Eurygaster integriceps Puton, 1881

Neimorovets, Vladimir, 2020, Review of the genus Eurygaster (Hemiptera: Heteroptera: Scutelleridae) of Russia, Zootaxa 4722 (6), pp. 501-539 : 516-521

Eurygaster integriceps Puton, 1881

Figs 7 , 15 , 22 , 28 , 34 , 40, 41 , 52 , 66–74 .

Eurygaster integriceps Puton, 1881: 119 .

Material examined. RUSSIA: Belgorod Prov.: 1♂, Krasnensky District, lake Krugloye , 16.v.1951, M. Belgovsky leg. 2♂, Shebekino , 25, 30.iii.1902, Serebryannikov leg. 3♂, 1♀, Borisovka , vii.1972, Vistavkina leg. 2♂, 3♀, Bor- isovka, vi–vii.2004, Unknown collector. 1♂, Borisovka , vi–vii.1999, Toktarova leg. Voronezh Prov.: 1♂, Grib- anovsky District, Gribanovkoye lesnichestvo, 29.vii.1953, Stark leg. 1♂, 3♀, Ternovsky District, Savalskoye lesn- ichestvo, Balash River , 11.v.1954, 19.vii.1954, 6, 7.vi.1955, Stark leg. 1♂, Ternovsky District, Savalskoye lesnichestvo, 30.v.1951, Stark leg. 21♀, Kalatch , 10.iv.1934, I. Gudim leg. > 1100 specimens, Talovsky District, settlement of Res. Inst. of agriculture of the Central Chernozem strip of a name of V. Dokuchaev, vi–vii.2007, vivii.2008, A. Shpanev leg. ( VIZR) . 1♂, Voronezhskoye lesnichestvo, “Tellermanova Rosha”, 6.viii.1953, Unknown collector . 1♂, Voronezhskoye lesnichestvo, 12.vi.1953, Moravskaya leg. Ulyanovsk Prov.: 9♂, 8♀, Staraya Ku- latka, x.1953, V. A. Kleshcheva leg. Saratov Prov.: 1♀, Saratov , 25.ix.1936, Lukyanovic leg. 1♀, Volsk , 1923, P.S. Kozlov leg. 7♂, 11♀, Engelssky District , 19.vii.2007, Unknown collector . 1♀, Zolotoye , 06.vii.1903, K. Demoki- dov leg . 4♂, 1♀, Khvalynsk , 27.vi.1952, 3.vii.1952, Lyubishchev leg. Samara Prov.: 1♂, Kamyshinsky forest belt, Bezenchuksky District , 19.ix.1949, Grunin leg. 3♂, 1♀, Zhiguli Nature Reserve , 25.viii.1941, 2.ix.1941, E. Novo- derezhkin leg . Orenburg Prov.: 1 nymph 5 th instar, Near Orenburg , 27.vii.1924, A. Ivanov leg. 5♂, 2♀, Buzuluksky Bor , 1.vi.1941, 6.x.1941, Chistovsky leg. Volgograd Prov.: 1♂, Saltovsky Les , 23–29.vi.1936, Vorontsov leg. 8♂, 13♀, Kamyshin , 9, 19.vi.1936, 25.vii.1936, 1–5.viii.1936, 5, 6, 8, 12.viii.1936, Unknown collector . Rostov Prov.: 4♂, 4♀, “Oblast Voyska Donskogo” [now Rostov Province], Sep , 1910, V. Kizericky leg. 4♂, 15♀, Salsk Area , vi–vii.1970, K. V. Kamenkova leg. 1♂, 1♀, Novocherkassk , 15.iii.1909, 25.v.1909, Kizeritsky leg. 1♀, Taganrog , 12.vii.1902, Unknown collector . 1♀, Taganrog , 26.ix.1907, V. F. Nikolaeva leg. Crimea Rep. : 1♂, Foros , 30.vi.1924, Kiritshenko leg. 1♀, Evpatoria , V. Jakovlev leg. 1♂, Feodosia, Koktebel , 31.viii.1926, Lukyanovich leg. 3♀, Sev- astopol, 11.iv.1903, 11.vi.1906, 27.iii.1907, W. Pliginski leg. 1♀, Bakhchisaray , 12.viii.1906, W. Pliginski leg. 1♀, Saky , 23–24.iv.1906, V. Jakovlev leg. 1♂, Kerch , 9.vi.1906, V. Jakovlev leg. 4♂, 1♀, Kerch, Unknown date, Kirit- shenko leg . 