Lithosia quadra subsp. quadra Linnaeus 1758
Creators: name: Dubatolov, Vladimir V.
name: Zolotuhin, Vadim V.
name: Witt, Thomas J.
Publisher: name: Zenodo
Year: 2016
Subjects: Biodiversity
Taxonomy
Animalia
Arthropoda
Insecta
Lepidoptera
Arctiidae
Lithosia
Lithosia quadra
Lithosia quadra quadra (linnaeus, 1758)

description: Lithosia quadra quadra (Linnaeus, 1758)
(Figs 1–5)
Phalaena Noctua quadra Linnaeus, 1758, Syst. Nat. (Edn. 10) 1: 511. Type locality: not stated, probably, Europe (Watson et al., 1980). Lectotype: female (LSL), designated by Mikkola & Honey, Zool. J. Linn. Soc. 108: 155. Mikkola & Honey (1993) noted one male as a paralectotype, but this designation is incorrect, because Linnaeus (1758) described this species after female(s) only (see below).
= Phal.[aena] Noctva flaua O. F. Müller, 1764; Favna insectorvm Fridrichsdalina: 46. TL: "[Fridrichsdal] … In nemore" [Germany].
= Phalaena Noctua deplana Linnaeus, 1771; Mant. Plant.: 539–540. TL: “Habitat - - -” (no reliable specimen in Linnaeus’ collection: Mikkola & Honey, 1993: 124). Type: male(s). The species was described by Linnaeus with a remark “D.[ominus] Fabricius. Prof.[essor] Haffn.[iae]”—seemingly the specimen is kept at HMUG in the collection of Fabricius (Fig. 2); it is designated here as a lectotype of Phalaena Noctua deplana Linnaeus, 1771.
=Lithosie quadrata Walckenaer, 1802; Faune parisienne. Insectes 2: 308. TL: [environs de Paris].
= Oeonistis quadra insolata Dannehl, 1929, Ent. Zeit. 42: 315. TL: “Terlan, Sigmundskron … Sattniß und den Karawankentalern” [S. Tyrol, Italy].
A lot of forms are known: albescens Lempke, 1961, atra Torstenius, 1956, confluens Dumont, 1903, depauperata Henriot, 1923, externa Closs, 1919, fasciata Spuler, 1906, flavescens Lempke, 1961, luteomarginata Lambillion, 1906, obscura Schawerda, 1921, pallida van Wisselingh, 1961, seminigra Dufay, 1954, triangularis Lempke, 1961, unipunctata Spuler, 1906, violagrisescens Daniel, 1952. However, all are infrasubspecific therefore not considered in this article.
Type material studied. EUROPE: 1 ♀ (lectotype of quadra), “ quadra ”, “ quadra 840.”; 1 ♀ (probably, a paralectotype of quadra), “ quadra faem. Angl. Jones”; ENGLAND: 1 ♂ (lectotype of deplana), “ Noct. Diplana / Fabr. p. 215 N o29”.
Material dissected. SPAIN: 1 ♂, gen. prep. 17.402, Prov. Gerona, Besalu, M.viii.[19]83, Pavlas leg. (MWM); BULGARIA: 1 ♂, gen. prep. 17.401, Albena, 21.vi.1971 (MWM); 1 ♀, gen. prep. 17.425, Pirin Mts., Bez. Sandanski Liljanovo, vi–vii.1985, W. Thomas leg. (MWM); CRIMEA: 1 ♂, gen. prep. 17.412, Karadag, 3.vii.1992, Z. Kljutschko leg. (MWM); TURKEY: 1 ♂, gen. prep. 17.403, Prov. Bolu, 11 km WSW Bolu, 900 m, 3.viii. [19]86, de Freina leg. (MWM); 1 ♂, gen. prep. 17.414, Camlibel-Paß, 1600 m, 27.vii.1977, W. Thomas leg.
