Alevonota (Liota) gracilenta ( ERICHSON , 1839)

Wunderle, Volker Assing Paul, 2008, On the Alevonota species of the Western Palaearctic region (Coleoptera: Staphylinidae: Aleocharinae: Athetini), Beiträge Zur Entomologie = Contributions to Entomology 58 (5), pp. 145-189 : 172-179

Alevonota (Liota) gracilenta ( ERICHSON, 1839) ( Figs 59-61 , 63 , Map 5 )

Homalota gracilenta ERICHSON, 1839: 94 .

Homalota venustula HEER, 1839: 340 .

Homalota splendens KRAATZ, 1856: 246 f.; synonymy confirmed.

Liota hypogaea MULSANT & REY, 1875: 175 ff.; synonymy confirmed.

Type material examined:

Lectotype  [damaged, head missing], here designated: "Cassel / coll. Kraatz / Syntypus / Lectotypus Homalota splendens Kraatz desig. V. Assing 2007 / Alevonota gracilenta (Erichson) det. V. Assing 2007 " ( DEI). Paralectotype : "Misdroy [= Poland: Międzyzdroje] / coll. Kraatz / Syntypus / splendens mihi = gracilenta Er., German. " ( DEI).

Comments:

Homalota gracilenta was described from an unspecified number of syntypes from Paris ("Habitat Lutetiae") collected by Aubé ( ERICHSON 1839).

The original description of Homalota splendens is based on an unspecified number of syntypes from Germany ("ueber ganz Deutschland verbreitet") ( KRAATZ 1856). Two syntypes, a male without head and a female, were found in the Kraatz collection at the DEI. The male is here designated as the lectotype.

Liota hypogaea was described from an unspecified number of syntypes from "Massane (Pyrénées- Orientales)" ( MULSANT & REY 1875). In the Rey collection, however, only a female labelled "Nice, Grouvelle" was found; this specimen is conspecific with A. gracilenta .

The genus Liota was described by MULSANT & REY (1874) to include two species, Homalota gracilenta ERICHSON and H. laeviceps BRISOUT. FENYES (1918) designated the former as the type species. Liota was later synonymised with Alevonota and has remained a synonym ever since (BERN- HAUER & SCHEERPELTZ 1926; FAUVEL 1895; SMETANA 2004). The present study revealed, however, that a revalidation as a subgenus appears justified for several reasons. Though somewhat similar in the general subparallel facies and in the morphology of the antennae, the type species of Liota , H. gracilenta , differs in various significant respects from other species of Alevonota , not only by external characters such as the distinctly transverse head, the more pronounced postgenal carinae, and the shape of the pronotum, but especially also in the the general morphology of the genitalia (aedeagus with long flagellum; spermatheca with twisted duct, somewhat resembling that of the genus Geostiba THOMSON ). Based on these observations, it does not seem unlikely that a thorough phylogenetic analysis, including also molecular data, may eventually reveal an even more distant relationship of Liota to Alevonota , so that both would have to be considered distinct genera.

Additional material examined:

Finland: 1 ex., Hanko, 11.V.-17.VI.1990, leg. Rutanen ( ZMH) .

Spain: Galicia: 1 ex., Pontevedra, Beloso ["Belus", 42°19'N, 08'°45'W], leg. Franz ( NHMW) .

France: Île-de-France: 1 ex., Paris, leg. Fauvel ( NHMW) . Provence: 1 ex., Alpes Maritimes, St. Augustin , IV.1949 ( MHNG) ; 1 ex., Alpes-Maritimes, Nice, leg. Grouvelle ( MGHNL) .

Switzerland: Vaud: 1 ex., Marchissy ( MHNG) .

Germany: Nordrhein-Westfalen: 1 ex., Marsberg, Dahlberg , mesobrometum, 5.VII.1992, leg. Lückmann (cFel) ; 1 ex., same data, but 3.VI.1995 (cFel) ; 1 ex., Münster-Kinderhaus , garden, V.2002, leg. Feldmann (cFel) ; 1 ex., same data, but 26.VI.2006 (cFel) ; 1 ex., Porta Westfalica, Wittekindsberg , car-net, 10.VI.1993, leg .

