Relationships among the tiphiid wasp subfamilies (Hymenoptera)

Abstract. The phylogenetic relationships among the subfamilies of Tiphiidae are discussed, with detailed analysis of characters, and presentation of a phylogenetic tree. The tribe Diammini Turner is elevated to subfamily status. A key to the subfamilies is included, and a brief synopsis of each subfamily is given.     

Systematics of the cyclostome subfamilies of braconid parasitic wasps (Hymenoptera: Ichneumonoidea): a simultaneous molecular and morphological Bayesian approach

Phylogenetic relationships among 95 genera collectively representing 17 of the 18 currently recognized cyclostome braconid wasp subfamilies were investigated based on DNA sequence fragments of the mitochondrial COI and the nuclear 28S rDNA genes, in addition to morphological data. The treatment of sequence length variation of the 28S partition was explored by either excluding ambiguously aligned regions and indel information (28SN) or recoding them (28SA) using the 'fragment-level' alignment method with a modified coding approach. Bayesian MCMC analyses were performed for the separate and combined data sets and a maximum parsimony analysis was also carried out for the simultaneous molecular and morphological data sets. There was a significant incongruence between the two genes and between 28S and morphology, but not for morphology and COI. Different analyses with the 28SA data matrix resulted in topologies that were generally similar to the ones from the 28SN matrix; however, the former topologies recovered a higher number of significantly supported clades and had a higher mean Bayesian posterior probability, thus supporting the inclusion of information from ambiguously aligned regions and indel events in phylogenetic analyses where possible. Based on the significantly supported clades obtained from the simultaneous molecular and morphological analyses, we propose that a total of 17 subfamilies should be recognized within the cyclostome group. The subfamilial placements of several problematic cyclostome genera were also established.     

A phylogeny of aphid parasitoids (Hymenoptera: Braconidae: Aphidiinae) inferred from mitochondrial NADH 1 dehydrogenase gene sequence

Wasps of the braconid subfamily Aphidiinae are solitary endoparasitoids of aphids. Several aspects of their biology have been the focus of intuitive evolutionary hypotheses which could be tested with a robust phylogeny. Phylogenetic hypotheses have been proposed previously for aphidiines based on morphology, embryology, and DNA sequences. However, many of them are based on a limited number of characters and/or taxa and lack congruence. In addition, many of the inferred phylogenies have not been based upon cladistic analysis. Therefore, a phylogenetic study of Aphidiinae was undertaken, utilizing 465 bp of DNA sequence of the mitochondrial NADH1 dehydrogenase gene. DNA sequences were obtained from 40 taxa, including 14 genera and three outgroups. It is suggested that in agreement with most of the previously proposed phylogenies, the aphidiines, each of the three recognized tribes (Praini, Ephedrini, Aphidiini), and most genera are monophyletic. In contrast to previously proposed phylogenies, the clade of Praon + Dyscritulus (=Praini), rather than Ephedrini, is basal among the aphidiines.     

A new enigmatic apterous cyclostome genus (Hymenoptera: Braconidae) from the Greek Islands

A new West Palaearctic genus Achaiabracon gen. n. with the type species A. aenigmatus sp. n. is described and illustrated from the Greek Islands Crete and Corfu. The position of this genus in the cyclostome group of subfamilies of Braconidae and its possible relation with rhyssaline genera are discussed. A new tribe Achaiabraconini trib. n. is proposed.     

The polyphyletic origin of endoparasitism in the cyclostome lineages of Braconidae (Hymenoptera)

Abstract. The evolution of endoparasitism in braconid wasps is demonstrably polyphyletic, not only within the family as a whole but even within the rogadine/exothecine assemblage of genera. A cladistic analysis of rogadine and exothecine genera, coupled with an optimization approach to estimating ancestral character states, is used to analyse the evolution of various aspects of endoparasitism within this assemblage. At least two independent origins of endoparasitism are hypothesized-one in the Rodagini and the other in an assemblage including the Gnaptodontini, Opiinae and Alysiinae.
Phylogenetic analyses and hypotheses can be useful not only in interpreting evolutionary trends, but in guiding and providing insights into more general comparative behavioural and physiological studies of parasitism.     

