Phylogenetic analysis of mealybugs (Hemiptera: Coccoidea: Pseudococcidae) based on DNA sequences from three nuclear genes, and a review of the higher classification

Abstract. Mealybugs (Hemiptera: Pseudococcidae) are small, plant‐sucking insects which comprise the second largest family of scale insects (Coccoidea). Relationships among many pseudococcid genera are poorly known and there is no stable higher level classification. Here we review previous hypotheses on relationships and classification and present the first comprehensive phylogenetic study of the Pseudococcidae based on analysis of nucleotide sequence data. We used three nuclear genes, comprising two noncontiguous fragments of elongation factor 1α (EF‐1α 5′ and EF‐1α 3′), fragments of the D2 and D10 expansion regions of the large subunit ribosomal DNA gene (28S), and a region of the small subunit ribosomal DNA gene (18S). We sampled sixty‐four species of mealybug belonging to thirty‐five genera and representing each of the five subfamilies which had been recognized previously, and included four species of Puto (Putoidae) and one species each of Aclerda (Aclerdidae) and Icerya (Margarodidae), using Icerya as the most distant outgroup. A combined analysis of all data found three major clades of mealybugs which we equate to the subfamilies Pseudococcinae, Phenacoccinae and Rhizoecinae. Within Pseudococcinae, we recognize the tribes Pseudococcini (for Pseudococcus, Dysmicoccus, Trionymus and a few smaller genera), Planococcini (for Planococcus and possibly Planococcoides) and Trabutinini (represented by a diverse range of genera, including Amonostherium, Antonina, Balanococcus, Nipaecoccus and non‐African Paracoccus), as well as the Ferrisia group (for Ferrisia and Anisococcus), some ungrouped African taxa (Grewiacoccus, Paracoccus, Paraputo and Vryburgia), Chaetococcus bambusae and Maconellicoccus. The ‘legless’ mealybugs Antonina and Chaetococcus were not closely related and thus we confirmed that the Sphaerococcinae as presently constituted is polyphyletic. In our analyses, the subfamily Phenacoccinae was represented by just Phenacoccus and Heliococcus. The hypogeic mealybugs of the Rhizoecinae usually formed a monophyletic group sister to all other taxa. Our molecular data also suggest that the genera Pseudococcus, Dysmicoccus, Nipaecoccus and Paracoccus are not monophyletic (probably polyphyletic) and that Phenacoccus may be paraphyletic, but further sampling of species and genes is required. We compare our phylogenetic results with published information on the intracellular endosymbionts of mealybugs and hypothesize that the subfamily Pseudococcinae may be characterized by the possession of β‐Proteobacteria (primary endosymbionts) capable of intracellular symbiosis with γ‐Proteobacteria (secondary endosymbionts). Furthermore, our data suggest that the identities of the secondary endosymbionts may be useful in inferring mealybug relationships. Finally, cloning polymerase chain reaction products showed that paralogous copies of EF‐1α were present in at least three taxa. Unlike the situation in Apis and Drosophila, the paralogues could not be distinguished by either the presence/absence or position of an intron.     

A molecular phylogeny of the tribe Aphidini (Insecta: Hemiptera: Aphididae) based on the mitochondrial tRNA/COII, 12S/16S and the nuclear EF1α genes

