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.     

A new genus and subfamily of Typhloplanidae (Platyhelminthes,Rhabdocoela) from Australia with a cladistic analysis of subfamily relationships

During a survey of marine interstitial Platyhelminthes of North Stradbroke Island, Queensland, Australia, a new taxon of Typhloplanidae (Rhabdocoela) was found, namely Cephalopharynx cannoni gen. nov. sp. nov. This new monotypic genus combines a suite of characters that set it apart from the remaining 63 genera present in the known eight subfamilies. It is unique in the lack of adenal and dermal rhabdites, the anterior position of the pharynx rosulatus, presence of a single anterior testis, and novel stylet structure. These character states substantiate the erection of a new subfamily, Cephalopharynginae, based on cladistic criteria. A key to the subfamilies of Typhloplanidae is provided.     

Generic classification of the family Pipunculidae (Diptera): a cladistic analysis

Classification of the higher taxa of the Pipunculidae is analysed by cladistic techniques. Twenty-five taxa are studied, and 117 character states are examined for these taxa. Based on the cladistic analysis a classification is proposed, recognizing three subfamilies: Chalarinae, Nephrocerinae, and Pipunculinae. The subfamily Pipunculinae is separated into five tribes: Pipunculini, Cephalopsini trib. nov; Microcephalopsini trib. nov., Eudorylini trib. nov., and Tomosvaryellini. Monophyletic genus groups within these tribes are recognized. The position of Claraeosphaera is unclear. Twenty-three taxa of generic status are recognized, and Wittella is considered synonymous with Cephalops. The genus Collinias is revalidated. Claraeomorpha is retained as a subgenus of Dorylomorpha. A presumed phylogenetic history of the taxa and evolutionary transformation series for certain characters are extrapolated from the analysis.     

V.—A note on the synonymy of three species of fossil crocodile of De Blainville

Summary

The Rhagionidae are reclassified on the basis of a review of all of the known genera. One new subfamily and 2 new genera are erected and keys are given for the separation of subfamilies and genera. Omphalophora is treated as a synonym of Ptiolina. It is almost certain that Archicera and Spatulina fall into synonymy with Spania and Ptiolina respectively. Four subfamilies and 21 genera are thus recognized and their supposed phylogenertic relationships are given in fig. 1. Desmomyia may be identical with Rhagio. If true Neorhagio is a synonym of Atherimorpha, ‘Neorhagio’ in this paper represents a new genus.

The female terminalia are here considered to be one of the most important characters in subfamily diagnoses in Rhagionidae, but it is not necessarily certain that Pseudoerinna belongs to Glutopinae and Bolbomyia and Alloleptis to Rhagioninae.

I believe that Austroleptis and Glutops belong to the family Rhagionidae based chiefly on the structure of genitalia in both sexes, although further study will be necessary to establish their systematic positions.     

Phylogenetic relationships of millipedes in the subclass Penicillata (Diplopoda) with a key to the genera

Bristly millipedes (subclass Penicillata, order Polyxenida) are minute diplopods characterised by uncalcified cuticle and a body covered with unique tufts of bristles. The order is found worldwide and comprises less than 200 described species divided into three families, with many of the species and genera being poorly known. The first evolutionary analysis of the order presented here utilises both molecular (COI, 18S rRNA and 16S rRNA) and morphological data to examine monophyly of the families and subfamilies and the evolutionary relationships between them. Maximum likelihood analysis was based on molecular data only, whereas parsimony analyses were based on molecular data as well as combined morphological and molecular data. The results of these analyses with two different optimality criteria were incongruent in many aspects. Unlike parsimony, the likelihood result found strong support for a basal position of the family Synxenidae and separation of the order into two monophyletic clades corresponding to the two superfamilies Synxenoidea, containing the family Synxenidae, and Polyxenoidea, containing the families Polyxenidae and Lophoproctidae. Parsimony results did not support the existence of the two superfamilies. Both analyses resolved the family Synxenidae as monophyletic and Polyxenidae as polyphyletic, whereas the family Lophoproctidae was shown to be paraphyletic in likelihood and monophyletic in parsimony analysis. The subfamilies Monographinae and Polyxeninae were found to be monophyletic in the likelihood tree but parsimony suggested paraphyly of both. The results suggest that further revision of the systematics of the Polyxenida may be necessary. However, a much larger molecular data set will be necessary to clarify and provide stronger nodal support for phylogenetic trees and to confirm the relationships, particularly of the families Polyxenidae and Lophoproctidae. Molecular identification is likely to be an important tool for this taxonomically challenging order in future. A simplified key to the genera is provided.     

