Phylogenetic analysis of the Malacostraca (Crustacea)

The Malacostraca comprises about 28 000 species with a broad disparity in morphology, anatomy, embryology, behaviour and ecology. The phylogenetic relationships of the major taxa are still under debate. Is the Leptostraca the sister group of the remaining Malacostraca, or is this taxon more closely related to other Crustacea? Does the Stomatopoda or the Bathynellacea represent the most basal taxon within the remaining taxa? Is the Peracarida monophyletic or are some peracarid taxa more closely related to other ‘caridoid’ taxa? Is the Thermosbaenacea part of the Peracarida or its sister group, and how much support is there for a taxon Amphipoda + Isopoda? To answer these questions a phylogenetic analysis of the Malacostraca combining different phylogenetic approaches was undertaken. In a first step, the monophyly of the Malacostraca including the Leptostraca is shown using the ‘Hennigian approach’. A computer cladistic analysis of the Malacostraca was carried out with NONA and PEE ‐WEE , based on 93 characters from morphology, anatomy and embryology. Nineteen higher malacostracan taxa are included in our analysis. Taxa whose representatives are exclusively fossils were not included. The Leptostraca was used as an operational out‐group. The present analysis supports the basal position of the Stomatopoda. Syncarida and Peracarida (including Thermosbaenacea) are supported as monophyletic, the Eucarida is not. Instead a sister‐group relationship is suggested between Euphausiacea and Peracarida (including Thermosbaenacea), with the Syncarida as the sister group to both taxa. Certain embryonic characters are interpreted as support for the monophyly of the Peracarida (without Thermosbaenacea) because convergences or reversals of these characters seem implausible. Within the Peracarida, the Mysidacea (Lophogastrida + Mysida) represents the sister group to the remaining taxa. A sister‐group relationship between Amphipoda and Isopoda is not supported.     

An updated phylogeny of Anisoptera including formal convergence analysis of morphological characters

Family interrelationships among Anisoptera (dragonflies) are unresolved. Molecular markers applied thus far have not been particularly useful for resolving relationships at the family level. Previous morphological studies have depended heavily on characters of wing venation and articulation which are believed to display considerable degrees of homoplasy due to adaptations to different flight modes. Here, we present a comprehensive anatomical dataset of the head morphology of Anisoptera focusing on muscle organization and endoskeletal features covering nearly all families. The characters are illustrated in detail and incorporated into an updated morphological character matrix covering all parts of the dragonfly body. Phylogenetic analysis recovers all families as monophyletic clades except Corduliidae, Gomphidae as sister group to all remaining Anisoptera, and Austropetaliidae as sister group to Aeshnidae (=Aeshnoidea). The position of Petaluridae and Aeshnoidea to each other could not be resolved. Libelluloidea is monophyletic with Neopetalia and Cordulegastridae as first splits. Chlorogomphidae is sister to monophyletic [Synthemistidae + (‘Corduliidae’ + Libellulidae)]. In addition, we applied a recently published formal approach to detect concerted convergence in morphological data matrices and uncover possible homoplasies. Analyses show that especially head and thorax characters may harbour homoplasies. After exclusion of possible homoplastic characters, Gomphidae is corroborated as sister group to all remaining Anisoptera.     

Characteristics of sperm ultrastructure in the gall midges Porricondylinae (Insecta,Diptera, Cecidomyiidae), with phylogenetic considerations on the subfamily

 Sperm ultrastructure was studied in ten genera of the Porricondylinae (Cecidomyiidae). Sperm structure is remarkably simplified by the absence of the acrosome and the accessory tubules, as happens in all the cecidomyiid flies. All genera of the Porricondylinae show a peculiar 9+3 axonemal model except Diallactes, which retains the plesiomorphic condition of a 9+2 axoneme, and Winnertzia, which appears to have secondarily acquired a 9+0 model. A cladistic analysis of relevant sperm characters (based on the axonemal model, the number of mitochondrial derivatives and the size and structure of the centriolar adjunct) was performed to infer phylogenetic relationships among six tribes of the Porricondylinae. In this cladogram, the Porricondylini are the sister group to the Asynaptini, Heteropezini and Winnertzini and these four taxa form the sister group to the Dicerurini. The tribe Diallactini are regarded as the group with the most plesiomorphic characters within the family. Accepted: 15 March 1996     

Morphological appraisal of Collembola phylogeny with special emphasis on Poduromorpha and a test of the aquatic origin hypothesis