Abstract A phylogeny of the tribe Aphidini (Hemiptera: Aphididae) was reconstructed from three gene fragments: two mitochondrial regions, partial tRNA‐leucine + cytochrome oxidase II (tRNA/COII), partial 12S rRNA + tRNA‐valine + 16S rRNA (12S/16S) and one nuclear gene, the elongation factor‐1 alpha (EF1α). Bayesian phylogenetic (BP) analyses were performed on each individual dataset of tRNA/COII, 12S/16S and EF1α, and maximum parsimony (MP), Bremer support test, maximum likelihood (ML) and BP analysis were performed on the combined dataset. After comparing our molecular phylogenetic results with the classic classification based on morphological and ecological data, we analysed three main issues: the monophyletic relationships among tribes and subtribes, the validities of the latest taxonomic positions of genera and species and the status of certain Aphis species groups. Our results indicate that 36 of the species analysed, with the exception of Cryptosiphum artemisiae, are clustered within the clade of Aphidini. Also, the 28 species representative of the subtribe Aphidina were separated from the eight species representative of Rhopalosiphina; each monophyletic subtribe was supported by significant P‐values in the combined analysis. According to our results, Cryptosiphum should be moved to Macrosiphini because it is more closely related to the genera Lipaphis and Brevicoryne. The genus Toxoptera was recovered as non‐monophyletic. In Rhopalosiphina, three genera, Hyalopterus, Rhopalosiphum and Schizaphis, were relatively closer to each other than to the genus Melanaphis. In the relationships between species‐groups among Aphis, most species were separated into two main lineages; the fabae group seemed to be more closely related to the spiraecola and craccivora group rather than to the gossypii group.     

Phylogenetics, biogeography and classification of, and character evolution in, gamebirds (Aves: Galliformes): effects of character exclusion, data partitioning and missing data

The phylogenetic relationships, biogeography and classification of, and morpho‐behavioral (M/B) evolution in, gamebirds (Aves: Galliformes) are investigated. In‐group taxa (rooted on representatives of the Anseriformes) include 158 species representing all suprageneric galliform taxa and 65 genera. The characters include 102 M/B attributes and 4452 nucleic acid base pairs from mitochondrial cytochrome b (CYT B), NADH dehydrogenase subunit 2 (ND2), 12S ribosomal DNA (12S) and control region (CR), and nuclear ovomucoid intron G (OVO‐G). Analysis of the combined character data set yielded a single, completely resolved cladogram that had the highest levels of jackknife support, which suggests a need for a revised classification for the phasianine galliforms. Adding 102 M/B characters to the combined CYT B and ND2 partitions (2184 characters) decisively overturns the topology suggested by analysis of the two mtDNA partitions alone, refuting the view that M/B characters should be excluded from phylogenetic analyses because of their relatively small number and putative character state ambiguity. Exclusion of the OVO‐G partition (with > 70% missing data) from the combined data set had no effect on cladistic structure, but slightly lowered jackknife support at several nodes. Exclusion of third positions of codons in an analysis of a CYT B + ND2 partition resulted in a massive loss of resolution and support, and even failed to recover the monophyly of the Galliformes with jackknife support. A combined analysis of putatively less informative, “non‐coding” characters (CYT B/ND2 third position sites + CR +12S + OVO‐G sequences) yielded a highly resolved consensus cladogram congruent with the combined‐evidence cladogram. Traditionally recognized suprageneric galliform taxa emerging in the combined cladogram are: the families Megapodiidae (megapodes), Cracidae (cracids), Numididae (guineafowls), Odontophoridae (New World quails) and Phasianidae (pheasants, pavonines, partridges, quails, francolins, spurfowls and grouse) and the subfamilies Cracinae (curassows, chachalacas and the horned guan), Penelopinae (remaining guans), Pavoninae sensu lato (peafowls, peacock pheasants and argus pheasants), Tetraoninae (grouse) and Phasianinae (pheasants minus Gallus). The monophyly of some traditional groupings (e.g., the perdicinae: partridges/quails/francolins) is rejected decisively, contrasted by the emergence of other unexpected groupings. The most remarkable phylogenetic results are the placement of endemic African galliforms as sisters to geographically far‐distant taxa in Asia and the Americas. Biogeographically, the combined‐data cladogram supports the hypothesis that basal lineages of galliforms diverged prior to the Cretaceous/Tertiary (K‐T) Event and that the subsequent cladogenesis was influenced by the break‐up of Gondwana. The evolution of gamebirds in Africa, Asia and the Americas has a far more complicated historical biogeography than suggested to date. With regard to character evolution: spurs appear to have evolved at least twice within the Galliformes; a relatively large number of tail feathers (≥ 14) at least three times; polygyny at least twice; and sexual dimorphism many times. © The Willi Hennig Society 2006.     

