The Tanyderidae are a small, ancient, poorly studied nematoceran family with 10 extant genera. А previous study found in the adults of Eutanyderus and Peringueomyina a full set of mouthparts (including mandibles) - a rare case among Diptera. The new study examined the mouthparts of adult Tanyderus, Araucoderus and Nothoderus using light and scanning electron microscopy. A full set of mouthparts, with no significant differences between sexes, was again found in all three genera. Three structural types were identified, suggesting as many feeding strategies among recent Tanyderidae. The genera form a series according to the degree of reduction of mandibles and other piercing structures. The well-developed toothed mandibles, hypopharynx, and laciniae were found only in both sexes of Tanyderus. Such a pattern has not previously been observed in males of any recent Nematocera. In the absence of live observations, the structure of these mouthparts suggests active penetration or laceration during feeding. Similar mouthparts, with both mandibles and hypopharynx toothed, are described, for the first time, in a female of Jurassic Protanyderus. For comparison, piercing mouthparts of two recent predaceous genera of Blephariceridae, Blepharicera and Liponeura, were examined. They showed numerous differences from the mouthparts of Tanyderus. Moreover, the mandibles, well-developed in the females, were absent in the males, suggesting sexual dimorphism in the feeding of these blepharicerids. Correlations between the structure of the mandibulate mouthparts, their sexual dimorphism, and the diet in lower Diptera are reviewed from the evolutionary perspective.

Immatures of Diptera are extremely rare in the Triassic fossil record and up to now they are only known from the early Anisian of Europe (Upper Buntsandstein of France and Spain). Herein, three immatures are described from the Norian (Upper Keuper) of Seinstedt, Germany: larva of Tanyderidae Larvaderus triassicus gen. et sp. nov. and pupa of Simuliidae Simulipupa curriei gen. et sp. nov., and Ramobuccina herrigi gen. et sp. nov., a pupa whose affinity is uncertain. Branched respiratory organs of both pupae suggest a rheophilic mode of life. The expansion of the order to (fast-) flowing waters no later than the Late Triassic is proven by the Norian immatures, which were strictly aquatic and could live without contact to the water surface in contrast to Anisian ones. The immature Tanyderidae and Simuliidae are the oldest records of these extant families. Simulipupa gen. nov. is the oldest record of a putative dipteran bloodsucker.

Four new species of Tega (Diptera: Anisopodidae) are described from the Jurassic of Eurasia: T. thuyi sp. nov. (early Toarcian Posidonia Shale Formation of Luxembourg), T. germanica sp. nov. (early Toarcian Posidonia Shale Formation and Grimmen Formation of Germany), T. storozhenkoi sp. nov. and T. sibirica sp. nov. (Bathonian Itat Formation of Central Siberia, Russia). These taxa demonstrate the stability of the wing venation of Tega during the Jurassic period, expanding the geographical and temporal framework of the genus significantly. A distribution map of Teginae and a key for genera and species are provided. The venation of the Jurassic Heterorhyphidae can be derived from the Tega venation. On the other hand, we consider the Mesozoic anisopodoid family Oligophrynidae with a revised composition, including Oligophryne, Austrorhyphus and Brachyrhyphus, as a probable ancestor of Anisopodidae.

First Upper Triassic dipterans (Nematocera) are described based on isolated wings from Fuchsberg near Seinstedt (Sevatian, uppermost Norian of Lower Saxony, Germany): Crivoptychoptera nebrias gen. et sp. nov. (Psychodomorpha, Ptychopteridae), Ptychopteridae incertae sedis, Dohloboyia triassica gen. et sp. nov. (Bibionomorpha, Boholdoyidae) and Chironomidae incertae sedis (Culicomorpha). All taxa under discussion present the oldest occurrences of their families. Geological background and composition of the Upper Triassic insect fauna of Seinstedt are briefly discussed.

