Evolution of green lacewings (Neuroptera: Chrysopidae): an anchored phylogenomics approach

A phylogeny of green lacewings (Neuroptera: Chrysopidae) using anchored hybrid enrichment data is presented. Using this phylogenomic approach, we analysed 137 kb of sequence data (with < 10% missing) for 82 species in 50 genera of Chrysopidae under Bayesian and maximum likelihood criteria. We recovered a strongly supported tree topologically congruent with recently published phylogenies, especially relationships amongst higher‐level groups. The subfamily Nothochrysinae was recovered as paraphyletic, with one clade sister to the rest of Chrysopidae, and the second clade containing the nominal genus (Nothochrysa Navás) as sister to the subfamily Apochrysinae. Chrysopinae was recovered as a monophyletic with the monobasic Nothancylini tribe n. sister to the rest of the subfamily. Leucochrysini was recovered sister to Belonopterygini, and Chrysopini was rendered paraphyletic with respect to Ankylopterygini. Divergence times and diversification estimates indicate a major shift in rate in ancestral Chrysopini at the end of the Cretaceous, and the extensive radiation of Chrysopinae, the numerically dominant clade of green lacewings, began in the Mid‐Paleogene (c. 45 Ma).     

The Status of Knowledge of the Genus Ecdyonurus in the Iberian Peninsula,with Description of Two New Species of the E. venosus Group from Spain (Ephemeroptera: Heptageniidae)

As a result of studies of material of Ecdyonurus from Spain, nymphs and adults of two new species, E. baeticus sp. n. and E. olgae sp. n., are described. Three species described by Navás must be considered as nomina dubia (E. diversus Navás, 1923, E. excelsus Navás, 1927 and E. fluminum var. speciosus Navás, 1915). Variability in adults of E. venosus is figured and discussed.     

Systematics of the dung beetle tribe Sisyphini Mulsant (Scarabaeidae: Scarabaeinae) inferred from a molecular phylogeny and biogeography of southern African species

The tribe Sisyphini Mulsant was recently redefined following the transfer of the endemic southern African genus Epirinus Dejean from the polyphyletic tribe Deltochilini Lacordaire. A molecular phylogeny of the southern African members of Sisyphini supports Epirinus as sister to Sisyphus Latreille and recovered three major clades in Sisyphus classified here as subgenera Sisyphus (Neosisyphus Müller) stat. rev. , Sisyphus (Parasisyphus Barbero, Palestrini & Zunino) stat.n. and Sisyphus (Sisyphus) stat.n. A molecular clock analysis suggests that Sisyphus and Epirinus diverged from their last common ancestor during the Lower to Middle Oligocene (c. 29.37 Ma). Biogeographical analysis indicated that southern African Sisyphus species are centred in the east and northeast in Highveld grassland and warmer savannah regions. By contrast, Epirinus species are largely restricted to the southwest and southeast in the cooler winter and bimodal rainfall regions plus arid highland Karoo and Highveld grasslands. Based on morphological and biogeographical differences between Epirinus and Sisyphus, we propose that the monogeneric Epirinus be placed in its own tribe, Epirinini van Lansberge stat. rev.     

Phylogeny and evolution of Mesozoic and extant lineages of Histeridae (Coleoptera), with discovery of a new subfamily Antigracilinae from the Lower Cretaceous