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.     

Is the Caprellidea a monophyletic group?

Cladistic analysis of caprellidean genera has revealed major divergence near the outgroup producing two major lines of evolution: Pseudoprotomima-Perotripus, and Caprogammarus-Caprella. The most plesiomorphic genus of each line, Pseudoprotomima and Caprogammarus, respectively, shows different plesiomorphic characters to each other. Pseudoprotomima has pereopods III and IV with 6 articles, antenna II flagellum with > 2 articles and three pairs of gills, while Caprogammarus has an abdomen 5-segmented with three pairs of pleopods. Pseudoprotomima and related genera are distributed from the temperate regions of the northern hemisphere to the South Indian Ocean, while Caprogammarus and related genera are recorded primarily from the boreal region of the northern hemisphere. That pereopods III and IV in Pseudoprotomima and its relatives have 6 articles indicates the functional gap from the Podoceridae (Gammaridea)-Caprogammaridae-Caprellidae gradation, since in the Podoceridae to the Caprellidae these pereopods are not used for clinging to substrata and, even in the Podoceridae, are reduced in size. This shows the possibility that the Caprellidea is not a monophyletic group. A new phylogeny derived from the present study is proposed for future study.     

A new species of Fabricinuda Fitzhugh, 1990 (Fabriciinae: Sabellidae: Polychaeta) from the Caribbean,with an emendation of the genus

During a survey of the infauna of seagrass meadows in Venezuela, several specimens of an unknown species of Fabriciinae were collected. The species could not be assigned to any genus of the subfamily, although it was most closely allied to Fabricinuda Fitzhugh, 1990. However, the absence of pseudospatulate inferior notochaetae on chaetiger 8 prevented assignment of the specimens to this genus since their presence on chaetigers 3 to 8 is one of its diagnostic apomorphies. In order to decide whether to establish a new genus for the species or to redefine Fabricinuda, a phylogenetic analysis of the subfamily was made on the basis of a pre‐existing character matrix. Twenty‐two anatomical characters were analysed for 59 species including the new one, and from the resulting consensus tree it was clear that the new species must be included within Fabricinuda and consequently an emendation of the genus is provided herein.     

Phylogenetic implications of ovipositor structure in Eucharitidae and Perilampidae (Hymenoptera: Chalcidoidea)

The external and cross-sectional morphology of the ovipositor was examined for 21 genera and 28 species of Philomidinae, Perilampidae and Eucharitidae. Eleven characters were scored and either mapped onto existing topologies reflecting the relationships of genera, or included in a larger data set focused on the generic relationships of Eucharitidae and analyzed using parsimony. The presence of a setiform flap in the ventral wall of the dorsal valve supported the monophyly of Perilampinae and possibly Chrysomalla (Chrysolampinae) + Perilampinae. Monophyly of Eucharitidae was supported by a thinning of the dorsal wall of the dorsal valve. One of two islands of most parsimonious trees that have recently been proposed was selected on the basis of fewer step changes as a result of both character mapping and parsimony analysis. Within Eucharitinae, this would support a clade that includes Kapala, Stilbula and Pseudochalcura, with Schizaspidia and Chalcura as its sister group. By mapping characters, an additional hypothesis of monophyly of Anorasema and Gollumiella was preferred, which on the basis of the pruned topologies was a single step shorter than the preferred islands of trees.     

Phylogenetic relationships among genera in the Calanidae (Crustacea: Copepoda) based on morphology

Cladistic analysis of Calanidae recovered two major clades: a tropical epipelagic clade composed of species without ontogenetic vertical migration (Canthocalanus + Cosmocalanus + Nannocalanus + Undinula); and a clade of ontogenetically migrating genera (Neocalanus + Calanoides + Calanus + Mesocalanus). The latter clade is least well supported although its Calanoides and Neocalanus lineages are well supported. Morphology-based topologies are largely congruent with published molecular trees, differing chiefly in the positions of Mesocalanus and Calanus. Independent homoplasious trends in character states in the male leg 5 include: reduction in segmentation and number of setae, increasing asymmetry as well as the probable retention of larval characteristics. The monophyly of Calanoides, Neocalanus and Calanus is well supported. It is postulated that Calanidae radiated during the mid to late Triassic from a bathypelagic megacalanid which was adapted to low oxygen conditions. High latitude locations may have been the first epipelagic environments to be recolonized.     