Diptera often dominate fossil insect assemblages since the Jurassic but are much less common in the Triassic deposits, especially in the southern Pangaea, where only five species have previously been described—from Australia and Argentina. A single indeterminate Triassic dipteran from Africa has earlier been mentioned without a description. Herein, three dipteran pupae are described from the later Carnian Molteno Formation (Karoo Basin) in South Africa. One of these represents a new species of Metarchilimonia (Limoniidae), the genus previously known from the Norian Cow Branch Formation in the USA. The two other, incomplete pupae are placed in Chironomidae inc. sed. and Culicomorpha inc. sed. A complete list and a distribution map of Triassic Diptera are compiled, and the apparent geographical homogeneity of the Triassic dipterofauna is discussed.

Immature Diptera are more diverse and abundant in fresh water than any other insect order. The question arises whether the earliest dipterans, known from the Upper Buntsandstein of Europe (early Anisian, Middle Triassic), already developed in water and whether such mode of life was inherited from the ancestor of the order or if, alternatively, the dipteran larvae were ancestrally terrestrial. Nematoceran immatures have become common and diverse in the Mesozoic fossil record since the Middle Jurassic, but the vast majority of them represent only two culicomorphan families, Chaoboridae and Chironomidae. Earlier records and records of immatures of other families from the Mesozoic are extremely rare. A total of four larvae and about 30 pupae have been described from Anisian assemblages of France and Spain. Among these, one larva clearly belongs to Culicomorpha, the infraorder most closely associated with fresh waters, and one larva to Bibionomorpha, the infraorder most closely associated with terrestrial habitats, while the rest are hard to classify. Nevertheless, most of the pupae are morphologically similar to the semi-aquatic pupae of extant Limoniidae and Ptychopteridae from wet habitats. The oligopneustic respiratory systems of the Anisian larvae and their apparently allochthonous burial also suggest their semi-aquatic development, which appears to have been the ancestral mode of life of Diptera. The absence of dipteran immatures in lacustrine Konservat-Lagerstätten of the Madygen and the Cow Branch Formations (Ladinian–Carnian and Norian, respectively) is explained by the aquatic conditions being unfavorable for insect immatures and invertebrates in general.

The richest assemblage of the Triassic Diptera has been described from the famous Konservat-Lagerstätte Grès à Voltzia (Upper Buntsandstein) in the northern Vosges Mountains in France, dated as Early Anisian. A re-examination of the holotypes and additional material from the type locality allows for the establishment of Vogerhyphus gen. nov. and erection of the Vogerhyphinae subfam. nov. for Vymrhyphus blagoderovi Krzemiński and Krzemińska, 2003 and Vogerhyphus krzeminskorum sp. nov. (Protorhyphidae), and a new monotypic family Galliidae fam. nov. for Gallia alsatica Krzemiński and Krzemińska, 2003, originally described as Rhagionidae based on its wing venation. Galliidae fam. nov. is characterized by its closed cua cell and long moniliform antenna with 14-segmented flagellum and is hypothesized to belong to the stem-group Brachycera, along with the Late Triassic Prosechamyiidae. The process of brachycerization in the Diptera evolution is briefly discussed.

Blephariceridae, or net-winged midges, are a small group of highly specialized flies with immatures adapted to torrential streams. The family is known since the Jurassic, but the fossil record is very poor, as is typical of rheophiles. Up to now, only four species of three Mesozoic genera have been described, all of them from Asia. This paper provides a description of a male of Zwickina minaevi gen. et sp. nov. (Blepharicerinae), from mid-Cretaceous Burmese amber of northern Myanmar, and a brief discussion of the phylogenetic position of the new taxon. A key for identification of adults of extinct blepharicerid genera is included.

Here we provide evidence for the presence of non-biting midges of the subfamily Chironominae in the Mesozoic, based on descriptions of exceptionally rare amber inclusions. The subfamily has already been reported, but based on a single and unspecified record from Late Cretaceous Taimyr amber (similar to 84 Mya). That record is here revised and confirmed. Moreover, a new find in Burmese amber locates the temporal boundary of the subfamily back to the mid-Cretaceous (probably similar to 100 Mya). We describe two new genera assigned to the tribe Pseudochironomini: Mesoacentron gen. nov. with the species Mesoacentron kaluginae sp. nov. (Taimyr amber) and Palaeocentron gen. nov. with the species Palaeocentron krzeminskii sp. nov. (Burmese amber), the oldest known representative of the subfamily. The systematic position of the new taxa is discussed, and a key to the identification of adult males of extinct and extant Pseudochironomini genera is presented. Insights in the phylogeny and diversification tempo of the Chironominae and Pseudochironomini in the past are also provided.

