Taxonomic status of two European sibling and barcode-sharing species of Brachypogon Kieffer, 1899 (Diptera: Ceratopogonidae).

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A new giant dung midge from Miocene Ethiopian amber (Diptera: Scatopsidae).

Fossil new genus and species Afroscatopse haennii gen. et sp. nov. from Miocene Ethiopian amber are described and illustrated and placed in the tribe Colobostematini. The gigantic adults of this scavenger fly, largest within the family, with dull body, long antennae and broad wings evidently were adapted to live on bark of trees. It is suggested that it was a myrmecophilous species with larvae living in the nests of tree ants.

New records of biting midges (Diptera: Ceratopogonidae) from the United Arab Emirates, with a description of a new species.

Biting midges (Ceratopogonidae) are a relatively well-studied family of nematocerous flies (Borkent Dominiak 2020), but the ceratopogonid fauna of the Middle East is not well known. To date, only 264 species from this family have been recorded in this entire region (Alwin et al. 2016a, b) and just 53 in the United Arab Emirates (UAE) (Szadziewski et al. 2011, Alwin et al. 2016a, b). Here, we report on three species of biting midges new to the fauna of the UAE.

A new species of Bruchomyiinae (Diptera: Psychodidae) from Baltic amber.

Bruchomyiinae is one of seven subfamilies of Psychodidae. This small group comprises fewer than 60 extant species distributed mainly in tropical and sub-tropical regions (Wagner Stuckenberg 2016). All life stages of these flies are closely tied to forest environment (Fairchild 1952; Stuckenberg 1962) and as a result, bruchomyiines are frequently preserved in fossil resins. The first fossil member of this subfamily was described from Baltic amber (45 Ma) (Meunier 1905) and Burmese amber (100 Ma) is the oldest fossil resin in which representatives of Bruchomyiinae have been found (Stebner et al. 2015; Wagner 2017; Skibinska et al. 2019). Due to the relative scarcity of this subfamily in Baltic amber, any new specimens are of great interest. Up till now, there are only 12 bruchomyiine species described from Baltic amber. Wagner (2017) completed a revision of fossil Bruchomyiinae and proposed the genus Hoffeinsodes to include the species having male genitalia with gonocoxites fused with hypandrium. In this genus, he classified six species reported exclusively from Eocene Baltic amber: H. bifida Wagner, 2017, H. cubicula Wagner, 2017, H. longicauda Wagner, 2017, H. obtusa Wagner, 2017, H. reducta Wagner, 2017 and H. hoffeinsi (Wagner, 2006) transferred from Nemopalpus (Wagner, 2006). Here, we describe a new species belonging to this genus.

Haematophagous biting midges of the extant genus Culicoides Latreille (Diptera: Ceratopogonidae) evolved during the mid-Cretaceous.

Four new fossil species of haematophagous biting midges of the genus Culicoides Latreille, from mid-Cretaceous Burmese amber are described and illustrated: C. bojarskii Szadziewski Dominiak sp. nov., C. burmiticus Szadziewski Dominiak sp. nov., C. ellenbergeri Szadziewski Dominiak sp. nov. and C. myanmaricus Szadziewski Dominiak sp. nov. These extinct species are assigned to the new subgenus, Groganomyia Szadziewski Dominiak subgen. nov. which also includes an extant species that inhabits European mountains, Culicoides cameroni Campbell Pelham-Clinton, 1960, the type species. These very old (99 Ma) haematophagous biting midges of the extant genus Culicoides from Burmese amber supports the hypothesis that most groups of modern biting midges evolved during the mid-Cretaceous greenhouse climate.

Biting Midges (Diptera: Ceratopogonidae) from Cambay Amber Indicate that the Eocene Fauna of the Indian Subcontinent Was Not Isolated.

India's unique and highly diverse biota combined with its unique geodynamical history has generated significant interest in the patterns and processes that have shaped the current distribution of India's flora and fauna and their biogeographical relationships. Fifty four million year old Cambay amber from northwestern India provides the opportunity to address questions relating to endemism and biogeographic history by studying fossil insects. Within the present study seven extant and three fossil genera of biting midges are recorded from Cambay amber and five new species are described: Eohelea indica Stebner & Szadziewski n. sp., Gedanohelea gerdesorum Stebner & Szadziewski n. sp., Meunierohelea cambayana Stebner & Szadziewski n. sp., Meunierohelea borkenti Stebner & Szadziewski n. sp., and Meunierohelea orientalis Stebner & Szadziewski n. sp. Fossils of species in the genera Leptoconops Skuse, 1889, Forcipomyia Meigen, 1818, Brachypogon Kieffer, 1899, Stilobezzia Kieffer, 1911, Serromyia Meigen, 1818, and Mantohelea Szadziewski, 1988 are recorded without formal description. Furthermore, one fossil belonging to the genus Camptopterohelea Wirth & Hubert, 1960 is included in the present study. Our study reveals faunal links among Ceratopogonidae from Cambay amber and contemporaneous amber from Fushun, China, Eocene Baltic amber from Europe, as well as the modern Australasian and the Oriental regions. These findings imply that faunal exchange between Europe, Asia and India took place before the formation of Cambay amber in the early Eocene.