In order to place a newly discovered species Antigracilus costatus gen. sp. n. from the Lower Cretaceous Yixian Formation (China) and to assess previously unplaced fossil taxa, we investigated the relationships of extant and extinct lineages of Histeridae based on three data sets: (i) 69 morphological characters belonging to 48 taxa (representing all 11 subfamilies and 15 of 17 tribes of modern Histeridae); (ii) partitioned alignment of 6030 bp from downloaded nucleotide sequences (28S, CAD, COI, 18S) of 50 taxa (representing 10 subfamilies and 15 of 17 tribes of modern Histeridae); and (iii) a combined morphological and molecular dataset for 75 taxa. Phylogenetic analyses of the morphology and combined matrices recovered the new Lower Cretaceous taxon as a sister group to remaining Histeridae and it is placed in †Antigracilinae subfam. n. †Antigracilinae constitutes the earliest record of Histeridae from the Lower Cretaceous Yixian Formation (∼125 Myr), backdating the minimum age of the family by 25 Myr from the earliest Cenomanian (~99 Myr) to the Barremian of the Cretaceous Period. Our molecular phylogeny supports Histeridae to be divided into seven different clades, with currently recognised subfamilies Abraeinae (sensu lato), Saprininae, Chlamydopsinae, and Histerinae (sensu lato) recovered as monophyletic, while Dendrophilinae, Onthophilinae, and Tribalinae are polyphyletic taxa. The Burmese amber species †Pantostictus burmanicus Poinar & Brown is placed as a sister group to the tribe Plegaderini (Abraeinae) and was assigned as a new tribe Pantostictini trib. n. Both molecular and combined phylogenies recovered the subfamilies Trypanaeinae and Trypeticinae deeply within the subfamily Abraeinae (sensu lato), and they are downgraded into Trypanaeini stat. n. and Trypeticini stat. n.     

Comprehensive phylogeny of the Cleroidea (Coleoptera: Cucujiformia)

The first thorough molecular phylogeny of the superfamily Cleroidea, represented by 377 taxa, and the first with an emphasis on Trogossitidae, was undertaken. Maximum likelihood and Bayesian analyses were performed on a four‐gene dataset (18S, 28S, cox1, cytb) of 395 taxa (along with 18 outgroups), including all 16 currently recognized families of Cleroidea and all current and formerly recognized tribes of Trogossitidae. The superfamily as a whole received strong support in Bayesian analyses. On the basis of phylogenetic results, 18 families in Cleroidea are recognized, including three taxa elevated to family for the first time and two reinstated families. The former tribe Rentoniini (Trogossitidae: Peltinae) was strongly supported as a monophyletic group apart from the remainder of Trogossitidae, and is herein elevated to family status, Rentoniidae stat.n. Protopeltis was also found to be an isolated lineage and becomes Protopeltidae stat.n. Peltini + Larinotini were recovered as a weakly supported sister grouping; Peltini (including only Peltis) becomes Peltidae stat.rest. The trogossitid subfamily Lophocaterinae, to the exclusion of Decamerini, formed a clade which is here designated Lophocateridae stat.rest. and sensu n. The Trogossitinae tribes Calityini, Egoliini (represented by Egolia) and Larinotini were recovered apart from core Trogossitidae but showed no strong affinities to other taxa or congruence between analyses; they are here conservatively retained in Trogossitidae as Calityinae stat.rest. , Egoliinae stat.rest. and Larinotinae stat.rest. The genus Thymalus of the peltine tribe Thymalini was indicated with moderate to strong support as the sister group of the Decamerini (Trogossitidae: Lophocaterinae); together these represent Thymalidae stat.n. and sensu n. with subfamilies Decamerinae stat.rest. ( new placement ) and Thymalinae stat.n. The remainder of Trogossitinae, the tribes Trogossitini and Gymnochilini, formed a well‐supported clade which comprises the Trogossitidae: Trogossitinae sensu n. The tribe Gymnochilini syn.n. is synonymized with Trogossitini. The monotypic family Phloiophilidae was recovered, contradicting a recent placement within Trogossitidae. The melyrid lineage was recovered with moderate (maximum likelihood) to strong (Bayesian analyses) support and includes the families Phycosecidae, Rhadalidae, Mauroniscidae, Prionoceridae and Melyridae (including Dasytidae and Malachiidae). The genus Dasyrhadus is tentatively transferred from Rhadalidae to Mauroniscidae. The genus Gietella, once proposed as a distinct family but recently placed within Dasytidae, was recovered as strongly sister to Rhadalidae sensu n. , and we transfer it to that family as Gietellinae new placement . Attalomiminae (formerly Attalomimidae) syn.n. is synonymized with Melyridae: Malachiinae: Lemphini sensu n. Melyridae sensu n. includes only Dasytinae, Malachiinae and Melyrinae. Metaxina is returned to the Chaetosomatidae sensu n. , of which Metaxinidae syn.n. becomes a junior synonym. Resolution within Cleridae was generally poor, but a broadly defined Korynetinae stat rest. + Epiclininae received high support (Bayesian analyses). Outside of Trogossitidae, the main focus of this study, major rearrangements of the classification of Cleroidea were not undertaken, despite evidence indicating such changes are needed.     

