Diversity in neuroblast number forming mushroom bodies of the higher dipterans (Insecta,Diptera, Brachycera Cyclorrhapha)

Neurogenesis in mushroom bodies is studied in 12 species of the higher dipterans. A significant difference in the number of neuroblasts forming mushroom bodies is found. In the majority of species studied, Kenyon cells are formed by four solitary neuroblasts. Among six calliphorid species, the number of neuroblasts increases up to 10–15 (mean 12.6) in each mushroom body in Calliphora vicina only. In young pupae of Muscina stabulans and M. livida, four polyneuroblastic prolipherative centers occur instead of solitary neuroblasts. These centers disintegrate later into numerous solitary neuroblasts. A hypothesis on the origin of the polyneuroblastic structure of mushroom bodies found in C. vicina and, earlier, in Musca domestica, is proposed.     

Sperm ultrastructure in Chironomoidea (Insecta, Diptera)

The fine structure of spermatozoa from several species of chironomids, of Culicoides sp. (Ceratopogonidae) and of Odagmia pontina (Simulidae) was studied. A synapomorphic feature, consisting of nine kidney-shaped structures forming the centriole adjunct, was found in the chironomid species. All members of Chironomoidea share a mono-layered acrosome and a flagellar axoneme, provided with accessory tubules with 15 protofilaments in their tubular wall. The axoneme has a 9+9+2 pattern, but in an unidentified species of chironomid, a 9+9+0 model was observed where the central complex and the spokes are missing. Sperm motility is, however, maintained in all the examined species. The spermatozoa of this taxon have the tendency to complete maturation during their progression along the deferent ducts. Thus, in the proximal region of these ducts, they often show remnants of the spermatid cytoplasm.     

Morphology and function of the proboscis in Bombyliidae (Diptera, Brachycera) and implications for proboscis evolution in Brachycera

Based on serial semithin sections and SEM photographs of representatives of European Bombyliinae and Anthracinae, the mouthparts of Bombyliidae are studied and compared with the relevant data from literature on other families of Diptera Brachycera. The three moving units of the proboscis (clypeo-cibarial region, haustellum-maxillary base region, and labella) and their structures and muscles are described. Functions and possible movements are inferred from the structures observed. Articulations both between the parts of the organ and to the head capsule enable the fly to retract its proboscis into a resting position. Proboscis movement from a resting to a feeding position encompasses the following submovements: rotating of the basal clypeo-cibarial region (= fulcrum) against the head capsule, folding of the haustellum-maxillary base region against the fulcrum, evagination and invagination of the labial base, and the labella movements. This is a novelty as compared to the rigid proboscis of Tabanidae and agrees largely with the conditions in the Cyclorrhapha. The evolution of these novelties and their functional significance are discussed. The fulcrum, as well as the haustellum-maxillary base, as the new moving units are deduced from the plesiomorphic state as present in Tabanidae by fusions of sclerites, shifts of musculature and formation of new articulations. Accepted: 5 April 2000     

Regional differences in a nematoceran retina (Insecta,Diptera)

Summary The compound eye of Psychoda cinerea comprises two types of ommatidia, arranged so as to divide the retina into distinct dorsal and ventral regions. The P-type ommatidium, in the ventral part of the eye, differs fundamentally from the other dipteran ommatidia so far described, and is regarded as a primitive ommatidium. The acone dioptric apparatus is the same in both types, with a spherical lens and four Semper cells, the processes of which expand below the rhabdom to form a ring of pigment sacs. Only the distal region of the rhabdom is surrounded by a continuous ring of screening pigment, formed by 2 primary and 12–16 secondary pigment cells. The highly pigmented retinula cells penetrate the basement membrane proximally at about the level of their nuclei; in this region they are separated from the hemolymph by glial elements. The rhabdomeres R1–6 are fused to form a tube. The two types of ommatidia are defined by the arrangement of the retinula cells R7/8: in the T type the central rhabdomeres are one below the other, in the usual tandem position, whereas in the P type only R8 is central, with R7 in the peripheral ring. In the proximal region of the retina, retinula cells with parallel microvilli in neighboring ommatidia are joined in rows by lateral processes from the R8 cells. All the rhabdomeres are short and not twisted, which suggests that the retinula cells are highly sensitive to direction of polarization. The eye can adapt by a number of retinomotor processes. These findings, together with observations of behavior, imply that the psychodids have well-developed visual abilities.     

