Phylogenetic analysis of the Blephariceromorpha,with special reference to mountain midges (Diptera: Deuterophlebiidae)

Abstract. A cladistic analysis of the Blephariceromorpha (here including the Nymphomyiidae, Deuterophlebiidae and Blephariceridae) and related Diptera provides a test of the phylogenetic hypotheses of Rohdendorf (1964, 1974), Hennig (1973), Wood & Borkent (1989) and Courtney (1990a). In particular, monophyly of the Blephariceroidea and Blephariceromorpha (sensu Wood & Borkent), and their relationship to other Diptera, is tested. Evaluation of larval, pupal and adult characters supports the hypothesis of Wood & Borkent, as modified by Courtney. Four larval features suggest that the Blephariceromorpha + Psychodomorpha form a monophyletic group, although an alternate hypothesis predicting that the Blephariceromorpha is the sister group of the Psychodomorpha + (Ptychopteromorpha + Culicomorpha), is discussed. Monophyly of the Blephariceromorpha (Nymphomyioidea + Blephariceroidea) is supported by one adult and five larval characters. Monophyly of the Blephariceroidea (Deuterophlebiidae + Blephariceridae) is supported by thirteen synapotypies, including features of the larva (six), pupa (three) and adult (four). Nineteen, nineteen and nine hypothesized synapotypies support monophyly of the Nymphomyiidae, Deuterophlebiidae and Blephariceridae, respectively.     

EVOLUTIONARY ORIGIN OF THE CYCLORRHAPHA (DIPTERA): TEST OF ALTERNATIVE MORPHOLOGICAL HYPOTHESES

Abstract— The higher flies, infraorder Cyclorrhapha [=Muscomorpha (McAlpine, 1989)], have undergone enormous radiation since the Cretaceous (∼100 Myr). Rapid morphological evolution in cyclorrhaphans has made their phylogenetic placement with respect to more primitive clades a long-standing problem in dipteran systematics. Of the two most plausible hypotheses, one treats the Cyclorrhapha as sister group to the orthorrhaphous superfamily Empidoidea [=Empidiformia (Hennig, 1948), Orthogenya (Brauer, 1883)], while the other places them within the empidoids. The debate over cyclorrhaphan origin has heretofore focused on homology interpretations for a few character systems, particularly the male genitalia. We provide the first attempt to assemble and quantify all of the available morphological evidence. By cladistic analysis of these data under alternative codings of genitalic features reflecting opposing homology theories, and then excluding these features altogether, we sought to judge which genitalic theory is better supported by the evidence as a whole, and how much the debate matters to resolving cyclorrhaphan origins. Using the analog of a factorial design, we also measured the effect of alternative transformation series in several other controversial characters, of outgroup choice and of successive weighting. Under all manipulations, including both genitalic codings, the Cyclorrhapha originate within the Empidoidea, near the family Atelestidae. However, trees in which the Empidoidea are constrained to be monophyletic are only 1-6 steps longer (out of ∼150), a fit not significantly worse under a permutation test for monophyly. Adult morphological data may not suffice to settle either the placement of Cyclorrhapha or the debate over genitalic homology. Moreover, the issue of genitalic homology does not appear critical to that of cyclorrhaphan origin.     

Morphological and other observations on Chonocephalus (Phoridae) and phylogenetic implications for the Cyclorrhapha (Diptera)

Chonocephalus blackithorum sp. nov . is described from Sulawesi. Biological observations on it and five other species are reported. In describing the female it proved necessary to re-evaluate the morphology of the female abdomen both in Chonocephalus and in the Phoridae as a whole. This adds to the growing evidence that the ground-plan of the Phoridae, and hence the Cyclorrhapha, is plesiomorphic with respect to the Empidoidea. The supposed affinity between these two groups is called into question and it is suggested that the Cyclorrhapha could even be an independent lineage from the Nematocera. The case for continuing to regard the Brachycera as monophyletic is also called into question by casting doubt on the rotation of the larval mandibles as a synapomorphy.     