27♂, 13♀, Kerch , 17.viii.1901, 21.v.1902, 4, 5.iii.1903, 20.v.1903, 1907, 18.v.1907, 2, 5, 9.ix.1907, 28.ix.1917, x.1917, 11.iii.1918, 10.iv.1918, 8.vii.1918, 3.viii.1918, Kiritshenko leg. 1♀, Kerch , 17.viii, V. Jakovlev leg. 1♂, Staroselie [former Salachik], 21.viii.1905, L. Bianki leg. 6♂, 9♀, Agarmysh , Kiritshenko leg. 1♀, 1♂, Agarmysh , 28.v.1906, 17.vi.1906, Kiritshenko leg. 4♂, 4♀, Agarmysh , 10, 11.vi.1948, 21.vii.1948, K. Arnoldi leg. 3♂, 3♀, Stary Krym , 21.x.1947, K. Arnoldi leg. 1♀, Stary Krym , 19–24.vii.1901, V. Muralevich leg. 1♂, 3♀, Near Simferopol , 1898, 3.v.1898, 19.v.1899, Bazhenov leg. 1♂, Simferopol , 26.vi.1907, Kiritshenko leg. 2♂, Simfero- pol, 1.iv.1899, Unknown collector. 1♂, Near Simferopol [former Post Station Tauman–Bazar], 9.vii.1907, W. Pli- ginski leg . 2♂, 1 nymph 5 th instar, Near Simferopol [former Post Station Tauman–Bazar], 7, 21.vi.1907, B. Grigo- ryev leg . 1♂, Near Petrovo [former Khan–Eli], 29.vi.1897, Unknown collector . 1♂, Alma River , 5.vi.1899, A. Bazhenov leg. 3♂, 4♀, Belbek, Sevastopol , 12, 14, 17.v.1897, 1–15.vi.1897, 10.vii.1897, N. Kuznetsov leg. 1♂, 1♀, Sevastopol , 3, 8.iv.1906, N. Zhitkov leg. 1♀, Near Alushta , “Korbekli”, 3.viii.1900, N. Kuznetsov leg. 1♂, 1♀, Oliva [former Mukhalatka], 23.vi.1900, 4.vii.1900, Ageenko leg. 1♀, “The south coast of Crimea”, Unknown date, Vidgalm leg. 1♀, Gaspra , 5.xii.1938, E. Kiritshenko leg. 2♂, 1♂, Opolznevoe [former Kikineiz], 14.vi.1927, 7, 10.vii.1927, Kiritshenko leg. Krasnodar Ter .: 1♂, 1♀, SW Caucasus, Unknown locality, Stark leg. 2♂, 8♀, Un- known locality [former Kuban Prov. ], 1886, 2, 16.iv.27, 28.v.1923, N. Vorobiev leg. 1♂, “Tsitsinskaya Karaulka” [ Former Kuban Prov. ], 15.viii.1910, A. Birulya leg. 2♂, Krasny Les , 1910, Bryansky leg. 1♀, Anapa , v–vi.1924, M. Belgovsky leg. 4♀, Novorossiysk , Stark leg. 4♂, 6♀, Novorossiysk , 24.v.1956, 4.vi.1956, K. Arnoldi leg. 11♀, Kabardinka, Markotkh Range , 3.vi.1956, 12.vii.1956, K. Arnoldi leg. 2♀, Gelendzhik, Tonky Mys , vi.1910, Un- known collector . 2♂, 2♀, Gelendzhik , 1, 27.v.1956, K. Arnoldi leg. 3♂, 1♀, Gelendzhik , N. Vorobiev leg. 4♂, 4♀, Gelendzhik, Markotkh Range , 15, 16.vii.1956, Kurcheva leg. 2♀, Dzhankhot , 23.v.1951, 16.v.1956, K. Arnoldi leg. 1♂, Mikhailovsky Pass near Pshada , 26.v.1956, K. Arnoldi leg. 1♀, Dzhubga , 17.vi.1871, Jakovlev leg. Dzhubga , 7.vi.1912, Bogdanov-Katkov leg. 1♀, Lvovskoye , 11.vi.1956, Kurcheva leg. 1♂, 1♀, Krasnodar [former Ekaterino- dar], 15.iv.1911, 1.vi.1911, Bogdanov-Katkov leg. 1♂, Krasnodar [former Ekaterinodar], 1906, Unknown collector . 1♂, 1♀, Krasnodar [former Ekaterinodar], 25.v.1906, 1905, V. Vorobiev leg. 2♂, 2♀, Durso , 1, 2, 6.vii.2002, V. Neimorovets leg. 6♂, 2♀, Krasnodar , 25.ii.1926, 28.ii.1926, 11.iii.1926, 19.v.1926, E. Stepanov leg. 7♂, Krasnodar , 20.x.1992, E. Rubanova leg. 42♂, 35♀, Anastasievskaya , 24 May, 15, 17, 27.vi.2007, V. Neimorovets leg. 10♂, 3♂, Near Ilsky , 3.xii.1972, 11.x.1992, 3.xii.1992, E. Rubanova leg. > 600 specimens, Near Ilsky , litter of oak forest, 20. viii.2000, 5, 9, 11, 22, 24.iv.2007, 3, 15.ix.2001, 17.ix.2007, 17.xii.2008, 24, 25, 26, 31.ii, 2, 3, 6, 8, 9, 10, 13, 20. iv.2009, 2.ix.2009, 15.vi.2011, V. Neimorovets leg. ( VIZR) . 