Eurygaster maura ( Linnaeus, 1758 )

Neimorovets, Vladimir, 2020, Review of the genus Eurygaster (Hemiptera: Heteroptera: Scutelleridae) of Russia, Zootaxa 4722 (6), pp. 501-539 : 522-525

Eurygaster maura ( Linnaeus, 1758)

Figs 17 , 24 , 30 , 36 , 43 , 54 , 75–85 .

Cimex maurus Linnaeus, 1758: 441 .

Eurygaster maura f. meridionalis Péneau, 1911: 175 (upgraded to species by Ribaut, 1926: 103; syn. China, 1927: 252, with Eurygaster maura ( Linnaeus, 1758)) .

Material examined. RUSSIA: Bryansk Prov.: 2♂, 2♀, Bryansk , 14, 23.vii.1927, V. Stark leg. 1♀, “Bezhitsa” [now is district of Bryansk ], 11.vi.1911, Unknown collector . Oryol Prov.: 1♂, Petrovskoye , vi–vii.1897, Favorsky leg. Ryazan Prov.: 2♀, “ Shatsky uyezd” [now is part of Moskow and Ryazan Provinces], Kiritshenko leg. 2♀, Ko- zlovka, 23.vi.1899, Unknown collector. 5♂, 5♀, Gremyachka , 15.vii.1883, 8.vii.1886, 23.vii.1892, 12.vii.1899, 20, 28.viii.1899, 6.v.1902, 25.v.1908, 6.vi.1911, A. Semenov leg. Nizhny Novgorod Prov.: 4♂, 7♀, Staraya Pustyn , 19, 21, 25, 31.viii.1938, 4, 5, 7, ix.1939, Kiritshenko leg. Kirov Prov.: 1♀, Malmyzhsky District, Savali , 5.vii.2013, Galimova leg. Mari El Rep .: 3♂, 3♀, Sernursky District, Mari El Branch of Federal State Budgetary Institution “Russian Agricultural Center”, 20.vi.2012, Unknown collector ( VIZR) . Udmurt Rep .: 3♂, 3♀, Alnashsky Dis- trict, 22.vii.2013, V. V. Chuvashova leg. Kursk Prov.: 3♂, 3♀, Zakharkovo, near Lgov , 9.vii.1898, 12.viii.1898, R. Schmidt leg. 1♀, Lgov , vii.1926, Pliginsky leg. Belgorod Prov.: 2♂, 2♀, Novoborisovka Station , 19–30.v.1927, Pesotskaya leg. 12♂, 16♀, Borisovka , 18.v.1907, 10, 13, 19, 29.vii.1907, S. Malyshev leg. 1♂, Borisovka , vii.1972, Jakovleva leg. 1♂, Borisovka , vii.1972, Kireeva leg. 1♀, Borisovka , vii.1972, Unknown collector . 1♂, Borisovka , 25.vi.1975 , 1♂, Shoshina leg. Near Grayvoron, 5.vii.1907, S. Malyshev leg. 2♂, 2♀, Shebekino , 16, 25.iii.1902, 29.vii.1907, 6.ix.1902, Serebryannikov leg. Voronezh Prov.: 3♂, 4♀, Polyanskiy District, Polyanskoye lesnich- estvo, 29.vii.1953, 30.vii.1954, 2.viii.1954, Stark leg. 2♂, 3♀, Borisoglebsk , 8, 15.vi.1915, Anosova leg. 12♂, 7♀, Kalatch , 7, 9, 10, 12, 13, 14, 16, 20, 23, 30.vi.1934, 15, 23.vii.1934, I. Gudim leg. 1♂, Olhovatka , 19–21.vii.1937, Lukyanovic leg. 4♂, 5♀, Kamennaya Steppe , 11 km S of Talovaya, 25, 30.v.1932, 24.iv.1932, Frolova leg. 1♀, Ka- mennaya Steppe, Talovaya , 30.vi.1933, Frolova leg. 3♀, 11 km S of Talovaya , 15.vii.1935 , 1♂, 21.vi.1949, V. Stark leg. 1♂, 3♀, Talovsky District, settlement of Res. Inst. of agriculture of the Central Chernozem strip of a name of V. Dokuchaev, vi–vii.2007, A. Shpanev leg. 3♂, 3♀, Talovaya , 12.ix.1933, Unknown collector . 