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

Abstract

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.     

Phylogenetic evaluation of the tropical Camptotypus genus-group (Hymenoptera: Ichneumonidae), with a key to the world genera

The phylogeny of the disjunct pantropical Camptotypus genus-group (Hymenoptera, Ichneumonidae, Pimplinae) was reconstructed using 40 morphological characters.The monophyly of many established genera (Parvipimpla, Clydonium, Odontopimpla) was found to be supported, but Hemipimpla was found to nest within Camptotypus and has thus been treated as a junior synonym of it (syn. n.). A group of undescribed Neotropical species were found to form a separate monophyletic group, herein referred to as Genus A. Species of the Neotropical genus Zonopimpla formed a basal paraphyletic grade which could not satisfactorily be resolved into discrete monophyletic units. Rather than subsuming all the more derived genera within one large group, or attempting to erect numerous new and probably ephemeral genera (for isolated species of Zonopimpla), we propose the retention of this grade-group as a separate genus at present. The host utilization patterns of the genus-group, and the biogeography of the genus-group are discussed. The group is hypothesized as having originated and radiated in the Neotropics, as ectoparasitoids of weakly concealed lepidopterous hosts. One lineage, Clydonium, has become specialized at attacking hosts in woody galls where it utilizes coleopterous as well as lepidopterous hosts. The Old World genera (Camptotypus and Parvipimpla) comprise a monophyletic group and are suggested to have originated from a single dispersal event across the widening Atlantic. No evidence for either a southern or northern circum-polar dispersal was observed in this entirely tropical genus-group. One lineage in the Afrotropical region has switched from attacking lepidopterous hosts, to attacking pre-pupal and pupal polistine vespids.     

Genus Baseodiscus (Nemertea: Heteronemertea): Molecular identification of a new species in a phylogenetic context

A new heteronemertean, Baseodiscus jonasii sp. nov., is described from Guadalcanal, Solomon Islands. It resembles B. delineatus in inner morphology but can be distinguished from this species by its different colour pattern and differences in the nucleotide sequence of the mitochondrial 16S rRNA gene. The monophyletic status of the genus is investigated by reconstructing the phylogeny of six specimens from four species assigned to this genus, together with 22 specimens from nine other heteronemertean genera, using parsimony and Bayesian analysis. The results imply that Baseodiscus is a monophyletic group while several other heteronemertean genera are non‐monophyletic.     

The Eucerotinae: a Gondwanan origin for a cosmopolitan group of Ichneumonidae?

A new genus and species of the formerly monobasic and predominantly amphitropical, ichneumonid subfamily, Eucerotinae, is described from Chile. A phylogenetic analysis of the redefined Eucerotinae is undertaken. Barronia araucaria gen. et sp. n. has a sister-group relationship with Euceros. The southern hemisphere species of the latter genus were found to comprise a monophyletic clade, the sister group to all other Euceros species. Similarly all north temperate Euceros species form a monophyletic clade, the sister group of which occurs in Madagascar. Within the former clade, the North American species form a monophyletic group, the sister species of which is Palaearctic. Several other groups of Palaearctic taxa have a sister-group relationship to this lineage. These results suggest the Eucerotinae had an ancient origin on Gondwanaland in the lower Cretaceous, prior to the separation of the Indo-Madagascar terrane from the Australian-Antarctic-South American tectonic plate. We hypothesize the Palaearctic fauna colonized Eurasia following the impaction of India on to its southern margin, and that the North American fauna is derived from a single circum-polar dispersal from Eurasia. Our results do not support an earlier hypothesis that the Eucerotinae arose as hyperparasitoids of Symphyta. Evidence suggests the group arose as parasitoids of ophioniform ichneumonids attacking lepidopterous larvae feeding on gymnosperms or anemophilous angiosperm trees.     

A key to Australasian and Oriental Woodiphora (Diptera: Phoridae), affinities of the genus and description of new species

The taxonomic value of the notopleural cleft in Phoridae is highlighted. A cladogram relating Gymnophora and Woodiphora to Megaselia is presented. Recognition of the genus Woodiphora is clarified, and the Old World species of Haplophleba transferred to Woodiphora. The evolution of the female abdominal sternite 9 is discussed. The tribe Megaseliini is proposed. A key to the Australasian and Oriental species of Woodiphora is provided. Twenty new species are described and two new synonyms proposed.     

