Ecologica Montenegrina 40: 164-175 (2021) This journal is available online at: www.biotaxa.org/em http://dx.doi.org/10.37828/em.2021.40.14

True bugs (: ) from the taiga zone of the mountainous Altai of : the first records and new data on rare species

VIKTOR B. GOLUB1*, NIKOLAY N. VINOKUROV2, NATALIA V. GOLUB3, VIKTORIA A. SOBOLEVA1 & EVGENIY V. AKSENENKO1

1*Voronezh State University, 394001 Voronezh, Universitetskaya sq. 1, Russia. E-mails: [email protected]; [email protected];[email protected] 2Institute for Biological Problems of Cryolithozone, Siberian Branch RAS, 677980 Yakutsk, 41 Lenin Av., Russia. E-mail: [email protected] 3Zoological Institute RAS, 199034 St. Petersburg, Universitetskaya emb.1, Russia. E-mail: [email protected]

Received 22 February 2021 │ Accepted by V. Pešić: 23 March 2021 │ Published online 26 March 2021.

Abstract Twelve species are recorded for the first time from the , Russia, based on the materials collected by authors in the vicinity of the northern part of Lake Teletskoye in the southern taiga of Western . (Dolichonabis) americolimbatus (Carayon, 1961) is recorded for the first time for the Western Siberia. Seven species, Pachycoleus pusillimus (J. Sahlberg, 1870) (and family Dipsocoridae Dohrn, 1859), Saldula orthochila (Fieber, 1859), Xylocoris cursitans (Fallén, 1807), Acalypta carinata (Panzer, 1806), vagabundus (Linnaeus, 1758), brunneus (R.F. Sahlberg, 1848), thomsoni Reuter, 1875 are recorded for the first time from the Russian Altai. Four species are recorded for the first time from the Altai Republic – Phytocoris longipennis Flor, 1861, Nysius helveticus (Herrich-Shaeffer, 1850), resedae resedae (Panzer, 1797), Cymus glandicolor Hahn, 1832. For each species the general distribution, the detailed distribution in the adjacent territories of Siberia, and the data on the biotopic confinement are provided. In addition, the data on the abundance and ecology of some other rare species are provided.

Key words: , hemipterans, West Siberia, Altai Republic, Lake Teletskoye, new records, distribution, ecology.

Introduction

Currently, about 350 heteropteran species are known from the Altai Republic. Most of them are recorded from steppe and semi-desert regions of southern Altai. The bugs fauna of northern taiga of mountain Altai has been less studied then the southern regions, and counts, approximately, two times less species. The data on the taxonomic composition of bugs of the Russian Altai and of Asian part of Russia as a whole is presented in the catalog by Vinokurov et al. (2010). Faunistic data on the Heteroptera of the Altai of eastern

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Kazakhstan were provided by Asanova (1974, 1975, 1996) and Esenbekova (2013). A considerable volume of information on the species composition of bugs of the Mongolian Altai was published by Kerzhner (1974), Kiritshenko & Kerzhner (1974) and others in their studies of Heteroptera of Mongolia. Present paper provides the first records of 12 species of terrestrial Heteroptera based on the materials collected by V.B. Golub, E.V. Aksenenko, V.A. Soboleva and N.V. Golub in the northern part of the Altai Republic in the vicinity of Artybash village in 2019 (Fig.1). The general distribution of the listed species is given according to the Catalogue of Palaearctic Heteroptera (1995–2006) with additions (Catalogue of Palaearctic Heteroptera, Supplement 2013). More detailed information on the distribution of the discussed species in Western Siberia and closest regions of Eastern Siberia are cited from the Catalogue of Hemiptera of the Asian part of Russia (Vinokurov et al. 2010). Furthermore, additional information on the distribution of these species in Siberia is given based on data published later than the mentioned Catalogues (Galich & Ivanov 2012; Ivanov 2015; Vinokurov & Golub 2016; etc.). The nomenclature and order of taxa arrangement correspond to those from the both mentioned Catalogues.

Material and methods

The material listed in this paper was collected in July-August, 2019 in the vicinity of Artybash village and in the vicinity of field station “Teletskaya” of the Institute of Systematics and Ecology of of the Siberian Branch of the Russian Academy of Sciences at the northern part of the Lake Teletskoye.

Figure 1. Taiga on the slopes of the at northern part of the Lake Teletskoye (Photo by V.A. Soboleva).