A phylogenetic analysis of relationships among genera of Conopidae (Diptera) based on molecular and morphological data

Members of the family Conopidae (Diptera) have been the focus of little targeted phylogenetic research. The most comprehensive test of phylogenetic support for the present subfamily classification of Conopidae is presented here using 66 specimens, including 59 species of Conopidae and seven outgroup taxa. Relationships among subfamily clades are also explored. A total of 6824 bp of DNA sequence data from five gene regions (12S ribosomal DNA, cytochrome c oxidase subunit I, cytochrome b, 28S ribosomal DNA and alanyl‐tRNA synthetase) are combined with 111 morphological characters in a combined analysis using both parsimony and Bayesian methods. Parsimony analysis recovers three shortest trees. Bayesian analysis recovers a nearly identical tree. Five monophyletic subfamilies of Conopidae are recovered. The rarely acknowledged Zodioninae is restored, including the genera Zodion and Parazodion. The genus Sicus is removed from Myopinae. Morphological synapomorphies are discussed for each subfamily and inter‐subfamily clade, including a comprehensive review of the character interpretaions of previous authors. Included are detailed comparative illustrations of male and female genitalia of representatives of all five subfamilies with new morphological interpretation.     

Phylogeny and taxonomy of the Prenolepis genus‐group of ants (Hymenoptera: Formicidae)

Abstract. We investigated the phylogeny and taxonomy of the Prenolepis genus‐group, a clade of ants we define within the subfamily Formicinae comprising the genera Euprenolepis, Nylanderia, gen. rev. , Paraparatrechina, gen. rev. & stat. nov. , Paratrechina, Prenolepis and Pseudolasius. We inferred a phylogeny of the Prenolepis genus‐group using DNA sequence data from five genes (CAD, EF1αF1, EF1αF2, wingless and COI) sampled from 50 taxa. Based on the results of this phylogeny the taxonomy of the Prenolepis genus‐group was re‐examined. Paratrechina (broad sense) species segregated into three distinct, robust clades. Paratrechina longicornis represents a distinct lineage, a result consistent with morphological evidence; because this is the type species for the genus, Paratrechina is redefined as a monotypic genus. Two formerly synonymized subgenera, Nylanderia and Paraparatrechina, are raised to generic status in order to provide names for the other two clades. The majority of taxa formerly placed in Paratrechina, 133 species and subspecies, are transferred to Nylanderia, and 28 species and subspecies are transferred to Paraparatrechina. In addition, two species are transferred from Pseudolasius to Paraparatrechina and one species of Pseudolasius is transferred to Nylanderia. A morphological diagnosis for the worker caste of all six genera is provided, with a discussion of the morphological characters used to define each genus. Two genera, Prenolepis and Pseudolasius, were not recovered as monophyletic by the molecular data, and the implications of this result are discussed. A worker‐based key to the genera of the Prenolepis genus‐group is provided.     

Large‐scale molecular phylogeny of Cryptorhynchinae (Coleoptera,Curculionidae) from multiple genes suggests American origin and later Australian radiation

The monophyly of the highly diverse weevil subfamily Cryptorhynchinae is tested with a dataset of 203 taxa representing 159 genera of Curculionoidea, 105 of them Cryptorhynchinae s.l. We construct a phylogeny based on an alignment of 5523 bp, consisting of fragments from two mitochondrial genes (two fragments of COI, 16S) and seven nuclear genes (ArgK, CAD, EF1α, enolase, H4, 18S, 28S). Analyses of maximum likelihood and Bayes inference recovered largely congruent results. Groups with different morphology of the rostral furrow (e.g. Aedemonini, Camptorhinini, Cryptorhynchini, Ithyporini) are not closely related to each other. However, most taxa with a mesosternal receptacle are monophyletic and here defined as Cryptorhynchinae s.s., comprising Cryptorhynchini, Gasterocercini, Torneumatini and Psepholacini, but also Arachnopodini and Idopelma Faust. The genus Phyrdenus LeConte is excluded from Cryptorhynchinae and transferred to Conotrachelini of Molytinae. Thus defined, the group still comprises several thousand species with centres of its diversity in South America and Australia. The early lineages we find in America and the Palearctic, while the extremely diverse faunas of Australia and neighbouring islands mainly belong to a more recent, species‐rich radiation. This also includes a clade comprising the majority of litter‐inhabiting species of New Zealand and the genus Miocalles Pascoe. Flightlessness was attained repeatedly and resulted in convergent evolution of a similar habitus in different zoogeographic regions, mainly exhibited by the polyphyletic genus Acalles Schoenherr.     