Bryophyte-Feeders in a Basal Brachyceran Lineage (Diptera: Rhagionidae: Spaniinae): Adult Oviposition Behavior and Changes in the Larval Mouthpart Morphology Accompanied with the Diet Shifts

Dipteran larval morphology exhibits overwhelming variety, affected by their diverse feeding habits and habitat use. In particular, larval mouthpart morphology is associated with feeding behavior, providing key taxonomic traits. Despite most larval Brachycera being carnivorous, a basal brachyceran family, Rhagionidae, contains bryophyte-feeding taxa with multiple feeding habits. To elucidate the life history, biology, and morphological evolution of the bryophyte-feeding rhagionids, the larval feeding behavior and morphology, and the adult oviposition behavior of four species belonging to three genera of Spaniinae (Spania Meigen, Litoleptis Chillcott and Ptiolina Zetterstedt) are described. Moreover, changes of the larval morphology associated with the evolution of bryophyte-feeding are traced by molecular phylogenetic analyses. Spania and Litoleptis (thallus-miners of thallose liverworts) share a toothed form of apical mandibular sclerite with an orifice on its dorsal surface, which contrasts to those of the other members of Rhagionidae possessing a blade-like mandibular hook with an adoral groove; whereas, Ptiolina (stem borer of mosses) exhibits a weak groove on the adoral surface of mandible and highly sclerotized maxilla with toothed projections. Based on the larval feeding behavior of the thallus-miners, it is inferred that the toothed mandibles with the dorsal orifice facilitate scraping plant tissue and then imbibing it with a great deal of the sap. A phylogeny indicated that the bryophyte-feeding genera formed a clade with Spaniopsis and was sister to Symphoromyia, which presumably are detritivores. This study indicates that the loss or reduction of adoral mandibular groove and mandibular brush is coincident with the evolution of bryophyte-feeding, and it is subsequently followed by the occurrence of dorsal mandibular orifice and the loss of creeping welts accompanying the evolution of thallus-mining.     

Phylogenetic relationships and the larval head of the lower Cyclorrhapha (Diptera)

We examined final‐stage larvae of all currently recognized lower cyclorrhaphan (= Aschiza) families, except Ironomyiidae and Sciadoceridae, and those of the higher cyclorrhaphan (= Schizophora) families Calliphoridae, Conopidae, Lonchaeidae, Muscidae, and Ulidiidae, and compared them with larvae of two out‐group families, Rhagionidae and Dolichopodidae, paying particular attention to structures of the head. A set of 86 morphological characters were analysed phylogenetically. The results show that the lower Cyclorrhapha is paraphyletic in relation to the higher Cyclorrhapha. The monophyly of the Cyclorrhapha is strongly supported. The lower Cyclorrhapha is resolved into two clades, based on the Lonchopteridae. Within the Syrphidae the traditional three‐subfamily system is supported, based on the Microdontinae. Within the lower Cyclorrhapha, the larval head is variable in form and arrangement of components. In Lonchopteridae, the mouth lies at the back of an open trough or furrow, comprising ventrally an elongate labium and laterally the maxilla. This arrangement of components appears to facilitate scooping food in water films. In Platypezoidea there is no furrow, and the dorsolateral lobes bearing the antennae are connected by a dorsal extension of the pseudocephalon. The main food‐gathering structure is the hooked apex of the labium, but in Phoridae the mandibles may also be important. In Eumuscomorpha the mandibles are at the apex of the head skeleton. The pseudocephalon is extended and infolded dorsally to form an oral pocket over the mouth. In the Pipunculidae, and the Microdontinae and Syrphinae of the Syrphidae, ventrally it forms a V‐shaped groove or guide along which the mandibles project. The labium is sclerotized apically, and forms a plate or tapered projection. This arrangement of components facilitates holding, piercing and extracting prey tissues. In Eristalinae the pseudocephalon is attached to the mandibles and is formed into a pair of cirri bearing mandibular lobes that lie either side of the mouth. Furthermore, the epipharynx is produced anteriorly in relation to the hypopharynx, and the labium is attached to the anterior part of the epipharynx to form a cavity or atrium. This arrangement is suited to fragmenting and imbibing solid food in Eristalinae with hooked mandibles, and when the mandibles are reduced and the mandibular lobes are inverted and sclerotized, these structures form a filter for separating fluid‐suspended particulate food. In higher Cyclorrhapha an atrium is present as in Eristalinae, but a connection between the pseudocephalon and the mandibles is absent. Instead, the pseudocephalon is bifurcate dorsally and forms a pair of cephalic lobes that ventrally ensheath each mandible. The surface of the sheath may be coated in cirri and other food‐gathering structures. The cephalic lobes, mandibular sheaths and the head skeleton are maneuverable and retractile to a higher degree than in lower Cyclorrhapha. This arrangement of components facilitates feeding on both solid food, in which the mouthooks may extend from the sheath to break the food up, and particulate and suspended food, in which the food‐gathering structures of the sheath scoop up the food. In many higher Cyclorrhapha, maneuverability is enhanced by a break between the labium and the basal sclerite, to which it is fused in all lower Cyclorrhapha. Intermediate characters and states for the structures of the higher cyclorrhaphan larval head are present in out‐groups, and lower Cyclorrhapha and homologies are discussed. Liquidity of the food is an important factor explaining the structure of the larval head in Cyclorrhapha. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 153 , 287–323.     

