Mitochondrial phylogeny of the Chrysis ignita (Hymenoptera: Chrysididae) species group based on simultaneous Bayesian alignment and phylogeny reconstruction (original) (raw)

A thousand and one wasps: a 28S rDNA and morphological phylogeny of the Ichneumonidae (Insecta: Hymenoptera) with an investigation into alignment parameter space and elision

Journal of Natural History, 2009

The internal phylogeny of the Ichneumonidae is investigated using parsimony analysis of a large data set including 1001 partial 28S ribosomal DNA sequences, 621 of which are newly reported, and a morphological data set of 162 characters scored variously at subfamily, tribe, genus group and genus levels and including only informative characters. The data set includes members of 630 named genera, representing all currently recognized subfamilies, all but four tribes and all but one of the taxa noted by Townes as being of uncertain placement. Sequences were aligned using clustal X, and a sensitivity analysis was performed varying gap-opening and gap-extension parameters. Alignments were appraised by reference to their ability to recover a range of traditional and morphologically recognized groups. Each alignment was analysed both independently and simultaneously with the morphological data set, and also with gap characters treated as both missing data and as informative. No single set of alignment parameters was found to be markedly better by this criterion, and different ranges of parameters led to the recovery of different recognized groups of taxa. Elision (combining all alignments into a single analysis) was therefore used, both with and without morphology and with both gap character treatments, to summarize the overall molecular signal. Analysis of the morphological matrix alone produced a number of results that are undoubtedly a consequence of convergence of morphological characters as the result of parallel evolution of similar life histories. Simultaneous analysis of the morphological data set with each of the 120 DNA alignments recovered most accepted subfamilies as monophyletic. Several currently recognized subfamilies are supported by most of the molecular analyses but some appear to be paraphyletic or polyphyletic. The Ctenopelmatinae are paraphyletic with respect to the Metopiinae. Robustly recovered results lead us to resurrect the Brachyscleromatinae to include Brachyschleroma and the Erythrodolius group of Phrudinae. The Neorhacodinae and the Phrudus group of Phrudinae are transferred to the Tersilochinae. Nonnus is transferred to the Nesomesochorinae. Hyperacmus is transferred to the Cylloceriinae. The major groupings of subfamilies that have recently been proposed (i.e. ichneumoniformes, pimpliformes and ophioniformes) were recovered as monophyletic, but their exact limits remain in question.

Testing the validity of Northern European species in the Chrysis ignita species group (Hymenoptera: Chrysididae) with DNA Barcoding

Zootaxa, 2014

Containing more than a hundred species, the Chrysis ignita species group is the largest and one of the most taxonomically challenging groups in its genus. It has not been possible to resolve the taxonomy of the group using traditional methods due to the lack of robust diagnostic morphological characters. Here we present the results of a molecular analysis designed to delimit species in the Chrysis ignita group for the first time; using mitochondrial sequence data for 364 in-group specimens consisting of all 18 species known to occur in Northern Europe. Two mitochondrial loci were analysed: a COI gene fragment, and a continuous DNA sequence consisting of 16S rRNA, tRNA Val , 12S rRNA and ND4. Two approaches were employed for delimiting species: (1) genetic distance analysis based on the standard COI barcode sequences and; (2) phylogenetic analysis of the COI fragment together with rRNA genes. Both analyses yielded trees with similar topology, but support values for nodes were higher using the second approach. Fifteen species were distinguished in all analyses: and C. vanlithi Linsenmaier, 1959. The specific status of C. mediata Linsenmaier, 1951 and C. solida Haupt, 1957 was not resolved. Included unidentified specimens grouped in three clusters, two of which are distinctly delimited and apparently represent cryptic species. The specific status of the unidentified samples in the third cluster remained unclear. Moreover, our data suggest the existence of additional cryptic species currently lumped under the names C. pseudobrevitarsis C. schencki Linsenmaier, 1968. In conclusion, our results derived from analysis of mitochondrial loci strongly support the specific status of the majority of currently recognised species in the Chrysis ignita species group, and suggest the existence of additional cryptic species in Northern Europe. Thus, considering the difficulties that often arise during species determination based on morphological characters, the mtDNA loci used here appear highly suitable for assisting species delimitation in this group as well as identification of specimens.

