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A review of leaf-miner tephritid flies (Diptera, Tephritidae) of the south-eastern part of West Siberia, Russia

Eleven leaf-miner tephritid fly species from 7 genera are reviewed for four regions in the south-eastern part of West Siberia, Russia, namely Tomskaya and Kemerovskaya Oblasts, Altaiskii Krai, and Republics of Altai and Khakassia. The share of leaf-miner species in the tephritid fly fauna of the region is 10.2%. Cornutrypeta svetlanae Richter & Shcherbakov, 2000, recorded from four regions, is the most numerous and widely distributed species preferring forest belt in the mountains of Kuznetskii Alatau (Kemerovskaya Oblast and Khakassia), North Altai (Republic of Altai) and West Altai (Altaiskii Krai). Two species are registered in only one region, namely Cornutrypeta spinifrons (Schroeder, 1913) in the forest belt of the Kuznetskii Alatau mountains (Kemerovskaya Oblast) and Hemilea dimidiata (Costa, 1844) in the forest meadows of Tomskaya Oblast. The rarest species in the region are Cornutrypeta spinifrons (Schroeder, 1913) and Trypeta zoe Meigen, 1826 known by single or several specimens only. Comments on the distribution and trophic relation of all the species are provided.


Introduction
Tephritid flies, characterized by their life-cycle that involves their pre-imaginal development in different parts of plants, include leaf-miner species which form a topological group within this family. Leaf-mining is recorded in both long-horned flies (Chironomidae, Sciaridae) and the short-horned flies (Agromyzidae, Anthomyiidae, Drosophilidae, Ephydridae, Psilidae, Scathophagidae, Syrphidae). The largest number are found in the family Agromyzidae, but other families include species representatives in several different genera; for example, the family Tephritidae includes several representatives of the tribe Trypetini which are known to be leafminers of herbal plants, mainly of the family Asteraceae.
Tephritidae is one of the largest dipteran family included about 4350 species from 480 genera worldwide, and about 850 species in Palearctic (Korneyev, Ovtshinnikova 2004). The tephritid fly fauna of Russia is still insufficiently known, only European part of Russia and the Russian Far East are relatively well studied. 190 species of tephritid flies from 53 genera were listed in the Key to insects of European Part of USSR (Richter 1970), of which 25 species from 16 genera were recorded for West Siberia. 250 species from 88 genera are registered for the Russian Far East, of which 39 species from 21 genera are occurring in West Siberia (Korneyev, Ovtshinnikova 2004), the territory which is carefully studying at present time and presumably inhabited by more than 100 species from 36 genera (Shcherbakov 2016).
The leaf-miner tephritid fly fauna in the south-eastern part of West Siberia have been studied in Tomskaya and Kemerovskaya Oblasts, Altaiskii Krai, Republics of Altai and Khakassia over the past 20 years. The first information regarding leafminer tephritid flies of West Siberia were presented in the review of species from Kuznetskii Alatau (Shcherbakov 2002) and the analysed data were published in an International Conference Proceedings (Shcherbakov 2015). To date, 10 leaf-miner tephritid fly species have been discovered in the south-eastern part of West Siberia, Russia. A list of these species is presented below and their taxonomic structure is given according to the Systematic Database of Tephritidae names (Norrbom et al. 1999). Data on trophic relation and a distribution of species are also provided. Most of host-plants are belonging to the family Asteraceae which is not noted in the list below, unless otherwise noted, name of plant family is provided.

Material and methods
The material was collected mainly by entomological net-sweeping, as imago-rearing from leaf-mines was not effective enough. Currently, trophic relations for a number of tephritid species in the region remains unknown.
Most of the material cited in the present article is housed in the collection of Tomsk State University, Russia (not noted in the list below), and the remainder is housed in the Zoological Institute, Saint Petersburg (ZIN); unless otherwise noted, specimens were collected by M. Shcherbakov. Trophic relations. Imagos of the species were collected from Saussurea latifolia, usually possessing leaf-mines, but rearing from the latter was not successful.