Assing (cAss). Rheinland-Pfalz: 1 ex., "Wattenh."[= Wattenheim] ( DEI) ; 1 ex., Eifel, Fischbachtal, Neuerburg [49°59'N, 06°57'E], car-net, 1.VI.1990, leg. Wunderle (cWun) ; 1 ex., Altenahr , pitfall, 21.VI.1988, leg. Büchs (cWun). Baden-Württemberg : 1 ex., Stuttgart , 9.V.1876, leg. Simon ( NHMW). Hessen : 1 ex., Dautphe- Buchenau, Katzenbachtal , car-net, 30.IV.1988, leg. Wunderle (cWun). Niedersachsen : 1 ex., Lüneburger Heide, Schneverdingen env., Tütsberg , grassy heathland, pitfall, V.1995 (cAss) ; 4 exs., same data, but VI.1996 (cAss) ; 1 ex., same data, but V.1997 (cAss) ; 1 ex., same data, but VI.1997 (cAss) ; 3 exs., same data, but VII.1998 (cAss) ; 1 ex., same data, but IV.1999 (cAss) ; 2 exs., same data, but V.1999 (cAss) ; 1 ex., same data, but IX.1999 (cAss) ; 3 exs., same data, but IV.2000 (cAss) ; 1 ex., same data, but V.2000 (cAss) ; 3 exs., same data, but V.2001 (cAss) ; 1 ex., Hannover , garden, pitfall, V.1987, leg. Assing (cAss) ; 1 ex., same data, but VIII.1987 (cAss) ; 1 ex., Hannover, bank of Leine river , pitfall, V.1990 (cAss) ; 1 ex., Hannover-Buchholz , school grounds, meadow, pitfall, IX.1997, leg. Assing (cAss) ; 1 ex., Hannover, Kugelfangtrift , grassland, pitfall, V.2002, leg. Sprick (cAss) ; 1 ex., same data, but V.2001 (cAss) ; 1 ex., same data, but VIII.2002 (cAss) ; 1 ex., Braunschweig , arable land, pitfall, VII.1988 (cAss) ; 1 ex., 2 km SE Wolfenbüttel, Öselberg , 150 m, mesobrometum, pitfall, V.1998, leg. Schmidt (cAss) ; 1 ex., same data, but V.1999 (cAss) ; 1 ex., Hameln env., Grossenwieden , arable land, pitfall, VI.1987, leg. Sprick (cAss) ; 1 ex., Hameln env., Düt , pitfall, VI.1988, leg. Sprick (cAss) ; 1 ex., same data, but V.1988 (cAss) ; 2 exs., Süntel, Rannenberg , arable land, limestone, pitfall, V.1987, leg. Sprick (cAss) ; 1 ex., same data, but VI.1987 (cAss) ; 4 exs., S Hildesheim, Steinberg , mesobrometum, pitfall, V.1997, leg. Schmidt & Sprick (cAss) ; 2 exs., same data, but VI.1997 (cAss) ; 1 ex., 3 km NE Alfeld / Leine, Wernershöhe , arable land, pitfall, V.1998, leg. Schmidt (cAss) ; 3 exs., same data, but VI.1998 (cAss) ; 1 ex., Northeim env., Fredelsloh , mesobrometum, pitfall, VII.1984 (cAss) ; 1 ex., Harz, St. Andreasberg env., Jordanshöhe , 700 m, pitfall, VII.1991, leg. Vowinkel (cAss) ; 1 ex., same data, but IX.1992 (cAss) ; 1 ex., same data, but V.1993 (cAss). Mecklenburg-Vorpommern : 1 ex., SE Stralsund, Devin , pitfall, 30.V.1967, leg. Zerche ( DEI). Sachsen- Anhalt : 2 exs., Halle , S Gimritz, fallow, pitfall, 12.V.1993, leg. Teichmann (cFel, cAss) ; 1 ex., same data, but 21.VII.1993 (cFel) ; 1 ex., same data, but 4.VIII.1993 (cFel) ; 1 ex., same data, but 1.IX.1993 (cFel) ; 1 ex., same data, but 15.IX.1993 (cFel) ; 1 ex., same data, but 8.VI.1994 (cFel) ; 1 ex., same data, but 18.VII.1996 (cFel) ; 1 ex., same data, but 9.VII.1996 (cFel). Brandenburg / Berlin : 1 ex., Mehrow , 1.V.1990, leg. Arnold (cSch) .

Thüringen: 1 ex., Kyffhäuser, Rottleben env., 1.VI.1996, leg. Schülke (cSch) .

