Contributions to the study of the subtribe Eremophlepsiina Dmitriev, 2002 (Hemiptera: Cicadellidae: Deltocephalinae: Opsiini).

Three species of Eremophlepsiina, Eremophlepsius rohdendorfi, E. sexnotatus, and Pseudophlepsius binotatus are redescribed, variability of their coloration and genitalia is illustrated, the status of some subspecific taxa is discussed. Oscillograms of the male calling signals of E. sexnotatus and P. binotatus are given. The synonymy E. sexnotatus Kusnezov, 1929 = E. parvulus Dlabola, 1961, syn. n. is established.

An enigma of Handianus Ribaut, 1942 (Hemiptera: Cicadellidae: Deltocephalinae: Athysanini): identical host preferences and male calling signals in two morphologically distinctive sympatric species.

Handianus eurotiae and H. fartilis distinctly differ in the forewing venation and the male genitalia shape, but have strongly overlapping ranges, do not differ in ecological preferences, feed on the same host plant, Krascheninnikovia ceratoides, and produce almost identical male calling signals. They were never found in the same biotope, although the mechanism of such vicariation is not clear. According to Mayr's biological species concept, such taxa are good species, but Paterson's species recognition concept is inapplicable in this case. Similar examples in Orthoptera and Auchenorrhyncha are considered.

Contributions to the study of the genus Mongolojassus Zakhvatkin, 1953 (Homoptera: Cicadellidae: Deltocephalinae: Paralimnini) with description of a new species from Northern Caucasus.

Mongolojassus caucasicus sp. n. from the alpine zone of Dagestan, Northern Caucasus is described, new data on distribution and illustrated descriptions of M. bicuspidatus (J. Sahlberg, 1871) and M. vinokurovi Emelyanov, 1976 are provided. Information on the type series of M. elpatjevskii (Zakhvatkin, 1953) is given. Ecological preferences and distribution patterns of species of Mongolojassus are discussed.

Macropsidius bogutensis (Mityaev, 1990) species group (Hemiptera: Auchenorrhyncha: Cicadellidae: Eurymelinae: Macropsini) in Kazakhstan: taxonomy, biology, and evolution.

Illustrated descriptions of species from the Macropsidius bogutensis group from Southern Kazakhstan with data on biology, male calling signals, and distribution are provided. M. karatavicus sp. n. from Syrdaryinskiy Karatau Range is described and the synonymy M. bogutensis Mityaev, 1990 = M. zhuravlevi Mityaev, 2014, syn n. is established. Conclusions concerning the evolution of M. bogutensis species group are presented.

Review of the Gargara genistae (Fabricius, 1775) species group (Hemiptera: Membracidae) from Russia and adjacent territories.

In Russia, Kazakhstan, and Central Asia the Gargara genistae species group includes three species differing from each other in the male calling signal patterns, but almost indistinguishable in external appearance and the male genitalia shape. Illustrated descriptions of morphology and the male calling signals, and data on host plants and distribution for all species are given. Considerations concerning possible modes of species divergence in this group are provided.

On the morphological variability of Limotettix (Limotettix) striola (Falln, 1806) (Homoptera: Auchenorrhyncha: Cicadellidae: Deltocephalinae: Limotettigini) with notes on the taxonomic status and distribution of L. striola ferganensis Dubovskiy, 1966.

Investigation of the male calling signals of Limotettix (Limotettix) striola (Falln, 1806) in populations from central and southern European Russia, Northern Caucasus, and West Tien Shan, Kyrgyzstan demonstrated that they belong to the same species. Males of L. striola from West Tien Shan have a somewhat different penis shape and belong to a separate subspecies, L. striola ferganensis Dubovskiy, 1966. L. striola ferganensis from West Tien Shan differs from L. ferganensis sensu Hamilton (1983, 1994) from the Nearctic in the shape of penis in dorsal view. Apparently, L. ferganensis sensu Hamilton (1983, 1994) is a separate, possibly, undescribed Nearctic taxon.

Review of the Neoaliturus fenestratus (Herrich-Schffer, 1834) species group (Homoptera: Auchenorrhyncha: Cicadellidae: Deltocephalinae: Opsiini) from Russia, Kazakhstan, and Central Asia.