5♂, 4♀, Near Ilsky , on wheat, 26.vi.1996, 21, 23. iv.2009, V. Neimorovets leg. 1♂, 2♀, Ubinskaya , 9, 10.vi.1956, Kurcheva leg. 2♂, 1♀, Mostovskoy District, near Psebay , 14.vi.2002, V. Neimorovets leg. 4♀, Caucasus Nature Reserve , kordon “Tretya Rota”, 5, 6.vi.2002, V. Neimorovets leg. 1♀, Armavir , v.1894, Groom–Grzhimailo leg. 5♂, 7♀, Near Novodmitrievskaya [the place former named “Shebskaya Voiskovaya Dacha”], 14.v.1907, 17, 26.v.1910, 7, 28.vi.1910, 2, 6.vii.1910, Anderson & Gurov leg. 1♂, 9♀, Goryachy Klyuch , 21.v.1956, 17.vii.1956, 16.vi.1956, K. Arnoldi leg. 1♂, 1♀, Goryachy Klyuch, Kotkh Range , 21.vi.1956, K. Arnoldi leg. 7♀, Novokubansk [former Kubanskaya], 17.iii.1934, 9, 26.v.1934, 31.iii.1935, G. Zimin leg. 1♂, Gulkevichi , 4.vii.1925, Y. Sakharov leg. 1♀, Girey near Gulkevichi , 12.x.1929, Y. Sakharov leg. 1♀. Tuapse , 1915, Unknown collector . 1♂, 1♀, Lazarevskoye , 5.viii.1952, Unknown collector . 21♂, 37♀, Sochi , 19.vii.1927, I. Gidim leg. 1♀, Psebay , 26.vi.1911, Shaposhnikov leg. Stavropol Ter .: 2♂, 1♀, Stav- ropol, 24.vii.1911, V. Luchnik leg. 1♀, Stavropol, Unknown date, I. Savalov leg. 1♂, Stavropol , 19.vii.1927, V. Belousov leg. 1♀, Stavropol , 6.vi.1907, 1 nymph 5 th instar, Anderson & Gurov leg. 1♀, Yessentuki , 21.vi.1922, Ryabov leg. 1♀, Budyonnovsk [former Prikumsk], 20.v.1925, Unknown collector . 2♂, 3♀, Stavropol Ter. , vii.1908, Borisov leg. 1♀, Stavropol , vi.1914, Uvarov leg. 3♂, 2♀, Stavropol , 1.vi.1911, 24.vii.1911, V. Luchnik leg. 1♂, Stavropol, Unknown date, Bryansky leg. 11♂, 10♀, Stavropol Ter. [khutor Sizova], 29.v.1899, Demokidov leg. 2♀, Pyatigorsk , 20.iv.1912, N.L. Pastukhov leg. 1♀, Mashuk Mount, Pyatigorsk , 28.v.1908, Kiritshenko leg. 2♀, Kislo- vodsk, vii.1905, vii.1909, V. Luchnik leg. 2♀, Kislovodsk , 1–14.vii.1951, Bey-Bienko leg. 1♂, 1♀, Esentuki , 3.ix.1950, V. Vinogradova leg. 1♀, Svetlograd [former Petrovskoye], Unknown collector . 1♀, Achikulak , 12.ix.1953, P. Rafes leg. North Ossetia – Alania Rep.: 1 nymph 5 th instar, Near Vladikavkaz [between stations Kosar–Chay and Hachmos], 2.vi.1906, Bianki leg. 2♀, Mozdoksky District , v.1923, Unknown collector . Chechen Rep .: 1♀, Naur- skaya [former Naur ], Unknown date, Unknown collector . 1♂, Borozdinovskaya [former Yaman–aul], 13.vii.1892, Kiritshenko leg. 3♂, 3♀, Paraboch , 12, 16.vii.1927, Kiritshenko leg. Dagestan Rep .: 1♂, 1♀, Sergokala , 28.v.1946, Ryabov leg. 1♀, Aleksndriyskaya , 7.vi.1927, I. Filipiev leg. 1♂, 1♀, Buynaksk , 18.iv.1904, Borodin leg. 2♀, 1 nymph 4 th instar, “Petrovsk” [now is dstrict of Makhachkala ], 2.v.1925, 4.vi.1912, Kiritshenko leg. 1♂, 2♀, Khasavy- urt, 30.ix.1950, Ryabov leg. 1♀, Khasavyurt , 3.vi.1901, Bekman leg. 24♂, 21♀, Makhachkala , 27.x.1943, 29.v.1941, 2.vi.1944, 4, 9, vii.1944, 19.v.1948, Ryabov leg. 1♂, 2♀, Derbent , 1.vii.1925, Kiritshenko leg. 1♂, Derbent , 23. iv.1926, Ryabov leg. 2♂, 4♀, Derbent, Unknown date, Jakovlev leg. 2♂, 2♀, 1 nymph 5 th instar, Derbent , 11.vi.1904, Satunin leg. 1♀, Near Derbent “Abbas–Ava”, 24.vi.1917, Olsufiev leg. 4♂, 4♀, Derbent , 1.vii.1925, Kiritshenko leg. 2♀, Derbent , 23.iv.1926, 22.v.1928, Ryabov leg. 2♀, Kidero , 24.vi.1914, Mlokosevich leg. 1♂, 1♀, Belidzhi , 25.iv.1904, E. Suvorov leg. 1♀, Manas , 7.vi.1928, Ryabov leg. 1 nymph 5 th instar, Okuztau Mount, Gimrinsky Range , 27.vii.1941, Ryabov leg. 1♀, Kumtorkala, Kumtorkalinsky District , 16.vii.1947, Ryabov leg. 4♂, 4♀, Novy Buyuzyak , 27.vii.1957, Vorobiev leg.

