearrangements of suprageneric taxa are evident from the phylogeny inferred. Therefore, the only clades recognized by current classifications and supported by our analysis are Planorbini and Segmentinini. The present study found that Ancylidae as traditionally understood, i.e. covering most freshwater limpet gastropods, is paraphyletic, as the genera of Burnupia and Protancylus have been shown to lie phylogenetically outside the Ancylini. Chromosome numbers and levels of polyploidy are discussed in the light of the new phylogeny. An earlier theory of shell shape evolution, i.e. that of patelliform taxa being most advanced, was not supported by this study; a limpet‐shaped taxon is most basal within Planorboidea. Although many taxa still remain to be studied, our results will hopefully contribute towards a better understanding of this very important group of freshwater organisms. Some taxonomic implications are discussed.     

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.     

Phylogeny and a new tribal classification of the Panicoideae s.l. (Poaceae) based on plastid and nuclear sequence data and structural data

? Premise of the study: The subfamily Panicoideae (Poaceae) encompasses nearly one-third of the diversity of grass species, including important crops such as maize and sugarcane. Previous analyses recovered strong support for a Panicoideae+Centothecoideae lineage within the diverse Panicoideae+Arundinoideae+Chloridoideae+Micrairoideae+Aristidoideae+Danthonioideae (PACMAD) clade, although support for internal relationships was inconsistent. The objectives of this research were to (1) further test the monophyly of each subfamily and previously recovered clades within the Panicoideae+Centothecoideae lineage, (2) establish phylogenetic relationships among these groups, and (3) propose a new tribal classification for this lineage based explicitly on the phylogeny. ? Methods: Maximum parsimony and Bayesian inference analyses of 37 taxa were based on previously published sequences (ndhF and rpl16 intron) and on new plastid and nuclear (rbcL and granule-bound starch synthase I) sequence data as well as structural data. ? Key results. The Panicoideae+Centothecoideae lineage and a majority of the clades identified in previous analyses continue to be robustly supported, but resolution along the backbone of the topology remains elusive. Support for the monophyly of both subfamilies was lacking although support values for some clades increased. The tribes Centotheceae and Arundinelleae were confirmed as polyphyletic. ? Conclusions: Subfamily Centothecoideae is formally submerged into the Panicoideae, and a new tribal classification for the expanded Panicoideae is proposed based explicitly on the phylogeny. This classification includes 12 tribes of which Chasmanthieae and Zeugiteae are segretated from the Centotheceae; Tristachyideae is segregated from Arundinelleae, and a new tribe, Cyperochloeae, is validated to accommodate two isolated genera. A key to the tribes is provided.     

Phylogeny of the cricket subfamily Eneopterinae (Orthoptera, Grylloidea, Eneopteridae) based on four molecular loci and morphology

The phylogenetic relationships of 39 species of Eneopterinae crickets are reconstructed using four molecular markers (16S rRNA, 12S rRNA, cytochrome b, 18S rRNA) and a large morphological data set. Phylogenetic analysis via direct optimisation of DNA sequence data using parsimony as optimality criterion is done for six combinations of weighting parameter sets in a sensitivity analysis. The results are discussed in a twofold purpose: first, in term of significance of the molecular markers for phylogeny reconstruction in Ensifera, as our study represents the first molecular phylogeny performed for this insect suborder at this level of diversity; second, in term of corroboration of a previous phylogeny of Eneopterinae, built on morphological data alone. The four molecular markers all convey phylogenetic signal, although variously distributed on the tree. The monophyly of the subfamily, that of three over five tribes, and of 10 over 13 genera, are recovered. Finally, previous hypotheses on the evolution of acoustic devices and signals in the Eneopterinae clade are briefly tested, and supported, by our new data set.     