In Russia, Kazakhstan, and Central Asia the Neoaliturus fenestratus species group includes eight species. Illustrated descriptions, oscillograms of male calling signals, and data on host plants and distribution for all species are given. One new species from European Russia and two new species from Kazakhstan and Central Asia are described, and the synonymy N. fenestratus (Herrich-Schffer, 1834) = N. lituratus Dubovskiy, 1966, syn. n. is established. Considerations concerning possible modes of diversification of species of this group are provided.

Review of the genus Taurotettix Haupt, 1929 (Homoptera: Cicadellidae: Deltocephalinae: Cicadulini): morphology, acoustic signals, and geographical variability.

The leafhopper genus Taurotettix includes two subgenera, Taurotettix (Taurotettix) and Taurotettix (Callistrophia), and three species, T. (T.) beckeri (Fieber, 1885), T. (C.) modesta (Mityaev, 1971), and T. (C.) elegans (Melichar, 1900). T. (C.) elegans is subdivided into two subspecies, T. (C.) elegans elegans and T. (C.) elegans subornata (Mityaev, 1971) stat. nov. Illustrated descriptions and data on biology and distribution for all taxa are given. Oscillograms of male calling signals of T. (T.) beckeri, T. (C.) modesta, and T. (C.) elegans elegans are provided. A hypothesis about speciation in Taurotettix (Callistrophia) is presented.

Review of the leafhopper genus Anaceratagallia Zachvatkin, 1946 (Homoptera: Auchenorrhyncha: Cicadellidae: Megophthalminae: Agalliini) from Russia, Kazakhstan, and Central Asia.

In Russia, Kazakhstan, and Central Asia the genus Anaceratagallia includes 14 species from two subgenera; all species except one belong to the nominotypical subgenus and can be classified into four groups according to the structure of male genitalia. The illustrated descriptions for all species are given, with male calling signal oscillograms provided for nine species. Ten new synonyms are established. Comparative investigation of morphological and acoustic characters of Anaceratagallia species showed that small differences in the structure of male genitalia and 2nd abdominal apodemes are not species-specific traits. All species studied in nature appeared to be polyphagous feeding on different species of herbaceous dicotyledons. It is assumed that closely related species within each group of the subgenus Anaceratagallia arose as a result of allopatric speciation and became secondarily sympatric due to subsequent range expansions.

Review of the genus Macropsis Lewis, 1836 (Homoptera: Auchenorrhyncha: Cicadellidae: Eurymelinae: Macropsini) of Southeastern Kazakhstan with description of new species.

Macropsis aselae Tishechkin sp. n. from Dzhungarsky Alatau Mtn. Range, Southeastern Kazakhstan is described. The synonymy M. iliensis Mityaev, 1971 = M. tuvensis Vilbaste, 1980 syn. n. is established. Descriptions, data on distribution and host plants, and an illustrated key to 20 Macropsis species from Southeastern Kazakhstan are given.

Macropsini (Homoptera: Auchenorrhyncha: Cicadellidae: Eurymelinae) from the vicinity of Ambo, Ethiopia.

Macropsis tincta Tisheсhkin sp. n., M. viridofulgida Tisheсhkin sp. n., M. nikippa Tisheсhkin sp. n., and M. antibia Tisheсhkin sp. n. from the vicinity of Ambo, Ethiopia are described. Redescriptions of M. simplex Jacobi, 1910, M. tuberculata (Linnavuori, 1978), and M. hippodameia Linnavuori, 1978 based on the material from Ambo are given. Apparently, Indian and African Acacia-dwelling Macropsis species represent a separate lineage evolved on Mimosoidea, and dispersed throughout subtropical and tropical regions of the Old World. Acacia-dwelling group of Macropsis demonstrate some morphological trends unknown in Palaearctic species; this makes it necessary to re-evaluate the diagnostic significance of some morphological traits for the taxonomy of Macropsini.

Taxonomic structure and variability of Selenocephalus obsoletus (Germar, 1817) (Auchenorrhyncha: Cicadellidae: Deltocephalinae: Selenocephalini).

Comparative investigation of morphology and calling signals of Selenocephalus obsoletus (Germar, 1817) from different localities in Western Europe, Bulgaria, Ukraine, and Russia showed that this species includes two subspecies. S. obsoletus obsoletus occurs in the Mediterranean, S. obsoletus rossicus Zachvatkin, 1945, stat. n. is distributed from the steppes of Eastern Europe up to Bulgaria; the boundary between two subspecies in the Balkans coincides with the boundary of the Mediterranean Region. Lectotypes of Selenocephalus rossicus Zachvatkin, 1945 and Selenocephalus rossicus tesquicola Zachvatkin, 1945 are designated; the synonymy S. obsoletus rossicus = S. rossicus tesquicola syn. n. is established.