Additional material examined. UZBEKISTAN: Xorazm Prov.: 3♂, 1♀, Khiva, 15.vi.1927, L. Zimin leg. ( VIZR) .

Diagnosis. Body length 10–13 mm. Punctures on dorsum distinct and deep. Apices of mandibular plates not protruding beyond apex of clypeus ( Fig. 22 ). Apices of mandibular plates and clypeus located in same plane ( Fig. 15 ). Buccula at middle almost as high as width of second segment of labium. Eye 4.2–5.1 times as high as buccula at middle ( Fig. 28 ). Pronotum 2.1–2.3 times as long as wide. Anterolateral margins of pronotum slightly convex, humeral angles widely rounded and not protruding beyond margin of corium. Ridge along median line of scutellum usually unclear. Pale oblong tubercles next to base of scutellum clearly visible. Aedeagus with two pairs of conjunctival appendages ( Figs. 33 , 40 A–C, 41A–C); appendages of basal pair almost straight; apical halves of distant appendages bent from midline. Blade of crown of paramere ⅓ as long as stem (dorsal view). Width of middle of stem clearly more than width of base of crown (dorsal view) ( Figs. 40 D–40F, 41D–41F). Laterotergite IX 2.5–2.7 times as wide as long, ⅔ as long as valvifer VIII and not reaching lateral excisions of abdominal sternite VII ( Fig. 52 )

