Local biodiversity and multi-habitat use in empidoid flies (Insecta: Diptera, Empidoidea)

The empidid fauna of four small adjacent biotopes bordering a pond was investigated for 2 consecutive years in Brittany (France). Adult activity was studied using yellow water traps, whereas suitable larval habitats were determined using emergence traps. While 24 species emerged from the soils, 45 flew above the four sites. The number of species emerging from each site was nearly identical. However, the highest number of individuals emerged from the heathland and numbers rapidly declined towards the pond banks. On the contrary, the greatest aerial activity occurred in the woodlot and near the pond banks. Fourteen times less flying activity was found above the dry heathland. The latter appeared to be a site of larval growth but mating and feeding of the adults took place in the woodlot. Reproduction sites and space used by the adults differed among the dominant species. The species assemblage could not be fully explained within the spatial limits of the four sites. Considering the species'behaviour, it is suggested that immigration of species and individuals from other sites should explain these differences. The study, which is supported by four other research works, emphasizes the role of key resource played by ecotonal zones between aquatic and terrestrial ecosystems in the persistence of species over a larger set of habitats. Considering the complementarity of habitats is essential to explain diversity patterns in species which need different space units to complete their life-cycle.     

Complete mitochondrial genome of Dacus vijaysegarani and phylogenetic relationships with congeners and other tephritid fruit flies (Insecta: Diptera)

Molecular Biology Reports - Tephritid fruit flies of the genus Dacus are members of the tribe Dacini, subfamily Dacinae. There are some 274 species worldwide, distributed in Africa and the...     

New contributions towards the understanding of the phylogenetic relationships among economically important fruit flies (Diptera: Tephritidae)

Fruit flies (Diptera: Tephritidae) are a species-rich and economically important group. The phylogenetic relationships among the many taxa are still to be fully resolved and the monophyly of several groups is still to be confirmed. This paper reports a study of the phylogenetic relationships among 23 economically important tephritid species (representing several major lineages of the family) which examines the sequence of a region of mitochondrial DNA encompassing the cytb, tRNA(Ser) and ND1 genes. Substitutions characteristic of particular taxa were found that could help classify members of the family at any developmental stage. The trees obtained by the maximum parsimony, neighbour joining and maximum likelihood methods were generally compatible with present morphological classification patterns. However, the data reveal some characteristics of the phylogenetic relationships of this family that do not agree with present classifications. The results support the probable non-monophyletic nature of the subfamily Trypetinae and suggest that Bactrocera cucurbitae (Coquillet) is more closely related to the genus Dacus than to other species of Bactrocera.     

Cryptic biodiversity and phylogenetic relationships revealed by DNA barcoding of Oriental black flies in the subgenus Gomphostilbia (Diptera: Simuliidae)

Understanding the medical, economic, and ecological importance of black flies relies on correct identification of species. However, traditional taxonomy of black flies is impeded by a high degree of morphological uniformity, especially the presence of cryptic biodiversity, historically recognized by details of chromosomal banding patterns. We assess the utility of DNA barcoding, based on cytochrome c oxidase subunit 1 (COI) sequences, for identifying 13 species of Oriental black flies in the subgenus Gomphostilbia. Samples of larvae fixed in Carnoy's solution were used to gather molecular and chromosomal data from the same individual. We found that larvae refrigerated in Carnoy's fixative for as long as 11 years can be used for DNA study. Levels of intraspecific genetic divergence, based on the Kimura-2 parameter, range from 0% to 9.28%, with a mean of 2.75%, whereas interspecific genetic divergence ranges from 0.34% to 16.05%. Values of intraspecific and interspecific genetic divergence overlap in seven species owing to incomplete lineage sorting and imperfect taxonomy, implying that DNA barcoding to identify these species will be ambiguous. Despite a low level of success, we found that DNA barcoding is useful in revealing cryptic biodiversity, potentially facilitating traditional taxonomy. Phylogenetic analyses indicate that species groups currently recognized on morphological criteria are not monophyletic, suggesting a need to reevaluate the classification of the subgenus Gomphostilbia.     