Phylogenetic relationships within Collembola were determined through the cladistic analysis of 131 morphological characters and 67 exemplar taxa representing the major families of the group, with special emphasis on Poduromorpha. The results show that the order Poduromorpha is monophyletic and the sister group to the remaining Collembola, with Entomobryomorpha monophyletic and the sister group to the clade Neelipleona + Symphypleona. In Entomobryomorpha, Actaletidae is the sister group of the remaining families. In Poduromorpha, Tullbergiinae is monophyletic as well as Onychiurinae and the group Tetrodontophorinae + Onychiurinae which is the sister group of the remaining Poduromorpha; Tetrodontophorinae is paraphyletic; Onychiuridae is polyphyletic; Isotogastruridae is not an intermediate between Poduromorpha and Entomobryomorpha, it is the sister group of Tullbergiinae; Odontellidae is monophyletic and the sister group to the clade Neanuridae + Brachystomellidae; in Neanuridae, Frieseinae and the group Pseudachorutinae + Morulinae + Neanurinae are monophyletic; Morulinae is the sister group of Neanurinae; Pseudachorutinae is paraphyletic; Hypogastruridae is polyphyletic; Podura aquatica (Poduridae) is not 'primitive', it clusters with the genera Xenylla and Paraxenylla in Hypogastruridae. On the basis of these relationships and the position of the aquatic species, the most parsimonious hypothesis is a terrestrial edaphic origin for the springtails.     

Cladistics and the comparative morphology of linyphiid spiders and their relatives (Araneae, Araneoidea, Linyphiidae)

This paper provides the first quantitative cladistic analysis of linyphiid morphology. Classical and novel homology hypotheses for a variety of character systems (male and female genitalia, somatic morphology, spinneret silk spigot morphology, etc.) are critically examined and studied within a phylogenetic context. Critical characters have been illustrated. A sample of linyphiid taxa (nine genera in four subfamilies), five species of Pimoa (Pimoidae), and two other araneoid families (Tetragnathidae and Araneidae, represented by Tetragnatha and Zygiella , respectively) were used to study the implications of the phylogeny of Pimoidae for the systematics of linyphiids. The phylogenetic relationships of these 16 exemplar taxa, as coded for the 47 characters studied, were analysed using numerical cladistic methods. In the preferred cladogram Pimoidae and Linyphiidae are sister groups, Stemonyphantinae are sister group to the remaining linyphiids, and Mynogleninae are sister group to the clade composed of Erigoninae plus Linyphiinae. These results agree with the relationships recently proposed by Wunderlich, except by finding erigonines as the sister group to linyphiines rather than to mynoglenines.     

Red & black or black & white? Phylogeny of the Araschnia butterflies (Lepidoptera: Nymphalidae) and evolution of seasonal polyphenism

Phylogeny of the butterfly genera Araschnia, Mynes, Symbrenthia and Brensymthia (Lepidoptera: Nymphalidae: Nymphalini) is reconstructed, based on 140 morphological and ecological characters. The resulting tree shows that Araschnia is a sister group of the clade including Symbrenthia, Mynes and Brensymthia (Symbrenthia is paraphyletic in the respect of remaining genera; Symbrenthia hippalus is a derived species of Mynes). The species-level relationships within Araschnia are robustly supported as follows: (A. davidis (prorsoides ((zhangi doris) (dohertyi (levana burejana))))). Analysis of the wing colour-pattern characters linked with the seasonal polyphenism in the Araschnia species suggests that the black and white coloration of the long-day (summer) generation is apomorphic. Biogeographically, the origin of polyphenism in Araschnia predates the dispersal of some Araschnia species towards the Palaearctic temperate zone, and the ecological cause of the polyphenism itself is then probably not linked with thermoregulation. The possible mimetic/cryptic scenarios for the origin of Araschnia polyphenism are discussed.     