14♂, 19♀, “Elan–Ko- lenkovsky disrikt” [now is part of Talovsky District ], 5.v.1937, 1, 7, 20.vii.1937, Frolova leg. 5♂, 7♀, Ternovsky District, Svalskoye lesnichestvo, 24.vi.1955, 2, 7, 17, 24.vi.1955, 1.vii.1955, Speranskaya leg. 19♂, 26♀, Ternovka, Svalskoye lesnichestvo, 30.v.1951, 21.vi.1951, 10.vi.1950, 25.vii.1950, 10, 27, 28.vii.1951, 25.v.1953, 12.vi.1953, 1.vii.1953, 4, 7.viii.1953, 25.v.1954, 7, 17.vi.1955, Stark leg. Tambov Prov.: 3♀, Chakino , 6.vi.1915, Unknown collector . 1♂, Rozhdestvenskoye to the east of Tambov , 12.vi.1864, Chekanovsky leg. 1♂, Michurinsk [former Ko- zlov], N. Longe leg. Penza Prov.: 1♂, 2♀, Penza , Unknown date, Olsufiev leg. Saratov Prov.: 1♂, Saratov “Lisaya Mount”, 21 Sept 1935, Lukyanovic leg. 1♀, Zolotoye , 06.vii.1903, K. Demokidov leg. Ulyanovsk Prov.: 3♂, 2♀, Ulyanovsk , 18, 20.vii.1952, 16.iii.1953, 13.vi.1953, 21.vii.1961, Lyubishchev leg. 1♂, 3♀, Cherdaklinsky District, Stary Bebeli Yar [former Bebeli Yar], 26.vi.1962, 9.vii.1962, A. Lyubishchev leg. 1♀, Ulyanovsk , [Nizhnye Polki], 7.viii.1962, A. Lyubishchev leg. Samara Prov.: 1♂, 1♀, Zhiguli , 1.vii.1926, 23.viii.1928, Dmitryev leg. 7♂, 8♀, Kinel , 21–22.viii.1918, 3.v.1927, 28.vii.1927, 24.v.1928, 5.vii.1928, 8, 30.vii.1929, 4.viii.1929 , 1♂, 6.vii.1930, Lyubishchev leg. 1♂, 3♀, Krasnaya Glinka , 10.viii.1920, 27.viii.1928, 1.viii.1929, 16.vi.1933, Unknown collector . 1♀, “ Dubovy Omet ”, 26–27.ix.1949, Grunin leg. Orenburg Prov.: 1♂, Dneprovka [former Verkhnya Dneprovka], 23.vii.1932, L. Zimin leg. Bashkortostan Rep .: 1♂, Zilim River , 24.vi.1925, Gekker leg. Kurgan Prov.: 1♀, Lebyazhyevsky District, Lisye , 10.vi.1901, V. Sorokina leg. Omsk Prov.: 2♂, 8♀, Omsky District , spring soft wheat, 15.vi.2017, Unknown collector . Altai Ter .: 3♂, 4♀, Belokurikha Mount , 2, 3, 8, 9.viii.2006, N.N. Vinokurov leg. 1♀, Tigereksiy Nat. Reserve , 23.vi.2005, A. Namyatova leg. 176♂, 190♀, Barnaul, Altai Research Institute of Agriculture , 15.viii.2014, Unknown collector ( VIZR) . 12♂, 12♀ Barnaul, Altai Research Institute of Agriculture , spring soft wheat, 10.viii.2015, 20.viii.2015, G.G. Sadovnikov leg. ( VIZR) . Crimea Rep .: 1♂, Ayu–Dag Mount , 6.vi.1894, K. Arnoldi leg. 1♀, Stary Krym , 29.v.1947, K. Arnoldi leg. 11♂, 21♀, Opolznevoe [former Kikineiz], 21.vii.1921, 14, 20, 25, 28, 30.vi.1926, 1, 8, 10, 11, 15, 22.vii.1926, 24.viii.1926 , 1♂, 6, 11, 16, 20, 27.vi.1927, 3, 5.vii.1927, Kiritshenko leg. 1♀, Zaprudnoe [Former Degermenkoy], 