The phylogenetic position of Thermophis (Serpentes: Colubridae), an endemic snake from the Qinghai‐Xizang Plateau,China

The snakes of the genus Thermophis, including two species (Thermophis baileyi and Thermophis zhaoermii), which are found only on the Qinghai‐Xizang Plateau, were studied to determine the phylogenetic position of the genus within the broad superfamily Colubroidea. In this study, T. baileyi from two different localities in Xizang and T. zhaoermii from western Sichuan formed a monophyletic group as determined using Bayesian and likelihood methods of phylogenetic inference. Based on the nucleotide sequence of the cyt b and c‐mos genes, Thermophis appeared to be a member of the subfamily Xenodontinae, which prior to this finding had been confined to a New World distribution. The molecular data also indicate a large genetic distance between T. baileyi and T. zhaoermii, which strengthens the validity of the new species: T. zhaoermii.     

Seed-feeding beetles of the weevil tribe Mecysolobini (Insecta: Coleoptera: Curculionidae) developing in seeds of trees in the Dipterocarpaceae

Species comprising two monophyletic groups of Alcidodes sensu lato (crassus group and dipterocarpi group) feed on fruits of the ecologically and economically important tropical timber family Dipterocarpaceae. Fifteen Alcidodes species are newly described from several thousand specimens of seed-feeding beetles reared from south-east Asian dipterocarp hosts (Dipterocarpoidae) and four other species are revised. Morphological keys are provided for identifying these 19 taxa. Weevil associations are provided for 70 species of Dipterocarpaceae in five genera (16 Dipterocarpus spp., four Dryobalanops spp., six Hopea spp., 39 Shorea spp. and five Vatica spp.). These records relate primarily to specimens reared from seeds in Borneo and in Peninsular Malaysia, but all known previous host records are also included (from India, Andaman Islands, Thailand, Laos, Vietnam and the Philippines) and assessed. These validated and documented host associations refute general assertions that (1) seed-eating beetles are host-specific in the tropics and (2) seed-eating beetles found on dipterocarp hosts are broad generalists across species in several genera of mast-fruiting Dipterocarpaceae. Most Alcidodes species exploit seeds from several dipterocarp species. Some Alcidodes species occur across broad geographic regions with several host species from a dipterocarp genus, other species have been found only on Bornean Hopea or Dipterocarpus species. Others, such as the two previously undescribed species feeding on Vatica species, may be restricted to separate sections of the host genus. Within an extensively sampled and diverse interspecific mast-fruiting dipterocarp community, no Alcidodes species was observed to feed on sympatric species across genera within the Dipterocarpaceae. Additional host records and specimens of Alcidodes spp. throughout the range of the Dipterocarpaceae in south and southeast Asia are required to further define these clades and the tribe Mecysolobini in general as well as to resolve the complex host associations and distributions documented.     

A synopsis of the subterranean mealybug genus Neochavesia Williams and Granara de Willink (Hemiptera: Pseudococcidae: Rhizoecinae)

The neotropical mealybug genus Neochavesia, associated with the ant genus Acropyga Roger, is discussed and placed in the tribe Xenococcini on the bases that it lacks dorsal ostioles, the distal end of the circulus is depressed and cup-shaped, and the third-instar female is a pupa. These are characters shared by Xenococcus Silvestri and Eumyrmococcus Silvestri, also in the tribe Xenococcini, and differ from those in the tribe Rhizoecini, the other tribe in the subfamily Rhizoecinae. Two new species, Neochavesia iwokramae sp. nov. and N. lapollai sp. nov., are described from Guyana, and the female pupal instar and adult male of N. lapollai and the adult male of N. caldasiae (Balachowsky) are described and illustrated. A key to adult females is provided.     

Phylogenetic analysis of the subgenus Lampetis (Spinthoptera) (Coleoptera: Buprestidae) of North and Central America,and the West Indies

Phylogenetic relationships between species of Lampetis (Spinthoptera) of North and Central America and the West Indies were analysed by cladistic methods, based on 65 characters from the external morphology and male genitalia. Eleven species of different generic groups of the subtribe Dicercina were considered as outgroups, including also species of Lampetis (Lampetis) and one South American species of L. (Spinthoptera). The monophyly of Lampetis (Spinthoptera) is supported by two synapomorphies, but this subgenus appears to be more closely related to species of Psiloptera rather than to Old World species of Lampetis (Lampetis), suggesting that Lampetis (Spinthoptera) may be segregated from Lampetis as a different genus. The species of Lampetis (Spinthoptera) from North and Central America and the West Indies do not represent a monophyletic taxon, because L. (S.) tucumana (South American outgroup) is nested within them. Comparison of these results with a previous panbiogeographic analysis indicate that several generalized tracks possess species from different clades, thus suggesting an ancient radiation of this taxon in Mesoamerica and the Mexican Transition Zone.     