Biting midges of the subfamily Forcipomyiinae (Diptera: Ceratopogonidae) from the Middle East, with keys and descriptions of new species.

Middle East biting midges of the genera Atrichopogon Kieffer and Forcipomyia Meigen, subfamily Forcipomyiinae Lenz, covering 41 species are reviewed. Two new species are described and illustrated: Forcipomyia (F.) siverekensis Alwin & Szadziewski sp. nov. and Forcipomyia (Microhelea) borkenti Alwin & Szadziewski sp. nov. The list includes 16 species of Atrichopogon and 25 of Forcipomyia. Nine species previously described by Vimmer and Kieffer from the Middle East are treated as nomina dubia and not included in the list.        Keys to identification of Atrichopogon and Forcipomyia species of the Middle East are also provided.

A fossil biting midge (Diptera: Ceratopogonidae) from early Eocene Indian amber with a complex pheromone evaporator.

The life-like fidelity of organisms captured in amber is unique among all kinds of fossilization and represents an invaluable source for different fields of palaeontological and biological research. One of the most challenging aspects in amber research is the study of traits related to behaviour. Here, indirect evidence for pheromone-mediated mating behaviour is recorded from a biting midge (Ceratopogonidae) in 54 million-year-old Indian amber. Camptopterohelea odora n. sp. exhibits a complex, pocket shaped structure on the wings, which resembles the wing folds of certain moth flies (Diptera: Psychodidae) and scent organs that are only known from butterflies and moths (Lepidoptera) so far. Our studies suggests that pheromone releasing structures on the wings have evolved independently in biting midges and might be much more widespread in fossil as well as modern insects than known so far.

Biting midges of the tribe Ceratopogonini (Diptera: Ceratopogonidae) from the Middle East, with keys and descriptions of new species.

Middle East predatory biting midges of the tribe Ceratopogonini, covering 22 species of 7 genera are reviewed. Three new species are described and illustrated: Allohelea israelensis Szadziewski & Alwin sp. nov., Kolenohelea levantica Szadziewski & Alwin sp. nov. and Serromyia galilaeae Szadziewski & Alwin sp. nov. The genus Boreohelea Clastrier & Delécolle, 1990 syn. nov. is recognized as a junior synonym of Allohelea Kieffer, 1917. Thysanognathus nilogenes Kieffer, 1925 syn. nov. from Egypt is a junior synonym of Alluaudomyia melanosticta (Ingram & Macfie, 1922). Keys to identification of subfamilies, tribes, genera and species of Ceratopogonini of the Middle East are also provided.

A redescription of Culicoides griseidorsum Kieffer, 1918, with comments on subgeneric position of some European taxa (Diptera: Ceratopogonidae).

Culicoides griseidorsum Kieffer, 1918 is formally redescribed, and the male described and illustrated for the first time. The species is placed within Sensiculicoides Shevchenko, 1977 the subgenus restored from the synonymy with Oecacta Poey, 1853. A checklist of European species placed in subgenera Sensiculicoides and Oecacta is provided.

All stages of the Palaearctic predaceous midge Palpomyia schmidti Goetghebuer, 1934 (Diptera: Ceratopogonidae).

All stages and the ecology of the Southern Palaearctic Palpomyia schmidti collected from the vicinity of the saline Lake Elton in Russia are described and illustrated. The morphology of larvae and pupae as well as the detailed ecology of the larvae are described for the first time. P. schmidti is a halobiontic biting midge, widely distributed in the steppes and deserts of the Palaearctic region. It is proposed that the Palpomyia schmidti group should include five Holarctic species. P. downesi Grogan & Wirth, 1979 from north-western North America is recognized as a new junior synonym of the Eastern Palaearctic P. tuvae Remm, 1972. New synonymy.

Two new species of predatory biting midges of the genus Alluaudomyia from Europe and the Canary Islands (Diptera: Ceratopogonidae).

Alluaudomyia canariensis Szadziewski & Dominiak sp. nov. from the Canary Islands and A. wyskokensis Szadziewski & Dominiak sp. nov. from Poland and Ukraine are described and illustrated. The genus Alluaudomyia is reported from the Canary Islands for the first time. The article is supplemented with a checklist and an identification key for the species so far recorded from Europe and the Canary Islands.

Afrostilobezzia, a new genus of predatory biting midges from the Afrotropical Region (Diptera: Ceratopogonidae).

A new genus Afrostilobezzia gen. nov. including A. clastrieri sp. nov. and A. ornatithorax (Clastrier, 1988) comb. nov. is described from West Africa (Nigeria and Guinea).