(MWM); 1 ♂, gen. prep. 17.415, Kars, Posof, 1400–1700 m, 10–19.vii. [19]80, Eckweiler leg. (MWM); 1 ♀, gen. prep. 17.426, Kars, Posof, 2 km NE, 1700 m, 30.vii.1981, Groß, Herbst, R. & A. Hoffman leg. (MWM); GEORGIA: 1 ♀, Tkvibuli, 6–31.vii. [18]95, Kisljakov leg. (ZISP); 1 ♀, Kobulety (Kobuleti), 24.vii. [19]10, N. Satunin leg. (ZISP); 1 ♂, 1 ♀, Chakva (Chakvi), 18.viii. [19]34, V. Nikolskii leg. (ZISP); 1 ♂, 1 ♀, Batumi, 5.viii, 20.viii.1976, S. Sinev leg. (ZISP); 1 ♂, Bakurian (Bakuriani), 8.viii. [18]80, Christoph leg. (ZISP); 2 ♂, 1 ♀, Borshom (now—Borzhomi), 6.vii. [18] 80, 7.vii. [18]98, Christoph & anonymous leg. (ZISP); 1 ♂, Tiflis (now—Tbilisi), 28.vi. [18]97, E. Koenig leg. (ZISP); 5 ♂, 5 ♀, Lagodekhi, 14.viii. [18] 80, 12.viii. [18] 90, 22.vi, 29.viii. [18] 93, 15.vi, 7.vii, 20–27.vii, 29.viii. [18]96, Mlokossevich leg. (ZISP); ARMENIA: 1 ♂, Gen. Präp. 17.418, Garni, E von Eriwan, 21–23.viii.1995, Kazarjan leg. (MWM); UZBEKISTAN: 1 ♂, Gen. Präp. 17.409, 50 km E Dargan-Ata, Amu Darja river, 10–16.v.1995, A. Karpov leg. (MWM); KAZAKHSTAN: 1 ♂, gen. prep. 17.410, Saur Mts., Saisan, 10.vi.1990, Schintlmeister coll. (MWM); RUSSIA: 1 ♂, gen. prep. 17.404, 1 ♀, Gen. Präp. 17.428, NW-Kaukasus, Pjatigorsk, 24.viii.1993, Schintlmeister coll. (MWM).
Other material studied. ENGLAND: 1 ♂, “ quadra mas / Angl. Jones” (BMNH); SWEDEN: 1 ♀, Baltic Sea near Gotland, 15.vii.1912, L. S. Bagrov leg. (ZISP); GERMANY: 1 ♂, Herrenalb, Schwarzwald, ix.[18]98, O. Adelung leg. (ZISP); 1 ♂, Kieshof, Greifswald, 24.viii. [19]02, Bundel coll. (ZISP); 1 ♂, 3 ♀, Berlin, Eversmann coll. (ZISP); CZECHIA: 1 ♀, Turnov, 22.vii. [19]40, Bundel coll. (ZISP); POLAND: 1 ♂, Stettin (now—Szczecin), 9.vii.1908, A. Djakonow leg. (ZISP); 1 ♂, 1 ♀, Preussen, Allenstein (now—Olsztyn), 7–13.vii. [18]98 (ZISP); LITHUANIA: 1 ♂, Mustula, 3.vi.1975, P. Ivinskis leg. (SZMN); 1 ♀, Oškiuiai, 26.vii.1976, P. Ivinskis leg. (SZMN); HUNGARIA: 1 ♂, Mikepércs, 29.ix.1964 (SZMN); 1 ♀, Ócsa, Nagywrdő, 17.vi.1952, Dr. Kovács leg. (SZMN); 3 ♂, 20 km W Kecskemét, Fülöpháza, 11.ix.1991, V. V. Dubatolov leg. (SZMN); UKRAINE: 1 ♂, Kiew, Meinhard coll. (ZISP); 2 ♂, Poltava, Dikan’ka, 15.vii, 27.vii.1928, E. Miljanovskij leg. (ZISP); CRIMEA: 9 ♂, 6 ♀, Bakhchisarai, 4 km from the town, 26.vi–8.viii.1993, S. V. Vasilenko leg. (SZMN); 1 ♂, Sebastopol, 7.ix. [19]06, W. Pliginski leg. (ZISP); 1 ♂, Taush. Bazar, 28.vi. [19]07, W. Pliginski leg. (ZISP); 1 ♂, Mukhalatka, viii.1902, N. Kusnezov leg. (ZISP); 2 ♀, distr. Jaltensis, loc. Mischor, 9.viii.1907, S. Tshetverikov leg. (ZISP); 1 ♂, 1 ♀, southern coast, 18.vii. [19]24, Bundel coll. (ZISP); 1 ♀, Yalta, Nature Reserve, 8.vii.1954, Yu. P. Korshunov leg. (SZMN); 1 ♂, 2 ♀, Sudak, 5– 7.vii.[19]71, V. Prasolov leg. (ZISP); 1 ♂, 15 km NE Sudak, Kurortnoe, 5.viii.1988, A. Barkalov leg. (SZMN); 3 ♂, 1 ♀, Kertsch, 25.vii. [19] 00, 25.vi, 2.vii.1916, Kiritschenko leg. (ZISP); TURKEY: 2 m $, 1 ♀, Prov. Artvin, 10 km S Zevtinlik, vic. Marlik, 350 m, 30–31.vii.1983, de Freina leg. (MWM); ABKHASIA: 3 ♂, 5 ♀, Sukhumi, 7.x.1936, 9.ix.1945, 8.xi.1949, 3.x.1953, 16.ix.1954, 22.ix.