Borneodessus zetteli,new genus and new species,notes on biogeography and a checklist of Dytiscidae from Borneo (Insecta: Coleoptera: Dytiscidae)

A new genus of Dytiscidae, Borneodessus gen. n., is described for Borneodessus zetteli zetteli sp. n. (Sabah, East Malaysia) and Borneodessus zetteli kalimantanensis ssp. n. (East Kalimantan, Indonesia). The new genus belongs to the tribe Bidessini because of the presence of two-segmented parameres of the aedeagus. A modified male clypeus is an apomorphy for Borneodessus zetteli. The species habitually resembles some species of Africodytes Biström, 1988 (Africa), Amarodytes Régimbart, 1900 (South America), and some Clypeodytes Régimbart, 1894 (Old World tropics). However, a sister group relation or closer affinities to any of the genera of Bidessini could not be established. Borneodessus is so far the first genus of Dytiscidae which is endemic to Borneo, but we suggest a certain probability of its occurrence in other Sundaic areas. The species is a rheobiont, being collected from forest streams. We briefly address biogeographical aspects and provide a checklist of Bornean Dytiscidae, indicating the currently known degree of endemism.     

Revision of the millipede genus Xystodesmus,with reference to the status of the tribe Xystodesmini (Diplopoda: Xystodesmidae)

On the basis of morphometrics and geographic distribution, we revise Xystodesmus martensii Peters and its related forms in Japan, which had been recognized as ten species belonging to five genera. In delimitation of species, first we sought the biggest pheno-geographic units based on the dendrogram from a cluster analysis and geographic distribution; a pheno-geographic unit is defined as an aggregation of populations being phenetically closely similar and geographically closely distributed to each other. Then, we decided whether each unit was warranted as a species. As a result, we propose that these forms constitute a single genus Xystodesmus comprising six species including one new species. The newly defined Xystodesmus is characterized by high variation in gonopod morphology and conservative coloration, body form and size. The status of some populations is still uncertain because of lack of material. The tribe Xystodesmini, to which Xystodesmus belongs, and the tribe Harpaphini are so similar that we could not warrant their separation. We propose the following new synonymies: the tribe Harpaphini with the tribe Xystodesmini; Phrurodesmus Takakuwa and Nikkonus Chamberlin and Wang with Xystodesmus; Xystodesmus scabra Verhoeff, Rhysodesmus tuberculatus Takakuwa, Rh. ikaoensis Takakuwa, and Rh. kitazawai Miyosi, with X. martensii; Phrurodesmus kinshaensis Murakami with X. gracilipes (Takakuwa). Riukiaria shirozui Takakuwa, Rh. serrulatus Miyosi, and P. gracilipes Takakuwa, are transferred to Xystodesmus. We describe X. tokaiensis as a new species.     

New species of Hygiella Mesnil (Diptera: Tachinidae), parasitoids of leaf insects (Phasmatodea: Phylliidae)

The Oriental genus Hygiella Mesnil is revised: three species are described as new, Hygiella angustifrons sp. nov., Hygiella luteipes sp. nov. and Hygiella proclinata sp. nov., and two known species are re-described, Hygiella nigripes Mesnil and Hygiella pygidialis Mesnil. A key to these five species is provided. The host of H. proclinata, Phyllium siccifolium (Linnaeus) (Phasmatodea: Phylliidae), is recorded for the first time as a host of the Tachinidae. The systematic position of Hygiella is discussed and the genus is placed in the tribe Acemyini (Exoristinae) based on the structure of the male terminalia.     

A review of the encyrtid wasp tribe Aenasiini,with descriptions of new Afrotropical taxa (Hymenoptera: Chalcidoidea)

The tetracnemine tribe Aenasiini is discussed and a key is provided for the separation of the nine genera that can be attributed to this tribe. Neodiscodes is synonymized with Aenasius and the following seven species, which were originally assigned to the former genus, are transferred to Aenasius as new combinations: abengouroui (Risbec), comperei (Kerrich), lepelleyi (Kerrich), martinii (Compere), indicus (Narayanan & Subba Rao), parvus (Kerrich) and subbaraoi (Kerrich). A key is given for the separation of the four Afrotropical species of Aenasius and the male of A. comperei is described for the first time. Incisencyrtus gen. n. is described for the following three new Afrotropical species: secus (the type-species), sirus and afer. Cladiscodes incisius sp. n. is described from South Africa and it represents the first species of this genus from Africa. New distribution records are given for three Alfrotropical species of Metaphaenodiscus.     

Deep-sea Tanaidacea (Crustacea) of the North-east Atlantic: the tribe Agathotanaini

The tanaidacean tribe Agathotanaini, which was formerly the family Agathotanaidae, is well-represented in the Rockall Trough and Bay of Biscay areas of the North-east Atlantic. Five species have been recorded from depths of 1160–4829 m. Agathotanais ingolfi is the most numerous and one of the most widely distributed tanaidacean species in the region, and is often predominant in epibenthic sled samples from 1500–2500 m. Less numerous and frequent are three new species of Paragathotanais: P. nanus, P. robustus and P. gracilis. The first two are bathyal in distribution, and the last is abyssal. A new monospecific genus is established for Metagathotanais insulcatus, which has been recorded from the deepest part of the study region (4600–4800 m). Males of this tribe are characterized by their rigid pleopods bearing short terminal setae, and are otherwise very similar to the females. A key is provided for the identification of agathotanaids from the North-east Atlantic, including Agathotanais hanseni, and the genera Allodaposia and Paranarthrura, which should also be classified in the Agathotanaini.     