The material collecting was carried out using the traditional methods: by entomological net, hand (manual) collecting from grassy and tree-shrub vegetation, soil, mosses, at the roots of plants, and by mosses analyzing (Golub et al. 2012). The species determination was verified using the materials of the Zoological Institute of the Russian Academy of Sciences, St. Petersburg (ZIN). The collected and studied material is stored in the collections of the Department of Zoology and Parasitology of Voronezh State University, Voronezh and the ZIN.

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Results

List of the Heteroptera recorded from the Altai Republic for the first time

Family Dipsocoridae Dohrn, 1859

Pachycoleus pusillimus (J. Sahlberg, 1870) Fig. 2

Material. Altai Republic, the vicinity of Artybash village: a swampy area on the margin of the forest, below the waterfall on the Oyrok (Third) River, 51º47'30.7" N, 87º19'8.2" E, 492 m a. s. l., on wet mosses, 30.07.2019 (V. Golub), 1 male. Distribution. European–West Siberian species. Northern and Central and the mountains of Southern Europe. In European part of Russia it was recorded only from Karelia, Voronezh and Belgorod Provinces (Prokin, 2008); in Asian part it was recorded only from Tyumen Province (Ivanov 2015). This family was recorded for the first time from the Altai Republic and Altai as a whole. Ecology. The species occurs in swampy habitats. In Western Europe, it almost always collected on Sphagnum mosses (Heiss & Péricart 2007). In the European part of Russia (Voronezh Province) (Golub 2004), and in Western Siberia (Tyumen Province) (Ivanov 2015) the species was also collected on wet or dried mosses, non Sphagnum. In the vicinity of Artybash village a single specimen was collected on mosses in a small swampy meadow at the edge of the forest, however, Sphagnum was not found at the collection point. Note. Until 2015, the species was known only from Europe. The previous finding of the species in Siberia (Ivanov 2015) and our record from the Altai Mountains are the evidences of its wide range of the boreal type.

Figure 2. Habitat of Pachycoleus pusillimus (J. Sahlberg, 1870) and Saldula orthochila (Fieber, 1859) in the environs of village Artybash, Altai Republic (Photo by V.A. Soboleva).

Family Amyot et Serville, 1843 Subfamily Saldinae Amyot et Serville, 1843 Tribe Saldoidini J.T. Polhemus, 1954

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Saldula orthochila (Fieber, 1859) Fig. 2

Material. Altai Republic, the vicinity of Artybash village, meadow on the right bank of the Oyrok (Third) River, 500 m below the waterfall, 51º47'30.7" N, 87º19'8.2" E, 492 m a. s. l., 30.07.2019 (Aksenenko), 1 male, 1 female. Distribution. The range is very wide: in the Palaearctic – Eurasian, eastward to the Yenisei River; Middle Asia; northwestern China. India (Kashmir). In Western Siberia it was previously recorded from the Tyumen Province. First record from the Altai Republic and Altai as a whole. The nearest localities in Eastern Siberia are the south of the Krasnoyarsk Territory and the Irkutsk Province. Ecology. In the vicinity of Artybash the species was collected in a humid habitat, on the river bank overgrown with grassy vegetation.

Family A. Costa, 1853 Subfamily Nabinae A. Costa, 1853 Tribe Nabini A. Costa, 1853

Nabis (Dolichonabis) americolimbatus (Carayon, 1961) Figs 3–5

Material. Altai Republic, the vicinity of Artybash village, 51º47'06–8.6" N, 87º19'45" E, 500–510 m a. s. l., margin of the forest on the southern slope, 26, 29.07, 03.08.2019 (V. Golub, V. Soboleva), 6 males; 2 km NNE of field station “Teletskaya”, 51º48'06" N, 87º18'18" E, 530 m a. s. l., glades in the taiga, on the grasses, 27.07, 01.08.2019 (V. Golub, V. Soboleva), 4 males. Distribution. Holarctic boreal species. In Palearctic it is known mainly from the eastern part (Eastern Siberia and the Far East of Russia, Mongolia, northeastern China and Japan); additionally, it was indicated from the Central part of the European Russia (Kerzhner 1996). The species is recorded from Western Siberia for the first time. The nearest localities in Eastern Siberia are the south of the Krasnoyarsk Territory and the Irkutsk Province (Kerzhner 1981; Vinokurov et al. 2010).