Review of species of Limotettix (Scleroracus) Van Duzee, 1894 (Homoptera: Auchenorrhyncha: Cicadellidae: Deltocephalinae) of the fauna of Russia with description of a new cryptic species from Sakhalin.

Illustrated descriptions of 10 species of Limotettix (Scleroracus) of the fauna of Russia with data on host plants, male calling signals, and distribution are given. L. (S.) falco sp. n. from Southern Sakhalin is described, the synonymy L. (S.) transversus (Fallén, 1826) = Scleroracus fasciatus Kuoh, 1985, syn. n. is established. Evolution of the subgenus in Palearctic, and acoustic communication channels segregation in L. (Scleroracus) are discussed.

Geographical variability of Lepyronia coleoptrata (Linnaeus, 1758) (Hemiptera: Cercopoidea) across Russia.

Comparative analysis of body size, coloration, penis shape, and male calling signals of Lepyronia coleoptrata from different localities in European Russia, Northern Kazakhstan, Siberia, and the Russian Far East showed that all populations studied belong to the same species. A clear boundary between European and the Far-Eastern subspecies can only be drawn based on body size and proportions. Light coloration is typical of European populations; on the contrary, in the Far East only dark specimens occur, but in Siberia a mixture of different forms presents. Penis shape and male calling signal pattern do not demonstrate geographical variability. The range of intraspecific variability of these traits is far less than interspecific differences between L. coleoptrata, L. koreana, and L. okadae.

Contributions to the study of Idiocerinae (Homoptera: Auchenorrhyncha: Cicadellidae) of Central Asia with notes on synonymy.

Investigation of morphological variability in three species of Central-Asiatic Idiocerinae showed that small differences in the shape of male genitalia and abdominal apodemes are not species-specific traits. Based on these data, the following synonymies are established: Populicerus ambigenus (Dubovskiy, 1966) = P. tenellus (Dubovskiy, 1966), syn. n., Idiocerus bilituratus Dubovskiy, 1966 = I. bipustulatus Mityaev, 1967, syn. n.; the synonymy Sahlbergotettix salicicola (Flor, 1861) = S. mesasiaticus Dubovskiy, 1966 = Idiocerus fulvius Dlabola, 1967 established by Anufriev (1978) is confirmed. Illustrated descriptions of three species considered and data on their male calling signals, biology, and distribution are provided.

On the taxonomic structure and variability of Macropsidius involutus Dlabola, 1963 (Homoptera: Cicadellidae: Macropsinae)-morphology and calling signal pattern.

Variability of genitalia shape and male calling signal pattern in Macropsidius involutus from 10 localities in South-eastern Europe (the Crimea, the Lower Volga Region), Kazakhstan, and Kyrgyzstan was investigated. Variability of pygofer processes, penis shape, and signal pattern clearly has a geographic component, but none of these traits clearly supports subdividing this species into different subspecies unequivocally. M. involutus was found at altitudes from 0 up to ca. 2000 m a. s. l. in steppes and deserts in the plains and in steppe zones of mountains. Apparently, different environmental conditions in different parts of its range led to divergence of local populations, but the absence of zoogeographical barriers between them makes possible gene exchange and thus prevents emergence of clearly delineated subspecific taxa.

On the taxonomy of Anaceratagallia Zachvatkin, 1946 (Homoptera: Auchenorrhyncha: Cicadellidae: Agalliinae) of Western Kyrgyzstan.

In Western Kyrgyzstan the genus Anaceratagallia includes four species: A. venosa (de Fourcroy, 1785), A. aciculata (Horvath, 1894), A. laevis (Ribaut, 1935) = A. acuteangulata Dubovskiy, 1966 nec Zachvatkin, 1946, and A. alabugensis Dubovskiy, 1966 = A. collicola Dubovskiy, 1966, syn. n. = A. turanica Dubovskiy, 1966, syn. n. = A. laevis Dubovskiy, 1966 nec Ribaut, 1935. Illustrated descriptions and male calling signal oscillograms for all species are given. The range of A. acuteangulata Zachvatkin, 1946 includes Italy, Greece, Turkey, and North Caucasus; records of this species from Ukraine, Kazakhstan and Central Asia are erroneous. Investigation of morphological and acoustic characters in Anaceratagallia showed that small differences in the shape of male genitalia and 2nd abdominal apodemes are not species-specific traits.