Phylogenetic relationships and biogeography of the Ipsiura cuckoo wasps (Hymenoptera: Chrysididae

Phylogenetic studies addressing relationships among chrysidid wasps have been limited. There are no hypotheses proposed for the Neotropical lineages of Chrysidini other than the classic cladogram published in the 1990s by Kimsey and Bohart. Herein we present a cladistic analysis based on 64 morphological characters coded for 54 species of Chrysidini, 32 of them being Ipsiura and 22 representing Caenochrysis, Chrysis, Exochrysis, Gaullea, Neochrysis, and Pleurochrysis. The species of Ipsiura were recovered as monophyletic and as the sister clade of Neochrysis in all most parsimonious trees. We discuss the high plasticity of some morphological characters as evidenced by their high homoplasy in the phylogenetic results, and we clarify the main morphological changes inferred on the phylogenetic tree for this genus. The effects of the inferred homoplasy were evaluated under an implied weighting cladistic analysis, and from a probabilistic perspective with Bayesian inference. Those alternative strategies did not alter the general conclusions about the monophyly of Ipsiura or the generic relationships in Chrysidini (changes were noticed in the species-level relationships within certain parts of Ipsiura, where low branch support was common across all approaches). Among the species groups proposed by Linsenmaier (1985), only the marginalis group was recovered as monophyletic. We also evaluated the convoluted biogeographic history of the group. The resulting historical reconstructions indicate a complicated scenario of diversification of these wasps in the Neotropics, and a close association with forested biomes is discussed.

Testing the validity of Northern European species in the Chrysis ignita species group (Hymenoptera: Chrysididae) with DNA Barcoding

Zootaxa, 2014

Containing more than a hundred species, the Chrysis ignita species group is the largest and one of the most taxonomically challenging groups in its genus. It has not been possible to resolve the taxonomy of the group using traditional methods due to the lack of robust diagnostic morphological characters. Here we present the results of a molecular analysis designed to delimit species in the Chrysis ignita group for the first time; using mitochondrial sequence data for 364 in-group specimens consisting of all 18 species known to occur in Northern Europe. Two mitochondrial loci were analysed: a COI gene fragment, and a continuous DNA sequence consisting of 16S rRNA, tRNA Val , 12S rRNA and ND4. Two approaches were employed for delimiting species: (1) genetic distance analysis based on the standard COI barcode sequences and; (2) phylogenetic analysis of the COI fragment together with rRNA genes. Both analyses yielded trees with similar topology, but support values for nodes were higher using the second approach. Fifteen species were distinguished in all analyses: and C. vanlithi Linsenmaier, 1959. The specific status of C. mediata Linsenmaier, 1951 and C. solida Haupt, 1957 was not resolved. Included unidentified specimens grouped in three clusters, two of which are distinctly delimited and apparently represent cryptic species. The specific status of the unidentified samples in the third cluster remained unclear. Moreover, our data suggest the existence of additional cryptic species currently lumped under the names C. pseudobrevitarsis C. schencki Linsenmaier, 1968. In conclusion, our results derived from analysis of mitochondrial loci strongly support the specific status of the majority of currently recognised species in the Chrysis ignita species group, and suggest the existence of additional cryptic species in Northern Europe. Thus, considering the difficulties that often arise during species determination based on morphological characters, the mtDNA loci used here appear highly suitable for assisting species delimitation in this group as well as identification of specimens.