Figure 3. One of habitats of Nabis (Dolichonabis) americolimbatus (Carayon) on the taiga edge near of Artybash village (Photo by E.V. Aksenenko).

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Ecology. The species inhabits swamps and damp meadows, dwells in the grass, occasionally – on bushes and trees (Kerzhner 1981). In the vicinity of Artybash village it was collected in damp biotopes, in areas of taiga with damp soil, and in drier but shaded biotopes Notes. 1. The presence of lamellar carina at the base of paramere (Fig. 5) and the absence of lamellar carinae on the basal hook of aedeagus are the characteristics of this species that ensures its reliable identification. 2. All the males of subgenus Nabis (Dolichonabis) studied from the vicinity of Artybash village belongs only to this species. 3. An expected presence of N. (D.) limbatus Dahlbom, 1851in this locality was not confirmed.

Family Fieber, 1836 Subfamily Anthocorinae Fieber, 1836 Tribe Xylocorini Herrich-Schaeffer, 1850

Xylocoris cursitans (Fallén, 1807) Fig. 6

Material. Altai Republic, southern outskirts of the Artybash villalge, 51º47'43" N, 87º19'32" E, 530 m a. s. l., under the bark of a sawn pin 03.08.2019 (E. Aksenenko, V. Golub, V. Soboleva), 1 male, 3 females. Distribution. Holarctic species. In Western Siberia it was previously recorded from the Tyumen Province (Vinokurov & Golub 2009; Vinokurov et al. 2010). First record from the Altai Republic and Altai as a whole. The nearest localities of the species in Eastern Siberia are the Krasnoyarsk Territory and the Irkutsk Province. Ecology. The specimens collected under the bark of a recently sawn pine with the other invertebrates, including the larvae of bark beetles (Coleoptera, Ipidae), that corresponds to the described ecological features of this species (Péricart 1972).

Family Hahn, 1833 Subfamily Baerensprung, 1860 Tribe Bryocorini Baerensprung, 1860

Bryocoris pteridis (Fallén, 1807) Fig. 7, 10

Material. Altai Republic, 2 NNE of field station “Teletskaya”, 51º48'06" N, 87º18'18" E, 550 m a. s. l., glade in a mixed forest, on a fern, 27.07.2019 (V. Golub), 1 female. Distribution. Transpalearctic species. In Western Siberia it was previously recorded from the Khanty-Mansi Autonomous Okrug – Yugra, Kemerovo Province, Altai Territory, Tyumen Province (Galich & Ivanov 2012) and the Altai Republic (Vinokurov & Golub 2016). In Eastern Siberia the species is recorded from Khakassia, the south of Krasnoyarsk Territory, Irkutsk Province and from Buryatia. Ecology. The species lives on a fern, as well as filicis (Linnaeus, 1758). However, in the taiga zone of the Altai Mountains, the species is obviously quite rare. In the vicinity of Lake Teletskoye a single specimen and only at one locality (Fig. 10) was collected together with numerous M. filicis.

Subfamily Mirinae Hahn, 1833 Tribe Mirini Hahn, 1833

Phytocoris longipennis Flor, 1861 Fig. 8

Material. Altai Republic, 2 km NNE of field station “Teletskaya”, 51º48'06 " N, 87º 18'18" E, 500 m a. s. l., forest edge, southern slope, 27.07.2019 (V. Golub, V. Soboleva), 1 male, 1 female; field station “Teletskaya”, 51º47'42.1" N, 87º18'11.8" E, 495–500 m a.s.l., on deciduous trees, 31.07.2019 (N. Golub), 1 male.

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Distribution. Transpalearctic species. In Western Siberia it was previously recorded from the Altai Territory. First record from the Altai Republic. The nearest localities in Eastern Siberia are the Khakassia and the Irkutsk Province. Ecology. In the vicinity of Artybash village the species was collected in deciduous trees species (, aspen, bird cherry). Note. Females of the genus Phytocoris Fallén, 1814, almost identical in appearance to those of Ph. longipennis, were collected together with males, however, they are not listed in the materials in order to avoid a possible misidentification.

Figures 4–9. Adult specimens and paramere of Heteroptera. 4, 5. Nabis (Dolichonabis) americolimbatus (Carayon) (4 – dorsal view, 5 – paramere, arrow indicates lamellar carina at its base), 6. Xylocoris cursitans (Fallén), 7. Bryocoris pteridis (Fallén), 8. Phytocoris longipennis Flor, 9. Gampsocoris culicinus Seidenstücker (Photo by V.A. Soboleva).