A model for the interaction of the G3‐subdomain of Geobacillus stearothermophilus IF2 with the 30S ribosomal subunit

Bacterial translation initiation factor IF2 complexed with GTP binds to the 30S ribosomal subunit, promotes ribosomal binding of fMet‐tRNA, and favors the joining of the small and large ribosomal subunits yielding a 70S initiation complex ready to enter the translation elongation phase. Within the IF2 molecule subdomain G3, which is believed to play an important role in the IF2‐30S interaction, is positioned between the GTP‐binding G2 and the fMet‐tRNA binding C‐terminal subdomains. In this study the solution structure of subdomain G3 of Geobacillus stearothermophilus IF2 has been elucidated. G3 forms a core structure consisting of two β‐sheets with each four anti‐parallel strands, followed by a C‐terminal α‐helix. In line with its role as linker between G3 and subdomain C1, this helix has no well‐defined orientation but is endowed with a dynamic nature. The structure of the G3 core is that of a typical OB‐fold module, similar to that of the corresponding subdomain of Thermus thermophilus IF2, and to that of other known RNA‐binding modules such as IF2‐C2, IF1 and subdomains II of elongation factors EF‐Tu and EF‐G. Structural comparisons have resulted in a model that describes the interaction between IF2‐G3 and the 30S ribosomal subunit.     

Phylogeny of Indo‐West Pacific pontoniine shrimps (Crustacea: Decapoda: Caridea) based on multilocus analysis

The phylogenetic relationships and evolutionary processes within the subfamily Pontoniinae, a speciose group of shrimps with diverse lifestyles (free living, semi‐symbiotic and symbiotic) inhabiting the coral reefs of tropical oceans, are an interesting and undeveloped subject of study. In this work, two mitochondrial ribosomal genes (12S rRNA and 16S rRNA) and two protein‐coding nuclear genes (Histone 3 and the sodium–potassium ATPase α‐subunit) were employed to reconstruct the phylogenetic relationships of 42 genera and 101 species within Pontoniinae. Compared to previous studies, ten additional genera were shown to be monophyletic groups, and the genera Dactylonia and Periclimenaeus were shown to be paraphyletic. The shallow‐water crinoid‐associated pontoniines were divided into several groups which were mostly consistent with the morphological analysis. The studied bivalve‐associated taxa exhibited ancestries that were traceable to different lineages, and two groups could be distinguished: Anchiopontonia + Conchodytes and Anchistus. The similar situation occurred in other echinoderm‐associated pontoniines. These results suggest that pontoniines sharing the same hosts may have different evolutionary origins resulting from multiple intrusions of their hosts by morphologically plastic ancestral groups.     

Major lineages of Nolidae (Lepidoptera,Noctuoidea) elucidated by molecular phylogenetics