Czech Republic: 2 exs., Praha, leg. Skalitzky ( NHMW) ; 1 ex., Brandýs nad Labem , leg. Skalitzky ( NHMW) ; 1 ex., Moravia, Černá Hora [50°50'N, 15°12'E], 12.-21.VIII.1926, leg. Stolz ( NHMW) ; 1 ex., Moravia, Brno-Hády , 1.V.1966, leg. Nohel ( NHMW) ; 1 ex., Moravia, Pálava [" Pollauer Berge "] ( MNHUB) .

Slovakia: 1 ex. [with worker of Tetramorium sp. glued on the label], Trenčin, Inovec, 5.V.1929, leg. Rambousek ( NMP) .

Italy: Trentino-Alto Adige: 1 ex., Biotopi Lago Pudro ( TN) , pasture, pitfall, 16.VI.-1.VII.1989, leg. Perini (cZan); 1 ex. [with 2 workers of Formica lemani attached to the pin], Lago di Molveno , 24.VII.1976, leg. Magrini (cZan) . Piemonte: 1 ex., Biella, Sagliano Micca [ca. 45°36'N, 08°02'E], 30.VII.1958, leg. Rosa ( MHNG) . Friuli-Venezia Giulia: 1 ex., Trieste, Opcina-Basovizza [45°39'N, 13°52'E], 21.-28.IV.1921, leg. Moczarski & Scheerpeltz ( NHMW) ; 1 ex., Maniago ( PN) , Magredi , 16.IV.1977, leg. Visentini (cZan) ; 1 ex., Pozzuolo del Friuli, 5.VII.1989 (cZan) . Emilia-Romagna: 1 ex., Monteriolo near Sarsina [43°55'N 12°09'E], 800 m, 15.IV.1976, leg. Sama (cZan) ; 1 ex., locality illegible, 6.VI.1901, leg. Fiori ( MNHUB) .

Toscana: 2 exs., Castelnuovo di Garfagnana , leg. Paganetti ( DEI) ; 1 ex., Passo della Futa [44°05'N, 11°17'E], 3.XI.1962, leg. Castellini (cBor) ; 1 ex., Pratolino (FI), IV.1971, leg. Bordoni (cBor) ; 1 ex., Mte. Morello (FI), 3.V.1925, leg. Andreini (cBor) ; 1 ex., Elba [island], leg. Moczarski ( NHMW) . Marche: 1 ex., Ancona, Monte Conero [43°32'N, 13°35'E], leg. Paganetti ( DEI) . Puglia: 1 ex., Gargano , San Giovanni Rotondo, leg. Holdhaus ( NHMW) . Basilicata: 2 exs., Policoro ( MT) , 12.V.1987, leg. de Marzo (cAss). Locality ambiguous: 1 ex., " Belvedere ", 3.VI.1894, leg. Solari ( NHMW) .

Austria: Vorarlberg: 2 exs., Feldkirch ( NHMW) ; 1 ex., locality illegible, 10.V.1908 ( NHMW) . Oberösterreich: 1 ex., Linz , Lichtenberg, leg. Mader ( NHMW) . Niederösterreich / Wien: 1 ex., Wien env., Sievering [48°15'N, 16°20'E], leg. Skalitzky ( NHMW) ; 2 exs., Wien env., leg. Breit, Winkler ( NHMW) ; 1 ex., Mödling ( NHMW) ; 1 ex., Langenzersdorf , leg. Luze ( NHMW) ; 1 ex., Hainburg, Hundsheimer Berg [48°06, 16°57'E], 20.V.1942, leg. Bischoff ( MNHUB) . Locality not specified : 1 ex., " Austria ", leg. Pipitz ( NHMW) .

Hungary: 1 ex., Villany , V.1981, leg. Sieber (cWun) ; 2 exs., Bugac National Park , grassland, pitfall, IV.1983, leg. Galle (cAss) ; 1 ex., same data, but V.1983 (cAss) ; 1 ex., same data, but IX.1986 (cAss) .

Romania: 1 ex., Munţii Rodnei [47°35'N, 24°40'E], leg. Deubel ( NHMW) .

Croatia: 1 ex., Črnica ["Cernizza", 45°27'N, 13°56'E], leg. Strupi (cAss) ; 1 ex., Mljet island , 1907, leg. Moczarski ( NHMW) ; 1 ex., " Istrien ", 30.VI.1984 (cWun) .

Bosnia-Herzegovina: 1 ex., Vlasic planina, 600-1700 m, car-net, 5.V.1990, leg. Wunderle (cWun) .

Bulgaria: 1 ex., Samokov, 1911, leg. Hilf ( NHMW) .