Family Laporte, 1832 Subfamily Tinginae Laporte, 1832 Tribe Acalyptaini Blatchley, 1926

Acalypta carinata (Panzer, 1806)

Material. Altai Republic, 2 km NNE of field station “Teletskaya”, 51º48'06" N, 87º 18'18" E, 500 m a. s. l., glade in a mixed forest, on mosses, 01.08.2020 (V. Golub), 2 males, 5 females, 1 larva. Distribution. Transpalearctic species. In Western Siberia the species was previously recorded from Khanty-Mansi Autonomous Okrug – Yugra and the Kemerovo Province. First record from the Altai

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Republic and Altai as a whole. The nearest localities of the species in Eastern Siberia are the Krasnoyarsk Territory, Tuva and the Irkutsk Province. Ecology. A. carinata, like all species of the genus Acalypta Westwood 1840, lives on mosses, on which it was collected in the vicinity of Lake Teletskoye. The species was found in forest glades together with Ceratocombus coleoptratus (Zetterstedt, 1819) ( Fieber, 1860) on mosses growing in moist soil.

Figure 10. Habitat of Bryocoris pteridis (Fallén) in taiga at the environs of Lake Teletskoye, Altai Republic (Photo by N.V. Golub).

Family Latreille, 1807 Subfamily EmesinaeAmyot et Serville, 1843 Tribe Van Duzee, 1916

Empicoris vagabundus (Linnaeus, 1758)

Material. Altai Republic, the vicinity of field station “Teletskaya”, 51º47'07" N, 87º19'45" E, 510 m a. s. l., taiga, on the fir (Abies sibirica Ledeb.), 03.08.2019 (N. Golub), 1 male, 1 fifth instar larva. Distribution. Holarctic species. In Western Siberia it was recorded from the Tyumen and Kemerovo Provinces. First record from the Altai Republic and Altai as a whole. The nearest localities in Eastern Siberia are in the south of the Krasnoyarsk Territory and the Irkutsk Province. Ecology. Both specimens were collected on Abies sibirica Ledebur, 1833 in a moist and shaded habitat that corresponds to the ecological requirements of this species (Putshkov & Moulet, 2010).

Family Schilling, 1829 Subfamily Orsiliinae Stål, 1872 Tribe Nysiini Uhler, 1876

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Nysius helveticus (Herrich-Shaeffer, 1850)

Material. Altai Republic, the vicinity of the Artybash village: glades on the slopes, mainly southern exposure, on the grass, 51º47'06" N, 87º19'45" E, 472 m a. s. l., 31.07.2019 (V. Soboleva), 1 female. Distribution. Holarctic species. In Western Siberia it was recorded from the Novosibirsk and Kemerovo Provinces and the Altai Territory. First record from the Altai Republic. The nearest localities in the Eastern Siberia are the south of the Krasnoyarsk Territory, Tuva, and the Irkutsk Province. Ecology. The open well-insulated habitat in the vicinity of Artybash where the species was collected, is fully consistent with its ecological requirements (Putshkov 1969; Péricart 1999a).

Subfamily Ischnorrhynchinae Stal, 1872 Kleidocerys resedae resedae (Panzer, 1797)

Material. Altai Republic, the vicinity of the Artybash village: glades on the slopes, mainly southern exposure, on grass, 51º47'06" N, 87º19'45" E, 472 m a. s. l., 31.07.2019 (V. Soboleva), 1 male; on fir (E. Aksenenko), 2 males, 1 female. Distribution. Holarctic species. The species is widely distributed in Western Siberia, including the Altai Territory. First record from the Altai Republic. The nearest localities in Eastern Siberia are the Irkutsk Province and Tuva. The species is recorded from Kazakhstan and Mongolia (Péricart 2001). Ecology. The species lives on many species of trees and shrubs (Betula spp., Alnus spp., etc.) (Putshkov 1969; Péricart 1999a), it is also found on grassy vegetation as a result of falling from trees. We collected the species both on the grass and by entomological net on the crown of trees.

