Anaulacaspis nigra ( GRAVENHORST, 1802 )

Assing, Volker, 2016, Revision of the Anaulacaspis species of the Palaearctic region (Coleoptera: Staphylinidae: Aleocharinae), Beiträge Zur Entomologie = Contributions to Entomology 66 (2), pp. 201-255 : 210-225

Anaulacaspis nigra ( GRAVENHORST, 1802)

( Figs 1–30 , Map 1 )

Aleochara nigra GRAVENHORST, 1802: 75 .

Aleochara picea GRAVENHORST, 1802: 75 .

Aleochara fracticornis GRAVENHORST, 1806: 152 .

Falagria pusilla HEER, 1839: 351 .

Falagria nigra var. jonica BERNHAUER, 1910: 78 ; syn. n. Melagria anatolica FAGEL, 1969: 9 ff.; syn. n.

Melagria corcyrana FAGEL, 1969: 8 f.; syn. n.

Type material examined: Aleochara nigra : Lectotype ♂, present designation: “Hist.-Coll. ( Coleoptera ), Nr. 5297, Falagria nigra Gyllh. , Germania, Zool. Mus. Berlin / Syntypus Aleochara nigra Gravenhorst, 1802 , labelled by MNB 2011 / Lectotypus ♂ Aleochara nigra Gravenhorst , desig. V. Assing 2016 / Anaulacaspis nigra (Gravenhorst) , det. V. Assing 2016” ( MNB) . Paralectotypes: 1 ♂, 4 ♀♀: same data as lectotype ( MNB) .

Aleochara picea : Lectotype ♂, present designation: “Hist.-Coll. ( Coleoptera ), Nr. 5297, Falagria nigra Gyllh. , Germania, Zool. Mus. Berlin / Syntypus Aleochara picea Gravenhorst, 1802 , labelled by MNB 2011 / Lectotypus ♂ Aleochara picea Gravenhorst , desig. V. Assing 2016 / Anaulacaspis nigra (Gravenhorst) , det. V. Assing 2016” ( MNB) . Paralectotype ♀: same data as lectotype ( MNB) .

Falagria jonica : Lectotype ♀: “Gr. Attica, Bang-Haas / jonica / v. jonica Brh, Cotypus / Chicago NHMus, M. Bernhauer Collection / Lectotypus Fagel 1969 / Anaulacaspis nigra (Gravenhorst) , det. V. Assing 2016” ( FMNH).

Melagria anatolica : Holotype ♂: “ Anatolie mér., Alanya (plage), VI.1968 G. Fagel / G. Fagel det., anatolica n. sp. / Type / Anaulacaspis anatolica (Fagel) , det. V. Assing 2016” ( IRSNB) . Paratypes: 1 ♀: same data as holotype ( IRSNB); 1 ♀: “ Anatolie mér., Alanya : Dim Irmak , VI.1968 G. Fagel / G. Fagel det., anatolica n. sp. / Paratype / Anaulacaspis nigra ( Gravenhorst) , det. V. Assing 2016” ( IRSNB) .

Melagria corcyrana : Holotype ♂ [dissected prior to present study; aedeagus damaged considerably]: “ Corfu , Reitter. / G. Fagel det., corcyrana n. sp. / Type / Anaulacaspis corcyrana (Fagel) , det. V. Assing 2016” ( IRSNB) . Paratype ♀: same data as holotype ( IRSNB)

.

Comment

: The original description of Aleochara nigra is based on an unspecified number of syntypes in “Mus. Prof. Knoch” ( GRAVENHORST 1802). The locality is not specified, but the species was described in a work on the Staphylinidae of the environs of Braunschweig (North Germany). Six syntypes, two males and four females, were found in the collections of the MNB. One of the males is designated as the lectotype.

Aside from the type series of A. nigra , four pins with syntype labels of Aleochara picea , two of them with mounting cards with only a single leg left, were located in the MNB. Two syntypes, a male and a female, are still in good condition. The male is designated as the lectotype. The previously established synonymy of A. picea with A. nigra is confirmed.