Greece: 1 ex., S Aristi, 850 m, 1.V.1973, leg. Löbl ( MHNG) ; 1 ex., Flórina , ca. 20 km SSW Flórina, Oros Vitsi, 40°39'N, 21°23'E, N-slope, 1850-1900 m, grass and moss sifted, 22.V.2005, leg. Wunderle (cWun).

Turkey: Rize: 1 ex., Cağlayan D. river valley, 1800-1900 m, 23.VI.1998, leg. Solodovnikov (cAss) . Mersin: 1 ex., Mut-Karaman, Sertavul Geçidi , 36°55'N, 33°16'E, 1570 m, 5.V.2004, leg. Brachat & Meybohm (cAss) ; 1 ex., Kırobası-Güzeloluk , 14 km W Güzeloluk, 36°45'N, 33°58'E, 1430 m, 8.V.2004, leg. Brachat & Meybohm (cAss) .

Armenia: 1 ex., Yerevan, 1.VI.1952 (cAss) .

Georgia: 1 ex., Khashuri ["Michailowo am Suramgebirge"; 41°59'N, 43°35'E], leg. Leder ( NHMW) .

Locality not specified or not identified: 1 ex., 13.VII.1993 ( ZMH) ; 1 ex., ["Hungar"] ( DEI) ; 1 ex., "Hungaria" ( NHMW) ; 2 exs. ( DEI) .

Diagnosis:

Extremely variable species, 1.8-3.4 mm; RL: 0.85-1.4 mm. Usual coloration: head and abdomen, except for the posterior margins of the segments and the apex, dark brown to blackish; pronotum brown to dark brown; elytra yellowish brown to brown; legs yellowish; antennae yellowish to yellowish brown. Occasionally, the whole body may be considerable darker or paler.

Head transverse (HW/HL: 1.10-1.25 [sic]), of transversely rectangular to distinctly wedge-like shape (i. e. lateral margins subparallel to distinctly diverging in dorsal view); puncturation very fine, often barely noticeable; microreticulation very shallow to pronounced. Eyes of very variable shape and size, in small-eyed specimens weakly protruding from lateral contours of head and little more than half the length of postocular region in dorsal view, in large-eyed specimens distinctly prominent and approximately as long as postocular region or nearly so. Postgenal carina distinct and longer than in the preceding species, extending below eyes or nearly so. Antenna variable; usual morphology: antennomere III approximately twice as long as wide; IV approximately 1.5 times as wide as long; V-IX increasingly transverse and of gradually increasing width; IX and X approximately twice as wide as long.

Pronotum of remarkably variable shape, slightly to distinctly wider than head (PW/HW: 1.12-1.25) and weakly to distinctly transverse (PW/PL: 1.05-1.30 [sic]); maximal width approximately in the middle, in posterior half only weakly tapering, i. e. posterior margin broader than anterior margin; puncturation similar to that of head; microreticulation pronounced to practically absent.

Elytra dimorphic, without sexual dimorphism; in macropterous morph slightly, in submacropterous morph distinctly shorter than pronotum (EL/PL: 0.82-0.95); puncturation as fine as or even finer than that of head and pronotum; microreticulation pronounced to practically absent. Hind wings fully developed or of somewhat reduced length.

Abdomen subparallel; tergite IV without anterior impression; puncturation fine and sparse, barely noticeable; microsculpture composed of isodiametric meshes everywhere and usually shallow, more distinct on tergites VII-VIII than on tergites III-VI; posterior margin of tergite VII with narrow palisade fringe; tergites VII-VIII without sexual dimorphism; posterior margin of tergite VIII broadly convex in both sexes.

: posterior margin of sternite VIII strongly convex; median lobe of aedeagus of variable size; ML: 0.32-0.38 mm; shape highly distinctive; internal sac with very long flagellum ( Figs 59-60 ).

: posterior margin of sternite VIII moderately convex, in the middle not concave; spermatheca with twisted duct ( Fig. 61 ).

Intraspecific variation:

The species is even more variable than its congeners, this enormous variation affecting especially size, coloration, the microsculpture of the forebody, eye size, the shape of the head and the pronotum, the length of the elytra and the hind wings, and the size of the aedeagus. Owing to this extreme variability, non-average specimens may be difficult to attribute to this species, or even to the genus. However, a reliable identification is always possible based on the highly distinctive primary sexual characters.