[19]62, Miljanowsky leg. (ZISP); ARMENIA: 2 ♂, 50 km NE Erevan, Tsakadzor, 40° 32' N, 44° 42' E, 1900 m, 5.ix.2012, V. Anikin leg. (SZMN); 9 ♂, 4 ♀, Kafan, 31.vii–6.viii.1969, Bundel leg. (ZISP); 1 ♀, Gedzhanan (now—Kadzhants), 7500–8000', 16.vii. [19]39, Rjabov leg. (ZISP); 2 ♂, Megri, 9.viii, 24.ix.1963, Azarjan leg. (SZMN); AZERBAIJAN: 1 ♀, Gerust, 3.vii. [18]81, Christoph leg. (ZISP); 1 ♂, Avrora, 3.vi.1980, M. Danilevsky leg. (MWM); 1 ♂, 1 ♀, Talysh Mts., Kogui, 900 m, 19.vii.1984, V. Lukhtanov leg. (SZMN); 1 ♂, 1 ♀, Talysh, 15 km S- W Masalli, h= 500 m, 2.vi.2005, V. Tikhonov leg. (MWM); IRAN: 1 ♂, Nord Elburs, Caspian Sea, Babolsal, 500 m, vi.2001, G. Müller leg. (MWM); KAZAKHSTAN: 3 ♂, Dzhungar, Boro-Horo Range, 35 km N Panfilov, 1800 m, 28–30.vi.2010, leg. S. Korb, D. Pozhogin, A. Shaposhnikov & L. Tzylin (coll. S. Korb); BELARUS: 1 ♂, 1 ♀, Pinsk, Meinhard coll. (ZISP); RUSSIA: Bryansk Province: 9 ♂, 3 ♀, 15 rv SW Bryansk, Poluzh’e, 14–19.vii.1985, V.G. Makhat (SZMN); Belgorod Province: 1 ♂, Borisovka, 13.vii.1984, Kokiev leg. (ZISP); Tver’ Province: 1 ♂, Udomelskii District, 6.viii. [20]06, A.G. Korobkov leg. (ZISP); Ul’yanovsk Province: 3 ♂, 2 ♀, Staraya Maina Distr., 10 km NE Staraya Maina, biostation, 24.vi.1993, V. Zolotuhin leg. (coll. Zolotuhin); 2 ♂, 2 ♀, Cherdakly Distr., vill. Staroe Eremkino, by lake, 25.vii.1998, V. Zolotuhin leg. (coll. Zolotuhin); 1 ♀, Melekessky Distr., ‘Fakel’ camp by Dimitrovgrad, coast of Cheremshan River, 20.vi.2006 ex p., S. Pugaev leg. (coll. Zolotuhin); 2 ♂, ♀, Novaya Malykla Distr., outsk. vill. Starya Besovka, coast of Cheremshan River, 27.vi.2010, V. Zolotuhin leg. (coll. Zolotuhin); Samara Province: 1 ♀, Zhigulevskii Nature Reserve, 9.vii.1937, Preobrazhenskii leg. (ZISP); Astrakhan Province: 1 ♂, Akhtubinskii District, a border between sands and river flood-plain, 3.vi.2006, I. Kamskov leg. (SZMN); Krasnodar Province: 1 ♂, Ust’-Labinskii District, stanitsa Voronezhskaya, 6.vi.2008, A. Matov leg. (ZISP); 1 ♀, Tuapse, 1914, N.S. Bryanskii leg. (ZISP); 1 ♂, Kropotkin, 45° 25' N, 40° 37' E, 20.vi.2004, Tikhonov leg. (MWM); 1 ♂, North Caucasus, Bolshoi Zelenchuk river, 11.vii.1939, Djakovov leg. (ZISP); 1 ♀, loc. Gelendshik, 2.ix.1915, A. Ohotnikova leg. (ZISP); 1 ♀, Sochi, Zarya sanatorium, 27.viii.1976, V.V. Dubatolov leg. (SZMN); 1 ♂, Sochi, Agura river, Agura waterfalls, 18.viii.1976, V.V. Dubatolov leg. (SZMN); 1 ♂, Sochi District, Nizhnee Loo, 43.72° N, 39.60° E, 7.vii.2008, Valerskiy leg. (ZISP); 3 ♂, 1 ♀, Sochi District, Belye Nochi, 43° 39' N, 39° 37' E, 23–29.vi.2008 Valerskiy leg. (ZISP); Stavropol’ Province: 1 ♂, 2 ♀, Stavropol, 14.vii, 22.vii, 16.viii. [1]920, Filipjev leg. (ZISP); 1 ♀, Stavropol, 21.vii.1984, O. Kosterin leg. (SZMN); 3 ♂, 3 ♀, Zheleznovodsk, without data and 23.vii, 28.vii.1895, 7.vii.