BERNHAUER (1910) described Falagria nigra var. jonica based on an unspecified number of syntypes from “Korfu, Attika und Kroatien ”, stating that this variety was distinguished from Central European populations by longer and broader elytra, but otherwise identical. FAGEL (1969) treated this taxon as a species, designated a female [sic] from “ Attica ” from the Bernhauer collection as the lectotype, recorded A. jonica from “différentes parties de la Grèce, de Dalmatie et ... Turquie d´Europe: Istanbul (Rumeli Hisar)”, and assigned the material studied from Corfu to a new species. A comparison of the lectotype of A. jonica with material from other regions studied in the course of the present revision, however, revealed that, regarding its external and the female secondary sexual characters, the specimen is within the range of intraspecific variation of A. nigra . Moreover, A. nigra appears to be the only species of the A. nigra group present in mainland Greece. Consequently, A. jonica is placed in synonymy with A. nigra .

The original description of Melagria anatolica is based on a male holotype from “Alanya (plage)”, one female from the same locality, one female of “même origine, sur plages du Dim Irmak”, and two females from the Rhodopes in Bulgaria ( FAGEL 1969). An examination of material from various regions revealed that the type material of M. anatolica is conspecific with that of A. nigra (see notes on intraspecific variation below).

Melagria corcyrana was described from a male holotype and seven paratypes from “Corfu” ( FAGEL 1969). Material from Corfu differs from many other populations of A. nigra by the modifications of the male pronotum (with an impression in posterior half rather than with a median sulcus) and by the absence of a pronounced semi-transparent carina at the base of the ventral process. However, similarly deviant character conditions were observed also in other populations, e.g. from the Pelopónnisos and Albania. Moreover, these differences do not correspond to a plausible distribution pattern, so that they are interpreted as intraspecific variation of A. nigra .

Additional material examined: Spain: Cataluña: 1 ♂, Olot [42°11'N, 2°28'E] env., pitfall, 14.VII.1977, leg. Hoffmann ( MHNG) .

France: Aquitaine: 1 ♀, Gironde, Lamothe [44°37'N, 0°59'W], 8.IX.1928, leg. Tempère ( MHNG) ; 1 ♀, Léognan [44°43'N, 0°35'W], leg. Gouin ( MHNG) ; 1 ♀, Beautiran [44°41'N, 0°26'W], VII.1944 ( MHNG) ; 1 ♀, Gironde, Gajac , 44°26'N, 0°08'W], 28.VI.1908 ( MHNG) . Midi-Pyrénées: 1 ♂, 2 ♀♀, Hautes-Pyrénées , Luz-Saint-Sauveur [“Luz”; 42°52'N, 0°00'E], IX.1911, leg. Tempère ( MHNG, cAss) . Picardie: 1 ♀, Becquigny [50°00'N, 3°27'E], near Bohain-en-Vermandois , 10.–19. IX.1918, leg. Salchert ( MNB) . Alsace: 1 ♀, Strasbourg , VIII.1953 ( MHNG) . Rhône-Alpes: 1 ♂, Megève [45°51'N, 6°36'E], X.1960 ( MHNG) . Provence: 1 ♂, Var, Agay [43°26'N, 6°52'E], 30.I.1955 ( MHNG) ; 1 ♂, Var, Fréjus [43°26'N, 6°44'E], VIII.1951 ( MHNG) ; 1 ♂, 1 ♀, Meailles [44°02'N, 6°38'E], 13.–17.IV.1949 ( MHNG) ; 1 ♂, La Cayolle [43°14'N, 5°23'E], 14.VII.1938 ( MHNG) ; 1 ♀, Caille [43°46'N, 6°43'E], IV.1959 ( MHNG) ; 1 ♂, 1 ♀, Le Plan-de-la-Tour [43°20'N, 6°33'E], flood, XII.1959 ( MHNG) ; 1 ♀, Saint-Martin-Vésubie [44°04'N, 7°15'E], leg. Buchet ( MHNG) ; 1 ♀, ex., Alpes-Maritimes , “ Pont du Pali ”, 20.III.1946 ( MHNG) .

