Anthobium fusculum (Erichson, 1839)

(Figs. 1, 6, 8, 10, 12, 14, 16, 18, 20, 22, 24, 26, 28–32, 38–43, 44)

Lathrimaeum fusculum Erichson, 1839: 626

Lathrimaeum fusculum: Erichson, 1840: 870; Redtenbacher, 1849: 829; Kelch 1852: 8; Fairmaire and Laboulbène, 1856: 637; Kraatz, 1857: 947; Lentz 1857: 57, 1879: 20; Gutfleisch & Bose 1859: 179; Clasen 1861: 188; Hochhuth 1862: 103; Preller 1862: 48; Stierlin & Gautard 1867: 106; Wilken 1867: 46; Fauvel, 1871: 120 [= 1872: 94]; Letzner 1871: 99, 1886: 142; Bertolini 1872 -1878: 242; Brüggemann 1873: 475; Redtenbacher, 1874: 268; Kellner 1876: 58; Seidlitz, 1875: 239; Dalla Torre 1879: 73; Mulsant and Rey, 1880: 131; Westhoff 1881: 98; Cornelius 1884: 25; Kittel 1884: 170; Cherkunov 1889: 164; Favre 1890: 129; Fauconnet 1892: 121; Ganglbauer, 1895: 722; Klima 1902: 71; Koltze 1901: 60; Stierlin 1900: 407; Fauconnet & Viturat 1903 -1904: 251; Weber 1903: 38; Luze 1905: 54, 58, 67; Stierlin 1906: 179; Linke 1907: 11; Jakobson 1908: 459; Reitter, 1909: 188; Bernhauer and Schubert, 1910: 64; Gerhardt 1910: 53; Prossen 1910: 174;

Röttgen 1911: 71; Łomnicki 1913: 65; Tenenbaum 1913: 17; Johansen 1914: 596; Ammann & Knabl 1922: 61; Porta

1926: 26; Fleischer 1927 -1930: 104; Zirk 1928: 6; Portevin, 1929: 442; Nowotny & Polentz 1933: 14; Rapp 1933 -1935:

223; Scheerpeltz, 1933: 1055, 1961: 79, 1968: 13; Sainte-Claire Deville 1935: 83; Walles 1936: 176; Borchert 1937: 150,

1938: 22; Lysholm 1937: 154; Hölzel 1946: 71; Lohse 1948: 199; 1964: 52; Palm 1948: 100, 102; Wörndle 1950: 127;

Borchert 1951: 70; Hansen 1952: 79, 1964: 98; Horion 1951: 109, 1961: 243, 1963: 107; Strand 1953: 65; Porta 1959: 49;

Friden 1960: 30; Kirschenblat 1965: 163; Koch 1968: 91, 1989: 229; Franz 1970: 286; Heiss 1971: 37; Medvedev &

Sosnina 1973: 825; Peez & Kahlen 1977: 129; Coiffait, 1978: 274; Focarile 1978: 27, 1981: 35; Bercio & Folwaczny

1979: 98; Burakowski et al. 1979: 55; Kofler 1980: 122; Uhlig & Zerche 1981: 150; Berstorf et al 1982: 24; Korge 1989:

30, 1991: 284, 2005: 4; Toth 1982: 89; Terlutter 1984: 27; Zanetti, 1987: 297; Schatz et al. 1990: 205; Schülke et al. 1992:

167; Meybohm 1994: 48; Gürlich et al. 1995: 26; Kahlen 1995: 21, 2011: 180; Zanetti, 1995: 8; Böhme 1996: 46; Schatz

1996: 257; Scholze 1999: 380, 2009: 229; Geiser 2001: 188 <?!> Lathrimaeum? fusculum: Baudi di Selve 1870: 401 Lathrymaeum [sic!] fusculum: Fröhlich 1897: 44 Lathrimaeum (Lathrimaeum) fusculum: Scheerpeltz, 1961: 79; Tichomirova 1973: 139 Olophrum fusculum: Redtenbacher, 1857: 249 Anthobium fusculum: Silfverberg 1979: 18; Rutanen 1983: 213; Lundberg 1988: 83; Zerche 1991: 314; Stoltze & Pihl 1998:

109; Kubisz et al 1998: 234; Derunkov 1999: 8; Direktoratet for Naturforvaltning 1999: 85; Herman 2001: 232; Andersen

& Olberg 2003: 61; Assing 2004: 676, 2005: 58, 2006: 226, 2007: 6, 2013: 106; Scholze et al. 2004: 275; Andersen &

Hanssen 2005: 3505; Barndt 2005: 81; Bohač 2005: 436, 660, 2016: 4; Gutowski et al. 2006: 129; Schatz 2007: 90;