The taxonomy and phylogenetic relationships of the Pollenia semicinerea species-group (Diptera: Calliphoridae)

Abstract. Within Pollenia Robineau-Desvoidy, a semicinerea species-group is defined. It consists of the redescribed P.semicinerea Villeneuve, P.grunini sp.n., P.mystica sp.n. and P.paragrunini sp.n., and P.pectinata Grunin, for which some new characters are given. A key is provided and the known terminalia of both sexes are illustrated. A lectotype is designated for P.semicinerea Villeneuve. P.semicinerea is known from the Middle East (Israel, Syria, Lebanon), P.grunini, P.mystica and P.paragrunini from the Caucasus and neighbouring areas in the U.S.S.R. and P.pectinata from the temperate parts of the Palaearctic Region except west of Poland and U.S.S.R. It is argued that the semicinerea- and racfe-groups are monophyletic sister-groups. The tenuiforceps-group is most probably the monophyletic sister-group of the composite monophyletic group ( rudis-group+semicinerea-group ). The intermedia-group and the griseo-tomentosa-group are both monophyletic. The composite group ( rudis- group + semicinerea-group + tenuiforceps-group + intermedia-group + griseotomentosa-group ) is probably a monophyletic subgroup of Pollenia. A partly resolved synapomorphy diagram is provided, which is compared to trees generated by various computer programs in the Phylogeny Inference Package (PHYLIP) of Felsenstein. Pollenia alajensis Rodendorf (= sytshevskajae Grunin) is assigned to the tenuiforceps species-group on the basis of male and female genitalia.     

The genera of Diopsidae (Insecta:Diptera)

Taxonomic characters which have been used in defining the genera of Diopsidae are discussed with special reference to the structure of the head and eyestalks. In consequence, the subdivisions of this family put forward by Hennig in 1965 are revised and extended, a new scheme of three subfamilies being proposed and a possible phylogenetic sequence of genera suggested.
A key for the identification of these subfamilies and current generic taxa is given.     

The adult head structures of Tipulomorpha (Diptera,Insecta) and their phylogenetic implications

Schneeberg, K. and Beutel, R.G. 2011. The adult head structures of Tipulomorpha (Diptera, Insecta) and their phylogenetic implications. —Acta Zoologica (Stockholm) 92 : 316–343. Head structures of adults of Tipula paludosa, Limonia sp. and Trichocera saltator were examined and described. The results are compared with conditions found in other dipterans and other antliophoran groups, notably Nannochoristidae. Several potential synapomorphies of a dipteran–nannomecopteran–siphonapteran clade are present in Tipuloidea and Trichocera, the labro‐epipharyngeal food channel, the loss of the galea and the postpharyngeal pumping apparatus. The sensorial field of the maxillary palpomere 3, a potential dipteran–nannomecopteran synapomorphy, is also present but modified. The presence of M. clypeolabralis, labellae and mandibular stylets are groundplan apomorphies of Diptera, with secondary loss of the mandibles in Tipuloidea, Trichoceridae and many other groups. Tipuloidea is supported by the origin of M. tentorioscapalis anterior on the head capsule, the reduction of M. frontobuccalis anterior and the loss of the ocelli. The reduced tentorium, the origin of two further antennal muscles on the head capsule, the maxillary sensorial field with sensilla in individual pits, the lacking dorsal prelabial concavity and the unpaired salivary channel entering the head are apomorphies of Tipulidae. Closer affinities of Tipulidae and Cylindrotomidae are suggested by pseudotracheae of the advanced type, which have evolved independently in this lineage. The results do neither support a basal placement of Tipuloidea nor close affinities with Brachycera.     

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.     

Evolution and phylogenetic utility of CAD (rudimentary) among Mesozoic-aged Eremoneuran Diptera (Insecta)

We sequenced nearly the entire carbomoylphosphate synthase (CPS) domain of CAD, or rudimentary, (ca. 4 kb) from 29 species of flies representing all major clades within Eremoneura, or higher flies, and several orthorrhaphous brachyceran outgroups. We compared these sequences with orthologs from Anopheles gambiae and Drosophila melanogaster to assess structure, compositional bias, and phylogenetic utility. CAD is large (6.6+ kb), complex (comprised of three major and myriad minor functional domains) and relatively free of introns, extreme nucleotide bias (except third codon positions), and large hypervariable regions. The CPS domain possesses moderate levels of nonsynonymous divergence among taxa of intermediate evolutionary age and conveys considerable phylogenetic signal. Phylogenetic analysis of CPS sequences under varying methods and assumptions resulted in well-resolved, strongly supported trees concordant with many traditional ideas about higher dipteran phylogeny and with prior inferences from 28S rDNA. The most robustly supported major eremoneuran clades were Cyclorrhapha, Platypezoidea, Eumuscomorpha, Empidoidea, Atelestidae, Empidoidea exclusive of Atelestidae, Hybotidae s.l., Microphoridae+Dolichopodidae, and Empididae s. str. Because CAD is ubiquitous, apparently single copy (at least within holometabolous insects), readily obtained from several insect orders using primers described herein, and exhibits considerable phylogenetic utility, it should have wide applicability in insect molecular systematics.     