A preliminary molecular phylogeny of the Sigalphinae (Hymenoptera: Braconidae), including Pselaphanus Szépligeti,based on 28S rDNA,with descriptions of new Afrotropical and Madagascan Minanga and Malasigalphus species

Minanga phoebea sp. n. Quicke, from Uganda, and Malasigalphus roa sp. n. Sharkey, from Madagascar, are described and illustrated. The new species of Minanga displays a distinct posterior metasomal carina and thus provides another example of the co‐occurrence of this feature with a metasomal carapace within the Braconidae. Malasigalphus roa is the second species of the genus recently described by Achterberg and Austin. A preliminary molecular phylogeny of the Sigalphinae is presented including the rare Neotropical genus Pselaphanus the placement of which has been debated. Sensitivity analysis to multiple alignment parameters was conducted and a single elided data set, based on the combined 21 separate alignments, was analysed. Strong support was obtained for the following relationships: Pselaphanus+(Malasigalphus+(other Sigalphinae)), (Minanga+(Sigalphus bicolor+(S. irrorator+S. gyrodontus))), and Earinini being basal (as a grade in these analyses) to other Agathidinae. The Mesocoelus group of genera (Mesocoelini in part) are shown to be derived agathidines rather than being associated with the Orgilini. The Microtypinae is shown be non‐monophyletic and Plesiotypus Achterberg is proposed as the sister group to the Agathidinae+Sigalphinae clade. The Acampsohelconinae does not appear as monophyletic; however, the placements of both of the two included genera, Afrocampsis and Canalicephalis, had less than 100% support in the elised analysis tree, and therefore monophyly of this recently redefined subfamily must be more rigorously tested.     

A review of Coelotinae epigynal teeth morphology,with descriptions of two species from China (Araneae: Amaurobiidae)

Five Exorista species from Malaysia and Indonesia are treated and diagnostic characters of the male postabdomen are illustrated. Three of them are described as new: Exorista globosa sp. nov., E. flaviventris sp. nov. and E. sabahensis sp. nov. Phylogenetic analysis is performed with previously published morphological data along with the five species treated here. Based on the generated tree, the five species treated here are assigned to Spixomyia Crosskey. Exorista ladelli (Baranov) is transferred to Spixomyia from the subgenus Podotachina Brauer and Bergenstamm. Evolutionary patterns of host use in the genus Exorista Meigen are discussed. Host information is optimized on the resultant tree under the most parsimonious reconstruction. The original host of the tribe Exoristini (except Ctenophorinia Mesnil and Phorinia Robineau-Desvoidy) was order Lepidoptera, and transitions to other orders (Hymenoptera, Mantodea and Orthoptera) occurred in subgenera Adenia Robineau-Desvoidy and Spixomyia, respectively.     

Contributions to the knowledge of the genera Haemerophygus Baudi, 1876 and Ceratanisus Gemminger, 1870 (Coleoptera: Tenebrionidae: Ceratanisini) in the Anatolian region

Previously known species of the genus Ceratanisus Gemminger, 1870 are studied, based on a comparison of specimens. This study is a first attempt to scrutinize the composition of this poorly known genus, as a preliminary approach to the tribe Ceratanisini Gebien, 1937. As a result of this study three previously described taxa are recognized as valid species and three new species from Turkey, Ceratanisus selimi sp. nov., Ceratanisus labriquei sp. nov. and Ceratanisus andiberti sp. nov., are described. A key for the separation of the genera of Ceratanisini is presented, including the new species described herein.     

Muhaka icipe,an enigmatic new genus and species of Kleidotomini (Hymenoptera: Figitidae: Eucoilinae) from an East African coastal forest

A remarkable new eucoiline genus and species, Muhaka icipe, is described herein. The genus is clearly a Kleidotomini, but is distinguished from other genera in the tribe by a unique head and scutellar morphology. The genus belongs to the ‘wedge-head’-syndrome group of species that, to date, is unique to Afrotropical eucoilines. The new genus and species is reminiscent of Stentorceps Quinlan and Nanocthulhu Buffington, but is readily distinguished from these genera. Muhaka was collected from a threatened kaya (sacred forest) of coastal Kenya. The biological importance of this and other kaya forests, as well as their protection, is discussed.

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