Germany: Niedersachsen: 1 ♂, 1 ♀, Northeim env., Weper , pitfall, 4.VIII.1989, leg. Joger (cAss); 1 ♂, same data, but 28.IV.1988 (cAss); 1 ♀, same data, but 5.VI.1988 (cAss); 1 ♂, 1 ♀, same data, but 6.VII.1987 (cAss); 1 ♀, same data, but 31.VIII.1989 (cAss); 7 ♂♂, 4 ♀♀, Lüchow-Dannenberg, Schreyahn, saline habitat, 16.VI.1987, leg. Wunderle (cWun, cAss) . Niedersachsen or Sachsen-Anhalt: 3 ♀♀, Harz ( MNB) .

Mecklenburg-Vorpommern: 2 ♂♂, 2 ♀♀, Schönberg [53°50'N, 10°56'E], 13.VII.1917, leg. Benick ( MHNG) .

Sachsen-Anhalt: 1 ex., Halle env., Gimritz , fallow, pitfall, 3.VIII.1994, leg. Teichmann (cFel); 1 ex., same data, but 11.V.1994 (cFel); 1 ♂, Halle, leg. Ruthe ( MNB) ; 1 ♂, Questenberg [51°28'N, 11°07'E], 25.VIII.1915, leg. Petry ( MNB) . Hessen: 1 ♂, Naumburg , 18.IV.1920, leg. Maertens ( MNB) ; 1 ♀, Marburg , 1.IV.1904 ( MNB) ; 1 ♀, Kühkopf [49°49'N, 8°26'E], 29.I.1955, leg. Vogt ( MHNG) . Brandenburg / Berlin: 1 ♀, Berlin, 2.IV.1990, leg. Göllner ( MNB) ; 1 ♀, Berlin-Lankwitz, Pädagogische Hochschule , fallow, 16.IV.1971 ( MNB) ; 1 ♂, Berlin, Bahnhof Heerstrasse , 6.V.1958, leg. Puthz ( MNB) ; 1 ♂, same data, but 19.VI.1958 ( MHNG) ; 1 ♀, Berlin, leg. Ruthe ( MNB) ; 1 ♂, Berlin-Lichtenrade, Richard-Tauberdamm , fallow, 14.VI.1980, leg. Korge ( MNB) ; 1 ♂, 1 ♀, Pankow, 1909 ( MNB) ; 1 ♂, Berlin env., Marienfelde ( MNB) ; 1 ♀, Schönefeld env. ( MNB) ; 1 ♀, Rüdersdorf , 17.VIII.1942 ( MNB) ; 1 ♀, Eberswalde ( MNB) ; 1 ♂, Döberitzer Heide , sand pits, 3.VII.2002, leg. Korge ( MNB) ; 1 ♂, Luckenwalde , IX.1903, leg. Delahon ( MNB) ; 1 ♂, same data, but V.1918 ( MNB) ; 1 ♀, same data, but VII.1914 ( MNB) ; 1 ♀, Schloss Güterfelde [“Gütergotz”; 52°22'N, 13°12'E] ( MNB) ; 1 ♀, Eisenhüttenstadt , 26.III.1982, leg. Pütz ( MNB) . BadenWürttemberg: 8 ♂♂, 4 ♀♀, Altshausen env. , 28.–30. VII.1987, leg. Wunderle (cWun, cAss) ; 1 ♂, Gruibingen [48°35'N, 9°38'E], 10.V.1964, leg. Lohse ( MHNG) .