12.v.1907, Kiritshenko leg. 2♂, 1♀, Yalta , 19.v.1907, Kiritshenko leg. 1♂, Gurzuf , 6.ix.1958, A. Lyubishchev leg. 2♂, Simeiz , 25, 30.vi.1918, Kiritshenko leg. 1♀, Agarmish , 20.vii.1948, K. Arnoldi leg. 5♂, 10♀, Agarmish , 30.iv.1906, 2, 20.v.1906, Kiritshenko leg. 2♂, Par- tizanskoye [former Sabli ], 17.v.1907, Kiritshenko leg. 1♂, 1♀, Kuybyshevo [former Albat], 26.vi.1924, Kiritshenko leg. 1♀, Bakhchysarai , 10.vii.1916, Kiritshenko leg. 2♂, 1♀, Bakhchysarai , 18, 25.vii.1936, Dombrovskaya leg. 1♀, Krasnogorskoye [former Neizats], 23.v.1927, E. Kuznetsov & V. Kuznetsov leg. 1♂, Simferopol , 13.vi.1947, K. Arnoldi leg. 1♂, 5♀, Simferopol , 10.v.1907, 27.vi.1907, 15.vii.1924, Kiritshenko leg. 1♂, 2♀, Near Simferopol , 1898, Bazhenov leg. 1♂, Near Simferopol , vi–vii.1898, Graftio leg. 1♀, Near Kerch , 1901, N. Kuznetsov leg. 1♂, Sevastopol , 2.vi.1907, V. Pliginsky leg. Belbek , 1♀, Sevastopol , 18.vi.1897, N. Kuznetsov leg. 1♂, Kastel Mount near Alushta , 3.vii.1900, N. Kuznetsov leg. 1♂, 1♀, Artek , 9.vii.1904, 28.v.1905, N. Kuznetsov leg. 1♂, Izobilnoe [former Korbek] near Alushta , 10.vi.1907, B. Grigoryev leg. 2♂, 3♀, Oliva [former Mukhalatka], 1900. 12.vi.1900, 6, 17.vii.1900, 23.vi.1901, Areenko leg. 1♀, Karatau Mount, Unknown date, Vigalm leg. 1♂, 2♀, Alma River , 26.v.1899, 3, 29.vii.1899, A. Bazhenov leg. 23♂, 24♀, Gurzuf , 22, 25.v.1947, 16, 21, 23, 27.vi.1947, 2, 4.vii.1947, 31.x.1947, 6, 15.vi.1948, 7, 25, 30.vii.1948, K. Arnoldi leg. 1♂, 1♀, Partenit , 24, 25.vi.1948, K. Arnoldi leg. 1♀, Yuhnoe [former Mshatka], 1.vi.1948, K. Arnoldi leg. Krasnodar Ter .: 1♀, Krasny Les , 1910, N. Bryansky leg. , 2♂, 1♀, Abinsk [former Abinskaya], 12.vii.1939, Risakov leg. 1♀, 3♀, Near Novodmitrievskaya [the place former named “Shebskaya Voiskovaya Dacha”], 26.vi.1910, 1, 6, 17.vii.1910, Andersonon leg. Adygea Rep .: 1♀, Abago Mount , 26.vi.1911, Volnukhin leg. 1♀, Umpir , 8–11 Sept 1891, Prikhodko leg. Stavropol Ter .: 1♀, Mashuk Mount, Pyatigorsk , 12.v.1908, Kiritshenko leg. 1♀, Podkumok , 13.vii.2010, D.A. Gapon leg. 2♀, Beshtau Mount near Pyatigorsk , 15.vii.2010, D.A. Gapon leg. Karachay – Cherkess Rep .: 1♀, Arkhyz, Psish River , 1.viii.1960, Sugo- nyaev leg . 1♀, Teberda , 4.viii.2013, V. V. Golub leg. 1♀, 2♀, Teberda , 13.vi.1952, 4, 30.viii.1953, L. Arens leg. 1♀, Teberda , vi–vii.1915, Kryzhanovskiy leg. 1♂, 2♀ Sofia River , viii.1935, Unknown collector . Dagestan Rep .: 1♀, Makhachkala , 6.vii.1944, Ryabov leg. 2♀, Sergokala , 27.v.1946, Ryabov leg. 1♀, Gunib , 7–9.vi.1982, Golovtch leg. 1♂, Derbent , 3.vii.1925, Kiritshenko leg.