Phylogeny of the plant bug subfamily Mirinae (Hemiptera: Heteroptera: Cimicomorpha: Miridae) based on total evidence analysis

The first comprehensive phylogenetic analyses of the most diverse subfamily of plant bugs, Mirinae, is presented in this study, for 110 representative taxa based on total evidence analysis. A total of 85 morphological characters and 3898 bp of mitochondrial (16S, COI) and nuclear (18S, 28S) sequences were analysed for each partitioned and combined dataset based on parsimony, maximum likelihood and Bayesian inference. Major results obtained in this study include monophyly of the tribe Mecistoscelini. The largest tribe, Mirini, was recovered as polyphyletic, and Stenodemini was recovered as paraphyletic. The clade of Stenodemini + Mecistoscelini is the sister group of the remaining Mirinae. The monophyly of two complexes composed of superficially similar genera were tested; the Lygus complex was recovered as nonmonophyletic, and the Adelphocoris–Creontiades–Megacoelum complex was confirmed to be monophyletic. The generic relationships of the main clades within each tribe based on the phylogeny, as well as their supported morphological characters, are discussed.     

Molecular evidence for the phylogeny of Australian gekkonoid lizards

Partial sequences of the mitochondrial 12S rRNA and nuclear c-mos genes were determined for 12 species of gekkonoid lizards representing the four major taxa of the Australian region, the Diplodactyhni and Carphodactylini (forming the subfamily Diplodactylinae), the Pygopodidae and the Gekkoninae. One further species represented a non-Australian gekkonoid lineage, the Eublepharinae. The combined sequence data were used to reconstruct the underlying molecular phylogeny. We used the molecular phylogeny to test the monophyly of the diplodactyline tribes and conflicting hypotheses of relationships of die pygopods and of the genus Oedura. Monophyly of the Diplodactylinae is supported, while pygopods form a monophyletic sister lineage to all Diplodactylinae. The molecular data support the monophyly of the Diplodactyhni, with Oedura firmly placed as a diplodactylin. Monophyly of the Carphodactylini is not supported. The four carphodactylin genera form a paraphyletic cluster at the base of the Diplodactyhni. Pygopods are nested within the traditional Gekkonidae and pygopods plus diplodactylines form a well-supported monophyletic group with respect to the remaining gekkonoids, the gekkonines and eublepharines.     

Several steps of lateral gene transfer followed by events of ‘birth-and-death’ evolution shaped a fungal sorbicillinoid biosynthetic gene cluster

Dung beetles (subfamily Scarabaeinae) are popular model organisms in ecology and developmental biology, and for the last two decades they have experienced a systematics renaissance with the adoption of modern phylogenetic approaches. Within this period 16 key phylogenies and numerous additional studies with limited scope have been published, but higher-level relationships of this pivotal group of beetles remain contentious and current classifications contain many unnatural groupings. The present study provides a robust phylogenetic framework and a revised classification of dung beetles. We assembled the so far largest molecular dataset for dung beetles using sequences of 8 gene regions and 547 terminals including the outgroup taxa. This dataset was analyzed using Bayesian, maximum likelihood and parsimony approaches. In order to test the sensitivity of results to different analytical treatments, we evaluated alternative partitioning schemes based on secondary structure, domains and codon position. We assessed substitution models adequacy using Bayesian framework and used these results to exclude partitions where substitution models did not adequately depict the processes that generated the data. We show that exclusion of partitions that failed the model adequacy evaluation has a potential to improve phylogenetic inference, but efficient implementation of this approach on large datasets is problematic and awaits development of new computationally advanced software. In the class Insecta it is uncommon for the results of molecular phylogenetic analysis to lead to substantial changes in classification. However, the results presented here are congruent with recent morphological studies and support the largest change in dung beetle systematics for the last 50 years. Here we propose the revision of the concepts for the tribes Deltochilini (Canthonini), Dichotomiini and Coprini; additionally, we redefine the tribe Sisyphini. We provide and illustrate synapomorphies and diagnostic characters supporting the new concepts to facilitate diagnosability of the redefined tribes. As a result of the proposed changes a large number of genera previously assigned to these tribes are now left outside the redefined tribes and are treated as incertae sedis. The present study redefines dung beetles classification and gives new insight into their phylogeny. It has broad implications for the systematics as well as for various ecological and evolutionary analyses in dung beetles.     

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.     

Molecular dating of the diversification of Phyllostominae bats based on nuclear and mitochondrial DNA sequences

Times of divergence among the three tribes included within the subfamily Phyllostominae were estimated using a Bayesian approach to infer dates of divergence based on mitochondrial and nuclear sequence data. The subfamily Phyllostominae is particularly attractive for such analysis, as it is one of the few groups of bats to have fossil specimens. Our molecular time analyses suggest that diversification among tribes and genera of phyllostomine bats occurred during the Early to Mid-Miocene, and was coincident with diversification events in two co distributed taxa: Caviomorph rodents and New World monkeys.     