Review of the genus Oncopsis Burmeister, 1838 (Homoptera: Auchenorrhyncha: Cicadellidae: Macropsinae) of Russia and adjacent countries with description of a new species from Central Asia.

Illustrated descriptions and data on host plants and distribution for 21 species of Oncopsis of Russia and adjacent countries are given, and O. abdykulovi sp. n. from Central Asia is described. Conspecificity of O. planiscuta from East Siberia and Sakhalin,of O. tristis from Moscow Area, Alati Mts., and Sakhalin, and of O. burjatica from East Siberia and Sakhalin is corroborated by male calling signal analysis.

Host plant shifts and transitions into new adaptive zones in leafhoppers: the example of Macropsinae (Homoptera: Auchenorrhyncha: Cicadellidae) of Russia and adjacent countries.

The modes of diversification of Palaearctic Macropsinae (Homoptera: Auchenorrhyncha: Cicadellidae) are reconstructed based on data on their host plants and distribution in Russia and the adjacent territories. Macropsinae (Homoptera: Auchenorrhyncha: Cicadellidae) is originally an Oriental group, which penetrated into the Palaearctic from Southeast Asia. The genus Pediopsoides and species of the genus Macropsis that feed on East Asian oaks have not dispersed beyond broadleaf forests of the Eastern Palaearctic. Apparently, Pediopsis and elm-feeding species of Macropsis initially dispersed throughout the entire broadleaf forest zone. Division of this zone into two widely separated parts in temperate areas of Europe and East Asia (nemoral disjunction), produced closely related vicariant pairs of sister species. The genus Oncopsis and species of Macropsis feeding on Salicaceae dispersed throughout the entire Palaearctic following their host plants. Both lineages penetrated into riparian forests of the foothills and midlands of Central Asia, where they produced endemic species. The Central Asian Macropsis lineage shifted from Salicaceae to trees and shrubs of unrelated families (wild roses, barberry, oleaster, and sea-buckthorn) growing in the same biotopes. Subsequent diversification on those plants produced several separate host-associated species-groups, some of which penetrated following their hosts from riparian forests into arid habitats. One such lineage apparently shifted from shrubs to wormwood species (Artemisia spp.) and thus gave rise to the genus Macropsidius. This genus underwent adaptive radiation on wormwood species in the plains of South Kazakhstan and Central Asia; advancing westward, it formed secondary centres of diversity in Transcaucasia and the Mediterranean. Finally, some lineage of Macropsidius (or its sister-group) switched from feeding on Artemisia to polyphagy, yielding the ancestral form of the genus Hephathus. In general, the evolution of the Macropsis-Macropsidius-Hephathus lineage in the Palaearctic closely followed the classical Simpsonian model: the group underwent diversification within a particular adaptive zone, then one lineage entered a new adaptive zone and secondarily diversified there, etc. Transitions into new adaptive zones by different Macropsinae lineages were probably caused by one of two factors: shift to a new plant unrelated to the original host (e.g., from Salicaceae to plants of other families) or adaptation to new microclimatic conditions (penetration from riparian forests into open arid habitats).

Contributions to the study of the genus Hephathus Ribaut, 1952 (Homoptera: Auchenorrhyncha: Cicadellidae: Macropsinae) of Russia and adjacent countries.

In Russia and the adjacent territories, the genus Hephathus includes three species, H. nanus (Herrich-Schäffer, 1835), H. freyi (Fieber, 1868) = H. tshakaranus Dlabola, 1957, syn. n., and H. achilleae Mityaev, 1967. They are indistinguishable in genitalia shape, but differ in male calling signal structure and black pattern of face. Photos of habitus and face, drawings of genitalia and male 2(nd) abdominal apodemes, signal oscillograms, and distribution maps for all species are provided. H. orientalis Linnavuori, 1953 is indistinguishable from H. freyi in coloration and genitalia shape; therefore, investigation of male calling signals is necessary for elucidation of its status. Macropsis fergusoni Evans, 1942 from Tasmania and Asmaropsis troilos Linnavuori, 1978 from Eritrea differ from Palaearctic Hephathus in the shape of head, pro-, and mesonotum and apparently belong to other genera.