Phylogenomics of the lepidopteran endoparasitoid wasp subfamily Rogadinae (Hymenoptera: Braconidae) and related subfamilies

Rogadinae are a cosmopolitan, species‐rich braconid wasp subfamily whose species are endoparasitoids that attack larvae of a number of lepidopteran families. Members of this subfamily are characterized by pupating within the mummified host larval skin. The subfamily contains six tribes whose relationships have only been partially clarified: Aleiodini, Betylobraconini, Clinocentrini, Rogadini, Stiropiini and Yeliconini. The limits and composition of the closely related subfamilies to the Rogadinae, Hormiinae and Lysiterminae, also remain unclear. Here, we generated ultraconserved element data to reconstruct an almost fully resolved phylogeny for the members of Rogadinae and related subfamilies. Based on our best estimate of phylogeny, we confirm the monophyly of Rogadinae including Betylobraconini, synonymize Xenolobus Fahringer and Bequartia Cameron within the species‐rich genus Aleiodes Wesmael ( syn.n. ) based on DNA, and synonymize Promesocentrus van Achterberg with Pilichremylus Belokobylskij ( syn.n. ) based on morphology. We also consistently recovered Hormiinae and Lysiterminae as not reciprocally monophyletic, and thus propose to unite their members under Hormiinae. The ancestral host preference for Rogadinae was probably attacking concealed lepidopteran larvae, with the occurrence of at least two main subsequent transitions to attack both concealed and exposed hosts, one within Rogadini and a second within Aleiodini. We highlight the importance of natural history collections as a source for conducting genomic‐based studies using techniques that allow to obtain a substantial amount of data from considerably old preserved insect specimens.     

Relationships among characiform fishes inferred from analysis of nuclear and mitochondrial gene sequences

Suprafamilial relationships among characiform fishes and implications for the taxonomy and biogeographic history of the Characiformes were investigated by parsimony analysis of four nuclear and two mitochondrial genes across 124 ingroup and 11 outgroup taxa. Simultaneous analysis of 3660 aligned base pairs from the mitochondrial 16S and cytochrome b genes and the nuclear recombination activating gene (RAG2), seven in absentia (sia), forkhead (fkh), and alpha-tropomyosin (trop) gene loci confirmed the non-monophyly of the African and Neotropical assemblages and corroborated many suprafamilial groups proposed previously on the basis of morphological features. The African distichodontids plus citharinids were strongly supported as a monophyletic Citharinoidei that is the sistergroup to all other characiforms, which form a monophyletic Characoidei composed of two large clades. The first represents an assemblage of both African and Neotropical taxa, wherein a monophyletic African Alestidae is sister to a smaller clade comprised of the Neotropical families Ctenolucidae, Lebiasinidae, and the African Hepsetidae, with that assemblage sister to a strictly Neotropical clade comprised of the Crenuchidae and Erythrinidae. The second clade within the Characoidei is strictly Neotropical and includes all other Characiformes grouped into two well supported major clades. The first, corresponding to a traditional definition of the Characidae, is congruent with some groupings previously supported by morphological evidence. The second clade comprises a monophyletic Anostomoidea that is sister to a clade formed by the families Hemiodontidae, Parodontidae, and Serrasalmidae, with that assemblage, in turn, the sistergroup of the Cynodontidae. Serrasalmidae, traditionally regarded as a subfamily of Characidae, was recovered as the sistergroup of (Anostomoidea (Parodontidae+Hemiodontidae)) and the family Cynodontidae was recovered with strong support as the sistergroup to this assemblage. Our results reveal three instances of trans-continental sistergroup relationships and, in light of the fossil evidence, suggest that marine dispersal cannot be ruled out a priori and that a simple model of vicariance does not readily explain the biogeographic history of the characiform fishes.     

Kauriphanes n. gen., a new genus of braconid parasitoid wasp (Hymenoptera: Braconidae: Doryctinae) from New Zealand

A new genus belonging to the braconid wasp subfamily Doryctinae, Kauriphanes n. gen. (type species K. khalaimi n. sp.), is described from New Zealand. This genus is placed within the doryctine subtribe Caenophanina. The extent of this subtribe is discussed and the phylogenetic relationships of three of its genera were investigated using one mitochondrial and one nuclear DNA sequence markers. Similar to previous studies, the Bayesian analyses performed significantly support a clade with the included members of Caenophanina as a sister group of a clade with the examined species of Spathiini sensu stricto. The placement of the Caenophanini within Doryctini, however, is left pendant to further exhaustive phylogenetic studies. A key to genera and subgenera belonging to Caenophanina is given.     