Eurygaster integriceps differs from other species of this genus by several features. The mandibular plates of E. austrica and E. laeviuscula are longer than the clypeus and converge in front of it; apices of mandibular plates of E. integriceps do not protrude beyond apex of clypeus and never converge in front of it ( Figs. 19, 20, 22 and 23 ). Eurygaster austriaca is larger while E. laeviuscula is smaller than E. integriceps . Eurygaster integriceps differs from E. dilaticollis in having a narrower pronotum: pronotum of E. dilaticollis is usually 2.4–2.5 times as wide as long whereas in E. integriceps this ratio usually equals to 2.1–2.3 times. The apices of the mandibular plates lay above the clypeus in E. testudinaria and located in the same plane in E. integriceps ( Figs. 15, 18 ). In E. maura the buccula at middle is usually 1.2–1.5 as high as width of second segment of the labium and the eye 3.9–4.1 times as high as buccula at middle ( Figs. 28, 30 ). The anterolateral margins of pronotum are slightly concave or straight in E. maura and slightly convex in E. integriceps . The aedeagus has one pair of conjunctival appendages in E. maura and two pairs of appendages in E. integriceps has ( Fig. 34 A–C, 40A–C and 41A–C). The laterotergite IX of the female of E. maura reaches to the border of abdominal sternite VII ( Fig. 54 ); the laterotergite IX of the female of E. integriceps usually does not reach the border of abdominal sternite VII ( Fig. 52 ). It is quite difficult to distinguish E. integriceps , E. maura , and E. testudinaria using a single external feature; a combination of the characters should be used.

Natural History and Bionomics. E. integriceps is a meso-xerophilic steppe species. It develops in one generation per year. The overwintering stage is imago.

The spring activity of overwintered bugs is started when the forest litter warms up to 6–7°С. When the aver- age daily air temperature reaches 12–13°C and the maximum daily temperature reaches 18°C, the bugs leave the wintering grounds and fly to the fields of wheat (second half of April in the south of Russia). In Krasnodar Territory, E. integriceps first appears in the fields of the foothills, they spend several days there, and then the bugs fly to the fields of the steppe areas ( Kamenkova 1957), where they feed intensively and their gonads develop during this time. The egg-laying starts about two weeks later depending on the weather. It is noteworthy that sometimes the first eggs are laid on weeds, and even on clumps of soil. The oviposition is stretched in time. The first instar nymphs are observed from early May to early June. Before beginning of the wheat flowering, the nymphs feed on the vegeta- tive parts of the plant. After the beginning of flowering, they start feeding on the generative organs. During the time when the grain reaches wax ripeness, most of the nymphs turn into imago. The adults of the new generation feed intensively about two weeks to gain weight. At this time reserve substances are intensively accumulated in the form of fat body and food masses in the anterior sections of the intestine ( Fedotov 1945, 1946, 1947, Strogaya 1950, 1954, Ushatinskaya 1953). The departure of imago to wintering grounds begins from mid-July in Russia and Ukraine. In Kazakhstan, Uzbekistan, Kyrgyzstan, and Tajikistan, the adults of the new generation fly from valleys to foothill areas with more comfortable temperature by the end of June. They stay there to accumulate food reserves, but their activities generally are reduced ( Fedotov 1944). By the end of September, the activity of the bugs increases again, and they fly to the wintering grounds, where they stay active until the end of October.

E. integriceps is a wide oligophytophage. In the mountains of Crimea and the Northwestern Caucasus, nymphs were observed feeding on Poaceae : Hordeum bulbosum L., Poa pratensis L., Dactylis glomerata L., Elytrigia intermedia (Host) Nevski and other species of the genus Elytrigia . In the steppe regions, the bugs were noted on other Poaceae : Elytrigia trichophora (Link) Nevski , Agropyron pectiniforme Roemer & Schultes , Aegilops cylindric a Host and many other wild cereals ( Putchkov 1961). Eurygaster integriceps is a dangerous pest of wheat; it also harms barley and oats, sometimes it is found on other cultivated cereals ( Putchkov 1972).

For hibernation, E. integriceps prefers elevations of relief in dry deciduous forests, forest belts, in tree stands along railroads and other similar places with sparse undergrowth, loose forest litter and optimum humidity. The bugs avoid the old densely packed forest litter with high humidity ( Taranukha 1955). Hibernation can also take place among the thickets of high grasses on the slopes of river terraces, under stubble and weeds in the fields. In the North Caucasus ( Russia), Kazakhstan, Uzbekistan, Kyrgyzstan, and Tajikistan, the bugs fly off for hibernating places in the mountain forests, sometimes located at a considerable distance-up to 100 km.