No certain representatives ofCulicoidea have yet been found in Triassic deposits.Chaoboridae, commonly known as "phantom midges," is a small relict family of Culicoidea.At present, the family is represented by only six genera, together including about fifty species [Borkent, 2014].In the past it was more diverse: 17 Mesozoic genera can be considered valid, although this number can be overestimated.For example, Jordanobotomus Kaddumi, 2005 and Libanoborus Azar et al., 2009, from Jordanic and Lebanese ambers, close in geography and age, are candidates for synonymization once the type material is revised [Lukashevich, 2011].Chaoboridae flourished in Jurassic and Early Cretaceous fresh waters of Asia; their immatures and/or adults often quantitatively dominate in taphocenoses [Kalugina, 1980;Kalugina, Kovalev, 1985; Lukashevich, 1996bLukashevich, , 2008]].However, such abundance and diversity are recorded only since the Middle Jurassic [Kalugina, Kovalev, 1985], whereas from the Lower Jurassic deposits no immatures and only two chaoborid wings have been described.Both of the latter are representatives of Rhaetomyia Rohdendorf, 1962, known from Asia (Dzhil Formation, Sinemurian; Sogyuty locality, Kyrgyzstan) and Europe (Toarcian "Grünen Serie"; Grimmen locality, Germany) [Rohdendorf, 1964;Ansorge, 1996; Lukashevich, 1996a].The systematic position of Rhaetomyia has been subject of debate: Ansorge [1996] separated it as a monotypic family, ancestral to all other Culicoidea, whereas Amorim [1993] included the genus in Axymyiomorpha, a monophyletic group additionally including Axymyiidae, Perissommatidae, and Pacheneuridae.Arguments in favor of the chaoborid affinity of the genus have been presented previously [Lukashevich, 1996a], while a photograph of a Rhaetomyia wing is being published here for the first time.The new genus, described herein from the Triassic Dzhailoucho locality (Madygen Formation, Ladinian-Carnian, Kyrgyzstan), is the oldest member of the family.ABSTRACT.The fossil record of Triassic Culicomorpha is extremely poor, with the oldest member of the infraorder having been described, as Chironomoidea incertae familiae, from the the 'Grès à Voltzia' Formation (Upper Buntsandstein, early Anisian, France).In this paper, the oldest member of another superfamily, Culicoidea, is described from the Madygen Formation (Ladinian-Carnian, Kyrgyzstan) based on an isolated wing.The new genus, Triassomyia, gen.n., is assigned to the family Chaoboridae.РЕЗЮМЕ.Триасовые находки Culicomorpha исключительно редки, а древнейший представитель инфраотряда описан как Chironomoidea incertae familiae из формации Гре-а-Вольция (верхи пестрого песчаника, ранний анизий, Франция).В данной работе древнейший представитель другого надсемейства, Culicoidea, описывается из формации Мадыген (ладин-карний, Киргизия) по изолированному крылу.Новый род, Triassomyia, gen.n., отнесен к семейству Chaoboridae.

A new species of the extinct genus Archizelmira Rohdendorf, 1962 is described based on an isolated wing from the Middle Jurassic of West Siberia (Itat Formation, Bathonian). Both previously known species are re-examined. Additional wing venation characters are found for the genus, which support its status as a sister group to the rest of the family. A new Cretaceous subfamily, Burmazelmirinae subfam. nov. is proposed.

The fossil record of Triassic Diptera is still poor, with the oldest dipteran assemblage described from the Upper Buntsandstein of the 'Grs Voltzia Formation (early Anisian, France). From the stratigraphically closest insect fauna of the Rt Formation of Lower Franconia, Germany, the first Diptera, Bashkonia franconica gen. et sp. nov. is described based on an isolated wing. The new genus is assigned to the family Nadipteridae, bridging the gap between two other genera included.