Phylogeny and character evolution in the Empidonax group of tyrant flycatchers (Aves: Tyrannidae): a test of W. E. Lanyon's hypothesis using mtDNA sequences

We sequenced mitochondrial DNA from four protein-coding genes for 26 taxa to test W. E. Lanyon's hypothesis of intergeneric relationships and character evolution in the Empidonax group of tyrant flycatchers. Three genera in this group (Empidonax, Contopus, and Sayornis) successfully occupy north temperate habitats for breeding, while the remaining genera (Mitrephanes, Cnemotriccus, Aphanotriccus, Lathrotriccus, and Xenotriccus) are restricted to neotropical latitudes. Lanyon hypothesized two major clades in the group based on differences in syringeal morphology and proposed relationships among genera using a combination of morphologic, behavioral, and allozymic characters. The mtDNA data strongly support Lanyon's division of genera into two clades. In addition, the molecular and nonmolecular data sets agree in uniting Aphanotriccus and Lathrotriccus as sister taxa, with Cnemotriccus as basal to these genera. Species of Aphanotriccus, Lathrotriccus, and Cnemotriccus form a clade that exploits a distinctive nesting niche relative to other members of the Empidonax group. Within the second major clade, mtDNA sequences support a reconstruction based on allozymes that places Contopus and Empidonax as sister taxa. This hypothesis contradicts that of Lanyon, who allied Contopus with Mitrephanes on the basis of similarity in foraging mode. Genera in the Empidonax group are members of a larger assemblage that radiated in South America. Occupancy of temperate habitats by certain genera in this group is coincident with their evolution of migratory behavior and with independent diversification in foraging modes that reduces potential competition in sympatry.     

A phylogenetic hypothesis for Asilidae based on a total evidence analysis of morphological and DNA sequence data (Insecta: Diptera: Brachycera: Asiloidea)

An hypothesis of phylogenetic relationships of Asilidae and its constituent taxa is presented, combining morphological and DNA sequence data in a total evidence framework. It is based on 77 robber fly species, 11 Asiloidea outgroup species, 211 morphological characters of the adult fly, and approximately 7300 bp of nuclear DNA from five genes (18S and 28S rDNA, AATS, CAD, and EF-1α protein-encoding DNA). The equally weighted, simultaneous parsimony analysis under dynamic homology in POY resulted in a single most parsimonious cladogram with a cost of 27,582 (iterative pass optimization; 27,703 under regular direct optimization). Six of the 12 included subfamily taxa are recovered as monophyletic. Trigonomiminae, previously always considered as monophyletic based on morphology, is shown to be non-monophyletic. Two of the three Trigonomiminae genera, Holcocephala Jaennicke, 1867 and Rhipidocephala Hermann, 1926, group unexpectedly as the sister taxon to all other Asilidae. Laphriinae, previously seen in the latter position, is the sister group of the remaining Asilidae. Five other subfamily taxa, i.e. Brachyrhopalinae, Dasypogoninae, Stenopogoninae, Tillobromatinae, and Willistonininae, are also shown to be non-monophyletic. The phylogenetic relationships among the higher-level taxa are partly at odds with findings of a recently published morphological study based on more extensive taxon sampling. The total evidence hypothesis is considered as the most informative one, but the respective topologies from the total-evidence, morphology-only, and molecular-only analyses are compared and contrasted in order to discuss the signals from morphological versus molecular data, and to analyze whether the molecular data outcompete the fewer morphological characters. A clade Apioceridae+Mydidae is corroborated as the sister taxon to Asilidae.     

Phylogeny of the plant bug subfamily Mirinae (Hemiptera: Heteroptera: Cimicomorpha: Miridae) based on total evidence analysis

The first comprehensive phylogenetic analyses of the most diverse subfamily of plant bugs, Mirinae, is presented in this study, for 110 representative taxa based on total evidence analysis. A total of 85 morphological characters and 3898 bp of mitochondrial (16S, COI) and nuclear (18S, 28S) sequences were analysed for each partitioned and combined dataset based on parsimony, maximum likelihood and Bayesian inference. Major results obtained in this study include monophyly of the tribe Mecistoscelini. The largest tribe, Mirini, was recovered as polyphyletic, and Stenodemini was recovered as paraphyletic. The clade of Stenodemini + Mecistoscelini is the sister group of the remaining Mirinae. The monophyly of two complexes composed of superficially similar genera were tested; the Lygus complex was recovered as nonmonophyletic, and the Adelphocoris–Creontiades–Megacoelum complex was confirmed to be monophyletic. The generic relationships of the main clades within each tribe based on the phylogeny, as well as their supported morphological characters, are discussed.     