To elucidate the evolutionary relationships of the major lineages within the moth family Nolidae, we analysed a molecular dataset comprising eight independent gene regions (6.4 kbp), cytochrome c oxidase subunit I (COI) from the mitochondrial genome, and elongation factor‐1α (EF‐1α), ribosomal protein S5 (RpS5), carbamoylphosphate synthase domain protein (CAD), cytosolic malate dehydrogenase (MDH), glyceraldehyde‐3‐phosphate dehydrogenase (GAPDH), isocitrate dehydrogenase (IDH) and wingless genes from the nuclear genome, using parsimony and model‐based evolutionary methods (maximum likelihood and Bayesian inference). Our analyses revealed a well‐resolved phylogenetic hypothesis, again recovering the six previously recognized families within Noctuoidea (i.e. Oenosandridae, Notodontidae, Euteliidae, Erebidae, Nolidae and Noctuidae), and monophyly of the quadrifid Noctuoidea (i.e. Euteliidae, Erebidae, Nolidae and Noctuidae). The family Nolidae is diagnosed and characterized by two synapomorphies from morphology: construction of a ridged boat‐shaped cocoon that bears a vertical exit slit at one end; and two other morphological character states: elongation of the forewing retinaculum into a bar‐like or digitate condition and possession of a postpiracular counter‐tympanal hood. We present a new phylogenetic hypothesis for Nolidae consisting of eight strongly supported subfamilies, two of which are erected here: Diphtherinae, Risobinae, Collomeninae subfam. nov., Beaninae subfam. nov., Eligminae, Westermanniinae, Nolinae and Chloephorinae. Where we are able, each monophyletic lineage is diagnosed by morphological autapomorphies and within each subfamily, monophyletic tribes and subtribes are circumscribed, most of which are also diagnosable by morphological apomorphies. We also describe two new taxa: Gelastocerini trib. nov. and Etannina subtrib. nov. The Neotropical subfamily Diphtherinae, here newly circumscribed, is considered to be the plesiomorphic sister lineage to the rest of Nolidae. Diphtherinae are characterized by loss of the proximal pair of metatibial spurs in males and by the presence of a frontal tubercle, which is presumably associated with a derived strategy of emergence from the cocoon.     

The taxonomic status of Digramma (Pseudophyllidea: Ligulidae) inferred from DNA sequences

The traditional classification of the ligulid tapeworms into 2 genera, Ligula Bloch, 1782 and Digramma Cholodkovsky, 1914, remains controversial. Molecular data of sequences for the 5' end of the nuclear 28S ribosomal ribonucleic acid (rRNA) gene, the mitochondrial cytochrome c oxidase subunit I (COI) gene, and the nicotinamide adenine dinucleotide dehydrogenase subunit 1 (ND1) gene, as well as the first internal transcribed spacer (ITS1) of the nuclear ribosomal deoxyribonucleic acid (DNA), were used to characterize Digramma and to investigate its relationship with Ligula. Digramma spp. exhibited identical sequences with Ligula intestinalis both in the 28S rRNA and the COI gene and differed from L. intestinalis by 0.7% in the ITS1 region and 7.4% in the ND1 gene, respectively. A high degree of genetic conservation within 28S ribosomal DNA, COI, ITS1, and even ND1 genes, was found in Ligula and Digramma. The low genetic divergence in the 4 genes between Ligula and Digramma indicates that Digramma is probably not an independent genus. Therefore, it is proposed that Ligula and Digramma should be considered as 2 species within the genus Ligula and the tapeworms of Digramma collected from diverse localities in China belong to the same species. The present study also suggests that ITS1 and ND1 sequences can act as useful genetic markers to distinguish Ligula and Digramma.     

Molecular phylogeny and systematics of the Pieridae (Lepidoptera: Papilionoidea): higher classification and biogeography