Taxonomy
Distribution. Only recorded from mountains of southern Siberia. Trophic relations. In Europe has been reared from leaf-mines on Hieracium, Lactuca and Taraxacum spp. (Merz 1994). Trophic relations in the region have not been investigated.
Distribution Trophic relations. Urtica dioica (Urticaceae) (Ellis 2020). A number of references provide data on the trophic relations of the species such as leaf-miners on stinging nettle, but in the Key to Tephritid Flies of Great Britain I, White (1988) commented on data by Beiger (1968) regarding mines located in the leaf petiole, but not the leaf lamina (White 1988). In A Handbook Plant Parasites of Europe devoted to leaf-miners, galls and fungal infection, this species is regarded as an inhabitant of stems lacking gall generation (Ellis 2020). A reference to the biology of Myoleja caesio was overlooked during the compilation of this Handbook. Ferrar (1987) noted that Beiger (1968) reared Philophylla caesio from mines in the petioles of an Urtica sp. (Urticaceae). The larva is described as having 19-20 anterior spiracle lobes, a number typical of stem-and leaf-mining Trypetini.
Distribution. Europe, Middle East, China; Russia: East and Central regions of European part, South Siberia, the Russian Far East (Korneyev, Ovtshinnikova 2004 (Ellis 2020). Trophic relations in the region have not been investigated, but leaf-mines were noted on Senecio nemorensis and Tussilago farfara.
Distribution. Europe, China, Japan, Russia: NW and Central regions of European part, Siberia, the Russian Far East (Korneyev, Ovtshinnikova 2004 (Ellis 2020). Trophic relations have not been investigated in the region.

Discussion
Analysis of leaf-miner tephritid fly species of the south-eastern part of West Siberia showed Cornutrypeta svetlanae Richter & Shcherbakov, 2000 to be the most numerous species, registered in four regions; it is the most widely distributed species preferring the forest belt in the Kuznetskii Alatau mountains (Kemerovskaya Oblast and Khakassia), North Altai mountain (Republic of Altai) and West Altai (Altaiskii Krai). Two species are registered in only one region, namely Cornutrypeta spinifrons (Schroeder, 1913) in the forest belt of Kuznetskii Alatau mountains (Kemerovskaya Oblast) and Hemilea dimidiata (Costa, 1844) in the forest meadows of Tomskaya Oblast.
Leaf-miner tephritid species are not large in number in comparison with the other representatives of the family which inhabit Asteraceae inflorescences. The rarest species, Cornutrypeta spinifrons (Schroeder, 1913) is known by a single specimen, and Trypeta zoe Meigen, 1826, also previously known from a single specimen, is currently found in several localities. According to A. Humala and А. Polevoy this species is rare and in Finland belongs to the Red Data NT category and therefore threatened with extinction in the near future, although it does not currently qualify for the threatened status (Humala, Polevoi 2015).
The share of leaf-miner species in the tephritid fly fauna of the region is 10.2% that corresponds with the data from other regions such as Great Britain (11%) (White 1988), Switzerland (8.3%) (Merz 1994) and the Ussuriisky State Reserve in the Asian part of Russia (9.6%) (Galinskaya, Ovtshinnikova 2014).
The study of trophic relations in leaf-miner tephritids is difficult because is not always possible to organize favourable conditions for larval and imago development. Up to date, only two species have been bred after regular collecting in different regions. Acidia cognata collected in Kuznetsky Alatau, Kemerovskaya Oblast was reared from leaf-mines on Petasites radiatus, and in Tomskaya Oblast from leaf mines on Tussilago farfara. Euleia heraclei was reared from leaf-mines of Heracleum dissectum in all collecting localities of the species. All other species have been collected strictly by the use of an entomological sweep net.
At present, the fauna of leaf-miner tephritid flies of the south-eastern part of West Siberia is almost completely revealed; the number of species could only be increased by a detailed study of the adjoining regions to the south. However, an improvement of imago rearing from leaf-mines could help in the determination of trophic relations and the distribution of the recorded species.