Cyminae Baerensprung, 1860 Tribe Cymini Baerensprung, 1860

Cymus glandicolor Hahn, 1832

Material. Altai Republic, the vicinity of the Artybash village: ruderal area, 51º47'43.4 "N, 87º18'31.6" E, 545 m a. s. l., 04.08.2019 (E. Aksenenko, V. Golub, V. Soboleva), 17 imago. Distribution. Transpalearctic species. In Western Siberia it was recorded from the Tyumen and Kemerovo Provinces and the Altai Territory. First record from the Altai Republic. The nearest localities in the Eastern Siberia are Khakassia, the south of the Krasnoyarsk Territory, the Irkutsk Province, and Tuva. The species is known from Kazakhstan and Mongolia. Ecology. The species inhabits both moist and relatively dry biotopes; lives mainly on different species of Cyperaceae and Juncaceae (Putshkov 1969; Péricart 1999a). On the territory close to the Lake Teletskoye, it was collected in single habitat – on the outskirts of an open and well insulated ruderal area with a fairly dry sandy soil.

Subfamily Amyot et Serville, 1843 Tribe Stål, 1872

Drymus (Sylvadrymus) brunneus brunneus (R.F. Sahlberg, 1848)

Material. Altai Republic, the vicinity of the Artybash village, bank of a stream flowing into Lake Teletskoye, 51º47'18.5" N, 87º19'10.6" E, 474 m a. s. l., on soil, 30.07.2019 (V. Golub), 1 male, 1 female; field station “Teletskaya”, 51º47'42.1" N, 87º18'11.8" E, 495 m a. s. l., in a sample of mosses with the topsoil, 31.07.2020 (V. Golub), 1 female. Distribution. Transpalearctic species. In Western Siberia it was previously recorded from the Yamalo-Nenets National District, Tyumen and Tomsk Provinces. First record from the Altai Republic and Altai as a whole. The nearest localities in Eastern Siberia are the south of the Krasnoyarsk Territory, the Irkutsk Province, and Buryatia.

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Ecology. The species inhabits moist biotopes, most often in forests, as well as the banks of rivers and lakes, streams and other water bodies (Putshkov 1969; Péricart 1999b). Inhabitant of mosses and forest floor. In the vicinity of Lake Teletskoye the species was found in its characteristic biotopes: on the bank of a stream and in moss samples together with Acalypta carinata (see above).

Lamproplax membranea Distant, 1883

Material. Altai Republic: field station “Teletskaya”, 51º47'42.1" N, 87º18'11.8" E, 495 m a. s. l., herbaceous vegetation, 31.07.2019 (E. Aksenenko, V. Golub), 1 male, 2 females, 1 larva; meadow near Teletskoye Lake with mixed grass vegetation, 51º47'18.1" N, 87º17'58" E, 482 m a. s. l., 01.08.2019, (Aksenenko), 1 female; ruderal area near vill. Artybash, 51º47'43.4" N, 87º18'31.6" E, 545 m a. s. l., 04.08.2019, (E. Aksenenko), 1 female. Distribution. East Palaearctic species with known western range border in Altai Republic (Vinokurov & Golub 2016). The nearest localities in Eastern Siberia are the south of the Krasnoyarsk Territory, and the Irkutsk Province. Ecology. All undisturbed natural habitats in which the species was collected were open mesophytic areas with meadow vegetation. One of the collection points is the margin of the forest at the base of the mountain slope. The rocky soil of this place was destroyed by machinery and overgrown mainly with ruderal vegetation. Thus, the extensive materials collected in 2019, compared to the materials of 2009 (Vinokurov & Golub 2016) has clarified the environmental features of the species.

Scolopostethus thomsoni Reuter, 1875

Material: Altai Republic, the vicinity of the Artybash village, meadow on the edge of a mixed forest, near a stream, 51º47'47.4" N, 87º17'17.7" E, 526 m a. s. l., 28.07.2019 (E. Aksenenko), 1 male, 2 females. Distribution. Holarctic species. In Western Siberia it was recorded from the Tyumen and Tomsk Provinces. First record from Altai as a whole. It was previousely recorded from Eastern Siberia without localities indications (Péricart 2001). Ecology. The species was collected in open moist biotope that corresponds to its ecological requirements as mesophilic and hygrophilic species (Putshkov 1969; Péricart 1999b).