Distribution and bionomics:

Based on the revised material, A. gracilenta is a widespread Ponto-Mediterranean species, its distribution ranging from southern Turkey, Armenia, and Georgia in the southeast and east to northwest Spain, western France, and the south of Great Britain in the west and northwest, and to southern Scandinavia in the north ( Map 5 ). It is here recorded from Spain, Croatia, Bosnia- Herzegovina, Bulgaria, Armenia, and Georgia for the first time. LUNDBERG (1995) reports the species from all Scandinavian countries; in Sweden, it is confined to the south. According to BAUDI DI SELVE (1870), it is present also in Cyprus, but this record requires confirmation. The species was only recently recorded from Greece and Turkey for the first time ( ASSING 2004, 2005). In Germany, it has been reported from all regions ( EISINGER 2006; KÖHLER & KLAUSNITZER 1998; material examined). In Italy, it is known from the north and south of the mainland, as well as from Sicily ( ZANETTI 1995). For additional literature records see ALLEN (1991), BENICK & LOHSE (1959), EISINGER (2006, KACHE & ZUCCHI (1993), KOCH (1968), KÖHLER & KLAUSNITZER (1998), KROKER & RENNER (1983), KORGE (1956, 1960, 1989), LINKE (1907), MÜLLER (1926), PEEZ & KAHLEN (1977), RENNER (2001), SAINTE-CLAIRE DEVILLE (1920), SCHOLZE & JUNG (1994), SEGERS (1986), STUMPF (1998), SZUJECKI (1968), TRONQUET (2006), VÁSÁRHELYI (1985), VOGEL (1978), and VOGEL & DUNGER (1980). Older records may be doubtful because the species was confounded with A. egregia by various authors (e.g. WÖRNDLE (1950).

Although records are comparatively rare, the species has been collected in a wide range of - usually unforested - biotopes, especially with pitfall traps (see material examined; ASSING 1988, 1992, 1994a, 1994b, 2001b; FELDMANN & LÜCKMANN 1998; KORGE 1960; KROKER & RENNER 1983; STUMPF 1998; ZERCHE 1980), occasionally also with car-nets (material examined), sweep-nets (material examined; ALLEN 1991; EISINGER 2006; KOCH 1968; VOGEL & DUNGER 1980), or by sifting (material examined; LINKE 1907; PEEZ & KAHLEN 1977). It has been found in xerothermous limestone grassland (material examined; ASSING 1994a, 2001b; FELDMANN & LÜCKMANN 1998), subalpine grassland (material examined; ASSING 2005), meadows (material examined; KOCH 1968, VOGEL & DUNGER 1980), pastures (material examined), arable land (material examined), fallow land (material examined), river banks (material examined), the leaf litter of old trees and the roots of grass in a sandpit ( PEEZ & KAHLEN 1977), sparsely vegetated heathlands on sandy soils (material examined; KORGE 1960), a chalkpit ( ALLEN 1991), a sparsely vegetated dump ( STUMPF 1998), abandoned railroad tracks with ruderal vegetation ( KORGE 1989), nests of ants ( Formica lemani , Tetramorium sp. , Lasius flavus ) (material examined; KORGE 1956), compost ( KOCH 1968), flood debris ( LINKE 1907), and various urban or suburban habitats such as gardens, lawns, and greens ( ALLEN 1991; ASSING 1988, 1992; KACHE & ZUCCHI 1993). The wide range of different habitats, as well as the fact that the species is collected rarely, accidentally, and usually in singles or very small numbers suggest that the true reproduction habitat is cryptic and unknown. While the records in Central Europe are generally from lower to intermediate elevations, the upper end of the known altitude range is at 1800-1900 m in Greece and Turkey (material examined).

The examined material was collected during spring and summer, from April through September, with a maximum in May and June. One specimen was found also in November ( Fig. 63 ). Since the habitat is probably subterranean, the ground and above-ground catches (mainly assessed by pitfall trapping and flight records) reflect dispersal activity. Flight records are confined to May and June. Teneral beetles were not observed.

Figures

Map 5: Distribution of Alevonota gracilenta (ERICHSON), based on examined records (filled circles) and selected literature records (open circles).

Figs 59-62: Alevonota gracilenta (ERICHSON) (59-61) and Tropimenelytron mirabilis (EPPELSHEIM) (62): median lobe of aedeagus in lateral view of male from Turkey (59) and of nanistic male from Italy (60); spermatheca (61); forebody (62). Scale bars: 0.1 mm.

Fig. 63: Alevonota gracilenta (ERICHSON): Seasonal distribution of the examined specimens (black bars) and samples/records (grey bars).