[19]15, Bramson, Keler et anonymous leg. (ZISP); 1 ♀, Pyatigorsk, 15.vii. [19]53 (ZISP); 1 ♂, 1 ♀, Pjatigorsk, 24.viii.1993, Schintlmeister coll. (MWM); 8 ♂, 7 ♀, Mashuk Mt., 20.vi, 5.vii, 25.vii. [19] 37, 17.vi, 23.vi, 27.vi, 4.viii, 27.viii. [19] 38, 23.vi, 5.viii. [19] 39, 22.viii, 27.ix.1940, 5.vii.1949, [Rjabov leg.] (ZISP); Karachaevo-Cherkesia: 6 ♂, 1 ♀, fl. et loc. Teberda, 11–29.vii.1912, S. Tschetwerikov leg. (ZISP); 2 ♂, 3 ♀, Teberda, 5.vii, 20.viii. [19]40, Bundel coll. (ZISP); 1 ♀, Teberda Nature Reserve, Late viii.[1]984, K. Gorodkov leg. (ZISP); 1 ♂, Dombai, 43º 18' N, 41º 38' E, 1590 m, 10–16.viii.2015, Anikin leg. (SZMN); Chechnya: 1 ♂, 1 ♀, loc. Groznyi, 23.vi, 15.vii.1906, N. Rodnenskij leg. (ZISP); Dagestan: 1 ♀, Tarki, 23.vii. [19]46, Rjabov leg. (ZISP); 1 ♂, Lake Ak-gjol, 4.x. [1]939, [Rjabov leg.] (ZISP); 1 ♀, Derbent, 1.ix. [19]33, [Rjabov leg.] (ZISP); 3 ♀, Derbent, 28.viii.1974 (SZMN); 1 ♂, 1 ♀, 8 km S Makhachkala, Talgi, h= 200 m, 42° 54' N, 47°, 31' E, 24.vi.2004, Kostjuk leg. (MWM); 1 ♀, Kapchugai, 22.viii. [19]40, Rjabov leg. (ZISP); village Arkas, 18.vi. [19]41, Rjabov leg. (ZISP).
Description. Male. Forewing length 16–22 mm. Forewing ground colour brownish-grey of different hues, orange at basal 1/5; basal part of costal margin black. Outer 1/4 significantly varied in coloration; typically it is dark brown, but sometimes yellow. Hindwings light yellow, with greyish costal area. Cilia coloration either likes the adjacent part of wing, or sometimes black from apex to Cu2 vein. Male genitalia (Figs 33–38). Uncus broad, downturned at apex, with a spine at the tip. Cucullus oval, sometimes slightly extended at apex, membranous, hairy. Sacculus sclerotized, constricted to apex; rounded at tip. Harpe short, broad, like a cockscomb, ventral spines heavier, longer. Juxta triangular. Saccus short, concave apically. Aedeagus short, only twice longer than uncus, cylindrical. Vesica very short, not longer and wider than aedeagus in diameter, with a large but thin single cornutus. Female. Forewing length 20–24 mm. Head and thorax light yellow. Abdomen yellow with a noticeable mixture of dark gray hair-like scales. Wing ground colour yellow. Forewing with two black spots: one between costa and fore angle of discal cell, another between discal cell and anal vein at middle part of the wing. Cilia coloration like the adjacent part of wing, except for the forewing apex, usually marked with black scales. Female genitalia (Figs 42–44). Antevaginal plate forming two simple semi oval folds on sides of vaginal sinus; these folds connected proximally with a sclerotized ring. Ductus covered with spines in distal half only.
Diagnosis. L. quadra can be separated from the similar L. yuennanensis in zones of sympatry only by genitalia. The characters separating L. quadra from L. yuennanensis are: in males, the harpe (shaped like a cockscomb) and the vesica (very short with a strong spine-like cornutus); in females, the two semi oval folds around vaginal sinus (without rugosity, connected proximally with a sclerotized ring) and the ductus bursae (covered with small spines in distal half only).