Phylogenetic relationships among genera of the tribe Cnephasiini (Lepidoptera: Tortricidae: Tortricinae) based on morphological characters of adults

A phylogenetic analysis was conducted on the moth tribe Cnephasiini based on 83 morphological characters of adults (63 binary and 20 multistate). The analysis included 26 species representing 20 genera, plus two outgroup representatives of the tribe Tortricini. The cladistic analysis resulted in the 10 most parsimonious trees. The tribe Cnephasiini was recovered as monophyletic; the genus Cnephasia was not recovered as monophyletic; the New World Decodes clustered with the primarily Old World Cnephasia; Oxypteron is best treated as monotypic (with the single species palmoni), and Amphicoecia adamana, Oporopsamma dunaria and O. wertheimsteini should be assigned to a different genus; Propriomorpha and Xerocnephasia were the most basal groups; Exapate duratella and Epicnephasia mongolica were recognised as sister-groups; and Kawabeia was closest to Tortricodes.     

A character analysis of the North American potter wasps (Hymenoptera: Vespidae; Eumeninae)

A cladistic analysis of the nearctic genera of Eumeninae is presented. The ground plan states of 43 character systems are discussed, and the first comprehensive cladogram for any significant portion of the subfamily is presented. At least eight of the 26 genera are apparently nonmonophyletic: Zethus, Montezumia, Euodynerus, Odynerus, Pterocheilus, Stenodynerus, Leptochilus and Microdynerus. A key to the nearctic genera accompanies the text.     

Observations on the morphology and classification of weevils (Coleoptera,Curculionoidea) with a key to major groups

A brief account of the present state of weevil taxonomy is followed by a detailed study of certain structures used in their classification, namely the venter, abdominal tergites, sternite 8 of the male, apex of the hind tibia and deciduous mandibular processes. A key to some 50 families and subfamilies of Curculionoidea is followed by a list of family-group taxa. The following changes are made: Brachyceridae, Erirhinidae, Cryptolaryngidae and Raymondionymidae are promoted to family rank from Curculionidae; Antliarhininae is demoted to a subfamily of Brentidae, and Allocoryninae to a subfamily of Oxycorynidae; Coptonotini is demoted to a tribe of Curculionidae-Scolytinae; Carinae, Subfam. n. is erected for Car Blackburn (genus incertae sedis) is Belidae; Dinomorphini is demoted to a tribe of Molytinae and Brachyceropsidinae is revived from synonymy with Dinomorphinae (Curculionidae); Brachyderini, Eremnini, Otiorhynchini and Sitonini are demoted to tribes of Entiminae; Desmidophorinae is transferred from Brentidae to Brachyceridae; Ocladiini is promoted to a tribe of Desmidophorinae (from Curculionidae-Cryptorhynchinae); Campyloscelini (including Phaenomerina) is transferred from Rhynchophoridae to Curculionidae-Zygopinae; Carphodicticinae is promoted to subfamily rank and transferred from Curculionidae-Scolytinae to Platypodidae; Perieges Schönherr is transferred from Curculionidae-Thecesterninae to Cryptolaryngidae and Agriochaeta Pascoe from Cryptorhynchinae to Hyperinae (Curculionidae); Schedlarius Wood and Mecopelmus Blackman are transferred from Coptonotidae to Platypodidae.     

Successful identification of the final instar nymph of Quintilia carinata (Thunberg) (Hemiptera: Cicadidae) by DNA extraction from the exuvium

Despite being taxonomically and phylogenetically informative, the morphology of the immature stages of cicadas has received comparatively superficial attention. One reason for this is the difficulty of positively identifying immature stages, particularly as these stages are fossorial. We present a method for identifying cicada exuviae using DNA sequence data and describe a set of characters and character states for the final instar nymph of Quintilia carinata (Thunberg). The identification of immature stages using molecular methods will increase our knowledge of African cicadas, allowing for the initiation of future phylogenetic and ecological comparisons.     