Rheinland-Pfalz: 1 ♂, 2 ♀♀, Bad Dürkheim, leg. Eppelsheim ( NHMW) ; 2 ♂♂, 2 ♀♀, Speyer , XI.1882 ( NHMW) ; 1 ♀, Rothaus [47°48'N, 8°14'E], 1908, leg. Riehn ( MNB) . Bayern: 1 ♂, 1 ♀, München , 3.IX.1907, leg. Ihssen ( MNB) ; 2 ♂♂, 7 ♀♀, München , 24.IV.1910, leg. Ihssen ( MNB) ; 1 ♀, München , 9.IV.1909, leg. Ihssen ( MNB) ; 1 ♂, München , 3.V.1910, leg. Ihssen ( MNB) ; 1 ♂, München , 24.VIII.1911, leg. Ihssen ( MNB) ; 1 ♂, Garmisch-Partenkirchen , 26.V.1942, leg. Ihssen ( MNB) ; 2 ♀♀, Kastl ( Oberpfalz ) [49°49'N, 11°53'E], leg. IV.1913, leg. Waegner ( MNB) ; 1 ♂, S Murnau, Eschenlohe [47°36'N, 11°10'E], 3.VIII.1937, leg. Ihssen ( MNB) ; 1 ♂, 2 ♀♀, Lenggries ( MNB) ; 1 ♀, Lenggries-Winkl , 11.V.1964, leg. Lohse ( MHNG) ; 1 ♀, Bamberg ( MNB) ; 1 ♂, “ Ob. - Bayern ” leg. Arnold ( MNB) . Thüringen: 3 ♀♀, Alter Stolberg [51°32'N, 10°55'E], 11.XI.1917, leg. Petry ( MNB) ; 1 ♂, same data, but 19.VIII.1910 ( MNB) ; 2 ♂♂, 2 ♀♀, Kyffhäuser, Kalktal , 26.IX.1958, leg. Dorn ( MHNG, MNB) . Sachsen: 2 ♀♀, Moritzburg [51°09'N, 13°41'E], leg. Minkwitz ( MNB) . German or Polish territory: 2 ♀♀, “Pommern”, leg. Schneider ( NHMW) .

Switzerland: 3 exs., locality not specified ( MHNG) .

Austria: Oberösterreich: 2 ♂♂, 2 ♀♀ [with workers of Formica pratensis attached to the pins], Niederranna, leg. Rupertsberger ( NHMW) ; 1 ♀, Wien, “ N. Donau ”, leg. Meschnigg ( NHMW) . Tirol: 1 ♂, Trins [47°04'N, 11°25'E; “F4”], leg. Franz ( NHMW) . Wien / Niederösterreich: 6 ♂♂, 4 ♀♀, Wien, Bisamberg, leg. Luze ( NHMW) ; 1 ♂, 1 ♀, Wechselgebirge , 1889, leg. Ganglbauer ( NHMW) ; 1 ♀, Hollenstein [47°48'N, 14°46'E], Krenngraben [“ X 1068 ”], leg. Franz ( NHMW) ; 2 ♂♂, 1 ♀, Mödling env. [“403”, “515”], leg. Franz ( NHMW) ; 1 ♀, Windisch Baumgarten [48°33'N, 16°43'E] ( MNB) ; 3 ♀♀, Lunz, Lunzberg , xerothermous slope , 10.IX.1962, leg. Puthz ( MHNG) ; 3 ♀♀, Theresienfeld [47°51'N, 16°15'E], 20.IV.1952, leg. Malicky ( MHNG) ; 1 ♂, 1 ♀, Theresienfeld , 15.III.1952, leg. Malicky ( MHNG) ; 1 ♂, Kollnitz [48°14'N, 15°08'E], 3.IV.1958 ( MHNG) ; 2 ♀♀, Fischauer Vorberge [47°49'N, 18°06'E], nest of mouse sifted, 16.III.1952, leg. Malicky ( MHNG) ; 1 ♀, locality illegible [“Feichsbach”?], 21.VI.1961 ( MHNG) ; 1 ♀, locality illegible, 10.III.1961 ( MHNG) ; 1 ♀, locality illegible, 1.XI.1960 ( MHNG) . Steiermark: 1 ♀, Weiz env. [“X 986”], leg. Franz ( NHMW) ; 1 ♀, Baierdorf [47°10'N, 14°06'E] near Schöder , 7.X.1949, leg. Franz ( NHMW) ; 4 ♂♂, 4 ♀♀, Frohnleiten ( MNB). Kärnten: 2 ♀♀, Maglern an der Gailitz , leg. Meschnigg ( NHMW) ; 1 ♂, 1 ♀, Villach, Seebach , leg. Meschnigg ( NHMW) ; 2 ♂♂, Heiligenblut [“G 370”], leg. Franz ( NHMW) ; 2 ♀♀, Mauthen [46°39'N, 12°59'E], 21.VII.1973 ( MHNG) ; 1 ♀, Rattendorf [46°36'N, 13°16'E], 2.VII.1966, leg. Lohse ( MHNG) ; 2 ♂♂, “ Kärnten ” ( MNB) . Burgenland: 3 ♀♀, Rohrbach bei Mattersburg, Marzer Kogel , nest of Formica sanguinea , 1.IV.1926, leg. Scheerpeltz ( NHMW) ; 6 ♂♂, 5 ♀♀, Neusiedlersee [“351”], leg. Franz ( NHMW, cAss) ; 9 ♂♂, 6 ♀♀, Zurndorf [“345”; “74-015”], leg. Franz ( NHMW, cAss) ; 1 ♂, 2 ♀♀, Illmitz , 22.VIII.1960, leg. Lohse ( MHNG) ; 7 exs., Breitenbrunn , pitfall, VII.– X.1965, leg. Malicky ( MHNG) ; 2 exs., same data, but V–VI.1965 ( MHNG) ; 1 ♀, Apetlon , pitfall, V.1967, leg. Malicky ( MHNG) .

