A remarkable new genus of stiletto flies from Egypt, with a key to Palaearctic genera of Phycinae (Diptera, Therevidae)

An unusual new genus (Salwaea burgensisgen. n., sp. n.) of phycine stiletto flies is described from Egypt. A key to Palaearctic genera of Phycinae is presented.     

Review of the stiletto fly genus Actenomeros Winterton & Irwin (Diptera, Therevidae, Agapophytinae)

The endemic Australian genus Actenomeros Winterton & Irwin, 1999b is reviewed. Three species are transferred from Nanexila Winterton & Irwin, 1999a: Actenomeros aureilineata (Winterton & Irwin, 1999a) comb. n., Actenomeros intermedia (Winterton & Irwin, 1999a) comb. n. and Actenomeros paradoxa (Winterton & Irwin, 1999a) comb. n. A new species (Actenomeros budawangsp. n.) is described and figured from New South Wales. A key to species is presented.     

Digestive system of larval black flies (Diptera: Simuliidae): structure and phylogeny

Abstract. Larval gut characters of 45 species of black flies, representing 17 subgenera and 15 genera, were studied for their utility in taxon diagnosis and phylogenetic inference. Phase-contrast and scanning electron microscopy revealed a wealth of new characters in the esophageal armature of the foregut. Larval gut morphology, particularly characters of the esophageal armature, varied significantly among taxa, permitting generic diagnoses. A cladogram inferred from 13 gut characters supported a sister-group relationship between Parasimulium and the remaining simuliids, but did not show a strong phylogenetic signal for the current, well-supported concept of tribal level monophyly.     

A large‐scale molecular phylogeny of flesh flies (Diptera: Sarcophagidae)

The available data for Sarcophagidae in GenBank were analysed in order to reconstruct the most comprehensive phylogeny to date. GenBank was explored for nine markers that are commonly used in various molecular and phylogenetic studies of flesh flies. We obtained data for 187 species and constructed an aligned dataset with 9241 characters. However, the matrix suffered from 74% missing data due to a low number of sequences for some markers and in most of the cases only short fragments of the analysed genes were available. The reconstructed tree was taxonomically biased towards the subfamilies Paramacronychiinae (12% of the described species) and Sarcophaginae (8.6% of the described species) and specifically the genus Sarcophaga. The third subfamily Miltogramminae was represented by only 0.7% of described species. Moreover, about half of the included species were of forensic importance, while the percentage of such species in the entire family was estimated at 7%. Many nodes had very low support, so in order to increase the support and thereby identify a ‘core topology’, we pruned ‘rogue’ taxa and applied different substitution models. Both strategies improved support considerably, although some nodes still were left unresolved. An analysis of the distribution of bootstrap values across chronograms showed that the weakest phylogenetic signal is restricted to that part of the tree which coincides with the onset of rapid radiations mainly within the genus Sarcophaga. Our study is concordant with phylogenies obtained by other authors, with the most noteworthy exception being the subfamily Paramacronychiinae emerging as paraphyletic with regard to the Miltogramminae, which is in strong conflict with morphological evidence. We discuss the new findings in the light of traditional taxonomical classifications of Sarcophagidae and recent molecular studies.     

Phylogenetic revision of Bonjeania Irwin & Lyneborg (Diptera: Therevidae)

The endemic Australian genus Bonjeania Irwin & Lyneborg is revised. Seven species are described as new: B. angelikae sp.n., B. clamosis sp.n., B. dynastis sp.n., B. flavofemoralis sp.n., B. irwinae sp.n., B. orphne sp.n. and B. trilineata sp.n. Three species, B. actuosa (White), B. nitidifrons (Macquart) and B. segnis (White), are redescribed. All species of Bonjeania were compared in a cladistic analysis with Agapophytus albobasalis Mann and five exemplars of the speciose sister genus Parapsilocephala Kröber, and the phylogenetic relationships of Bonjeania discussed. The male genitalic musculature of B. clamosis sp.n. is described and figured, and discussed with respect to previous studies on other Therevidae.     