Campaniform sensilla of Calliphora vicina (Insecta,Diptera)

Summary A classification scheme of campaniform sensilla using morphological criteria was developed. All variations of the two most important outer structural elements, the cuticular cap and the cuticular collar, were taken into consideration: (a) the external shape of the cuticular cap; (b) the position of the cuticular cap in relation to the remaining cuticle; (c) the position of the cuticular collar in relation to the cuticular cap. This resulted in a classification of campaniform sensilla into 24 types. This typology was applied to the campaniform sensilla of Calliphora, which show considerable variations in their outer structures. According to SEM (scanning electron microscope) pictures and TEM (transmission electron microscope) sections we found only 9 out of 24 different types of campaniform sensilla in the fly.     

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.     

New taxa of snipe flies (Diptera: Brachycera: Rhagionidae) in the Daohugou Biota,China

Three impressions of snipe flies from the uppermost Middle Jurassic or lowermost Upper Jurassic Daohugou Formation in northeastern China are described as Protorhagio parvus sp. nov. and Lithorhagio megalocephalus get. et sp. nov. Protorhagio parvus is the first record of the genus Protorhagio Rohdendorf, 1938 outside the Karatau-Mikhailovka locality (Kazakhstan), which has implications for biostratigraphic correlation of the formations, in which they are found. The taxonomic position of Lithorhagio gen. nov. is discussed. Although Orsobrachyceron Ren, 1998 demonstrates some resemblance in form to the new genus, it probably belongs to Xylomyidae rather than Rhagionidae.     

Mechanisms of wing rotating regulation in Calliphora erythrocephala (Insecta,Diptera)

Summary The blowfly Calliphora erythrocephala rotates its wings, i.e. changes the geometrical angle of attack, generating forces and moments for flight steering. There are two possibile ways to regulate this angle. The mechanisms for these movements are described. (1) The leading edge and the anterior part of the wing — between the costal vein and radial vein 4 — are pronated automatically due to the interaction of the moving parts during the downstroke. They are supinated during the upstroke. This is basic automatic regulation. (2) The posterior part of the wing — behind the anterior cross vein — is pronated and supinated independently of wing-drive. This is wing-drive independent additional regulation.Abbreviations a.c anterior cross vein - a.n anal veins - a.t.l anterior tergal lever - a.w anterior part of the wing - b.z bending zone - co costal vein - cr crossing of the tendons of the posterior notal wing process - c.s cross section - cu cubital vein - f fit or turning point of ventral radial vein 1 - h.a horizontal axis of pterale III - h.c humeral cross vein - h.co head of costal vein - h.r head of radial vein - k Klöppel - l.a longitudinal axis - me median vein - mp middle plate - ms mesoscutum - p anterior process of the anal veins - p.c posterior cross vein - pl pleurum - p.n.w.p posterior notal wing process - p.n.w.p 1–4 muscles 1–4 of the posterior notal wing process - pt I–III pterale I–III - p.t.l posterior tergal lever - p.w. posterior part of the wing - p.w.j pleural wing joint - r 1–4 radial veins 1–4 - r.s. ring stiffenings - sc subcostal vein - s.p semicircular part of the middle plate - s.t subalar tendon - t.c tip cross vein - te tegula - t.st thin strips - t.v.r tooth of ventral radial vein - v.a. vertical axis of pterale III - w wing - III 1–4 muscles 1–4 of pterale III     

Formation,properties and degradation of the peritrophic membranes of larval and adult fleshflies,Parasarcophaga argyrostoma (Insecta,Diptera)

Summary Parasarcophaga argyrostoma larvae continuously secrete a single, tube-like peritrophic membrane (PM), which has an electron-dense layer on the lumen side and a thicker chitin-containing electron-lucent part on the epithelium side. In the adult fleshfly, the secretion of PMs starts immediately after emergence. The initial part of the PMs is twisted and tight. The formation zone is folded with two separate secretory pads in which two tube-like PMs are formed continuously. The PMs are different, morphologically and with respect to their peripheral carbohydrate residues. The latter could be demonstrated with lectin gold conjugates. PM 1 consists of an electron-dense, chitin-free layer on the lumen side and a thicker part which contains chitin microfibrils in the matrix. PM 2 appears fluffy and has chitin microfibrils in its matrix, too. Chitin could be localized with WGA gold. Incubation of isolated PM 1 with lectin gold resulted in a peculiar pattern of bound lectins and gaps on the electron dense layer which otherwise appeared to be homogenous. Degradation of peritrophic membranes takes place in the hindgut. The cuticle of the anterior hindgut is studded with small teeth, which seem to be responsible for mechanical degradation of the peritrophic membranes into frayed pieces. This may be completed by the teeth on the rectal pads. From the appearance of the remnants of the peritrophic membranes it can be inferred that chemical degradation takes place in the hindgut.Supported by the Deutsche Forschungsgemeinschaft     

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.     