Revision of the genus Heliomantis Giglio-Tos 1915 (Insecta: Mantodea: Hymenopodidae)

Summary. The little-known flower mantis genus Heliomantis Giglio-Tos 1915 (Hymenopodidae) distributed in South and South-Eastern Asia is revised. All species are redescribed. New data on the morphology, distribution and ecology are given, and all known specimens are listed. The lectotype of H. elegans (Navás 1904) is designated. Due to highly divergent morphology, H. latipennis Werner 1930 from Sarawak, Borneo, is transferred to Werneriana n. gen., leaving the genus Heliomantis with the only species, H. elegans (Navás, 1904).     

Phylogeny of a new supertribe Cephenniitae with generic review of Eutheiini and description of a new tribe Marcepaniini (Coleoptera: Staphylinidae: Scydmaeninae)

Genera of Eutheiini are reviewed and Eutheimorphus is removed from this tribe of ant‐like stone beetles (Scydmaeninae) and transferred to Cephenniini. A monogeneric Marcepaniini trib.n. is described to accommodate Marcepania gen.n. from Malaysia, with five species: M. semengohensis sp.n. (the type species of Marcepania), M. tuberculata sp.n. , M. seramaensis sp.n. , M. minutissima sp.n. and M. elongata sp.n. A phylogenetic analysis of all genera of Cephenniini, Eutheiini and Marcepaniini based on adult morphological characters resulted in recovering a well‐supported monophyletic clade Eutheiini + (Marcepaniini + Cephenniini) and these tribes are included in Cephenniitae stat.n. (Eutheiini and Cephenniini are therefore removed from Scydmaenitae). Only a weak support for monophyly of Eutheiini was found, but morphological characters allow for maintaining this presumably relic group as a separate tribe. Previously proposed monophyletic groups within Cephenniini were recovered as such, but after inclusion of Eutheimorphus, a sister taxon to the ‘Cephennomicrus group’, the latter lineage gained weak statistical support. The evolutionary history of Cephenniitae is discussed, with focus on known northern hemisphere fossils classified in Scydmaenitae and Hapsomelitae, but possibly closely allied to Cephenniitae. Establishing the supertribe Cephenniitae is the first step toward a profound reclassification of Scydmaeninae on a robust phylogenetic basis. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:B0E1B12D-9587-4C4F-A908-A12A0C424A8C .     

Phylogenetic systematics of the gondwanan Empis macrorrhyncha group (Diptera,Empididae, Empidinae)

The Empis macrorrhyncha group (Diptera: Empididae) from cool to warm temperate areas of South America and Australia is diagnosed and cladistically analysed, and five new species, Empis animosa sp.n. , E. austera sp.n. , E. maculosa sp.n. , E. occidentalis sp.n. and E. pedivillosula sp.n. , are described. Cladistic analysis of 23 adult morphological characters for 14 species of the group generated a single tree of 28 steps (CI = 0.82; RI = 0.93). Monophyly was established on the basis of a single apomorphy, possession of a bilobed cercus of the male hypopygium. Three main clades were inferred: clade 1 included three Patagonian and a single southwestern Australian species; clade 2 included two species from southeastern Australia; clade 3 included a large Patagonian group of five species and a single southeastern Australian species. The E. fulvicollis complex (clade 1) is a sister‐group of the E. macrorrhyncha complex (clades 2 + 3). A provisional historical biogeographic hypothesis is advanced correlating the appearance of the South American and Australian sister lineages with the timing of the break‐up of Gondwana.     

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

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

Hecabolomorpha n. gen., a new Asian genus from the tribe Hecabolini (Hymenoptera: Braconidae: Doryctinae)

A new Oriental genus Hecabolomorpha n. gen. with the type species H. asiatica n. sp. is described and illustrated from India and Central China. The position of this genus in the tribe Hecabolini and its relations with hecaboline genera are discussed.     