A phylogeny of robber flies (Diptera: Asilidae) at the subfamilial level: molecular evidence

We present the first formal analysis of phylogenetic relationships among the Asilidae, based on four genes: 16S rDNA, 18S rDNA, 28S rDNA, and cytochrome oxidase II. Twenty-six ingroup taxa representing 11 of the 12 described subfamilies were selected to produce a phylogenetic estimate of asilid subfamilial relationships via optimization alignment, parsimony, and maximum likelihood techniques. Phylogenetic analyses support the monophyly of Asilidae with Leptogastrinae as the most basal robber fly lineage. Apocleinae+(Asilinae+Ommatiinae) is supported as monophyletic. The laphriinae-group (Laphriinae+Laphystiinae) and the dasypogoninae-group (Dasypogoninae+Stenopogoninae+Stichopogoninae+ Trigonomiminae) are paraphyletic. These results suggest that current subfamilial classification only partially reflects robber fly phylogeny, indicating the need for further phylogenetic investigation of this group.     

Molecular phylogeny of black flies (Diptera: Simuliidae) from Thailand, using ITS2 rDNA

The sequences of the second internal transcribed spacer (ITS2) of ribosomal DNA (rDNA) were determined for 40 black fly species from Thailand, belonging to 4 subgenera of the genus Simulium, namely Gomphostilbia (12 species), Nevermannia (5 species), Montisimulium (1 species), Simulium sensu stricto (21 species), and an unknown subgenus with one species (Simulium baimaii). The length of the ITS2 ranged from 247 to 308 bp. All black fly species had high AT content, ranging from 71 to 83.8%. Intraindividual variation (clonal variation) occurred in 13 species, ranging from 0.3 to 1.1%. Large intrapopulation and interpopulation heterogeneities exist in S. feuerboni from the same and different locations in Doi Inthanon National Park, northern Thailand. Phylogenetic relationships among 40 black fly species were examined using PAUP (version 4.0b10) and MrBAYS (version 3.0B4). The topology of the trees revealed two major monophyletic clades. The subgenus Simulium and Simulium baimaii were placed in the first monophyletic clade, whereas the subgenera Nevermannia + Montisimulium were placed as the sister group to the subgenus Gomphostilbia in the second monophyletic clade. Our results suggest that S. baimaii belongs to the malyschevi-group or variegatum-group in the subgenus Simulium. The molecular phylogeny generally agrees with existing morphology-based phylogenies.     

Higher-level phylogeny of the Therevidae (Diptera: insecta) based on 28S ribosomal and elongation factor-1 alpha gene sequences

Therevidae (stilleto flies) are a little-known family of asiloid brachyceran Diptera (Insecta). Separate and combined phylogenetic analyses of 1200 bases of the 28S ribosomal DNA and 1100 bases of elongation factor-1alpha were used to infer phylogenetic relationships within the family. The position of the enigmatic taxon Apsilocephala Kr?ber is evaluated in light of the molecular evidence. In all analyses, molecular data strongly support the monophyly of Therevidae, excluding Apsilocephala, and the division of Therevidae into two main clades corresponding to a previous classification of the family into the subfamilies Phycinae and Therevinae. Despite strong support for some relationships within these groups, relationships at the base of the two main clades are weakly supported. Short branch lengths for Australasian clades at the base of the Therevinae may represent a rapid radiation of therevids in Australia.     

Evolutionary history of Stratiomyidae (Insecta: Diptera): the molecular phylogeny of a diverse family of flies