Characteristics of the sperm flagellum in fungus gnats (Insecta,Diptera, Mycetophiloidea)

Spermatozoa from several members of the closely related Mycetophilidae and Keroplatidae were examined by electron microscopy using a fixative that contains glutaraldehyde and tannic acid, followed by a post-fixative that consists of uranyl acetate rather than osmium tetroxide. With this fixative, the detailed architecture of the flagellar axoneme and its various microtubules could be resolved. The so-called accessory tubules, which surround the central 9+2 unit of the sperm axoneme, were found to have 16 protofilaments in several examined Mycetophiloidea, but in no other Diptera. As 16 is the common number in holometabolic insects, it is presumably the plesiomorphic condition in Diptera. Other fungus gnats have accessory tubules with 15 or 14 protofilaments. The intertubular material situated between the accessory tubules is smaller in the examined members of the Mycetophilidae than in the Keroplatidae. The acrosome consists of an apical vesicle, which in one species, Macrorhyncha ancae, has three microtubular doublets in its anterior part and two large and three small extensions which extend posteriorly along the sperm axoneme.     

A monoclonal antibody against an adult-specific cuticular protein of Tenebrio molitor (Insecta, Coleoptera)

To study the sequential expression of the epidermal program in the mealworm Tenebrio molitor, monoclonal antibodies were prepared against the water-soluble proteins from preecdysial adult cuticle. Among the 16 clones obtained, one of them (named K2F6) recognized a 20-kDa antigen, found only in adult extracts but not in the larval or pupal ones, as revealed by immunoblot analysis. Our results strongly suggest an epidermal origin for this protein. The monoclonal antibody K2F6 fails to react with water-soluble proteins from fat body and hemolymph taken during the deposition of the 20-kDa antigen. Electron microscopic immunogold localization of this antigen showed that it is secreted, just after epicuticle deposition, in the 30 first-deposited preecdysial lamellae of sternal and elytral cuticles only. The sclerotizing process, which modifies the physicochemical properties of these cuticles, does not prevent the immunoreaction. When the expression of the adult program was inhibited by application of a juvenile hormone analog (ZR 515), the water-soluble proteins from different pupal-adult intermediates were never recognized by the monoclonal antibody K2F6 using immunoblot analysis. These results support the conclusion that this 20-kDa antigen is a protein specific for the sclerotized cuticle of the adult stage.     

Morphological phylogeny of the variable fly family Stratiomyidae (Insecta,Diptera)

Brammer, C. A. & von Dohlen, C. D. (2010). Morphological phylogeny of the variable fly family Stratiomyidae (Insecta, Diptera). —Zoologica Scripta, 39, 363–377. Stratiomyidae is a dipteran family distributed worldwide and containing 2800 species classified into 12 subfamilies. Previous phylogenetic work on the Stratiomyidae consisted of a 20‐character morphological analysis of the subfamilies [ World Catalog of the Stratiomyidae (Insect: Diptera). Leiden: Backhuys Publishers, 2001 ], and a molecular study using 69 taxa and two gene regions [ Molecular Phylogenetics and Evolution, 43, 2007, 660 ]. In this study, we present an expanded morphological cladistic analysis using 92 characters and 80 taxa, representing 36 of 39 described genera and all 12 Stratiomyidae subfamilies, as well as Xylomyidae and Pantophthalmidae outgroups. Data are analysed under maximum parsimony with all characters unordered and weighted equally; nodal support is assessed with the bootstrap and Bremer index. The strict consensus of all shortest trees is well resolved, and many of the deeper nodes are supported, although the root is ambiguous. Antissinae, Stratiomyinae, Sarginae and the diverse Clitellariinae are not monophyletic. Clitellariinae are positioned across several lineages, with most species grouped into a single, unsupported clade. Many of the well‐supported relationships are consistent with several aspects of the previous studies. The position of Exodontha remains elusive. Character support for subfamilies and other major clades is discussed.     