Host range of braconid species (Hymenoptera: Braconidae) that attack Asphondyliini (Diptera: Cecidomyiidae) in Japan

We reared six idiobiont braconids, Bracon asphondyliae, B. sunosei, B. tamabae, Simplicibracon curticaudis, Testudobracon longicaudis and T. pleuralis from 22 identified species and 11 unidentified segregates of Asphondyliini (Diptera: Cecidomyiidae) in Japan. A total of 22 cecidomyiid species and segregates were newly recorded as hosts of the braconids. Analysis of cytochrome oxidase subunit I (COI) did not show any evidence of host races among the braconids. Bracon sunosei, which was synonymized with B. asphondyliae, is restored to a valid species. The host range of the braconid species seemed to be related to the lineage of host genera within Asphondyliini.     

A new, putatively primitive Cretaceous fossil braconid subfamily from New Jersey amber (Hymenoptera, Braconidae)

Protorhyssalus goldmani gen. n., sp. n., in a new subfamily of braconid wasps, the Protorhyssalinae, is described from Late Cretaceous amber fossils from New Jersey, USA. The Protorhyssalinae appears to be cyclostome and shows a similar set of plesiomorphic characters to the extant Rhyssalinae. However, it possesses hindwing vein 2-CU, a feature only found among the cyclostome braconids in the rare and putatively primitive Chilean subfamily Apozyginae.     

Phylogenetic relationships among genera of Aphidiinae (Hymenoptera: Braconidae) based on DNA sequence of the mitochondrial 16S rRNA gene

Phylogenetic relationships among forty‐nine taxa representing twenty‐four genera of Aphidiinae (Hymenoptera: Braconidae) were investigated using DNA sequence of a portion of the mitochondrial 16S rRNA gene and parsimony analysis. Seven species in six other subfamilies of Braconidae were used as outgroup. The results suggested that members of Aphidiinae are monophyletic. The basal lineage of Aphidiinae was Aclitus in weighted and unweighted parsimony analyses and Praini was basal relative to Ephedrini. With the exception of Pauesia and Aphidius, all genera were monophyletic. The results support generic status for Euaphidius, but not for Lysaphidus. Diaeretus leucopterus was internal to a clade composed of three Pauesia species, suggesting that the latter genus may be paraphyletic. A combined analysis that included DNA sequence of 16S rRNA, NADH1 dehydrogenase and 28S rRNA resulted in more robust cladograms with topologies similar to those inferred from the 16S rRNA gene sequence alone. The results are compared to previously proposed phylogenies of Aphidiinae based on morphological and molecular characters.     

Relationships among several species of the subfamily Gymnelinae (Zoarcidae) inferred from mitochondrial DNA sequence analysis

Nucleotide sequences of the mtDNA COI and cytochrome b genes were determined in Magadanichthys skopetsi, a member of the new monotypic genus Magadanichthys, endemic to the northern coast of the Sea of Okhotsk. Comparison of this species with other representatives of the subfamily Gymnelinae (family Zoarcidae) revealed high genetic similarity of M. skopetsi to Hadropareia middendorffii and considerable differences between these species and Gymnelopsis ochotensis.     

The complete mitochondrial genome of Taeniogonalos taihorina (Bischoff) (Hymenoptera: Trigonalyidae) reveals a novel gene rearrangement pattern in the Hymenoptera

The family Trigonalyidae is considered to be one of the most basal lineages in the suborder Apocrita of Hymenoptera. Here, we determine the first complete mitochondrial genome of the Trigonalyidae, from the species Taeniogonalos taihorina (Bischoff, 1914). This mitochondrial genome is 15,927 bp long, with a high A + T-content of 84.60%. It contains all of the 37 typical animal mitochondrial genes and an A + T-rich region. The orders and directions of all genes are different from those of previously reported hymenopteran mitochondrial genomes. Eight tRNA genes, three protein-coding genes and the A + T-rich region were rearranged, with the dominant gene rearrangement events being translocation and local inversion. The arrangements of three tRNA clusters, trnY–trnM–trnI–trnQ, trnW–trnL2–trnC, and trnH–trnA–trnR–trnN–trnS–trnE–trnF, and the position of the cox1 gene, are novel to the Hymenoptera, even the insects. Six long intergenic spacers are present in the genome. The secondary structures of the RNA genes are normal, except for trnS2, in which the D-stem pairing is absent.     