Stratiomyidae is a cosmopolitan family of Brachycera (Diptera) that contains over 2800 species. This study focused on the relationships of members of the subfamily Clitellariinae, which has had a complicated taxonomic history. To investigate the monophyly of the Clitellariinae, the relationships of its genera, and the ages of Stratiomyidae lineages, representatives for all 12 subfamilies of Stratiomyidae, totaling 68 taxa, were included in a phylogenetic reconstruction. A Xylomyidae representative, Solva sp., was used as an outgroup. Sequences of EF-1alpha and 28S rRNA genes were analyzed under maximum parsimony with bootstrapping, and Bayesian methods to recover the best estimate of phylogeny. A chronogram with estimated dates for all nodes in the phylogeny was generated with the program, r8s, and divergence dates and confidence intervals were further explored with the program, multidivtime. All subfamilies of Stratiomyidae with more than one representative were found to be monophyletic, except for Stratiomyinae and Clitellariinae. Clitellariinae were distributed among five separate clades in the phylogeny, and Raphiocerinae were nested within Stratiomyinae. Dating analysis suggested an early Cretaceous origin for the common ancestor of extant Stratiomyidae, and a radiation of several major Stratiomyidae lineages in the Late Cretaceous.     

The bee‐killing flies,genus Melaloncha Brues (Diptera: Phoridae): a combined molecular and morphological phylogeny

The phylogeny of the bee‐killing flies, genus Melaloncha Brues (Diptera: Phoridae) is analysed using six genes –cytochrome oxidase I, 16S ribosomal DNA, 12S ribosomal DNA, NADH1 dehydrogenase, 28S ribosomal DNA and CAD– plus 47 morphological characters. A total of 91 specimens, including eight out‐groups and 83 Melaloncha (representing 70 species) were included in the analyses. Parsimony analysis of the combined data set produced a single most parsimonious tree with varied Bremer and bootstrap support of interior nodes. Bayesian analysis of molecules only and of morphology + molecules produced trees largely in agreement with parsimony results, although with a few differences. Supported groups included subfamily Metopininae, genus Melaloncha, and subgenera Melaloncha s.s. and Melaloncha (Udamochiras) Enderlein. Within the subgenera, the previously recognized Melaloncha furcata, Melaloncha cingulata, Melaloncha ungulata and Melaloncha stylata groups were recovered, as well as some new groupings. The M. furcata group was placed as the sister group of other Melaloncha s.s., which is consistent with known host‐attacking behaviour.     

The aquatic drift patterns of black flies (Diptera: Simuliidae)

Larval (5th-final instar) drift patterns of five species of black flies (Prosimulium mixtum, Stegopterna mutata, Simulium tuberosum FG, S. vittatum IIIL-1, and S. vittatum IS-7) were analyzed. The diel drift patterns were not species-specific, but consistently exhibited a marked increase after sunset, preceded by a period of minimal drift. The drift minimum generally occurred during or shortly after the period of maximum water temperature. Larvae showed significantly more nocturnal than diurnal drift. Pupal drift was attributed to accidental detachment from the substrate. Adult drift was over 90% diurnal and usually composed primarily of teneral individuals. Upstream larval movements of the S. vittatum and S. tuberosum complexes were each less than 6.5% of the total upstream and downstream larval movements. The proportion of parasitized larvae of the S. vittatum and S. tuberosum complexes was generally higher in the benthos than in the drift. Based in part on data included in a thesis submitted by the senior author to The Pennsylvania State University in partial fulfillment of the requirements for the Ph. D. degree.     

Taxon sampling to address an ancient rapid radiation: a supermatrix phylogeny of early brachyceran flies (Diptera)

Early diverging brachyceran fly lineages underwent a rapid radiation approximately 180 Ma, coincident in part with the origin of flowering plants. This region of the fly tree includes 25 000 described extant species with diverse ecological roles such as blood‐feeding (haematophagy), parasitoidism, predation, pollination and wood‐feeding (xylophagy). Early diverging brachyceran lineages were once considered a monophyletic group of families called Orthorrhapha, based on the shared character of a longitudinal break in the pupal skin made during the emergence of the adult. Yet other morphological and molecular evidence generally supports a paraphyletic arrangement of ‘Orthorrhapha’, with strong support for one orthorrhaphan lineage – dance flies and relatives – as the closest relative to all higher flies (Cyclorrhapha), together called Eremoneura. In order to establish a comprehensive estimate of the relationships among orthorrhaphan lineages using a thorough sample of publicly available data, we compiled and analysed a dataset including 1217 taxa representing major lineages and 20 molecular markers. Our analyses suggest that ‘Orthorrhapha’ excluding Eremoneura is not monophyletic; instead, we recover two main lineages of early brachyceran flies: Homeodactyla and Heterodactyla. Homeodactyla includes Nemestrinoidea (uniting two parasitic families Acroceridae + Nemestrinidae) as the closest relatives to the large SXT clade, comprising Stratiomyomorpha, Xylophagidae and Tabanomorpha. Heterodactyla includes Bombyliidae with a monophyletic Asiloidea (exclusive of Bombyliidae) as the closest relatives to Eremoneura. Reducing missing data, modifying the distribution of genes across taxa, and, in particular, removing rogue taxa significantly improved tree resolution and statistical support. Although our analyses rely on dense taxonomic sampling and substantial gene coverage, our results pinpoint the limited resolving power of Sanger sequencing‐era molecular phylogenetic datasets with respect to ancient, hyperdiverse radiations.     