Family Fieber, 1851 Subfamily Gampsocorinae Southwood et Leston, 1959 Tribe Gampsocorini Southwood et Leston, 1959

Gampsocoris culicinus Seidenstücker, 1948 Figs 9, 11

Material. Altai Republic, the vicinity of the Artybash village, 51º47 '06" N, 87º19'45" E, 472 m a. s. l., 29, 31.07, 03.08.2019, forest edge on the slope of the southern exposure, partially anthropogenically disturbed, on inflorescences of single Asteraceae species, 39 imago, 9 larvae. (V. Golub, V. Soboleva). Distribution. West–Central Palearctic species. In Western Siberia it was previously recorded from the Kemerovo Province and the Altai Republic, from Eastern Siberia – southern Yakutia. Ecology. All the material was collected in an open, well-insulated habitat, on the rocky slope of the southern exposure at the margin of the forest, with anthropogenically disturbed soil cover. Note. The first indication of the species from the Altai Republic (Vinokurov & Golub, 2016) was based on only three specimens collected in the vicinity of Artybash. A relatively high population numbers of this species in 2019 in the same locality allowed us to establish that the species prefers the open habitats in the conditions of the southern taiga of Western Siberia.

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Figure 11. Habitat of Gampsocoris culicinus Seidenstücker at the environs of Lake Teletskoye, Altai Republic (Photo by V.A. Soboleva).

Discussion

The first records of 12 Heteroptera species from the Altai Republic significantly clarify their general distribution. The finding of Nabis americolimbatus in Altai eliminates a huge gap in the distribution of this species between Eastern Europe and Eastern Siberia. The western boundary of the range for this species has significantly moved westward, in relation to the previous western records from Eastern Siberia (Irkutsk; Kultuk) and the Khentei highlands in the north-eastern Mongolia (Kerzhner 1981). The eastern boundary of the Pachycoleus pusillimus range has moved south-east nearby to 1400 km from the former record from Western Siberia (Tyumen Province). The additional information about the ranges of six species eliminated the extensive disjunction between Western and Eastern Siberia in the characteristics of their distributions. It concerns three species with Holarctic ranges (Xylocoris cursitans, Scolopostethus thomsoni, ), two species with Transpalaearctic ranges (Acalypta carinata, ), and one species with Eurasian distribution (Saldula orthochila), in the east reaches the Yenisei River. The first records of four species (Phytocoris longipennis, Nysius helveticus, Kleidocerys resedae, Cymus glandicolor) from the Altai Republic, with the closest known localities in the adjacent Altai Territory, supported data on their ranges in Southern Siberia. In addition, these records enhance the understanding of the ecological possibilities of these species inhabiting the southern taiga in the Altai Mountains. Ranges of all of twelve species mentioned are confined to the humid territories of the boreal and subboreal areas of the Holarctic. Ecologically, nine from twelve listed species (N. americolimbatus, X. cursitans, Ph. longipennis, A. carinata, E. vagabundus, N. helveticus, K. resedae, D. brunneus, S. thomsoni) are confined to mesophytic conditions, and 2 species (S. orthochila and C. glandicolor) – to mesohygrophytic and hygrophytic ones. On the basis of counting the number of individuals of Bryocoris pteridis and Monalocoris filicis collected together on a fern in the vicinity of Lake Teletskoye, it was found that the first species about 50 times less abundant than the second. The clear confinement of membranea to meadow habitats, including those with stony soil, was confirmed. A fairly high abundance of Gampsocoris culicinus in open habitat with rocky soil was established on Asteraceae species.

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Forty-five species have been revealed in the steppe zone of the Altai Republic, including 15 species recorded from this territory for the first time (Vinokurov & Rudoi 2020). Most species listed in mentioned paper (Nabis punctatus mimoferus Hsiao, Lygus rugulipennisus Poppius, fulvicollis Jakovlev, Kleidocerys resedae resedae Panzer, Myrmus calcaratus Reuter etc) are predominantly xerophilus and mesoxerophilus in their ecological characteristics and mostly confined to steppe biotopes in their habitat. In general, the rich and diverse fauna of the true Heteroptera of Altai is based on a wide variety of landscapes, steppe and taiga biotopes of this vast territory.

Acknowledgements We thank two reviewers, Dr. Prokin and anonymous referee, for their thorough revision of the manuscript and useful comments and corrections. The study was funded by the grant from the Russian Foundation for Basic Research no. 18-04-00464 (2018–2020) and by state research project no. AAAA-A19-119020790106- 0 (to N.V. Golub).

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