The morphology of the larval head of Tipulidae (Diptera, Insecta) - The dipteran groundplan and evolutionary trends

External and internal head structures of the larva of Tipula montium are described in detail. The results are compared to conditions found in other representatives of Tipuloidea and other dipteran and antliophoran lineages. Despite of the conceivably basal position of Tipulomorpha within Diptera, the larvae are mainly characterised by derived features. The partially retracted head, the specific hemicephalic condition and several other derived character states support the monophyly of Tipuloidea. A clade comprising Tipuloidea excluding Pediciidae is suggested by the strongly retracted head, by deep dorsolateral incisions of the head capsule, by a distinctly toothed anterior premental margin, by the loss of the second extrinsic maxillary muscle, and possibly by the loss of the pharyngeal filter. Eriopterinae and Hexatominae are characterised by a tendency towards an extreme reduction of the head capsule. Limoniinae, Cylindrotomidae, and Tipulidae form a clade supported by the presence of a premaxillary suture. This implies the non-monophyly of Limoniidae. A feature shared by Cylindrotomidae and Tipulidae is the presence of a movable lacinia mobilis. However, this is arguably a plesiomorphic feature, as it also occurs in Nannochoristidae. Features of the larval head of Trichoceridae, which were included in Tipulomorpha, do not show affinities with those of Tipuloidea. Trichocerid larvae share a specialised subdivided mandible with larvae of psychodomorph groups. Tipuloidea are a highly specialised group. The characters examined did not reveal plesiomorphic features supporting a basal position, and features suggesting closer affinities with Brachycera are vague. The evolution of dipteran larval head structures was apparently strongly affected by the loss of legs and the tendency to live in cryptic habitats. Diptera are the group of Endopterygota with the highest number of apomorphic features of the larval head. The appendages are generally simplified and the muscular apparatus is strongly reduced. Specialised features evolving within dipteran lineages include specifically arranged brushes of hairs on the labrum and epipharynx, movable messores, subdivided mandibles, different mandibular brushes, and a far-reaching reduction of labial parts.     

The male postabdomen and reproductive system of Bibio marci Linnaeus, 1758 (Hexapoda: Diptera: Bibionidae)

The male postabdomen and the internal parts of the male genital system of Bibio marci (Bibionomorpha) were examined and reconstructed 3‐dimensionally. Several features differ from the presumptive dipteran groundplan. The bases of the gonopods are fused with each other and with tergite IX. The penis is not tube‐shaped and only sclerotized on the ventral side. The vasa deferentia are S‐shaped, and two pairs of accessory glands are present. In contrast to these characteristics, the arrangement of the internal parts is probably close to the ancestral condition. With its specific shape, the penis is well suited for the transfer of a spermatophore. The dorsal sclerite of the copulatory organ probably represents the medially fused parameres. A cladistic analysis of 27 characters of the postabdomen yielded two most parsimonious trees, with the strict consensus as follows: Nannochoristidae (outgroup) + (Culicidae [Culicomorpha] + ((Nymphomyiidae + (Tipulidae + Trichoceridae)) + (Tabanidae [Brachycera] + (Bibionidae, Anisopodidae, Axymyiidae [Bibionomorpha])))). Potential synapomorphies of Bibionomorpha (including Axymyiidae) and Brachycera are the fusion of sternum IX with the gonocoxites, the fusion of the parameres forming the dorsal sclerite and the presence of an entire series of postabdominal muscles (M4, M20, M23, M26, M27, M31, M35 and M37). The results of the analysis are preliminary as it is based on a single‐character system with a limited taxon sampling. However, the main result – a clade Bibionomorpha + Brachycera – is fully compatible with current hypotheses on dipteran phylogeny.     