The systematic relationships of the butterfly family Pieridae are poorly understood. Much of our current understanding is based primarily on detailed morphological observations made 50–70 years ago. However, the family and its putative four subfamilies and two tribes, have rarely been subjected to rigorous phylogenetic analysis. Here we present results based on an analysis of molecular characters used to reconstruct the phylogeny of the Pieridae in order to infer higher‐level classification above the generic level and patterns of historical biogeography. Our sample contained 90 taxa representing 74 genera and six subgenera, or 89% of all genera recognized in the family. Three complementary approaches were employed: (1) a combined analysis of a 30 taxon subset for sequences from four gene regions, including elongation factor‐1 alpha (EF‐1α), wingless, cytochrome oxidase subunit I (COI), and 28S (3675 bp, 1031 parsimony‐informative characters), mainly to establish higher‐level relationships, (2) a single‐gene analysis of the 90 taxon data set for sequences from EF‐1α (1066 bp, 364 parsimony‐informative characters), mainly to establish lower‐level relationships, and (3) an all available data analysis of the entire data set for sequences from the four genes, to recover both deep and shallow nodes. Analyses using maximum parsimony, maximum likelihood and Bayesian inference provided similar results. All supported monophyly for the four subfamilies but not for the two tribes, with the Anthocharidini polyphyletic and the Pierini paraphyletic. The combined and all available data analyses support the following relationships among the subfamilies: ((Pseudopontiinae + Dismorphiinae) + (Coliadinae + Pierinae)), corroborating Ehrlich’s 1958 phenetic hypothesis. On the basis of these analyses, and additional morphological and life history evidence, we propose a reclassification of the subfamily Pierinae into two tribes (Anthocharidini s.s., Pierini s.s.) and two informal groups (Colotis group, Leptosia), with the tribe Pierini s.s. subdivided into three subtribes (Appiadina, Pierina, Aporiina) and three genera (Elodina, Dixeia, Belenois) of uncertain status (incertae sedis). The combined and all available data analyses support the following relationships among the Pierinae: (Colotis group + Anthocharidini s.s. + Leptosia + (Elodina + ((Dixeia + Belenois) + Appiadina + Pierina + Aporiina))). Application of a molecular clock calibrated using fossil evidence and semiparametric rate smoothing suggests that divergence between the Pierina and Aporiina occurred no later than the Palaeocene (> 60 Myr). The minimum estimate for the age of the crown‐group of the Pieridae was 112–82 Myr, with a mean of 95 Myr. A historical biogeographical hypothesis is proposed to explain the present‐day distribution of the clade Pseudopontiinae + Dismorphiinae, which argues for an origin of the two subfamilies in western Gondwana (Africa + South America) during the Late Cretaceous. © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society, 2006, 147 , 239–275.     

Molecular and chemical characters to evaluate species status of two cuckoo bumblebees: Bombus barbutellus and Bombus maxillosus (Hymenoptera,Apidae, Bombini)

Many methods, based on morphological, molecular or chemical characters, have been used to address the question of species taxonomic status. Integrative taxonomy aims to define stronger supported taxonomic hypotheses by considering complementary datasets from different characters. By following an integrative approach, the present study includes molecular, chemical and morphological criteria to establish the taxonomic status of two rare and doubtful cuckoo bumblebee taxa: Bombus (Psithyrus) barbutellus and Bombus (Psithyrus) maxillosus. These two sympatric taxa are discriminated by few morphological criteria (mainly wing darkness and hair length). We used these morphological character diagnoses to establish an a priori status of our samples (23 specimens). We developed a combined molecular dataset from one nuclear gene, elongation factor 1α (EF‐1α), and one mitochondrial gene, cytochrome c oxidase subunit I (COI), spanning 1623 bp, and a chemical dataset of sexual marking pheromones (73 compounds). The molecular data were subjected to maximum‐likelihood and Bayesian phylogenetic inference under partitioned model and maximum parsimony. The chemical data were analysed by clustering and the two‐group k‐means method to test divergences between the two species. The resulting phylogenetic trees show no consistent divergence between the two taxa. Moreover, we found no divergence in the sexual marking pheromones in the clustering and two‐group k‐means analyses. These converging results support the conspecificity of both taxa. Nonetheless, our determinations using the traditional morphological criteria separated our samples into two taxa. We conclude that the morphological criteria seem to relate to intraspecific variations: B. maxillosus is regarded as a syn.n. of B. barbutellus.     