Czech Republic: 3 ♀♀, Brandys nad Labem, leg. Skalitzky ( NHMW).

Slovakia: 1 ♀, Plešivec [48°32'N, 20°23'E] plateau, 5.V.1967, leg. Lohse ( MHNG) .

Italy: Trentino-Alto Adige: 1 ♀, Val di Non ( TN) , Castelfondo [48°27'N, 11°06'E], torbiera d. Regole , 1230 m, 2.VIII.1982, leg. Zanetti (cZan); 1 ♀, Bolzano ( MNB) ; 1 ♀, Bressano [46°42'N, 11°39'E], 4.VI.1965, leg. Lohse ( MHNG) . Lombardia: 1 ♂, NE Sondrio, Montagna in Valtellina , 6.IV.1972, leg. Dioli (cZan); 1 ♂, 1 ♀, Bergamo , Dalmine [45°39'N, 9°36'E], Brembo river , 30.V.1979, leg. Kiener ( MHNG) . Veneto: 3 ♂♂, 4 ♀♀, Venezia, San Giuliano, Ponte sul Oselin , 26.XII.1954, leg. Zecchini (cZan); 1 ♂, 3 ♀♀, Venezia, San Giuliano , 21.XI.1954, leg. Zecchini (cZan, cAss) . Emilia-Romagna: 1 ♂, 1 ♀, Montone, leg. Brenske ( NHMW) . Lazio: 1 ♂, 1 ♀, Roma, Aniene river , II.1902, leg. Fiori ( MHNG, cAss); 5 exs. [with worker of Formica sp. attached to the pin], Acquapendente , 14.VIII.1922, leg. Botto ( MHNG) .

Hungary: 1 ♀, Bugac National Park, grassland, pitfall trap, 30.VII.1986, leg. Galle (cAss).

Romania: 1 ♀, Băile Herculane, 1895, leg. Ganglbauer ( NHMW).

Croatia: 1 ♂, 1 ex. without head and pronotum, locality not specified ( NHMW); 1 ♀, Josipdol – Karlovac, car-net, 9.V.1990, leg. Wunderle (cWun) .

Bulgaria: 1 ♂, Macedonia , Sandanski–Struma , 16.VII.1986, leg. Pütz (cAss); 1 ♂, Sandanski–Struma , 28.IV.1985, leg. Jäger ( MNB); 1 ♂, Sandanski , vineyard, under stones, 5.VI.1984, leg. Pütz (cPüt); 2 ♀♀, Sandanski, 16.–23.VII.1985, leg. Schülke ( MNB); 1 ♂, 3 ♀♀, 1 ♂, 3 ♀♀, Rupite [41°26'N, 23°15'E] near “General Todorow”, 7.V.1984, leg. Hieke ( MNB); 1 ♂, Pimorsko, 16.IX.1977, leg. Hieke & Uhlig (cAss) .

Bosnia-Herzegovina: 3 ♂♂, 2 ♀♀, Travnik , leg. Brandis ( NHMW, cAss); 1 ♀, Fatničko polje, entrance of Lepirnica cave, 17.–22.VI.2003, leg. Hlaváč (cTro); 4 ♀♀, Trebinje ( MHNG) .