Two remarkable new species of scuttle-fly (Diptera: Phoridae) that parasitize termites (Isoptera) in Sulawesi

ABSTRACT. Diplonevra mortimeri sp.nov. and D.watsoni. sp.nov. are described from specimens caught at Nasutitermitinae termite colonies in Sulawesi. The female of both species has a remarkable pre-oviposition behaviour in which she lures a worker termite into following her away from the colony. She then oviposits in the termite's abdomen. Pupation takes place within the remains of the host's abdomen. Both species only mature a single egg at a time. The female probably guards the comatose host termite during the period of larval development. The morphology of the male hypopygia of the new species allows reinterpretation of the peculiar anal tube in typical male Diplonevra. Furthermore the suggested homologies reinforce the view that the epandrium (tergite 9) has not been replaced by a periandrium (fused gonocoxites) in the Cyclorrhapha and consequently the latter cannot be derived from the Empidoidea.     

Evolution of the middle leg basal articulations in flies (Diptera)

The morphology of the coxa and trochanter was studied in 205 species from 68 fly families to compare these structures with respect to ability to fly in a streamlined posture, with the middle legs pointing forward and pressed to the thorax. Only Brachycera are able to attain this posture. The forward turn of the coxa at this position is hindered by the junction of the coxa with the pleuron. Recovery of mobility is gained in two ways. (1) By reduction of the contact zone between coxa and pleurite, as in Asiloidea, Bombyloidea, and Empidoidea. Within these flies, the streamlined posture was recorded in Bombyliidae and in a robber-fly, Laphria flava . Others fly with their middle legs straddled laterally or trailing backwards. (2) Longitudinal splitting of the coxa into three coxites provides intracoxal mobility in most Tabanoidea and Cyclorrhapha. The hind and medial coxites rotate about the front coxite and change the coxo-trochanteral axis, thus compensating for restricted protraction. Separation of the hind coxite appears in primitive Tabanoidea, and a separate middle coxite was found in several families among the Nematocera. The streamlined posture was recorded in horse-flies, stratiomyids, and in many Cyclorrhapha except Micropezidae and Hippoboscidae. There is morphological evidence for a possible secondary fusion of coxites at least in Dolichopodidae and Opetidae as well as for the origin of Cyclorrhapha from a miniature ancestor.     

An interesting new species of Megaselia from Sulawesi, the ground plan of the Phoridae (Diptera) and phylogenetie implications for the Cyclorrhapha

Megaselia biarticulata sp.n. is described from Sulawesi. Its distinctly two-segmented palp reinforces the view that a two-segmented palp is part of the ground plan of the family. The tibial hair palisades are also postulated as part of the ground plan and serve to link the Phoridae to the Platypezidae. The median furrow on the frons is considered as part of the ground plan. Furthermore it is postulated that it is homologous with the frontal vitta in the Schizophora, and that its invaginated lower end gave rise to the ptilinum. The Empidoidea cannot be ancestral to the Cyclorrhapha and the case for regarding these two taxa as sister groups seems highly tenuous in the light of the inferred ground plan for the Phoridae.     

骆驼斯氏副柔线虫病传播媒介西方角蝇和截脉角蝇的幼虫龄期划分

【目的】本研究旨在找出区分西方角蝇Haematobia irritans和截脉角蝇H.titillans幼虫龄期划分标准,为准确鉴定两种角蝇各龄幼虫,研究斯氏副柔线虫在角蝇体内的发育过程,以及制定防控骆驼斯氏副柔线虫病的有效措施等奠定基础。【方法】采用实验室人工孵育两种角蝇幼虫的方法,分别测量不同发育阶段幼虫的虫体长、咽骨体长和咽骨体宽3项指标,利用SPSS Statistics 19.0统计软件对数据进行处理,结合Crosby生长法则和线性回归的方法进行分析,比较两种角蝇幼虫之间差异,以确定两种角蝇幼虫最佳龄期划分标准。【结果】结果表明,两种角蝇的幼虫均分为3龄,咽骨体是两种角蝇幼虫龄期划分的特征性结构,两种角蝇各龄幼虫相同指标的测量值随龄期的增长呈现出相同的增长规律。咽骨体长是划分两种角蝇幼虫龄期的最佳测量指标,咽骨体宽可作为分龄的辅助指标;两种角蝇相邻龄期幼虫的体长变化范围存在相互重叠,不能准确划分角蝇幼虫龄期。【结论】研究表明通过西方角蝇和截脉角蝇幼虫咽骨体的形态特征可简便、快速和准确地鉴定两种角蝇幼虫的龄期。     