Some authors ( Arnoldi 1947, Arnoldi & Bocharova 1952) suggested that the original distributional area of E. integriceps was limited to mountainous areas from Turkey to Tajikistan. The spreading to the European steppe zone occurred in historical time, following the spreading of wheat and other cultivated cereals. In some areas of Uzbekistan and Tajikistan, feeding of E. integriceps was observed almost exclusively on wild barley ( Fedotov 1947). Arnoldi (1947, 1955) concluded that there are three ecological types of E. integriceps . The 1 st— “the non-migratory mountain type ” (the bugs live in the mountains without flying to the plain); the 2 nd —“the migrating type ” (the bugs hibernate in the forests of the mountains, from where they fly to the fields with crops in spring, which are next to these mountains) and the 3 rd —“the non-migrating plain type ” (the bugs live in the plain districts far from the mountains and hibernate in the forest belts and other suitable places near to agricultural fields; they do not fly far from the fields where they develop).

Natural entomophages of E. integriceps . Some egg parasitoids are: Hymenoptera : Scelionidae : Trissolcus simoni (Mayr, 1879) , T. vassilievi (Mayr, 1903) , T. simoni (Mayr, 1879) , T. alpestris (Kieffer 1909) , Dissolcus rufiventris Rubstov, 1944 , Telenomus chloropus (Thomson, 1861) , T. politus (Thomson, 1861) , Gryon pedestre (Nees, 1834) , G. monspeliense (Picard, 1924) ; Encyrtidae : Ooencyrtus telenomicida (Vassiliev, 1904) , Some parasitoids of nymphs and imago are: Diptera : Tachinidae : Cylindromyia brassicaria ( Fabricius, 1775) , Rhodogyne clavatum Rohdendorf, 1947 , Elomya lateralis (Meigen, 1824) , Phasia subcoleoptrata (Linnaeus, 1767) , Ectophasia crassipennis (Fabricius, 1794) , Eliozeta helluo (Fabricius, 1805) ( Putchkov 1961) .

Economic importance. An extensive literature is devoted to the economic importance of the species and to the control of its population density (for example: Vodjdani 1954, Putchkov 1961, Putchkov 1972, Golub 1980, Areshnikov & Starostin 1982, Critchley 1998, Alekhin 2002, Gul et al. 2006, Pavlushin et al. 2008, Darkoh et al. 2010, Pavlushin et al. 2010, Neimorovets & Protsenko 2012, Neimorovets & Protsenko 2013, Vilkova et al. 2014, Pavlushin et al. 2015). This species is included in the ”List of pest organisms being highly dangerous to plant production” (VIZR working group, 2010) for agriculture of Russia.

There are studies showing that different geographic populations of E. intericeps occurring on the territory of the former USSR noticeably differ from each other in their physiological responses ( Doronina & Makarova 1976). Fasulati (2005) divided all types of colouration of E. integriceps into five morphotypes: 1 st— scutellum with a clear contrasting pattern; 2 nd— scutellum with an unclear low-contrast pattern; 3 rd— the coloration of the upper body is monotonously grey-brown dark without a pattern; 4 th— the coloration of the upper body is monotonously greyyellow-orange light; 5 th— the melanistic forms. Based on this division into morphotypes some authors concluded that the frequency of occurrence of coloration types varies in different geographic regions of the former USSR. In different regions, the percentage of these morphotypes in the samples is different. As a result, five ecotypes were identified by the proportion of these morphotypes. ( Pavlyushin et al. 2008, 2010, 2015, Vilkova et al. 2014). These authors concluded that above-mentioned morphotypes have different resistance to insecticides. The first morphotype showed the most resistance.

However, in my opinion, it is very difficult to denote a clear boundary between the morphotypes, since there is a continuous series of transitions in coloration, especially between the 1 st and the 2 nd, and also the 3 rd and the 4 th ones. The age of the collected material, the storage conditions and the substances, with which the bugs were fixed, are very important for the preservation of the original colouration of the collected material. Often the colouration of the collected material becomes darker during storage.