Diagnosis. Body length: 9–11 mm. Apex of head widely rounded. Mandibular plates as long as clypeus and not extending beyond apex of clypeus (dorsal view, Fig. 24 ). Apices of mandibular plates and apex of clypeus laying on same plane (anterior view, Fig. 17 ). Buccula at middle usually 1.2–1.5 as high as width of second segment of labium, very rarely as high as width of second labial segment. Eye 3.9–4.1 times as high as buccula at middle ( Fig. 30 ). Lateral margins of pronotum slightly concave or straight. Humeral angles of pronotum not protruding or slightly protruding beyond margin of corium. Aedeagus with one pair of conjunctival appendages, their apical parts bluntly bent in dorsal direction ( Figs. 36 , 43 A–43C). Blade of crown of paramere ⅓ as long as stem (dorsal view) ( Figs. 43 D–43F). Width of middle of stem clearly more than base of crown (dorsal view). Laterotergite IX 1.7–2.0 times as wide as long, ½ as long as valvifer VIII and reaching to border of abdominal sternite VII ( Fig. 54 ).

Eurygaster maura is very similar to E. testudinaria , but the latter species differs in the apex of clypeus located below the level of the apices of mandibular plates (anterior view, Fig. 18 ) the aedeagus with two pairs of conjunctival appendages ( Fig. 44–49 A–C), the laterotergite IX of the female not reaching margin of sternite VII ( Fig. 55 ). Eurygaster maura differs from E. integriceps in having straight or concave anterolateral margins of the pronotum. The anterolateral margins of E. integriceps are convex. Eurygaster maura differs from E. austriaca and E. laeviuscula in mandibular plates, which are of the same length as the clypeus and are not convergent in front of it. The mandibular plates of E. austriaca and E. laeviuscula are longer than the clypeus and they often converge (or almost converge) in front of it. Eurygaster maura differs from E. dilaticollis in the width of the pronotum, which in E. maura is usually 2.1–2.3 times as wide as long, but in E. dilaticollis this ratio is usually equals to 2.4–2.5. The height of eye is 3.9–4.1 times the height of the middle of the buccula in E. maura ; this character is 4.2–5.1 times in E. integriceps and 4.7–5.4 times in E. testudinaria ( Tab. 3 , Figs. 28, 30, 31 ).

Natural History and Bionomics. E. maura is a typical mesophilic species and it often inhabits dry meadows, forest edges and forest glades, and fields of grain crops. In the steppe zone this species is less common. In the steppe regions of Ukraine, E. maura inhabits floodplains and areas of low relief and the northern slopes of the hills. In mountainous areas, it prefers open xerotopic cenoses, which are well warmed and illuminated by the sun. E. maura was recorded at an altitude of 1900 m in Karachay–Cherkess Republic ( Russia). In the Carpathians ( Ukraine), it occurs from the foothills to mountain meadows at altitudes up to 1000 meters above sea level ( Roshko 1953).

E. maura is a wide-oligophagous species, it breeds successfully only on cereal grasses ( Poaceae ): Elytrigia repens (L.) Nevski, Agropyron pectinatum (M. Bieb.) P. Beauv. , Arrhenatherum elatius (L.) J. Presl & C. Presl, Alopecurus pratensis L., Phleum pratense L., Poa pretensis L., P. palustris L., P. annua L., Agrostis gigantea Roth , A. tenuifolia M. Bieb. , Festuca pratensis Huds. , F. rubra L., F. pallens Host , Bromus inermis Leyss , Dactylis glomerata L., Lolium perenne L., Milium spp. and other wild and cultivated cereals. Last instar nymphs and young adults are known to feed on seeds of many Asteraceae ( Carduus , Cirsium , Sonchus , Matricaria , Centaurea , Scabiosa , etc.). Eurygaster maura was also noted on Lycopus (Lamiaceae) , Salvia (Lamiaceae) , Verbascum (Scrophulariaceae) , Poterium (Rosaceae) , Epilobium (Onagraceae) , Eriophorum (Cyperaceae) ( Putchkov 1961).