Feeding efficiency of Chaoborus flavicans (Insecta, Diptera) under turbulent conditions

Turbulence can affect predator–prey interactions. The effect of turbulence on the feeding efficiency of an ambush predator was tested with laboratory experiments. The experiments were conducted in 100-L aquaria in which ten individuals of fourth instar Chaoborus flavicans larvae were placed as predators. Two prey densities (3 and 10 ind. of Daphnia pulex L^?1) and two durations (30 and 120 min) were tested in a nonturbulent treatment and five different turbulence levels [average root-mean-square (RMS) velocities ranging from 0 to 7.3 cm s^?1, corresponding dissipation rates from 7.2 × 10^?7 to 1.3 × 10^?3 m² s^?3]. We hypothesized that the feeding rate of C. flavicans would be enhanced by turbulence due to increasing encounter rates up to a turbulence level above which a disturbance in post-encounter processes would lead to reduced feeding efficiency. However, the results showed no significant increase in the feeding rate of C. flavicans at intermediate turbulence. At high turbulence we found the expected significant negative response in the feeding rate of Chaoborus larvae. The feeding rate declined below the rates at nonturbulent and intermediate turbulence conditions as the average RMS velocity exceeded 3.1 cm s^?1 (dissipation rate 9.9 × 10^?5 m² s^?3, respectively).     

Macromycete fruit bodies as a habitat for dipterans (Insecta, Diptera)

The complex of Diptera (except for Sciaroidea) breeding in the fruit bodies of fungi is analyzed. The larvae of some species, such as Cecidomyiidae, Platypezidae, Heleomyzidae (Suilliinae), Anthomyiidae (Pegomya), Sphaeroceridae, and possibly Phoridae and Syrphidae feed on fresh fungal tissues. The ecological groups of flies are distinguished with respect to the type of feeding and the way of using fungal fruit bodies as habitat. The larvae of most groups developing inside the fruit bodies are saprobionts, typically occurring in decaying organic substrates of various origins.     

Monoclonal antibodies recognizing larval- and pupal-specific cuticular proteins of Tenebrio molitor (Insecta,Coleoptera)

To study the sequential expression of insect epidermal cells during metamorphosis, a library of monoclonal antibodies (MABs) was prepared against the water-soluble proteins from preecdysial pupal cuticle of Tenebrio molitor. Six selected MABs recognizing only larval and pupal cuticular proteins (CPs) in immunoblot analysis were classified into three types. Type 1 recognized a 21.5 and a 22 kDa polypeptide, type 2, a 26 kDa polypeptide, and type 3, three polypeptides of 18.5, 19.5 and 21.5 kDa. They did not immunoreact with any protein of fat bodies or haemolymph from pharate pupae, suggesting that the antigens originate from the epidermis. The stage-specificity was confirmed by electron microscopic immunogold labelling. Type 1 and 3 MABs recognized antigens characterizing larval and pupal preecdysial sclerotized cuticles, while the antigens recognized by type 2 were localized in the first few lamellae of unsclerotized postecdysial cuticle. When the expression of the adult programme was inhibited by application of a juvenile hormone analogue, the larval-/pupal-specific CPs were detected in the supernumerary pupal cuticle. These results suggest that the genes encoding these proteins are juvenile hormone dependent. These MABs should be useful tools to isolate pupal-specific genes whose regulation sems to be different from that of the adult-specific ones.     

Development of labellar taste hairs in the blowfly,Calliphora vicina (Insecta,Diptera)

Summary The early development of taste sensilla has been studied with special emphasis on cilia, dendrite, and pore formation.In the 39-h stage (the first stage investigated) differentiation of sensilla is already under way. The mechanisms of differentiation of dendrites (39–48 h) deviate from the mechanisms described for differentiation of true cilia. In taste hairs the centrioles meet in the tip of the narrowed apical region of the sensory neuron. Together they sink deeper into this region and line up coaxially, thus forming the basal body complex. Thereafter, lateral contacts between this complex and the plasma membrane of the neuron are established. Formation of open connections between the trichogen lumen of the hair and the environment, or the dendrite lumen, was not observed.Electrophysiological data indicate that the sensilla become functional from 3 days before emergence onwards.List of Abbreviations B Basal body complex - bc Basal compartment - bs Basal sheath - bl Basement lamina - D Dendrite(s) - EPC Non-sensillar epithelial cells - ES Ecdysial sheath - GERL Granular endoplasmatic reticulum-lysosomes - HS Hemolymph space - P Protrusions - PRN Prospective receptor neuron - RER Rough endoplasmatic reticulum - RN Receptor neuron - n Neck - THC Thecogen cell - TOC Tormogen cell - TRC Trichogen cell - TRS Trichogen sprout