Distribution. The species is known from the Southern Europe, the northern border passes through the southern France, Italy, Slovenia, Croatia, Romania, Moldavia, and Ukraine; in Asia the species is distributed from Turkey and Jordan to Kyrgyzstan and northern China (but not occurred in Mongolia); also known from Egypt ( Putchkov 1961, Vinokurov & Kanyukova 1995, Göllner-Scheiding 2006, Neimorovets et al. 2006, Syromyatnikov et al. 2017). In Russia, the northern border of distribution passes through Bryansk Province, almost reaches Moscow Province, passes through the south of Nizhny Novgorod Province, reaches the southern border of Republic of Mari El, passes through Republics of Tatarstan and Bashkortostan (south of Ufa) and through the Chelyabinsk Province (some north of Magnitogorsk); the southern border runs along the foothills of the Greater Caucasus.

Notes. The record from the Altai Territory ( Kapustkina & Nefedova 2015) is a mistake. I have studied all the material mentioned in this article, more than 500 specimens in total from Altai Territory. Most of them belong to E. maura , 4 specimens belong to E. testudinaria , and no specimen belongs to E. integriceps . However, the distribution of the species has a tendency to expand ( Neimorovets et al. 2006, Pavlyushin et al. 2008, Aljaryian et al. 2016).

Figures

FIGURES 1–7. Scutelleridae of Russia. 1—Odontoscelis fuliginosa (Linnaeus, 1760), female; 2—Irochrotus sibiricus Kerzhner, 1976, female, paratype; 3—Psacasta exanthematica (Scopoli, 1763), female; 4—Phimodera lapponica (Zetterstedt, 1828), female; 5—Poecilocoris lewisi (Distant, 1883), female; 6—Odontotarsus robustus Jakovlev, 1884, male; 7—Eurygaster integriceps Puton, 1881, male.

FIGURES 13–18. Head of Eurygaster, anterior view. 13—E. austriaca (Schrank, 1776), male; 14—E. dilaticollis Dohrn, 1860, female; 15—E. integriceps Puton, 1881, male; 16—E. laeviuscula Jakovlev, 1886, male; 17—E. maura (Linnaeus, 1758), male; 18—E. testudinaria (Geoffroy, 1785), female.

FIGURES 19–25. Head of Eurygaster, view from above. 19—E. austriaca (Schrank, 1776), male (apices of mandibular plates converge); 20—E. austriaca (Schrank, 1776), male (apices of mandibular plates don’t converge); 21—E. dilaticollis Dohrn, 1860, female; 22—E. integriceps Puton, 1881, male; 23—E. laeviuscula Jakovlev, 1886, male; 24—E. maura (Linnaeus, 1758), male; 25—E. testudinaria (Geoffroy, 1785), female.

FIGURES 26–31. Head of Eurygaster, left lateral view (A—photo; B—scheme). 26—E. austriaca (Schrank, 1776), female; 27—E. dilaticollis Dohrn, 1860, male; 28—E. integriceps Puton, 1881, male; 29—E. laeviuscula Jakovlev, 1886, female; 30—E. maura (Linnaeus, 1758), male; 31—E. testudinaria (Geoffroy, 1785), male.

FIGURES 32–37. Schemes of aedeagus of Eurygaster (A—dorsal view, B—dorsolateral view, C—left lateral view). 32—E. austriaca (Schrank, 1776); 33—E. dilaticollis Dohrn, 1860; 34—E. integriceps Puton, 1881; 35—E. laeviuscula Jakovlev, 1886; 36—E. maura (Linnaeus, 1758); 37—E. testudinaria (Geoffroy, 1785).

FIGURES 38–41. Male genitalia of Eurygaster (aedeagus: A—dorsal view, B—dorsolateral view, C—left lateral view; parameres: D—right paramere, E—left paramere, F—left paramere, dorsal view). view from abow. 38—E. austriaca (Schrank, 1776); 39—E. dilaticollis Dohrn, 1860; 40, 41—E. integriceps Puton, 1881.

FIGURES 50–55. Female’s postgenital segments of Eurygaster. 50—E. austriaca (Schrank, 1776); 51—E. dilaticollis Dohrn, 1860; 52—E. integriceps Puton, 1881; 53—E. laeviuscula Jakovlev, 1886; 54—E. maura (Linnaeus, 1758); 55—E. testudinaria (Geoffroy, 1785).

FIGURES 66–74. Eurygaster integriceps Puton, 1881, variability of coloration. 66, 68 and 73—female, Russia, Krasnodar Territory; 67, 69–72 and 74—male, Russia, Krasnodar Territory.