New braconid wasps from French Cretaceous amber (Hymenoptera, Braconidae): synonymization with Eoichneumonidae and implications for the phylogeny of Ichneumonoidea

Two new fossils of Braconidae are described from Albian-Cenomanian amber of south-western France, Protorhyssalodes arnaudi gen. n., sp. n., and Aenigmabracon capdoliensis gen. n., sp. n. The former appears superficially similar to the type genus and species of the extinct sub-family Protorhyssalinae, from Turonian New Jersey amber specimens, and the latter both to Protorhyssalus and to members of the extinct family Eoichneumonidae. However, both new taxa display unique combinations of wing venation characters making confident assignment to sub-family impossible. Indeed, they are the first braconids ever known to possess both vein 2-CU and a distinct trace of vein 2-1A on hindwing. The new fossil taxa are incorporated into a morphological analysis of extinct and extant ichneumonoids. As a result of the analyses we synonymize the Eoichneumonidae with the Braconidae.     

Mendesellinae, a new subfamily of braconid wasps (Hymenoptera, Braconidae) with a review of relationships within the microgastroid assemblage

Abstract. A new subfamily of braconid wasps, the Mendesellinae, is proposed and described. It is represented by two genera and nine species, all newly described: Mendesella gen.n., type species M.amazonica sp.n., M.braziliensls , sp.n., M.magna sp.n., Epsilogaster gen.n., type species E.bicolor sp.n., E.brazMensis sp.n., E.dureno sp.n., E.palenque sp.n., E.tico sp.n. and E.panama sp.n. The phylogenetic relationships among the subfamilies of the 'microgastroid assemblage' of Braconidae are reviewed and analysed, to clarify the status and relationships of the new subfamily.     

绿眼赛茧蜂线粒体基因组全序列测定和分析

【目的】测定绿眼赛茧蜂Zele chlorophthalmus线粒体基因组全序列,分析其基因组结构及茧蜂科(Braconidae)部分类群的系统发育关系。【方法】利用Illumina MiSeq二代测序技术对绿眼赛茧蜂的线粒体基因组进行测序,对基因组序列进行拼装、注释,分析其结构特点和碱基组成;基于22种茧蜂科昆虫的COX1蛋白编码基因序列,应用最大似然法(ML)和邻接法(NJ)构建系统发育树,分析绿眼赛茧蜂与茧蜂科其他昆虫的系统发育关系。【结果】绿眼赛茧蜂线粒体基因组全长16 661 bp(GenBank登录号: MG822749),包含13个蛋白质编码基因、22个tRNA基因和2个rRNA基因,共37个基因,以及1个控制区。线粒体基因组有明显的核苷酸组成的偏倚,AT偏正,GC偏负,其A+T含量为82.83%。基因排列顺序与推测的昆虫祖先的序列不完全一致,tRNA基因7处发生重排。13个蛋白质编码基因均以ATN为起始密码子,以TAA为终止密码子。在22个tRNA基因二级结构中,除tRNA^His(H)缺失TΨC环和tRNA^Cys(C)仅剩二氢尿嘧啶(DHU)臂和反密码子臂外,其余tRNA基因均能形成典型的三叶草结构。基于COX1蛋白编码序列的系统发育分析结果显示,与绿眼赛茧蜂亲缘关系最近的是同属于赛茧蜂属的雪跗赛茧蜂Z. niveitarsis。【结论】本研究首次获得绿眼赛茧蜂线粒体基因组全序列。结果表明绿眼赛茧蜂隶属于优茧蜂亚科(Euphorinae)赛茧蜂属,并支持赛茧蜂属的单系性。     

Venom gland and reservoir morphology in the Doryctinae and related braconid wasps (Insecta, Hymenoptera, Braconidae)