Phylogeny of ensign scale insects (Hemiptera: Coccoidea: Ortheziidae) based on the morphology of Recent and fossil females

The Ortheziidae (ensign scale insects) is a morphologically well‐defined family. The morphology and occurrence in the fossil record suggests a probable early origin of the family in scale insect evolution. The present phylogenetic analysis – based on 69 morphological characters of female ortheziids, using 39 exemplar Recent species – provides the first analytical assessment of relationships among living genera of the family, as well as the relationships of eight fossil species, based on complete, well‐preserved specimens in amber. Monophyly of the subfamilies Newsteadiinae, Ortheziinae and Ortheziolinae is supported, but Nipponortheziinae is found to be paraphyletic by inclusion of the Ortheziolinae. Thus, the subfamily Ortheziolinae is reduced in rank to tribe Ortheziolini stat.n. , which now includes Matileortheziola Kozár & Foldi, Ortheziolacoccus Kozár, Ortheziolamameti Kozár and Ortheziola?ulc. Consequently, the tribes Matileortheziolini, Ortheziolacoccini and Ortheziolamametini are synonymized ( syn.n. ) here under Ortheziolini. Five new species and one new genus of fossil ensign scales are described from three amber deposits: Burmorthezia gen.n. with type species Burmorthezia kotejai sp.n. and also B. insolita sp.n ., both in mid‐Cretaceous Burmese amber (98 Ma) and Arctorthezia baltica sp.n. in Eocene Baltic amber (c. 43 Ma) based on second‐instar nymphs; Mixorthezia kozari sp.n . and M. dominicana sp.n . in Miocene Dominican amber (c. 17 Ma) based on adult females. Fossil placements are unambiguous, with Burmorthezia forming a stem to crown‐group (Recent and Tertiary) Ortheziidae. A summary of described fossil ortheziids is provided.     

Molecular systematics of the butterfly tribe Preponini (Nymphalidae: Charaxinae)

The nymphalid butterfly tribe Preponini includes some of the Neotropical region's most spectacular and familiar butterflies, but the taxonomy of the group nevertheless remains unstable. Several recent studies of Nymphalidae phylogeny have suggested that both the tribe itself and several genera might not be monophyletic, but to date taxon sampling has not been sufficiently comprehensive to allow informed revision of the group's systematics. We therefore conducted the first complete species‐level phylogenetic study of the tribe to establish a firm higher classification. We used DNA sequence data from three genes, the two mitochondrial genes cytochrome oxidase subunits I and II (COI and COII), and the nuclear gene elongation factor‐1α (EF‐1α), to reconstruct the phylogeny of the tribe using maximum likelihood (ML), maximum parsimony (MP) and Bayesian inference (BI). We included 48 individuals representing the 22 recognised Preponini species, and an additional 25 out‐group taxa to explore taxonomic limits at different levels. Firstly, we found that Anaeomorpha splendida Rothschild never grouped with remaining Preponini, so that maintaining monophyly of the tribe requires the taxon to be excluded, and we thus reinstate the tribe Anaeomorphini stat.rev. Secondly, we investigated generic limits, in particular the relationship of Noreppa Rydon to Archaeoprepona Fruhstorfer, and that of Agrias Doubleday to Prepona Boisduval. The molecular results coupled with previous morphological studies suggest that Noreppa syn.n should be synonymised with Archaeoprepona, and that Agrias syn.n should be synonymised with Prepona. We found Prepona pheridamas (Cramer) to be sister to all other Prepona, and markedly divergent from them in both morphology and DNA sequences, suggesting the possibility that it should be placed in a separate genus. We also found a number of cases of significant DNA sequence divergence and paraphyly or polyphyly within putative species that require further taxonomic attention, including Prepona claudina (Godart) stat.n. and Prepona narcissus (Staudinger) stat.n., Prepona pylene Hewitson and Prepona deiphile (Godart). Future research should focus on a broader population sampling of widespread, polymorphic Preponini species to thoroughly revise the current species‐level taxonomy, thus creating a solid foundation for studies in ecology and conservation.     