Resolving Phylogeny at the Family Level by Mitochondrial Cytochrome Oxidase Sequences: Phylogeny of Carrion Beetles (Coleoptera, Silphidae)

Molecular Phylogenetics and Evolution, 2000

We investigated the phylogenetic relationships of carrion beetles (Coleoptera, Silphidae) using 2094 bp of their mitochondrial cytochrome oxidase subunit I and II and tRNA leucine gene sequences. Shorter fragments of this gene region previously have been used to establish generic relationships in insects. In this study, they provided more than sufficient resolution, although the third positions of the protein-coding sequences reached saturation for the deeper divergences. This first published phylogeny for the Silphidae comprises 23 species from 13 genera sampled across the geographic range of the family. In addition, we included species from three related families as outgroups. One of these families, the Agyrtidae, was, until recently, included in the Silphidae, but its resolution here justifies its current position as a separate family. The silphid subfamilies Nicrophorinae and Silphinae are monophyletic in all analyses. All genera for which several species were sampled are supported as monophyletic groups, with the exception of the genus Silpha. European and North American representatives of two Nicrophorus species described from both continents are supported as each others' closest relatives. The lineage that colonized Gondwanaland and that most likely originated in the Palearctic is the most basal within the Silphinae.

Molecular phylogeny of Diabrotica beetles (Coleoptera: Chrysomelidae) inferred from analysis of combined mitochondrial and nuclear DNA sequences

The phylogenetic relationships of thirteen Diabrotica (representing virgifera and fucata species groups) and two outgroup Acalymma beetle species (Coleoptera: Chrysomelidae) were inferred from the phylogenetic analysis of a combined data set of 1323 bp of mito-chondrial DNA (mtDNA) cytochrome oxidase subunit 1 (COI) and the entire second internal transcribed spacer region (ITS-2) of nuclear ribosomal DNA of 362 characters. Species investigated were D. adelpha , D. balteata , D. barberi , D. cristata , D. lemniscata , D. longicornis , D. porracea , D. speciosa , D. undecimpunctata howardi , D. u. undecimpunctata , D. virgifera virgifera , D. v. zeae , D. viridula , and outgroup A. blandulum and A. vittatum. Maximum parsimony (MP), minimum evolution (ME), and maximum likelihood (ML) analyses of combined COI and ITS-2 sequences clearly place species into their traditional morphological species groups with MP and ME analyses resulting in identical topologies. Results generally confer with a prior work based on allozyme data, but within the virgifera species group, D. barberi and D. longicornis strongly resolve as sister taxa as well as monophyletic with the neotropical species, D. viridula , D. cristata and D. lemniscata also resolve as sister taxa. Both relationships are not in congruence with the prior allozyme-based hypothesis. Within the fucata species group, D. speciosa and D. balteata resolve as sister taxa. Results also strongly supported the D. virgifera and D. undecimpunctata subspecies complexes. Our proposed phylogeny provides some insight into current hypotheses regarding distribution status and evolution of various life history traits for Diabrotica .

Phylogeny and classification of the extant basal lineages of the Hymenoptera (Insecta)

Zoological Journal of the Linnean Society, 2001

The phylogeny of the basal hymenopteran lineages, including representatives of all 'symphytan' families, is analysed. In total, 236 morphological characters were scored for 44 exemplars, including six outgroup, two xyelid, 16 tenthredinoid, five pamphilioid, three cephoid, five 'siricoid', one orussid, and six apocritan taxa. The dataset is analysed with parsimony under equal weights and under implied weights. The monophyly of the Hymenoptera is strongly supported but the sistergroup of the Hymenoptera cannot be identified with confidence. The relationships of the 'symphytan' lineages are Xyeloidea + (Tenthredinoidea + (Pamphilioidea + (Cephoidea + (Anaxyelidae + (Siricidae + (Xiphydriidae + (Orussoidea + Apocrita))))))). Many of the relationships between the superfamilies, especially in the basal branching pattern, are only weakly corroborated. The monophyly of most superfamilies is well supported, and all may be monophyletic except the 'Siricoidea', which is clearly paraphyletic. It is difficult to decide whether the Siricidae or the Anaxyelidae are the closest relatives of Xiphydriidae + (Orussoidea + Apocrita). The support for the sistergroup relationship between the Orussoidea and the Apocrita is substantial, putative synapomorphies being provided by most character systems. There is also good evidence in favour of the monophyly of the Apocrita. The internal phylogeny of the Tenthredinoidea differs considerably from the results of earlier analyses. The Blasticotomidae are the sistergroup of the Tenthredinoidea S.S. Relationships at the base of the Tenthredinoidea S. S. are weakly supported. It is uncertain whether the Tenthredinidae are monophyletic or comprise a basal paraphyletic grade within the Tenthredinoidea S.S. The Diprionidae may be the sistergroup to the Cimbicidae + (Argidae + Pergidae). Most relationships within the Cimbicidae + (Argidae + Pergidae) clade are well corroborated, with the exception of the monophyly of the Argidae. It is proposed to elevate the Anaxyelidae and the Xiphydriida both to superfamily status. The family-level classification of the Tenthredinoidea will probably also have to be changed, but this must await further clarification of the phylogeny of this superfamily.