Higher level phylogeny of Satyrinae butterflies (Lepidoptera: Nymphalidae) based on DNA sequence data

We have inferred the first empirically supported hypothesis of relationships for the cosmopolitan butterfly subfamily Satyrinae. We used 3090 base pairs of DNA from the mitochondrial gene COI and the nuclear genes EF-1alpha and wingless for 165 Satyrinae taxa representing 4 tribes and 15 subtribes, and 26 outgroups, in order to test the monophyly of the subfamily and elucidate phylogenetic relationships of its major lineages. In a combined analysis, the three gene regions supported an almost fully resolved topology, which recovered Satyrinae as polyphyletic, and revealed that the current classification of suprageneric taxa within the subfamily is comprised almost completely of unnatural assemblages. The most noteworthy findings are that Manataria is closely related to Melanitini; Palaeonympha belongs to Euptychiina; Oressinoma, Orsotriaena and Coenonympha group with the Hypocystina; Miller's (1968). Parargina is polyphyletic and its components group with multiple distantly related lineages; and the subtribes Elymniina and Zetherina fall outside the Satyrinae. The three gene regions used in a combined analysis prove to be very effective in resolving relationships of Satyrinae at the subtribal and tribal levels. Further sampling of the taxa closely related to Satyrinae, as well as more extensive sampling of genera within the tribes and subtribes for this group will be critical to test the monophyly of the subfamily and establish a stronger basis for future biogeographical and evolutionary studies.     

Molecular phylogeny of the horse flies: a framework for renewing tabanid taxonomy

Horse flies, family Tabanidae, are the most diverse family‐level clade of bloodsucking insects, but their phylogeny has never been thoroughly explored using molecular data. Most adult female Tabanidae feed on nectar and on the blood of various mammals. Traditional horse fly classification tends towards large heterogeneous taxa, which impede much‐needed taxonomic work. To guide renewed efforts in the systematics of horse flies and their relatives, we assembled a dataset of 110 exemplar species using nucleotide data from four genes—mitochondrial CO1, and nuclear 28S, CAD and AATS. All commonly recognized tribes in Tabanidae are represented, along with outgroups in Tabanomorpha. The phylogeny is reconstructed using Bayesian inference, and divergence times are estimated using Bayesian relaxed clock methods with time constraints from tabanid fossils. Our results show Athericidae strongly supported as the lineage most closely related to Tabanidae, and Pangoniinae and Tabaninae as monophyletic lineages. However, Chrysopsinae is nonmonophyletic, with strong support for both a nonmonophyletic Bouvieromyiini and for Rhinomyzini as sister to Tabaninae. Only the tribes Philolichini, Chrysopsini, Rhinomyzini and Haematopotini are recovered as monophyletic, although Scionini is monophyletic with exclusion of the peculiar genus Goniops Aldrich. Mycteromyia Philippi and Adersia Austen, two enigmatic genera sometimes placed in separate family‐level groups, are recovered inside Pangoniini and Chrysopsini, respectively. Several species‐rich genera are not recovered as monophyletic, including Esenbeckia Rondani, Silvius Meigen, Dasybasis Macquart and Tabanus L. Tabanidae likely originated in the Cretaceous, and all major extant groups were present by the early Palaeogene. This newly revised phylogenetic framework for Tabanidae forms the basis for a new assessment of tabanid diversification and provides context for understanding the evolution of trophic specialization in horse flies.     