Antlions and owlflies (Neuroptera: Myrmeleontidae,Ascalaphidae) of Kyrgyzstan

The myrmeoleontoid Neuroptera fauna of Kyrgyzstan comprises 34 species-group taxa, among which 23 are recorded from the republic for the first time. Epacanthaclisis alaica Krivokhatsky, 1998 is known only from the holotype collected in the Alai Mt. Range; Kirghizoleon cubitalis Krivokhatsky et Zakharenko, 1994 is known from Kyrgyzstan and from the Hissar Mt. country in Tajikistan; and Mesonemurus vartianorum Hölzel, 1972 is distributed from the Tien Shan to Afghanistan. 26 species are common in Kyrgyzstan and are fairly evenly distributed over its territory. New synonymies are established: Macronemurus persicus (Navás, 1915) = Barreja persica Navás, 1915, = Barreja amoena Hölzel, 1972, syn. n.; Myrmecaelurus acerbus (Walker) = Myrmeleon acerbus Walker, 1853; = Myrmecaelurus varians Navás, 1913, syn. n.; = Myrmecaelurus aequans Navás, 1913, syn. n.; = Myrmecaelurus nematicus Navás, 1932, syn. n. The polymorphous structure of the widely variable Macronemurus persicus is shown, and three its color morphs are distinguished: Macronemurus persicus var. persica (Navás, 1915), var. graciosa var. n., and var. amoena (Hölzel, 1972). Data on the antlion and owlfly fauna of some other Palaearctic countries are supplemented.     

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.     

New and little known crickets of the subfamily Phalangopsinae (Orthoptera,Gryllidae): 6. Neotropical taxa of the tribes Phalangopsini and Paragryllini

Two new genera, ten new species, and two new subspecies: Uvaroviella izerskyi sp. n., U. morona sp. n., U. ucayali sp. n., U. pastaza sp. n., U. affinis sp. n., U. bolivia sp. n., U. bora atalaya subsp. n., Kevanacla orientalis contraria subsp. n., Peruacla solitaria gen. et sp. n., Ecuadoracla propria gen. et sp. n., Adelosgryllus spurius sp. n., and A. phaeocephalus sp. n. are described. The tribe Paragryllini is divided into three subtribes: Paragryllina Desutter, 1988 stat. n., Neoaclina Desutter, 1988 stat. n., and Strogulomorphina Desutter 1988, stat. n. The composition of the genera Uvaroviella Chop. and Neoacla Desutter is discussed. The species from Costa Rica and the Antilles are supposed to belong to these genera and subgenera. New synonymy [U. trinidadi Gorochov, 2007 = U. enodos Otte et Perez-Gelabert, 2009, syn. n. and N. multivenosa (Chopard, 1937) = Selvacla choreutes Otte, 2006, syn. n.] is proposed.     

A review of the genus Eristena Warren in China (Lepidoptera: Crambidae: Acentropinae)

The larvae of three known species of Callibaetis are described here for the first time: C. gonzalezi (Navás), C. pollens Needham &; Murphy and C. sellacki (Weyenbergh). The larva of C. willineri Navás is resdescribed and the larva of C. guttatus Navás is revised and discussed. The diagnoses of the adults are also provided.     

Combined molecular and morphological phylogeny of Eulophidae (Hymenoptera: Chalcidoidea), with focus on the subfamily Entedoninae

A new combined molecular and morphological phylogeny of the Eulophidae is presented with special reference to the subfamily Entedoninae. We examined 28S D2–D5 and CO1 gene regions with parsimony and partitioned Bayesian analyses, and examined the impact of a small set of historically recognized morphological characters on combined analyses. Eulophidae was strongly supported as monophyletic only after exclusion of the enigmatic genus Trisecodes. The subfamilies Eulophinae, Entiinae (=Euderinae) and Tetrastichinae were consistently supported as monophyletic, but Entedoninae was monophyletic only in combined analyses. Six contiguous bases in the 3e′ subregion of the 28S D2 rDNA contributed to placement of nominal subgenus of Closterocerus outside Entedoninae. In all cases, Euderomphalini was excluded from Entiinae, and we suggest that it be retained in Entedoninae. Opheliminae n. stat. is raised from tribe to subfamily status. Trisecodes is removed from Entedoninae but retained as incertae sedis in Eulophidae until its family placement can be determined new placement . The genera Neochrysocharis stat. rev. and Asecodes stat. rev. are removed from synonymy with Closterocerus because strong molecular differences corroborate their morphological differences. Closterocerus (Achrysocharis) germanicus is transferred to the genus Chrysonotomyia n. comb. based on molecular and morphological characters.