Zoogeographical affinities and faunal relationships of bee flies (Diptera: Bombyliidae) in Egypt

The distributions of 229 beefly (Diptera: Bombyliidae) species across the eight Egyptian ecological zones, together with their faunal affinities to the main zoogeographical regions, were used to test the suggestion of Holt et al. (2013) that the SaharoArabian is a distinct region rather than a subregion of the Palaearctic. All Egyptian ecological zones but one have greater affiliation to the Palaearctic and SaharoArabian than to the Afrotropical region; the Gebel Elba ecological zone, the southeastern triangle of Egypt, has greater affinities with the Afrotropics. Affinities to the Saharo-Arabian region were not different from those to the Palaearctic. From its bombyliid fauna, therefore, the Saharo-Arabian region is so closely allied to the Palaearctic as to constitute merely a subregion of it. Sinai shows a high level of endemism reflecting its isolation from other parts of Egypt.     

Review of the systematics and phylogenetic relationships of the Sabethini (Diptera: Culicidae)

This paper reviews the historical treatment of the tribe Sabethini and genus-group taxa and examines the unusual life histories associated with the group. Although recognized by taxonomists as distinct, the taxonomic position of sabethines has been questioned and their rank within the family Culicidae unstable. In order to evaluate the current status of the classification of the tribe a cladistic analysis is performed. Thirty-seven taxa are selected from within the Sabethini and two outgroups were chosen from the tribe Aedini. Exemplars are selected from genus-group taxa world-wide and new and traditional character systems examined in larval, pupal and adult life stages. The results firmly establish the sabethines as a monophyletic group. However, the genera Runchomyia , Tripteroides and Wyeomyia are not demonstrably monophyletic. In addition, the data support the New World taxa as a monophyletic group to a paraphyletic assemblage of Old World taxa. The pattern displayed by the cladogram suggests the ability to vector arboviruses has arisen more than once in mosquitoes.     

On the head morphology of Phyllium and the phylogenetic relationships of Phasmatodea (Insecta)

Friedemann K., Wipfler B., Bradler S. and Beutel R.G. 2011 . On the head morphology of Phyllium and the phylogenetic relationships of Phasmatodea (Insecta). —Acta Zoologica (Stockholm) 00 : 1–16. External and internal head structures of Phyllium siccifolium are described in detail. The findings are compared with conditions found in other phasmatodeans and members of other neopteran lineages. The compiled 125 characters were analysed cladistically. A clade Eukinolabia (Phasmatodea + Embioptera) was confirmed. Synapomorphies of these two taxa are the shift of the origin of M. tentorioparaglossalis to the hind margin of the prementum, the presence of M. tentorioscapalis medialis, and antennal muscles that originate exclusively on the anterior tentorial arms. Within Eukinolabia, the position of Timema remains somewhat ambiguous because of missing anatomical data. However, it was confirmed as sister group of Euphasmatodea in a monophyletic Phasmatodea. Apomorphic groundplan features of Euphasmatodea are salivary ducts with separate external openings, apically rounded glossae, the presence of the galealobulus, and the reduction of the antennifer. The monophyly of Neophasmatidae was confirmed. Autapomorphies are the loss of M. frontobuccalis posterior, the anteriorly or dorsally directed maxillary palps, and the reduction of the mandibular incisivi. The analysis of characters of the head yielded three new autapomorphies of Phylliinae, the presence of a protuberance on the attachment site of the dorsal tentorial arms, dorsoventrally flattened maxillary‐ and labial palps, and possibly the narrow and U‐shaped field of trichomes on the apical part of the galea.     

Phylogeny of Oestridae (Insecta: Diptera)

The phylogeny of Oestridae was analysed at the generic level using 118 characters from all developmental stages and including morphology, ontogeny, physiology and behaviour. Four major clades were given subfamilial rank with the phylogenetic relationship (Cuterebrinae (Gasterophilinae (Hypodermatinae + Oestrinae))). The subdermal parasites of the African elephant, Neocuterebra squamosa Grünberg and Ruttenia loxodontis Rodhain, had their most probable affiliation subordinate to the clade of stomach parasites, although their exact position needs further investigation. Genus Ochotonia Grunin, which is known from a single third‐instar larva only, was the probable sister group of all other Hypodermatinae. Twenty‐five oestrid genera were recognised as valid and those containing more than one species were defined through lists of autapomorphies. Cuterebra Clark was proposed as a senior synonym of Alouattamyia Townsend, Andinocuterebra Guimarães, Pseudogametes Bischof and Rogenhofera Brauer. The clade of hypodermatine ungulate parasites (Hypoderma Latreille +Pallasiomyia Rubtzov +Pavlovskiata Grunin +Przhevalskiana Grunin +Strobiloestrus Brauer) remained largely unresolved, and genus Przhevalskiana emerged without defining characters.     

Phylogeny of Ptychopteroidea (Insecta: Diptera)

A phylogenetic tree is proposed for the superfamily Ptychopteroidea, reconstructed taking into account both extinct and extant taxa and based mainly on characters of wing venation.     