Ultrastructure of mediodorsal setae in biting midge larvae of the genus Atrichopogon Kieffer with notes on their biological significance (Diptera: Ceratopogonidae).

The ultrastructure of the strong mediodorsal setae in terrestrial stage IV larvae of Atrichopogon (Meloehelea) oedemerarum and A. (M.) meloesugans was examined using light, scanning and transmission electron microscopy. Serrated setae placed on prominent processes are distributed in pairs on all thoracic and abdominal segments. Setae are innervated by a single dendrite and their surface has no pores. The trichogen cell is not retracted from the setal lumen on completion of the hair-forming process but fills the mediodorsal seta also when the larval cuticle is fully sclerotised. Such a phenomenon was previously reported in terrestrial larvae of the genus Forcipomyia. We suggest that the mediodorsal setae described in Atrichopogon are plesiotypic mechanoreceptors for the subfamily Forcipomyiinae. They are preserved in the truly terrestrial larvae of Atrichopogon, but modified to secretory setae in the genus Forcipomyia. Both genera bearing distinct mediodorsal setae have developed functional tracheal gills, unknown in other biting midges.

First description of polytene chromosomes in biting midges (Diptera: Ceratopogonidae).

Polytene chromosomes are described from secretory cells in larvae of Forcipomyia nigra (Winnertz). They are present in large glandular-trichogen cells at the bases of secretory setae and in midgut cells that were observed by transmission electron microscopy and light microscopy. Polytene chromosomes, isolated from the glandular-trichogen cells using aceto-orcein squash technique, measure 50-200 microm, have braid-like strands of chromatin and no bands, features that are unique within the Culicomorpha.

Composition and antimicrobial activity of fatty acids detected in the hygroscopic secretion collected from the secretory setae of larvae of the biting midge Forcipomyia nigra (Diptera: Ceratopogonidae).

The hygroscopic secretion produced by the secretory setae of terrestrial larvae of the biting midge Forcipomyia nigra (Winnertz) was analysed using gas chromatography coupled with mass spectrometry (GC-MS). The viscous secretion is stored at the top of each seta and absorbs water from moist air. GC-MS analyses (four independent tests) showed that the secretion contained 12 free fatty acids, the most abundant of which were oleic (18:1), palmitic (16:0), palmitoleic (16:1) and linoleic (18:2). Other acids identified were valeric (5:0), enanthic (7:0), caprylic (8:0), pelargonic (9:0), capric (10:0), lauric (12:0), myristic (14:0) and stearic (18:0). Two other compounds, glycerol and pyroglutamic acid, were also found. The antibacterial activity of the fatty acids and pyroglutamic acid was tested using the agar disc diffusion method and targeted Gram positive (Bacillus cereus, Bacillus subtilis, Enterococcus faecalis) and Gram negative bacterial strains (Citrobacter freundii, Pseudomonas aeruginosa, Pseudomonas fluorescens). The antifungal activity was tested by determining minimal inhibitory concentration (MIC) of examined compounds. Fatty acids were tested against enthomopathogenic fungi (Paecilomyces lilacinus, Paecilomyces fumosoroseus, Lecanicillium lecanii, Metarhizium anisopliae, Beauveria bassiana (Tve-N39), Beauveria bassiana (Dv-1/07)). The most effective acids against bacterial and fungal growth were C(9:0), C(10:0) and C(16:1), whereas C(14:0), C(16:0,) C(18:0) and C(18:1) demonstrated rather poor antifungal activity and did not inhibit the growth of bacteria. The antimicrobial assay investigated mixtures of fatty and pyroglutamic acids (corresponding to the results of each GC-MS test): they were found to be active against almost all the bacteria except P. fluorescens and also demonstrated certain fungistatic activity against enthomopathogenic fungi. The hygroscopic secretion facilitates cuticular respiration and plays an important role in the antimicrobial protection of F. nigra larvae living in moist terrestrial habitats.

Morphology and histology of secretory setae in terrestrial larvae of biting midges of the genus Forcipomyia (Diptera: Ceratopogonidae).

Apneustic larvae of the genus Forcipomyia possess unique secretory setae located on the dorsal surface along the body in two rows, one pair on each thoracic and abdominal segment and two pairs on the head. Morphological and histological studies of secretory setae in fourth instar larvae of Forcipomyia nigra (Winnertz) and Forcipomyia nigrans Remm indicate they are modified mechanoreceptors (sensilla trichodea) in which the trichogen cell is a glandular cell producing a hygroscopic secretion. The cytoplasm of the glandular trichogen cell fills the lumen of a secretory seta, which shows one or more pores on the apex. The cytoplasm contains numerous microtubules responsible for transportation of proteinaceous vesicles, and an extremely large polyploid nucleus typical of gland cells. The main role of the hygroscopic secretion is to moist the body and thus facilitate cuticular respiration.