The orientation behaviour of horse flies and deer flies (Tabanidae, Diptera)

The Manitoba Fly Trap is illustrated and described and its uses in behaviour studies, surveys, insect control and perhaps population studies are enumerated. Its advantages over other traps for diurnal biting flies are discussed.

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Zusammenfassung Die Manitoba-Fliegenfalle wird abgebildet und beschrieben. Ihre Verwendungsmöglichkeiten bei der Verhaltensforschung, Sammlung und Bekämpfung von Insekten und vielleicht bei Massenwechselstudien werden aufgezählt und ihre Vorteile vor anderen Fallen für den Fang diurnaler Stechfliegen diskutiert.

     

Molecular phylogeny of flat‐footed flies (Diptera: Platypezidae): main clades supported by new morphological evidence

The molecular phylogeny of flat‐footed flies is inferred from analysis of DNA sequence data from the five mitochondrial genes 12S, 16S, COI, COII and CytB, and the nuclear gene 28S and discussed with the recent systematics based on morphological features. The Bayesian inference, maximum likelihood and maximum parsimony analyses included 42 species of 18 genera, representing all four extant subfamilies (Microsaniinae, Melanderomyiinae, Callomyiinae and Platypezinae) and all known genera except one (Metaclythia). Representatives of the brachycerous taxa Lonchopteridae, Phoridae, Sciadocerinae (Phoridae) and Opetiidae are used as outgroups, and Lonchoptera was used to root the trees. Our results show Platypezidae consisting of two well‐supported clades, the first with the subfamilies Melanderomyiinae + Callomyiinae and the second formed by subfamily Platypezinae. Genus Microsania was resolved as a separate lineage distant from Platypezidae which clustered with Opetiidae as its sister group, both together forming a sister group to Platypezidae. At the generic level, the genus Agathomyia proved not to be monophyletic in any of the analyses. The species Chydaeopeza tibialis is sister to Agathomyia sexmaculata, and consequently, the genus Chydaeopeza Shatalkin, 1992 is a new junior synonym of Agathomyia Verrall, 1901. Bifurcated setae on legs of adult Platypezidae are documented as a new synapomorphy of the family, exclusive of Microsania. Outstretched wings and only a small overlap of their surfaces at resting position are considered a new synapomorphy for the subfamily Platypezinae. Other phylogenetically important characters defining main clades are documented, and their relevance/validity in phylogenetic studies is discussed. The current systematic concept of Platypezidae is discussed, and new phylogenetic hypotheses are proposed.     

Allozyme variation in stable flies (Diptera: Muscidae)

Polyacrylamide gel electrophoresis was used to resolve allozymes in the cosmopolitan blood-feeding stable fly,Stomoxys calcitrans (L.). Nineteen of 38 loci were polymorphic (53%). Mean heterozygosities among all loci and among only polymorphic loci were 0.096 and 0.182, respectively. These gene diversity measures are about half those among other muscid Diptera. Variation in gene frequencies was examined in 10 natural stable fly populations from Iowa and Minnesota. Gene frequencies were homogeneous at five of eight loci among six populations in 1990 and eight of eight loci among four populations in 1992. Wright'sF statistics showed no significant departure from random mating among stable flies. It was concluded that gene flow compensates for any local differentiation in stable fly populations.     