Molecular phylogenetics of Braconidae (Hymenoptera: Ichneumonoidea), based on multiple nuclear genes,and implications for classification

This study examined subfamilial relationships within Braconidae, using 4 kb of sequence data for 139 taxa. Genetic sampling included previously used markers for phylogenetic studies of Braconidae (28S and 18S rDNA) as well as new nuclear protein‐coding genes (CAD and ACC). Maximum likelihood and Bayesian inference of the concatenated dataset recovered a robust phylogeny, particularly for early divergences within the family. This study focused primarily on non‐cyclostome subfamilies, but the monophyly of the cyclostome complex was strongly supported. There was evidence supporting an independent clade, termed the aphidioid complex, as sister to the cyclostome complex of subfamilies. Maxfischeria was removed from Helconinae and placed within its own subfamily within the aphidioid complex. Most relationships within the cyclostome complex were poorly supported, probably because of lower taxonomic sampling within this group. Similar to other studies, there was strong support for the alysioid subcomplex containing Gnamptodontinae, Alysiinae, Opiinae and Exothecinae. Cenocoeliinae was recovered as sister to all other subfamilies within the euphoroid complex. Planitorus and Mannokeraia, previously placed in Betylobraconinae and Masoninae, respectively, were moved to the Euphorinae, and may share a close affiliation with Neoneurinae. Neoneurinae and Ecnomiinae were placed as tribes within Euphorinae. A sister relationship between the microgastroid and sigalphoid complexes was also recovered. The helconoid complex included a well‐supported lineage that is parasitic on lepidopteran larvae (macrocentroid subcomplex). Helconini was raised to subfamily status, and was recovered as sister to the macrocentroid subcomplex. Blacinae was demoted to tribal status and placed within the newly circumscribed subfamily Brachistinae, which also contains the tribes Diospilini, Brulleiini and Brachistini, all formerly in Helconinae.     

Molecular phylogeny and divergence times of Hormaphidinae (Hemiptera: Aphididae) indicate Late Cretaceous tribal diversification

The aphid subfamily Hormaphidinae is a good candidate for the study of the evolution of insect – plant relationships. Most hormaphidine species depend on woody primary host plants and woody or herbaceous secondary host plants, and represent high host specificity, especially to their primary hosts. No detailed molecular phylogeny of Hormaphidinae has been reported, and the taxonomic positions of some taxa in this group remain unclear. To reconstruct major phylogenetic relationships and to understand the evolution of host association patterns for major lineages, we present the first detailed molecular phylogeny of Hormaphidinae, as inferred from nuclear and mitochondrial DNA sequences, using maximum parsimony, maximum likelihood, and Bayesian methods. The monophyly of Hormaphidinae and its three traditional tribes was supported, and a sister relationship between Hormaphidini and Nipponaphidini was suggested. Most inner relationships within tribes were also supported, and some novel relationships were revealed. Two subtribes of Cerataphidini are proposed. Divergence times estimated using a Bayesian approach indicate that tribal diversifications occurred during the Late Cretaceous and were coincident with the appearance of their primary host plants. The current pattern of secondary host association for the three tribes may have evolved in different time ranges. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 165 , 73–87.     

Molecular phylogeny of the beetle tribe Oxypodini (Coleoptera: Staphylinidae: Aleocharinae)

This is the first study to comprehensively address the phylogeny of the tribe Oxypodini Thomson and its phylogenetic relationships to other tribes within the staphylinid subfamily Aleocharinae. Using the hitherto largest molecular dataset of Aleocharinae comprising of 4599 bp for representatives of 22 tribes, the Oxypodini are recovered as non‐monophyletic. Members of the tribe belong to three distantly related lineages within the Aleocharinae: (i) the Amarochara group as sister clade to the tribe Aleocharini, (ii) the subtribe Tachyusina within a clade that also includes the tribes Athetini and Hygronomini, (iii) all other Oxypodini in a clade that also includes the tribes Placusini, Hoplandriini and Liparocephalini. Based on the inferred phylogeny, five subtribes of the Oxypodini are recognized: Dinardina Mulsant & Rey, Meoticina Seevers, Microglottina Fenyes, Oxypodina Thomson and Phloeoporina Thomson. The following changes in the classification of the Aleocharinae are proposed: (i) Amarochara Thomson is removed from the Oxypodini and placed in the tribe Aleocharini; (ii) the subtribe Taxicerina Lohse of the Athetini is reinstated as tribe Taxicerini to include Discerota Mulsant & Rey, Halobrecta Thomson (both removed from the Oxypodini) and Taxicera Mulsant & Rey; (iii) the subtribe Tachyusina Thomson is excluded from the Oxypodini and provisionally treated as tribe Tachyusini; (iv) the oxypodine subtribe name Blepharhymenina Klimaszewski & Peck is placed in synonymy with the subtribe name Dinardina Mulsant & Rey.