Distribution (Fig. 56). Typical Amphipalaearctic species with widely disjunctive range. The Atlantic part of the distribution area occupies most parts of Europe, excluding the central and northern parts of Fennoscandia and Estonia, the northern part of European Russia, south-eastern Spain and southern Greece (Peloponnesus). In Russia, the northern distributional range is following: Darwin Nature Reserve in Tver Province, southern regions of Kirov (Vyatka) Province, central part of Udmurtia. The most eastern records are from the Katav-Ivanovsk Region in Chelyabinsk Province in Ural Mts. (50 km W from the headwater of Ural River), the central and southern parts of Orenburg Province in Russia and the western regions of Kazakhstan. However, the species is not known from arid regions in the south of European Russia (north from the Caucasus) but is not rare in humid localities of the Volga’s low valley and delta. In South-West Asia, L. quadra is known from the northern and central provinces of Turkey, the whole Caucasus and Transcaucasia as well as the north-western parts of Iran and the northern slope of Elburs Mts.; the most eastern locality in Iran is Siaret in the NNW of Shirvan south from the Kopetdagh Mts. A presence of the species in the Saur Mts. at the border between Kazakhstan and China was affirmed by Mr. S. Korb. Sporadic records from Amu-Darja River valley (Uzbekistan), Tomsk Province in West Siberia, Baikal, Kamchatka (Sedykh, 1979) have not been confirmed by recent collectors and appear doubtful although not impossible, at least from Uzbekistan and Kamchatka, the latter might be caused by vagrant specimens. One female from MWM is labeled “ Israel ” (N Dead Sea, zw. Beit HaArava & 5 km N Kalya junction, 12.v 2002, - 60 m, leg. A. Sulak)—this southernmost locality is extremely untypical for the species’ distribution and therefore has to be confirmed. The Pacific part of the distributional area occupies the eastern regions of Transbaikalia, the Amur basin north to Tukuringra Mts., 54° N (Dubatolov et al., 2013, 2015), Bureja Mts., 51° N (Koshkin, 2013), and the north-eastern border of broad-leaved forests along the lower valley of Amur, up to 51° 30' N (Dubatolov, 2009). In Sakhalin, the species occurs up to 52° N (Tshistjakov, 2012), which corresponds to the distribution along the river Amur. Overall the species is distributed in Primorskii Krai in SE Russia; it is also known from Dunbei provinces of China (Heilongjiang, Jiling, Liaonin) (Fang, 2000), North and South Korea, Japan (Hokkaido, Honshu, Shikoku, Kyushu, Yaku) (Kishida, 2011a). A record from Shandong (Fang, 2000) should be studied, as it may belong to the following species.
Witt et al. (2011: 175) cited L. quadra as “A Trans-Palaearctic species, its range extends from the Atlantic coast of Europe to the Pacific Region (Russian Far East, Korea, Japan, eastern China and Taiwan)”. This is partially wrong: the members of the genus are absent from Taiwan, and in eastern and central China L. quadra is replaced by a sibling species, L. yuennanensis (Dubatolov, 2010), see below.
Bionomics. L. quadra is monovoltine; moths are on the wings from mid June up to early September in the temperate part of Europe. The presence of a full second generation has not been confirmed (Ebert, 1997), but some moths can be occasionally collected in autumn. In subtropical regions (like in West Transcaucasia) moths can fly during autumn till early November (see material). The larvae feed on different tree-lichens; they were found on Peltigera canina and Parmelia spp., and they were also reported feeding on moist leaf litter, as well as living leaves and flowers, and feeding on other caterpillars, such as Lymantria monacha (Linnaeus, 1758) (Ebert, 1997). Caterpillars hibernate in mid and last instars. Pupation occurs in a weak cocoon under bark and stones. This species prefers locations with high humidity.
Taxonomic remarks. The species was described based on a female only accordingly to the original description given by Linnaeus (1758) as “alis depressis luteis; superiorinus punctis dubious atris”. The name levyi Silbernagel, 1944, was mistakenly attributed to Lithosia by LepIndex [http://www.nhm.ac.uk/research-curation/ research/projects/lepindex/]. It was originally introduced as a form of Lithosia deplana (Esper, [1787]), now— Katha depressa (Esper, [1787]).
The populations of the northern Levante and Transcaucasia are separated here as different subspecies.
description: Published as part of Dubatolov, Vladimir V., Zolotuhin, Vadim V. & Witt, Thomas J., 2016, Revision of Lithosia Fabricius, 1798 and Conilepia Hampson, 1900 (Lepidoptera, Arctiidae), pp. 175-196 in Zootaxa 4107 (2) on pages 177-181, DOI: 10.11646/zootaxa.4107.2.3, http://zenodo.org/record/260857