Phylogeny of Leafhopper Subgenus Errhomus (Erronus) (Hemiptera: Cicadellidae) Based on Mitochondrial DNA Sequences

Annals of the Entomological Society of America, 2005

The Nearctic leafhopper genus Errhomus Oman (Hemiptera: Cicadellidae) exhibits high host-plant selectivity and female brachyptery, which may limit gene ßow. Two subgenera, Errhomus (Errhomus) and Errhomus (Erronus), are recognized based on morphological and behavioral differences. We used mitochondrial DNA sequencing to elucidate the phylogenetic relationships among members of the subgenus Errhomus (Erronus). Portions of the NADH dehydrogenase subunit 1 (ND1) and the leucine (CUN) transfer RNA (tRNA-Leu) genes were sequenced and were analyzed with parsimony and maximum likelihood. Our results generally veriÞed the Þndings of Hamilton and Zack (1999); however, both optimality criteria revealed a paraphyletic Errhomus (Erronus). Errhomus (Erronus) calvus does not seem to be a member of the subgenus Errhomus (Erronus) but seems to be more closely aligned with the subgenus Errhomus (Errhomus). We tested two a priori null hypotheses: 1) that the phylogeny estimated from the molecular data matched a previous phylogeny based on morphology and biogeography from Hamilton and Zack (1999) and 2) that diagnostic characters of the male genitalia (serrated aedeagal processes and triangular aedeagal shaft tip) were synapomorphies that deÞned major clades. We rejected these hypotheses with both Bayesian tests and parametric bootstraps.

Simultaneous analysis of the basal lineages of Hymenoptera (Insecta) using sensitivity analysis

Cladistics, 2002

The first simultaneous analysis of molecular and morphological data of basal hymenopterans that includes exemplars from all families is presented. DNA sequences (of approximately 2000–2700 bp for each taxon) from the nuclear genes 18S and 28S and the mitochondrial genes 16S and CO1 have been sequenced for 39 taxa (four outgroup taxa, 29 symphytans, and six apocritans). These DNA sequences and 236 morphological characters from Vihelmsen [Zool. J. Linnean Soc. 131 (2001) 393] were analyzed separately as well as simultaneously. All analyses were performed on unaligned sequences, using the optimization alignment (¼direct optimization) method. Sensitivity analysis sensu Wheeler [Syst. Biol. 44 (1995) 321] was applied by analyzing the data under nine different combinations of analysis parameter values. The superfamily level relationships of basal hymenopterans as proposed by Vilhelmsen [Zool. J. Linnean Soc. 131 (2001) 393] and Ronquist et al. [Zool. Scr. 28 (1999) 13] are mostly confirmed, except that Pamphilioidea is the sister group to Tenthredinoidea s.l. and that Anaxyelidae (i.e., Syntexis libocedrii) and Siricidae are supported as a monophyletic group, partly reestablishing the traditional concept of Siricoidea. The resulting hypothesis that best represents the combined evidence from morphology and DNA sequences is (Xyeloidea (Tenthredinoidea s.l. Pamphilioidea) (Cephoidea (Siricoidea (Xiphydrioidea (Orussidae Apocrita))))), with Siricoidea¼Anaxyelidae + Siricidae. The phylogenetic system within Tenthredinoidea s.l., derived from the combined evidence, is (Blasticotomidae (Tenthredinidae including Diprionidae (Cimbicidae (Argidae Pergidae)))).