Albania: 1 ♂, 5 ♀♀, Vlorë [“Avlona”], leg. v. Oertzen ( MNB, NHMW, cAss).

Greece: mainland: 1 ♀, Katerini, Agios Dimitrios , 800 m, 19.–21.VII.1990, leg. Schuh & Zabransky (cAss); 1 ♀, Thessalia, Volos ( MNB) ; 1 ♂, 1 ♀, Evzoni [41°06'N, 22°32'E], 1.IX.1964, leg. Puthz ( MHNG) ; 1 ♀, Serrai [41°05'N, 23°33'E], river bank , 16.VIII.1964, leg. Puthz ( MHNG) ; 1 ♀, “ Attica ” ( NHMW) . Evvoia: 1 ♀, Karystos , leg. v. Oertzen ( MNB) . Pelopónnisos: 4 ♂♂, 2 ♀♀, Killini Oros, Lake Stimfalia , 500 m, 3.IV.1992, leg. Frisch ( MNB, cAss); 1 ♀, Killini , 37°55'N, 22°22'E, 1350 m, car-net, 3.IV.2016, leg. Schülke ( MNB) ; 1 ♂, Kalávrita , leg. Holtz ( MNB) ; 1 ♂, 1 ♀, Koumani , leg. Brenske ( NHMW, cAss); 4 ♂♂, 4 ♀♀, Taygetos, leg. Brenske ( MHNG, cAss) .

Corfu: 1 ♂, 2 ♀♀, NE-Corfu , Archaravi , 20.–25.X.1991, leg. Katschak (cWun, cAss); 9 exs., locality not specified, leg. Reitter, Winkler ( MHNG, NHMW, cAss). Ikaría: 2 ♂♂, 1 ♀, locality not specified, leg. v. Oertzen ( MNB, cAss). Locality not specified: 1 ♂, 1 ♀, locality not specified [“ Graecia ”], leg. Emge ( NHMW) ; 1 ♂, 1 ♀, “Graecia” ( MNB).

Ukraine: 1 ♂, Transcarpathia, Vel’ke Berezne , sandy river bank, 17.V.2012, leg. Gontarenko (cAss) .

Turkey: Bolu: 10 ♂♂, 8 ♀♀, 26 km S Bolu , N Seben, 40°30'N, 31°36'E, 1410 m, grassy slope with scattered pine trees, grass, moss, and shrub litter sifted, 27.III.2010, leg. Assing (cAss) . Kastamonu: 1 ♂, 5 km NW Kastamonu , 5 km NW Ağlı, 41°44'N, 33°30'E, margin of calcareous fir forest with interspersed pine trees, litter and moss sifted, 22.III.2010, leg. Assing (cAss); 1 ♀, same data, but floated from stream bank (cAss); 1 ♀, 40 km N Kastamonu , S Kayneak geçidi, 41°42'N, 33°41'E, 1250 m, calcareous slope, under stones, 10.IV.2009, leg. Assing (cAss) .

Sinop: 2 ♀♀, 30 km NNE Boyabat, Dıranaz geçidi, exit SE tunnel, 41°38'N, 34°52'E, calcareous grassland, under stones, 5.IV.2009, leg. Wunderle (cWun) . Ordu: 1 ♂, 1 ♀, 25 km S Ordu, S Kabaduz , 40°49'N, 37°54'E, 990 m, roadside, roots of grass and herbs, moss, 30.VII.2006, leg. Assing (cAss) . Muğla: 1 ♀, N Fethiye, Çaliş , 36°40'N, 29°06'E, 10 m, 1.X.2002, leg. Assing (cAss) . Aydin: 1 ♂, 25 km SSW Çine, Gökbel Dağı , N-slope at peak, 37°28'N, 28°00'E, 1420 m, litter of pine and grass sifted, 6.IV.2006, leg. Assing (cAss) . Konya: 1 ♀, Beyşehir [37°40'N, 31°44'E], lake shore , 13.III.1979 ( MHNG) . Antalya: 5 ♂♂, 3 ♀♀, ca. 70 km NE Fethiye, Gülübeli geçidi, east side, 36°50'N, 29°46'E, 1525 m, north slope, litter of cushion plants, grass roots, and moss between rocks sifted, 29.III.2002, leg. Assing & Wunderle (cAss, cWun) ; 1 ♀, same data, but grassland, under stones and sifted (cAss); 1 ♂, Alanya , 12.III.2000, leg. Esser (cAss) ; 2 ♀♀, Taşkesigi , 17.III.2000, leg. Esser (cAss).