Detailed venom reservoir and venom gland intima morphology has been investigated in reprsentatives of 84 genera and 10 subfamilies of Braconidae with particular reference to the Doryctinac, basal cyclostomes and related groups including the Opiinae, Alysiinae and Rogadinae sens. lat . Several new phylogenetically significant characters are described and illustrated. Extcnsivc secretory ductules on the primary venom duct is suggested as a synapomorphy for the subfamilies Braconinac, Doryctinae, Opiinae, Alysiinae and Gnamptodontinae. The presence of two separate venom gland insertions and their position beyond the spiral sculpture of the reservoir are suggested as synapomorphies for the subfamilies Doryctinae and Ypsistocerinac. An anterior, weakly sclerotized, probably glandular. expansion of the reservoir characterizes the Histeromerinae and Rhyssalinae and also the enigmatic genera Thoracoplites Fischer and Doryctomorpha Ashmead which are hcre transferred to the Rhyssalinae. Within the Doryctinac, venom reservoir morphology suggests three generic groups, one comprising the genera of the Odontohraconini and possibly also the genera Acanthodoryctes , Binareu and Monarea , all of which have an essentially undivided reservoir with extremely fine and uniform spiral sculpture. the second comprising the genera Doryctophasmus , Euscelinus, Gildoria, Halycea and Schlettereriella , which share horn-like processes at the base of the venom duct insertions, and a third including Acrophasmus. Dendro- soter, Heterospilus, Megaloproctus, Rutheia, Paraspathius, Schlettereriella. Spathius and Syngaster based on the presence of two markedly different densities of annulation of the reservoir. The degree to which external and other characters support these groups is discusscd.     

Phylogenetic relationships among the Ithomiini (Lepidoptera: Nymphalidae) inferred from one mitochondrial and two nuclear gene regions

Abstract.  A phylogenetic hypothesis for the tribe Ithomiini (Lepidoptera: Nymphalidae: Danainae) is presented, based on sequences of the mitochondrial cytochrome oxidase subunits I and II (COI–COII) region and regions of the nuclear genes wingless and Elongation factor 1-alpha . Branch support for each clade is assessed, and a partition congruence index is used to explore conflict among gene regions. The monophyly of the clade is strongly supported, as are many of the traditionally recognized subtribes and genera. The data imply paraphyly of some genera and tribes, but largely support recent classifications and phylogenetic hypotheses based on morphological characters.     

Phylogenetic relationships among the Braconidae (Hymenoptera: Ichneumonoidea) inferred from partial 16S rDNA, 28S rDNA D2, 18S rDNA gene sequences and morphological characters

Phylogenetic relationships among the Braconidae were examined using homologous 16S rDNA, 28S rDNA D2 region, and 18S rDNA gene sequences and morphological data using both PAUP* 4.0 and MRBAYES 3.0B4 from 88 in-group taxa representing 35 subfamilies. The monophyletic nature of almost all subfamilies, of which multiple representatives are present in this study, is well-supported except for two subfamilies, Cenocoelinae and Neoneurinae that should probably be treated as tribal rank taxa in the subfamily Euphorinae. The topology of the trees generated in the present study supported the existence of three large generally accepted lineage or groupings of subfamilies: two main entirely endoparasitic lineages of this family, referred to as the "helconoid complex" and the "microgastroid complex," and the third "the cyclostome." The Aphidiinae was recovered as a member of the non-cyclostomes, probably a sister group of Euphorinae or Euphorinae-complex. The basal position of the microgastroid complex among the non-cyclostomes has been found in all our analyses. The cyclostomes were resolved as a monophyletic group in all analyses if two putatively misplaced groups (Mesostoa and Aspilodemon) were excluded from them. Certain well-supported relationships evident in this family from the previous analyses were recovered, such as a sister-group relationships of Alysiinae+Opiinae, of Braconinae+Doryctinae, and a close relationship between Macrocentrinae, Xiphozelinae, Homolobinae, and Charmontinae. The relationships of "Ichneutinae + ((Adeliinae + Cheloninae) + (Miracinae + (Cardiochilinae + Microgastrinae)))" was confirmed within the microgastroid complex. The position of Acampsohelconinae, Blacinae, and Trachypetinae is problematic.     

The position of the Hymenoptera within the Holometabola as inferred from the mitochondrial genome of Perga condei (Hymenoptera: Symphyta: Pergidae)

We sequenced most of the mitochondrial genome of the sawfly Perga condei (Insecta: Hymenoptera: Symphyta: Pergidae) and tested different models of phylogenetic reconstruction in order to resolve the position of the Hymenoptera within the Holometabola, using mitochondrial genomes. The mitochondrial genome sequenced for P. condei had less compositional bias and slower rates of molecular evolution than the honeybee, as well as a less rearranged genome organization. Phylogenetic analyses showed that, when using mitochondrial genomes, both adequate taxon sampling and more realistic models of analysis are necessary to resolve relationships among insect orders. Both parsimony and Bayesian analyses performed better when nucleotide instead of amino acid sequences were used. In particular, this study supports the placement of the Hymenoptera as sister group to the Mecopterida.