Molecular phylogeny of Geoplaninae (Platyhelminthes) challenges current classification: proposal of taxonomic actions

Despite likely being the most diverse group within the Tricladida, the systematics of land planarians (Geoplanidae) has received minor attention. The most species‐rich ingroup, the subfamily Geoplaninae, is restricted to the Neotropics. The systematics of Geoplaninae remains uncertain. Unique features supporting the genera are scanty; moreover, parts of the known species have been poorly described, making comparative studies difficult. Likewise the evolutionary relationships among land planarians remain insufficiently understood. In the present study, a phylogenetic hypothesis for selected taxa of Geoplaninae based on the molecular data is presented and discussed in the light of morphological features. Our phylogenetic inference is based on the fragments of three nuclear regions (18S, 28S rDNA and EF‐1α) and a mitochondrial marker (cytochrome oxidase I) for which we considered three optimality criteria (parsimony, maximum likelihood and Bayesian inference). Although our data provide little support for most basal nodes, our phylogenetic trees show a number of well‐supported clades, unveiling morphologically homogeneous groups. According to these results, we propose to separate Geoplana into Barreirana (formerly considered a subgenus), Cratera gen. n., Imbira gen. n., Matuxia gen. n., Obama gen. n. and Paraba gen. n., emend the diagnoses of Barreirana, Geoplana, Notogynaphallia, Pasipha and Xerapoa and review the classification of the species within these genera. For Geoplana goetschi sensu Marcus, (1951), a new name is proposed.     

Further insights into the highly derived haptorids (Ciliophora,Litostomatea): Phylogeny based on multigene data

Evolutionary relationships of taxa within the ciliate subclass Haptoria are poorly understood. In this study, we broaden the taxon sampling by adding 14 small subunit ribosomal RNA gene sequences, 13 large subunit ribosomal RNA gene sequences and 13 ITS1‐5.8S‐ITS2 gene sequences of haptorians. This includes the first molecular data from two genera, Pseudotrachelocerca Song, 1990, and Foissnerides Song & Wilbert, 1989. Phylogenies inferred from the three individual genes and concatenated data sets show that: (i) the subclass Haptoria could be a multiphyletic complex with about up to four main clades while “interrupted” by some intermingled with the related subclasses Rhynchostomatia, Trichostomatia and some incertae sedis; (ii) the genus Pseudotrachelocerca Song, 1990, is clearly separated from Litostomatea and clusters within an assemblage comprising the classes Prostomatea, Colpodea and Plagiopylea; (iii) both morphological evidence and molecular evidence indicate that the genus Foissnerides should be transferred from family Trachelophyllidae to Pseudoholophryidae; (iv) the validity of the order Helicoprorodontida Grain, 1994, and its monophyly is strongly supported; (5) the family Chaeneidae does not belong to the order Lacrymarida but represents a distinct clade in the subclass Haptoria.     

Phylogenetic analysis of the minute brown scavenger beetles (Coleoptera: Latridiidae), and recognition of a new beetle family,Akalyptoischiidae fam.n. (Coleoptera: Cucujoidea)

We infer the first phylogenetic hypothesis for Latridiidae Erichson (Coleoptera: Cucujoidea). Portions of seven genes (18S ribosomal DNA, 28S ribosomal DNA, 12S ribosomal DNA, 16S ribosomal DNA, cytochrome c oxidase I and II and histone III) were analysed. Twenty‐seven latridiid species were included, representing both subfamilies and more than half of the currently recognized genera. Eight outgroup taxa from other families of Cucujoidea were included. Parsimony and partitioned Bayesian analyses were performed on the combined dataset. In both phylogenetic analyses, the enigmatic Akalyptoischion Andrews (Latridiinae) was recovered outside of Latridiidae. The subfamilies Corticariinae and Latridiinae (without Akalyptoischion) were each recovered as monophyletic in both analyses. A new family, Akalyptoischiidae fam.n. is erected based on the results of the phylogenetic study and further support from adult morphology, key features of which are illustrated.     