Armenia: 2 ♂♂, 20 km Kapan , W Tsav, 39°03'N, 46°26'E, 1170 m, stream valley, litter and flood debris near stream sifted, 9.VII.2016, leg. Assing & Schülke (cAss, MNB) .

Georgia: 1 ♀, Kasbek ( NHMW).

Russia: 1 ♂, 1 ♀, Rjasan, leg. Reitter ( NHMW); 1 ♀, Samara, leg. Eocrsmann [?] ( NHMW) .

Kyrgyzstan: 1 ♂, Issyk-Kul , 25 km W Balykchy, 2 km S Kek Mojnok Vtoroe, 42°27'N, 75°51'E, 1670 m, 18.VI.2011, leg. Frisch ( MNB); 2 ♀♀, Tienshan, Ala Tau: Ala Artsha, 2000 m, 9.VII.1984, leg. Wrase ( MNB) .

Locality not specified: 3 ♂♂, 3 ♀♀, “ Styria ”, “Styr.”, etc., leg. Pipitz, etc. ( MNB, NHMW); 1 ♀, “Silesia” ( MNB) ; 2 ♂♂, 1 ♀, “Kaukas, Leder” ( NHMW, cAss) .

Redescription: Body length 1.7–2.5 mm; length of forebody 0.9–1.1 mm. Coloration: body blackish, with the posterior portion of the elytra more or less extensively and more or less distinctly paler, narrowly yellowish to broadly brown; legs brown to dark-brown; antennae blackish-brown to black.

Head approximately 1.1 times as broad as long; with weakly pronounced sexual dimorphism. Eyes approximately as long as postocular region, or slightly shorter, or slightly longer (in specimens with long elytra). Antenna 0.65–0.80 mm long; antennomere IV weakly transverse or as long as broad; antennomeres V–X of increasing width and increasingly transverse, X slightly more than 1.5 times as broad as long. Mouthparts as in Figs 1–3 .

Pronotum approximately 1.1 times as broad as long and approximately as broad as head, strongly convex in crosssection; punctation subject to a more or less pronounced sexual dimorphism.

Elytra of variable length, 0.8–1.1 times as long as pronotum; punctation dense, fine, and distinct; interstices without microsculpture and glossy. Hind wings fully developed.

Abdomen approximately as broad as elytra in specimens with short elytra and narrower than elytra in specimens with long elytra; punctation denser and coarser in anterior than in posterior portions of tergites; posterior margin of tergite VII with palisade fringe; posterior margin of tergite VIII ( Figs 4–9 ) usually broadly and weakly, rarely deeply concave, with unmodified long marginal setae, in postero-lateral portions with dense long and thin setae.

♂: head often more or less distinctly depressed in posteromedian portion, punctation fine, but distinct; pronotum with very dense and often somewhat asperate punctation, with more or less pronounced median sulcus (this sulcus more distinct posteriorly than anteriorly), and sometimes with depression or shallow median impression in posterior half; sternite VIII ( Figs 10–12 ) strongly produced posteriorly, in the middle weakly convex (nearly truncate) and with numerous long thin setae; median lobe of aedeagus ( Figs 13–30 ) of rather variable size, 0.23– 0.28 mm long, at base of ventral process usually with, rarely without, more or less pronounced membranous median projection (lateral view); ventral process apically weakly incised and laterally weakly angulate (ventral view).

♀: head not depressed in postero-median portion, punctation finer and less distinct than in male; pronotum with shallower, or without, median sulcus, punctation moderately dense, moderately fine, not asperate; sternite VIII with broadly convex posterior margin; spermatheca small and shaped like a comma, not distinctive.