New species of Vomerina Winterton (Diptera, Therevidae, Agapophytinae) from Australia

Two new species of Vomerina Winterton, 2007 (Vomerina comapenissp. n. and Vomerina micorasp. n.) are described from New South Wales (Australia).     

Predation of Thereva nobilitata (Fabricius) (Diptera: Therevidae) on Agriotes obscurus L. (Coleoptera: Elateridae)

Little is known about the natural enemies of wireworms (Coleoptera: Elateridae), but there are frequent anecdotal reports of (usually unnamed) stiletto fly larvae (Diptera: Therevidae) preying on various species. We observed larvae of Thereva nobilitata (Fabricius) feeding on larvae of the dusky wireworm, Agriotes obscurus L., during the summer of 2011, in Agassiz, British Columbia. This finding is of interest as: both the predator and the wireworm are introduced species to this area from Europe; T. nobilitata is uncommon in North America; and this predator has not been associated with any wireworm species previously. We observed that larvae of male and female T. nobilitata will feed on various sizes of A. obscurus larvae, most feeding being carried out by the smallest T. nobilitata larvae. These findings suggest future work should assess the potential for therevid larvae as top‐down regulators of Agriotes larvae under field conditions.     

Gender-specific bacterial composition of black flies (Diptera: Simuliidae)

Although hematophagous black flies are well-known socioeconomic pests and vectors of disease agents, their associated bacteria are poorly known. A systematic analysis of the bacterial community associated with freshly emerged adult black flies of four North American species, using cultivation-independent molecular techniques, revealed 75 nonsingleton bacterial phylotypes. Although 17 cosmopolitan phylotypes were shared among host species, each fly species had a distinct bacterial profile. The bacterial composition, however, did not correlate strongly with the host phylogeny but differed between male and female flies of the same species from the same habitat, demonstrating that a group of insects have a gender-dependent bacterial community. In general, female flies harbor a less diverse bacterial community than do males. The anatomical locations of selected bacteria were revealed using fluorescence in situ hybridization. Understanding the physiological function of the associated bacterial community could provide clues for developing novel pest-management strategies.     

The phylogenetic relationships of flies in the superfamily Empidoidea (Insecta: Diptera)

We conducted a molecular phylogenetic study of the Empidoidea, a diverse group of 10,000 species of true flies, with two major goals: to reconstruct a taxonomically complete and robustly supported phylogeny for the group and to use this information to assess several competing classifications for the clade. We amassed 3900+ nucleotides of coding data from the carbamoylphosphate synthase domain of the rudimentary locus (CAD) and 1200+ nucleotides from the large nuclear ribosomal subunit (28S) from 72 and 71 species, respectively, representing several orthorrhaphan and cyclorrhaphan families and all previously recognized empidoidean subfamilies. Independent and combined phylogenetic analyses of these data were conducted using parsimony, maximum likelihood, and Bayesian criteria. The combined matrix included 61 taxa for which both CAD and 28S sequences were obtained. Analyses of CAD first and second codon positions alone and when concatenated with 28S sequences yielded trees with similar and largely stable topologies. Analyses of 28S data alone supported many clades although resolution is limited by low sequence divergence. The following major empidoid clades were recovered with convincing support in a majority of analyses: Atelestidae, Empidoidea exclusive of Atelestidae, Hybotidae sensu lato, Dolichopodidae+Microphorinae (including Parathallassius), and Empididae sensu lato (including Brachystomatinae, Ceratomerinae, Clinocerinae, Empidinae, Hemerodromiinae, Oreogetoninae, and Trichopezinae). The branching arrangement among these four major clades was Atelestidae, Hybotidae, Dolichopodidae/Microphorinae, Empididae. Previously recognized subclades recovered with robust support included Hybotinae, Brachystomatinae, Tachydromiinae, Clinocerinae (in part), Hemerodromiinae, Empidinae, and Empidiini.