The phylogeny of stiletto flies (Diptera: Therevidae)

The therevoid clade represents a group of four families (Apsilocephalidae, Evocoidae, Scenopinidae and Therevidae) of lower brachyceran Diptera in the superfamily Asiloidea. The largest of these families is that of the stiletto flies (Therevidae). A large‐scale (i.e. supermatrix) phylogeny of Therevidae is presented based on DNA sequence data from seven genetic loci (16S, 18S and 28S ribosomal DNA and four protein‐encoding genes: elongation factor 1‐alpha, triose phosphate isomerase, short‐wavelength rhodopsin and the CPSase region of carbamoyl‐phosphate synthase‐aspartate transcarbamoylase‐dihydroorotase). Results are presented from Bayesian phylogenetic analyses of approximately 8.7 kb of sequence data for 204 taxa representing all subfamilies and genus groups of Therevidae. Our results strongly support the sister‐group relationship between Therevidae and Scenopinidae, with Apsilocephalidae as sister to Evocoidae. Previous estimates of stiletto fly phylogeny based on morphology or DNA sequence data, or supertree analysis, have failed to find significant support for relationships among subfamilies. We report for the first time strong support for the placement of the subfamily Phycinae as sister to the remaining Therevidae, originating during the Mid Cretaceous. As in previous studies, the sister‐group relationship between the species‐rich subfamilies Agapophytinae and Therevinae is strongly supported. Agapophytinae are recovered as monophyletic, inclusive of the Taenogera group. Therevinae comprise the bulk of the species richness in the family and appear to be a relatively recent and rapid radiation originating in the southern hemisphere (Australia + Antarctica + South America) during the Late Cretaceous. Genus groups are defined for all subfamilies based on these results.     

The classification of genus Anopheles (Diptera: Culicidae): a working hypothesis of phylogenetic relationships

The internal classification of genus Anopheles is updated to reflect taxonomic actions published since the classification was last reviewed in 1994. Both formal and informal taxa are included. The classification is intended to aid researchers and students who are interested in analysing species relationships, making group comparisons and testing phylogenetic hypotheses. The genus includes 444 formally named and 40 provisionally designated extant species divided between six subgenera: Anopheles, Cellia, Kerteszia, Lophopodomyia, Nyssorhynchus and Stethomyia. Subgenera Anopheles, Cellia and Nyssorhynchus are subdivided hierarchically into nested informal groups of morphologically similar species that are believed to represent monophyletic lineages based on morphological similarity. Changes to the classification include additional species, eliminated species and changes to the hierarchical organization and composition of supraspecific groups, some as a result of molecular studies.     

Utility of the white gene in estimating phylogenetic relationships among mosquitoes (Diptera: Culicidae)

The utility of a nuclear protein-coding gene for reconstructing phylogenetic relationships within the family Culicidae was explored. Relationships among 13 species representing three subfamilies and nine genera of Culicidae were analyzed using a 762-bp fragment of coding sequence from the eye color gene, white. Outgroups for the study were two species from the sister group Chaoboridae. Sequences were determined from clone PCR products amplified from genomic DNA, and aligned following conceptual intron splicing and amino acid translation. Third codon positions were characterized by high levels of divergence and biased nucleotide composition, the intensity and direction of which varied among taxa. Equal weighting of all characters resulted in parsimony and neighboring-joining trees at odds with the generally accepted phylogenetic hypothesis based on morphology and rDNA sequences. The application of differential weighting schemes recovered the traditional hypothesis, in which the subfamily Anophelinae formed the basal clade. The subfamily Toxorhynchitinae occupied an intermediate position, and was a sister group to the subfamily Culicinae. Within Culicinae, the genera Sabethes and Tripteroides formed an ancestral clade, while the Culex-Deinocerites and Aedes- Haemagogus clades occupied increasingly derived positions in the molecular phylogeny. An intron present in the Culicinae- Toxorhynchitinae lineage and one outgroup taxon was absent in the basal Anophelinae lineage and the second outgroup taxon, suggesting that intron insertions or deletions may not always be reliable systematic characters.      

The Linnaean species of the family Tephritidae (Insecta: Diptera)

Linnaeus described nine species of Musca now placed in the Tephritidae. The Linnaean Collection was examined; a syntype of Tephritis hyoscyami and a possible syntype of Urophora solstitialis were identified. Syntypes of Tephritis arnicae, Urophora cardui and Rhagolelis cerasi were not found; specimens illustrated by Aldrovandi and de Reaumur are designated as lectotypes. No syntypic material of Euleia heracleii, Oxyna parietina, Terellia serratulae or Ensina sonchi could be found.