Intraspecific variation: This species is subject to pronounced, partly clinal variation. While in populations from West and Central Europe the elytra are shorter than the pronotum and approximately as broad as the abdomen, they tend to be longer than the pronotum and broader than the abdomen in Southeast Europe and Asia. Also, external characters such as the punctation and the modifications of the male head and pronotum may vary considerably. Moreover, in one examined male from Bulgaria, the posterior margin of the male tergite VIII is much more strongly concave than is usually the case ( Fig. 9 ). Finally, the membranous carina at the base of the ventral process of the aedeagus may be of various shapes ( Figs 13–25 ). In material from southern Turkey, this carina is more pronounced and more obliquely truncate than in populations from West and Central Europe and from most of the Balkans. On the other hand, it may be nearly completely reduced in material from Corfu, the Pelopónnisos, Albania, and mainland Greece. However, transitional conditions were observed for all these characters, suggesting that these differences should be interpreted as intra- rather than interspecific variation.

Distribution and natural history: Anaulacaspis nigra is the most widespread species of the genus, its confirmed distribution ranging from Northeast Spain and France across Central and South Europe, Russia, Turkey, and Armenia eastwards to Kyrgyzstan ( Map 2 ). Plausible records are known also from Scandinavia and the Baltic states ( SCHÜLKE & SMETANA 2015). The species appears to be absent from the British Isles. Based on currently available evidence, the distribution is of the PontoMediterranean type sensu LATTIN (1967). According to SCHÜLKE & SMETANA (2015), the species has been recorded also from Portugal; GAMARRA & OUTERELO (2005) report it from Madrid and Mallorca. However, no material from these regions was seen in the course of the present revision, and based on the currently known distribution, these records are most likely incorrect and may well refer to A. iberica .

The examined material was collected in various habitats such as different types of grassland (steppe, calcareous grassland, mountain slopes), fallows, sandy river banks, an inland saline habitat, a cave entrance, a vineyard, and nests of mice and of wood ants ( Formica pratensis RETZIUS, 1783 , F. sanguinea LATREILLE, 1798 ). In the north of its range, A. nigra is usually confined to xerothermic habitats (sandy soils, calcareous slopes, urban fallows, etc.). Numerous specimens were sifted from grass roots, moss, and litter, some were found under stones. The altitudes range from 10 to 1670 m. Adult specimens were found throughout the year. One specimen was collected with a car-net in May, suggesting that dispersal on the wing may take place in spring. According to HORION (1967), A. nigra is mainly found on sandy soils and teneral adults have been observed in summer (from the end of July). LOKAY (1905), ROUBAL (1932), and LAPEVA-GIONOVA & ILIEFF (2012) recorded it from the nests of various ant species: Formica pratensis , F. rufa LINNAEUS, 1761 , Lasius fulginosus (LATREILLE, 1798) , L. niger (LINNAEUS, 1758) , and Tetramorium caespitum (LINNAEUS, 1758) .

Figures

Map 1: Distribution of Anaulacaspis in the Palaearctic region, based on revised records (all records pooled).

Map 2: Distributions of Anaulacaspis nigra (black circles), A. pseudonigra (white circle), A. reticulata (white triangle), A. libanotica (black triangles), A. caucasica (white diamonds), A. truncata (black diamonds), and A. sinuata (stars) in the West Palaearctic and Middle Asia, based on revised records.

Figs 1–12: Anaulacaspsis nigra. Maxilla (1); labium (2); labrum (3); male tergite VIII (4–9); male sternite VIII (10–12). Scale bars: 1–2, 4–12: 0.1 mm; 3: 0.05 mm.

Figs 13–30: Anaulacaspsis nigra. Median lobe of aedeagus in lateral (13–25) and in ventral view (26–30) of the lectotype of A. nigra (13) and of males from Bosnia-Herzegovina (14), Bulgaria (15–16, 29), Ukraine (17, 30), mainland Greece (18), Turkey (19–20), Albania (21), Korfu (22, 28), and the Pelopónnisos (23–27). Scale bar: 0.1 mm.