Phylogenetic relationships of Moxostoma and Scartomyzon (Catostomidae) based on mitochondrial cytochrome b sequence data

A recent phylogenetic study based on morphological, biochemical and early life history characters resurrected the genus Scartomyzon (jumprock suckers, c . eight−10 species) from Moxostoma (redhorse suckers, c . 10–11 species) and advanced the understanding of relationships among species in these two genera, and the genealogical affinities of these genera with other evolutionary lineages within the tribe Moxostomatini in the subfamily Catostominae. To further examine phylogenetic relationships among moxostomatin suckers, the complete mitochondrial (mt) cytochrome b gene was sequenced from all species within this tribe and representative outgroup taxa from the Catostomini and other catostomid subfamilies. Phylogenetic analysis of gene sequences yielded two monophyletic clades within Catostominae: Catostomus + Deltistes + Xyrauchen + Erimyzon + Minytrema and Moxostoma + Scartomyzon + Hypentelium + Thoburnia . Within the Moxostomatini, Thoburnia was either unresolved or polyphyletic; Thoburnia atripinnis was sister to a monophyletic Hypentelium . In turn, this clade was sister to a monophyletic clade containing Scartomyzon and Moxostoma . Scartomyzon was never resolved as monophyletic, but was always recovered as a polyphyletic group embedded within Moxostoma , rendering the latter genus paraphyletic if ' Scartomyzon ' continues to be recognized. Relationships among lineages within the Moxostoma and' Scartomyzon 'clade were resolved as a polytomy. To better reflect phylogenetic relationships resolved in this analysis, the following changes to the classification of the tribe Moxostomatini are proposed: subsumption of' Scartomyzon 'into Moxostoma ; restriction of the tribe Moxostomatini to Moxostoma ; resurrect the tribe Erimyzonini, containing Erimyzon and Minytrema , classified as incertae sedis within Catostominae; retain the tribe Thoburniini.     

Phylogeny and higher classification of Hoplandriini (Coleoptera: Staphylinidae: Aleocharinae)

Abstract The monophyly of the aleocharine beetle tribe Hoplandriini is established and the phylogenetic relationships of sixteen genus-group taxa are resolved. Three primary lineages are recognized: the Ligulata + (( Platandria + Tetrallus ) + ( Paroplandria + ( Ditropandria + ( Omoplandria + ( Nosora + Tinotoma ))))) clade forms a sister group to the (( Hoplandria + undescribed species 5, 4, 6) + undescribed species 1, 2, 3) clade, and together form the sister group to Pseudoplandria . On the basis of phylogenetic inference, three subtribes are proposed within Hoplandriini. Hoplandriina is composed of Hoplandria s.l and two as yet undescribed genera from the New World. Platandriina, subtr.n., is composed of Ditropandria , Paroplandria , Platandria , Tetrallus , Omoplandria , Nosora and Tinotoma from the Nearctic, Neotropical, Oriental and Palearctic Regions. Pseudoplandriina, subtr.n., is composed of genus Pseudoplandria , primarily from the Oriental Region with one species from the Palearctic Region.     

A new Late Cretaceous family of Hymenoptera, and phylogeny of the Plumariidae and Chrysidoidea (Aculeata)

The taxonomic placement of an enigmatic species of wasp known from two specimens in Late Cretaceous New Jersey amber is investigated through cladistic analyses of 90 morphological characters for 33 terminals ranging across non-Aculeata, non-Chrysidoidea, most subfamilies of Chrysidoidea and all genera of Plumariidae (the family to which the fossils were initially assigned), based on use of exemplars. The fossil taxon is apparently basal in Chrysidoidea, most likely sister to Plumariidae, but perhaps sister to the remaining chrysidoids, or even sister to Chrysidoidea as a whole. It is described as representing a new family, Plumalexiidaefam. n., containing a single species, Plumalexius rasnitsynigen. et sp. n. Previous estimates of relationships for the genera of Plumariidae and for the higher taxa of Chrysidoidea are mostly confirmed. The importance of outgroup choice, and additivity and weighting of characters are demonstrated.     

Phylogenetic analysis of the family Ariidae (Ostariophysi: Siluriformes), with a hypothesis on the monophyly and relationships of the genera

Ariid monophyly and intrafamilial relationships are investigated based on cladistic analysis of 230 morphological characters. Terminal taxa examined include whenever possible type‐species, or the most morphologically similar species to the type‐species of the nominal genera, and the largest possible number of species, including cleared and stained specimens, available in zoological collections. Previous hypotheses about monophyly of the Ariidae are strongly corroborated by new synapomorphies discovered in the present study. The subfamily Galeichthyinae and the remaining ariids are strongly supported by new morphological characters. The monotypic subfamily Bagreinae is recognized as the sister group to all nongaleichthyin ariids, supported by a large series of exclusive synapomorphies. A new concept of Ariinae is presented: the subfamily is found to be unequivocally monophyletic and includes all ariid genera, except Galeichthys and Bagre. New data supporting the monophyly of the genera included in the Ariinae are introduced and previous hypotheses of monophyly, species composition, morphological definition, and relationships are reviewed and discussed.     