Phenetic and cladistic relationships among tenebrionid beetles (Coleoptera)

Abstract. The higher classification of Tenebrionidae is analysed using numerical phenetic, numerical cladistic and traditional Hennigian methods. In all, eighty characters are examined for about 335 taxa; definitive analyses are made on combinations of eighteen to seventy characters for thirty-three OTUs. At lower levels of relationship (genera and closely related tribes) phenetic and cladistic classifications are shown to be congruent, but at higher levels (tribes and subfamilies) there is marked discordance with phenetic results being more stable. A consensus classification is more similar to the Hennigian cladogram than is any single computer generated cladogram. Two main tribal groups – the Lagrioid and Tenebrionoid groups – are suggested which differ in defensive glands, female anatomy, wing and mouthpart morphology, larval characters and other features. The Tenebrionoid group consists of three main subdivisions – the tenebrionine, coelometopine and diaperine lineages. Changes in classificatory position are recommended for eighty-seven genera and tribes (listed in Appendix E) and implied for numerous others.     

Phytogeny of the nematocerous families of Diptera (insecta)

The relationships of the nematocerous families of Diptera are cladistieally analysed using the parsimony programs PAUP and Hennig86. An extensive review, as well as a data matrix, is presented for 98 almost exclusively morphological characters (larva, 56; pupa, 6; adult, 36). Four infraorders are recognized, viz , Ptychopteromorpha, Culicomorpha, Blephariceromorpha, Bibionomorpha, and a clade containing the 'higher Nematocera' and Brachycera. Traditionally the family Nymphomyiidae or the infraorder Tipulomorpha (=Tipulidae, with or without Trichoceridae) are considered the most basal clade of the extant Diptera. On the basis of our cladistic analysis it is suggested that the Ptychopteromorpha-Culicomorpha clade is the sister-group of all other extant Diptera. We provide evidence that the Axymyiidae are part of a monophyletic Bibionomorpha. The latter infraorder is proposed as the sister-group of the higher Nematocera and Brachycera. We transfer the Tipulidae (Tipulomorpha) to the higher Nematocera, at a position next to Trichoceridae and near the Anisopodidae-Brachycera lineage. Previous hypotheses concerning nematocerous relationships are reviewed.     

恒温条件下盐酸吗啡对肥须亚麻蝇幼虫的影响及法医学意义

研究石家庄地区优势蝇种肥须亚麻蝇Parasarcophaga crassipalpi(Macquart)在不同浓度的盐酸吗啡作用下的生长发育情况,探索毒物对肥须亚麻蝇幼虫口钩和咽骨的影响,积累石家庄地区尸源性蝇类的法医昆虫毒理学(forensic toxicologic entomology)资料,从而为死后间隔时间(postmortem interval,PMI)的推断提供科学依据。     

CLADISTIC TESTS OF ADAPTATIONAL HYPOTHESES

Abstract— A cladistic viewpoint provides an historical definition of adaptation and an operational ecological test for evolutionary adaptations. Adaptation is apomorphic function promoted by natural selection, as compared with plesiomorphic function. Adaptation is thus a conditional, hierarchical, comparative term, like homology. Hypotheses of adaptation that do not specify levels of apomorphy are weak; they should refer to and explain the function at the level at which it is apomorphic with respect to the plesiomorphic (outgroup) condition. The adaptational hypothesis serves as a prior prediction in the comparison of the apomorphic function of the derived trait with the plesiomorphic function of the plesiomorphic trait serving as the null hypothesis. It is useful to distinguish whether hypotheses about characters identify selection as facilitating: 1) the origin of a character; 2) its maintenance; 3) neither; or 4) both. The latter two are uniformitarian and testable in a strong sense. The former two possibilities use ancillary arguments to protect the hypothesis of the role of natural selection in one way or another, but might still be tested by the weak criterion of plausibility. Given an hypothesis of both origin and maintenance due to selection, the test of adaptation may still be thwarted because only certain kinds of cladistic structure allow feasible tests. Few of the really classic and common examples of supraspecific adaptation survive this kind of cladistic test.     