This issue of Palaeoentomology is dedicated to the foremost Russian entomologist, palaeontologist, and evolutionary biologist, Alexandr Pavlovich Rasnitsyn, who will be 85 on September 24 this year. The authors and those numerous colleagues, who could not, for various reasons, participate, wish a happy birthday to the undisputed worldwide leader of palaeoentomology!

An Erratum to this paper has been published: https://doi.org/10.1134/S0031030121340013

The paper overviews the fossil record of insects with long mouthparts and rostra adapted to feeding on floral nectar and pollination drops of extinct gymnosperms. The presence of suctorial mouthparts is demonstrated for the first time for the Permian mecopterans Permochoristidae and Permotanyderidae. The long-proboscid scorpionflies Mesopsychidae are recorded for the first time from the Upper Jurassic of Kazakhstan. A new finding of a detached head of a long-proboscid nectar-feeding brachyceran fly is reported from the Lower Cretaceous of Transbaikalia. Three major radiations of long-proboscid nectar feeders are identified: the Paleozoic, the Mesozoic, and the Cenozoic one; they were related to the Paleozoic seed ferns, the Bennettitales and other Mesozoic entomophilous gymnosperms, and the flowering plants, respectively. The earliest long-proboscid nectar feeders, found in the Lower Permian deposits of the Cis-Urals, belong to Protomeropidae (stem-Amphiesmenoptera). The few other Paleozoic insects specialized to nectarivory probably also included some long-proboscid Permochoristidae. The diversity of long-proboscid nectar feeders shows a dramatic increase since the Middle Jurassic. About 70 Mesozoic species with preserved long mouthparts and rostra are known to date; they belong to 12 families and 3 orders (Mecoptera, Neuroptera, and Diptera) and can be clustered into three morphogroups. With the beginning of the Cenozoic the long-proboscid Mecoptera and Neuroptera were supplanted by Hymenoptera and Lepidoptera in the nectar-feeding niche, while Diptera on the whole retained this specialization. Considerable abundance of long-proboscid nectar feeders before the appearance of flowers with hidden nectar indicates that complex pollination systems first evolved in gymnosperms. Therefore, insect pollination cannot be considered the key novelty in flowering plants crucial for their evolutionary success.

The Bibionidae or march flies are a dipteran family with a worldwide distribution, presently including over 750 species in the extant faunas (Hesperininae sometimes treated as a separate family) (Pape et al., 2009). Emergence of adults is markedly seasonal, and adults are often observed visiting flowers, apparently feeding on nectar, pollen, and honeydew (Fitzgerald, 2009). Larvae are mainly phyto-saprophagous, found on superficial soil layers, decaying organic matter, leaf litter, dung, and rotten wood (Krivosheina & Mamaev, 1967; Pinto & Amorim, 2000; Fitzgerald, 2009). Larvae of Bibionidae usually occur in dense aggregations in suitable habitats and adults emerge synchronously in huge numbers and often form dense mating aggregations (Skartveit, 2017).

This issue of Palaeoentomology is dedicated to the foremost Russian entomologist, palaeontologist, and evolutionary biologist, Alexandr Pavlovich Rasnitsyn, who will be 85 on September 24 this year. The authors and those numerous colleagues, who could not, for various reasons, participate, wish a happy birthday to the undisputed worldwide leader of palaeoentomology!

Mouthparts of adult Tanyderidae are examined in detail for the first time, with emphasis on scanning electron microscopy. A full set of mouthparts, devoid of conspicuous dentation or serration and showing no differences between sexes, was found in both examined species, Eutanyderus wilsoni Alexander, 1928 and Peringueyomyina barnardi Alexander, 1921. The mouthparts are elongated into a proboscis in E. wilsoni and are short, attached to the tip of a long rostrum, in P. barnardi. In the absence of direct field observations, nectar-feeding is hypothesized for both genera.