Larvae of Trichophya and phylogeny of the tachyporine group of subfamilies (Coleoptera: Staphylinidae) with a review, new species and characterization of the Trichophyinae

Abstract. Larvae of the staphylinid subfamily Trichophyinae are described for the first time based on larvae of a new species of Trichophya from the southwestern United States. Adults and larvae of the new species, Trichophya texana Ashe & Newton (type locality Texas, Brewster Co., Big Bend National Park), are described and illustrations of both provided. Also given are a key for separation of the Nearctic species of Trichophya , a checklist of the known World fauna of the Trichophyinae (including first report of the genus from Mexico and Guatemala), and a characterization of the subfamily Trichophyinae based on both larvae and adults. The relationships of major genera and higher taxa in the tachyporine group of staphylinid subfamilies are analysed cladistically using larval characters. No larval characters were found that provide evidence for the monophyly of the tachyporine group; no evidence was found for the monophyly of the Tachyporinae; Charhyphus, Olisthaerus and Phloeocharis (Phloeocharinae + Olisthaerinae) form a monophyletic group; the Trichophyinae and Habrocerinae are sister groups and together probably are the sister group to the Aleocharinae; the Aleocharinae are confirmed to be monophyletic based on larval characters; and Gymnusa + Deinopsis form the sister group to the remainder of the Aleocharinae.     

Phylogenetic relationships among superfamilies of Hymenoptera

The first comprehensive analysis of higher‐level phylogeny of the order Hymenoptera is presented. The analysis includes representatives of all extant superfamilies, scored for 392 morphological characters, and sequence data for four loci (18S, 28S, COI and EF‐1α). Including three outgroup taxa, 111 terminals were analyzed. Relationships within symphytans (sawflies) and Apocrita are mostly resolved. Well supported relationships include: Xyeloidea is monophyletic, Cephoidea is the sister group of Siricoidea + [Xiphydrioidea + (Orussoidea + Apocrita)]; Anaxyelidae is included in the Siricoidea, and together they are the sister group of Xiphydrioidea + (Orussoidea + Apocrita); Orussoidea is the sister group of Apocrita, Apocrita is monophyletic; Evanioidea is monophyletic; Aculeata is the sister group of Evanioidea; Proctotrupomorpha is monophyletic; Ichneumonoidea is the sister group of Proctotrupomorpha; Platygastroidea is sister group to Cynipoidea, and together they are sister group to the remaining Proctotrupomorpha; Proctotrupoidea s. str. is monophyletic; Mymarommatoidea is the sister group of Chalcidoidea; Mymarommatoidea + Chalcidoidea + Diaprioidea is monophyletic. Weakly supported relationships include: Stephanoidea is the sister group of the remaining Apocrita; Diaprioidea is monophyletic; Ceraphronoidea is the sister group of Megalyroidea, which together form the sister group of [Trigonaloidea (Aculeata + Evanioidea)]. Aside from paraphyly of Vespoidea within Aculeata, all currently recognized superfamilies are supported as monophyletic. The diapriid subfamily Ismarinae is raised to family status, Ismaridae stat. nov. © The Will Henning Society 2011.     

Limb myology and phylogenetic relationships in the superfamily Dipodoidea (birch mice, jumping mice, and jerboas)

Limb muscles were dissected in seven genera, representing all six superfamilies, of dipodoid rodents and myoloic characters were used to construct a phylogenetic hpothesis of relationships within this cfade. Mologic differences among genera suported tie monophyly of the superfamily Dipodoidea reLtive to the outrou taxon and reveaEd thatSicista is the sister group to all other zapodid and dipodid enera. Tkis picement of Sicista differs markedly from its position in previous classifications where it has been regarded merely as a primitive zapodid genus. The phlograrn based on rnyologic characters also indicated that Cardiocranius is not a rimitive dipodid genus; it is the sister group to the subfamily Dipodinae. Although myologic differences among taxa were not sufficient to warrant the continued separation of zaodids and dipodids into two families, a new classification that places Sicista in its own family, ficistidae, and places the remaining zaodids and dipodids in the family Dipodidae, is proposed. Differences in karyology, genitaP morholoy, and postcranial osteological characters among dipodoid rodents are discussed in light or this pjylogeny.     