A new genus and two new species of Philotarsidae. (Insecta: Psocoptera) from East Africa

Summary

Genus Tarsophilus and its species lineatus and aequabilis (Insecta: Psocoptera: Philotarsidae) are described as new. The specimens, all males, were collected in highland areas of East Africa. The new genus is placed in Philotarsidae rather than Pseudocaeciliidae on characters of hypandrium, phallosome, and lacinial tip. Its position in the subfamily Philotarsinae is justified and it is compared to the other genera of this group on both male and on trans-sexual characters.     

A new species of Nebalia (Crustacea,Leptostraca) from Unguja Island (Zanzibar), Tanzania,East Africa,with a phylogenetic analysis of leptostracan genera

Nebalia brucei sp. nov. is described from specimens taken in traps from Unguja Island (Zanzibar), Tanzania, East Africa. The species is characterized by the following unique combination of characters: a short and broad rostrum; a carapace that is large compared to overall body size and that covers pleonite 4; a basally narrow compound eye; and acutely pointed denticles at the dorsal margin of pleon segments 6 and 7. Cladistic analyses of the relationships of the seven recognized leptostracan genera resulted in two hypotheses. One tree has Nebaliopsis as sister group to the remaining leptostracans and supports the monophyly of Nebaliidae. The other tree has Paranebalia as sister group to the remaining leptostracans and Nebaliidae as paraphyletic with respect to Nebaliopsis. Both trees contain a Nebalia/Dahlella/Sarsinebalia clade.     

Ancestrular morphology in cyclostome bryozoans and the quest for phylogenetically informative skeletal characters

Cyclostomes are an ancient order of bryozoans with more than 500 living species. Traditionally, cyclostome taxonomy has depended almost entirely on characters of the calcareous skeletons of mature colonies. However, the taxonomic distribution of these characters is largely incongruent with well-supported molecular phylogenetic trees, implying the need for major revisions if cyclostome taxonomy is to reflect phylogeny. While far more taxa must be sequenced to ascertain their molecular relationships, there is also a need to discover morphological characters that can be used to support molecular clades, not least to allow these clades to be inferred among fossil cyclostomes for which molecular data will never be available. Here we survey the skeletal morphology of the first-formed part of the skeleton – the protoecium of the founding zooid (ancestrula) – in a variety of Recent and well-preserved fossil cyclostomes, motivated by the recent discovery of a morphologically disparate molecular clade that can be distinguished by protoecial characters. The distribution of pseudopores on the protoecium is shown to vary between species, with eight patterns recognizable, offering a new source of skeletal characters in both living and fossil taxa. However, the patterns seem not to be fully congruent with the cyclostome molecular tree and more data and analyses are required before these patterns can be used to recognize cyclostome clades.     

Mastacembelidae (Teleostei; Synbranchiformes) subfamily division and African generic division: an evaluation

The family Mastacembelidae is currently divided into two subfamilies, the Mastacembelinae and the Afromastacembelinae. Within the African Afromastacembelinae two genera, Caecomastacembelus and Aethiomastacembelus, are recognized. Both the subfamily and the African generic division were critically evaluated mainly through X‐ray analyses and osteological research. There is no supporting evidence for a split into two subfamilies. In addition, the present diagnosis of the African genera proved to be unworkable, and no convincing phylogenetic evidence supporting this classification is available. Therefore, pending new detailed phylogenetic research, the use of a subfamily classification within the family Mastacembelidae is not recommended. Furthermore, Caecomastacembelus and Aethiomastacembelus are placed in synonymy with the genus Mastacembelus.     

The most primitive Scoliidae (Hymenoptera)

Summary

The subfamily Proscoliinae Rasnitsyn, 1977, (Scoliidae) was based on a single male from Armenia. Following discovery of a second species known from both sexes, the Proscoliinae and Scoliinae are redefined as subfamilies of Scoliidae, and keyed. The new species is described and figured; biological observations are given. The relationships of Scoliidae are discussed in the light of recent work. Taxonomic observations are made on the genera Colpa Dufour, 1841 (Scoliidae; Scoliinae) and Paramyzine Berg, 1898 (= Menius Buysson, 1898) (Tiphiidae; Anthoboscinae); new generic and specific synonymies are recorded.