The developmental cycle of E. maura is longer than in E. integriceps . In the forest-steppe zone, E. maura appears from the wintering shelters when the soil and litter are heated to 12–15°C ( Taranukha 1954, Pavlov 1954). The bugs fly from wintering grounds to open places, preferring areas with wild cereals or cereal crops. When the daytime temperatures rise up to 16–18°C, the bugs begin to feed and mate. Mass egg-laying occurs in June. The eggs are laid on many species of wild cereals, in the lower or middle tier, depending on the height of the grass stand. The eggs develop in 1–1.5 weeks at average temperature of 20–24°C. Hatching of nymphs is observed from the first half of June. Adult bugs of the new generation appear from the middle of July. This happens later than in E. integriceps when harvesting grain is finishing. Probably this circumstance does not allow E. maura to accumulate enough reserve substances using cultivated cereals. Perhaps this is one of the main factors constraining the number of this species in areas where crops are cultivated ( Bey-Bienko 1969). The period of accumulation of reserve substances is short. Flying to the overwintering places starts at the end of July. The bugs do not fly to hibernate over long distances. They remain not far from breeding sites in local forests, forest belts or among high grass. The wintering grounds of E. maura are very similar to those of E. integriceps ( Putchkov 1961) .

Natural entomophages of E. maura . Hymenoptera : Scelionidae : Trissolcus vassilievi (Mayr, 1903) , T. cultratus (Mayr, 1879) ; Diptera : Tachinidae : Cistogaster globosa ( Fabricius, 1775) , Elomya lateralis (Meigen, 1824) , Cylindromyia auriceps (Meigen, 1838) , C. intermedia (Meigen, 1824) , Gymnosoma rotundatum ( Linnaeus, 1758) ( Putchkov 1961) .

Economic importance. The number of E. maura in cultivated fields in European part of Russia is usually low. This species damages rye, wheat, barley, grass, corn, and sometimes millet. The negative effect of E. maura saliva on flour quality is the same as in E. integriceps ( Putchkov 1972) . In some years, there was a significant increase in numbers of E. maura . Severe damage of cultivated cereals was recorded at the end of the 1930s in Hungary and Austria. Also damage of cultivated cereals was observed in Bulgaria, former Yugoslavia, partly in Zakarpattia Province ( Ukraine) and Belarus in the 1960–70 s. As a pest of cereals it was noted in Yaroslavl and Nizhny Novgorod Prov- inces of Russia ( Taranukha 1952, Pogorelov 1965, Putchkov 1972). E. maura does not cause much damage to crops in northern Kazakhstan ( Azhbenov 2004). It is an important pest in Iran, especially in the Gorgan region (Golestan province), but the harmfulness of this species was not noted in other parts of Iran ( Radjabi 2000).

Distribution. The species is known from North Africa (northern Morocco, Algeria, and Tunisia), all Europe except Norway and Finland and from Turkey and Egypt (Sinai) to Kazakhstan and Turkmenistan in Asia. ( Petrova 1975, Vinokurov & Kanyukova 1995, Kerzhner 2003, Göllner-Scheiding 2006, Neimorovets et al. 2008, Vinokurov et al. 2010, Syromyatnikov et al. 2017). In Russia, the species is distributed to the Kemerovo Province in the east; the northern distributional border lays through the Pskov, Tver, Yaroslavl and Kirov Provinces, along the northern border of Republic of Bashkortostan, through Kurgan, south of Tyumen, Omsk, and Novosibirsk Provinces.

Figures

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 42–45. 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). 42—E. laeviuscula Jakovlev, 1886 (holotype); 43—E. maura (Linnaeus, 1758); 44, 45—E. testudinaria (Geoffroy, 1785).

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 75–85. Eurygaster maura (Linnaeus, 1758), variability of coloration.75, 76 and 85—female, Russia, Altai Territory; 77, 79, 82 and 84—female, Russia, Omsk Province; 78, 80 and 81—male, Russia, Altai Territory; 83—male, Russia, Belgorod Province.