Molecular phylogeny of Allograpta (Diptera, Syrphidae) reveals diversity of lineages and non-monophyly of phytophagous taxa

Phylogenetic relationships of genera Allograpta, Sphaerophoria and Exallandra (Diptera, Syrphidae) were analyzed based on sequence data from the mitochondrial protein-coding gene cytochrome c oxidase subunit I (COI) and the nuclear 28S and 18S ribosomal RNA genes. The three genera are members of the subfamily Syrphinae, where nearly all members feed as larvae on soft-bodied Hemiptera and other arthropods. Phytophagous species have recently been discovered in two subgenera of Allograpta, sg Fazia and a new subgenus from Costa Rica. Phylogenetic analyses of the combined datasets were performed using parsimony, under static alignment and direct optimization, maximum likelihood and Bayesian inference. Congruent topologies obtained from all the analyses indicate paraphyly of the genus Allograpta with respect to Sphaerophoria and Exallandra. Exallandra appears embedded in the genus Sphaerophoria, and both genera are placed within Allograpta. The distribution of phytophagous taxa in Allograpta indicates that plant feeding evolved at least twice in this group.     

Molecular phylogeny of the higher taxa of Odonata (Insecta) inferred from COI, 16S rRNA, 28S rRNA,and EF1‐α sequences

In this study, we sequenced both two mitochondrial genes (COI and 16S rRNA) and nuclear genes (28S rRNA and elongation factor‐1α) from 71 species of Odonata that represent 7 superfamilies in 3 suborders. Phylogenetic testing for each two concatenated gene sequences based on function (ribosomal vs protein‐coding genes) and origin (mitochondrial vs nuclear genes) proved limited resolution. Thus, four concatenated sequences were utilized to test the previous phylogenetic hypotheses of higher taxa of Odonata via Bayesian inference (BI) and maximum likelihood (ML) algorithms, along with the data partition by the BI method. As a result, three slightly different topologies were obtained, but the BI tree without partition was slightly better supported by the topological test. This topology supported the suborders Anisoptera and Zygoptera each being a monophyly, and the close relationship of Anisozygoptera to Anisoptera. All the families represented by multiple taxa in both Anisoptera and Zygoptera were consistently revealed to each be a monophyly with the highest nodal support. Unlike consistent and robust familial relationships in Zygoptera those of Anisoptera were partially unresolved, presenting the following relationships: ((((Libellulidae + Corduliidae) + Macromiidae) + Gomphidae + Aeshnidae) + Anisozygoptera) + (((Coenagrionidae + Platycnemdidae) + Calopterygidae) + Lestidae). The subfamily Sympetrinae, represented by three genera in the anisopteran family Libellulidae, was not monophyletic, dividing Crocothemis and Deielia in one group together with other subfamilies and Sympetrum in another independent group.     

Phylogenetic relationships of leptophlebiid mayflies as inferred by histone H3 and 28S ribosomal DNA

Abstract Leptophlebiidae is among the largest and most diverse groups of extant mayflies (Ephemeroptera), but little is known of family‐level phylogenetic relationships. Using two nuclear genes (the D2 + D3 region of 28S ribosomal DNA and histone H3) and maximum parsimony (MP), maximum likelihood (ML) and Bayesian inference (BI), we inferred the evolutionary relationships of 69 leptophlebiids sampled from six continents and representing 30 genera plus 11 taxa of uncertain taxonomic rank from Madagascar and Papua New Guinea. Although we did not recover monophyly of the Leptophlebiidae, monophyly of two of the three leptophlebiid subfamilies, Habrophlebiinae and Leptophlebiinae, was recovered with moderate to strong support in most analyses. The Atalophlebiinae was rendered paraphyletic as a result of the inclusion of members of Ephemerellidae or the Leptophlebiinae clade. For the species‐rich Atalophlebiinae, four groups of taxa were recovered with moderate to strong branch support: (i) an endemic Malagasy clade, (ii) a Paleoaustral group, a pan‐continental cluster with members drawn from across the southern hemisphere, (iii) a group, uniting fauna from North America, southeast Asia and Madagascar, which we call the Choroterpes group and (iv) a group uniting three New World genera, Thraulodes, Farrodes and Traverella. Knowledge of the phylogenetic relationships of the leptophlebiids will aid in future studies of morphological evolution and biogeographical patterns in this highly diverse and speciose family of mayflies.     

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).