The larval hyobranchial apparatus of discoglossoid frogs: its structure and bearing on the systematics of the Anura (Amphibia: Anura)

The morphology of the larval hyobranchial apparatus of discoglossoid frog species representing the genera Ascaphus, Alytes, Bombina, and Discoglossus is described and the resulting characters were analysed cladistically. Seven species representing seven major lineages of frogs were included in the cladistic analysis of characters. Several changes in the terminology of the musculature are introduced, and a new interpretation of the subarcualis-muscle system is presented. The phylogenetic analysis suggest that the hyobranchial apparatus was substantially altered in the lineages leading to and within the Pipanura. This notably involved fusion, reduction and loss of skeletal structures and muscles, and splitting of certain muscles into muscle groups. The result confirm previous hypotheses based on the study of adults: discoglossoid species retain the most numlerous plesiomorphic characters among extant ianurans. The larval hyobranchial apparatus is in many features structrually similar to that of urodeles. Many of their character states were most likely present in the most recent common ancestor of all living forgs. The cladistic analysis of 31 characters of ithe larval hyobranchial apparatus supports major clades: Anura, Bombinanura, Pipanura, and Pelobatoidea + Neobatrachia. The cladiostic analysis and interpretation of larval characters is in part compatible with phylogenetic hypotheses based on characters of adults and rRNA sequences, but is in conflict with the Mesobatrachia and Archaeobatrachia concepts of other authors.     

The male abdominal,genital and pregenital sclerites and musculature in <Emphasis Type="Italic">Neria commutata</Emphasis> (Czerny, 1930) (Diptera,Micropezidae)

The muscles of the male abdomen and genitalia of Micropezidae were studied for the first time by the example of Neria commutata (Czerny, 1930). Based on analysis of the sclerites and musculature of the male genitalia of Micropezidae as compared to those of the previously studied Acalyptratae and Aschiza, we revealed several apomorphies of this group. The hypandrial complex is characterized by the presence of the phallic retractors and protractors M1 and M2, and the epandrial complex, by the presence of muscles M3 of the subepandrial sclerite, muscles M4 of the surstyli, muscles M7 of the cerci, and also the tergosternal muscles M5; all these muscles correspond to the ground plan of Cyclorrhapha. The following characters are considered apomorphic: the splitting of intersegmental sternal muscles ISM5–6 into 4 pairs that ensure the functioning of the forcipate appendages of sternite V; development of syntergosternite VII and reduction of muscles ISM6–7; the splitting of muscles M3 of the subepandrial sclerite into 4 pairs, enhancing the function of this sclerite; the appearance of pregonites with the associated muscles M42, which probably occurred independently several times in the evolution of different groups of Cyclorrhapha; asymmetry of syntergosternites VII and VIII and their muscles. The sclerites and muscles of the epandrium and hypandrium are characterized by complete symmetry.     

THE NATURE OF CLADISTIC DATA

Abstract— Cladistic data are the characters of organisms. Character is defined as a feature that can be evaluated as a variable with two or more mutually exclusive and ordered states. Cladistic characters must be treated as multistate variables, and coded as sequential numbers or in additive binary fashion. Any other interpretation and handling of cladistic data will introduce error into analysis. Character states cannot be treated independently as present or absent, i.e., as nominal variables, because redundancy is introduced into the data and information content is sacrificed. Non-additive binary coding demonstrates that treating cladistic variables as nominal data will lead to multiple, equally parsimonious solutions. Defining characters found universally in a group of organisms, but unknown outside those organisms have no alternative state that can be designated as absent. Absence, however, is valid as a character state if it can be shown to be apomorphic. When two or more character states occur within a taxon, that taxon must be coded as having an unknown state for that character, or the taxon must be split in two or more taxa. Continuously varying quantitative data are not suitable for cladistic analysis because there is no justifiable basis for recognizing discrete states among them. Quantitative data are questionable even when they exhibit mutually exclusive states because the states can be interpreted only in reference to an archetype, i.e., as implied homologies not subject to test.     

On the fauna of the Diptera Brachycera of Samarskaya Luka: 1. Brachycera Orthorrhapha; Cyclorrhapha: Aschiza

The presumptive data on results of the investigation of the Diptera Brachycera in the territory of Samarskaya Luka, the largest refuge in eastern European Russia, are given. Based on the published data and on examination of 18 000 specimens, a faunal list for the region is compiled, including 897 species of 395 genera of 62 families. The annotated list of Brachycera Orthorrhapha, Cyclorrhapha: Aschiza includes data on the flight periods, habitats, and number of examined specimens.