Phylogeny of Branchiopoda (Crustacea) based on a combined analysis of morphological data and six molecular loci

The phylogenetic relationships of branchiopod crustaceans have been in the focus of a number of recent morphological and molecular systematic studies. Although agreeing in some respects, major differences remain. We analyzed molecular sequences and morphological characters for 43 branchiopods and two outgroups. The branchiopod terminals comprise all eight “orders”. The molecular data include six loci: two nuclear ribosomal genes (18S rRNA, 28S rRNA), two mitochondrial ribosomal genes (12S rRNA, 16S rRNA), one nuclear protein coding gene (elongation factor 1α), and one mitochondrial protein coding gene (cytochrome c oxidase subunit I). A total of 65 morphological characters were analyzed dealing with different aspects of branchiopod morphology, including internal anatomy and larval characters. The morphological analysis resulted in a monophyletic Phyllopoda, with Notostraca as the sister group to the remaining taxa supporting the Diplostraca concept (“Conchostraca” + Cladocera). “Conchostraca” is not supported but Cyclestheria hislopi is the sister group to Cladocera (constituting together Cladoceromorpha) and Spinicaudata is closer to Cladoceromorpha than to Laevicaudata. Cladocera is supported as monophyletic. The combined analysis under equal weighting gave results in some respects similar to the morphological analysis. Within Phyllopoda, Cladocera, Cladoceromorpha and Spinicaudata + Cladoceromorpha are monophyletic. The combined analysis is different from the morphological analysis with respect to the position of Notostraca and Laevicaudata. Here, Laevicaudata is the sister group to the remaining Phyllopoda and Notostraca is sister group to Spinicaudata and Cladoceromorpha. A sensitivity analysis using 20 different parameter sets (different insertion–deletion [indel]/substitution and transversion/transition ratios) show the monophyly of Anostraca, Notostraca, Laevicaudata, Spinicaudata, Cladoceromorpha, Cladocera, and within Cladocera, of Onychopoda and Gymnomera under all or almost all (i.e., 19 of 20) parameter sets. Analyses with an indel‐to‐transversion ratio up to 2 result in monophyletic Phyllopoda, with Laevicaudata as sister group to the remaining Phyllopoda and with Spinicaudata and Cladoceromorpha as sister groups. Almost all analyses (including those with higher indel weights) result in the same topology when only ingroup taxa are considered. © The Willi Hennig Society 2007.     

Preliminary morphological analysis of relationships between the spider wasp subfamilies (Hymenoptera: Pompilidae): revisiting an old problem

No qualitative cladistic analysis has been performed previously for the subfamily classification of Pompilidae (Hymenoptera). In 1994 Shimizu proposed six subfamilies, but their validity and relationships remain inconclusive. The objective of this study was to perform a quantitative analysis of phylogenetic relationships of the Pompilidae, with emphasis on testing the validity of proposed subfamilies. Two cladistic analyses were performed based on morphological evidence. First, a maximum-parsimony analysis of Shimizu's original morphological data matrix (72 taxa by 54 characters) was conducted, with the data subjected to a heuristic search for the first time with phylogenetic software. The resulting strict-consensus cladogram yielded a monophyletic Ceropalinae that was sister group to a large polytomy containing members of the remaining five subfamilies. In a second analysis, several of Shimizu's characters were re-examined, and new characters and more taxa were added to the data set. Terminal taxa were coded as species rather than as generic abstractions, and 20 additional morphological characters were introduced. The analysis was based on 77 morphological characters derived from the adults of 84 taxa. This second analysis suggested that Notocyphinae sensu Shimizu (1994) was nested within Pompilinae and that Epipompilinae sensu Shimizu (1994) was nested within Ctenocerinae; neither should retain their status as a separate subfamily. Lastly, Chirodamus s .s., which historically has been a member of the Pepsinae, is placed within the Pompilinae with reservations rather than erecting a new subfamily. After these allowances were made, a strict consensus tree gave the following relationships: (Ceropalinae + (Pepsinae + (Ctenocerinae + Pompilinae))).