Schülke 2007: 139; Strobl 2007: 24; Bäse 2008: 130; Gontarenko 2008: 22; Szujecki 2008: 44, 116; Esser 2009: 27, 2017:

12; Semenov 2009: 24; Piryugin 2010: 281; Derunkov & Bacal 2011: 106; Samin et al. 2011: 1236; Solodovnikov 2012:

67; Zanetti 2012: 56; Vogel 2013: 37; Petrenko et al. 2014: 6; Odegaard et al. 2015: 21; Hlavač et al., 2016: 2 <?!> Anthobium fusculum: Bacal et al. 2013: 423 Anthobium (Anthobium) fusculum: Gamarra & Outerello 2008: 212; Smetana 2004: 238; Schülke & Smetana 2015: 306 Lathrimaeum melanochromum Iablokoff-Khnzorian, 1961: 140 Lathrimaeum (Lathrimaeum) melanochromum: Tichomirova, 1973: 139 Lathrimaeum melanochromum: Herman 2001: 235 Anthobium (Anthobium) melanochromum: Smetana 2004: 239; Lathrimaeum mesasiaticum Kirschenblat, 1961: 363 syn. nov. Lathrimaeum (Lathrimaeum) mesasiaticum: Tichomirova 1973: 139 Anthobium mesasiaticum: Herman, 2001: 235; Hlavač et al., 2016: 2 Anthobium (Anthobium) mesasiaticum: Smetana, 2004: 239; Schülke & Smetana, 2015: 306 Olophrum kondarense Iablokoff-Khnzorian, 1964: 157 Olophrum kondarense: Herman 2001: 349; Smetana 2004: 249 Lathrimaeum kashmiricum Coiffait, 1982: 150 Anthobium kashmiricum: Herman, 2001: 233 Anthobium (Prionothorax) kashmiricum: Smetana, 2004: 239; Schülke & Smetana, 2015: 307 Deliphrosoma ivanovi Shavrin, 2012: 17 Deliphrosoma kirgizicum Shavrin, 2012: 19 Deliphrosoma sp.: Shavrin, 2012: 19

Type material examined: Holotype of Lathrimaeum mesasiaticum Kirschenblat 1961 ♀ [specimen without right antennomeres 3–11 and left antennomeres 10–11]: TAJIKISTAN: ‘Cтaлинaбaд [Stalinabad (= Dushanbe)] | Taджик. [Tajik[istan].] | Киpичeнкo [Kirichenko] 28 X [handwritten] 943’ <rectangular label, printed>, ‘ Lathrimaeum | mesasiaticum sp. n. | Type | J Kirschenblatt det.’ <rectangular label, handwritten in black Indian ink>, ‘ HOLOTYPE | Lathrimaeum | mesasiaticum | Kirschenblatt J.D. det. 1961’ <red rectangular label, printed>, ‘ Anthobium | fusculum (Erichson, 1839) | Shavrin A.V. det. 2016’ <rectangular label, printed> (ZIN).

Paratypes of L. mesasiaticum Kirschenblat 1961: 1 ♀ [apical part of the abdomen is glued under the specimen]: TAJIKISTAN: ‘Пeнджикcит [Пeнджикeнт] | 10.XII. [1]943. | Киpичeнкo. [Pendzhiksit [Panjakent] 10.XII.1943 Kirichenko.]’ <rectangular label, handwritten>, ‘ Lathrimaeum | mesasiaticum sp.n. | JKirschenblatt det.’ <rectangular label, handwritten in black Indian ink> (ZIN); 1 ♀ [specimen without right antennomeres 3–11, left elytron and left middle leg]: UZBEKISTAN: ‘cт Гoлoднaя Cтeпь | Cp[eднe-] Aз[иaтcкaя].ж[eлeзнaя].д[opoгa]., Xoдж[eнтcкий].y[eзд] | Г Якoбcoн 5.[handwritten]IV.[19]03 [Golodnaya Step`st. Middle-Asian Railway, Khodzhentskiy uezd [Samarkand Area of Turkestan Territory of the Russian Empire] G. Yakobson 5.IV.03]’ <rectangular label, printed>, ‘ Lathrimaeum | mesasiaticum sp. n. | J Kirschenblatt det.’ <rectangular label, handwritten in black Indian ink> (ZIN). Paratypes with additional labels: ‘ PARATYPE | Lathrimaeum | mesasiaticum | Kirschenblatt J.D. det. 1961’ <red rectangular label, printed by the senior author>, ‘ Anthobium | fusculum (Erichson, 1839) | Shavrin A.V. det. 2016’ <rectangular label, printed>.

Additional material: UKRAINE: 2 ♂: Odessa obl., Berekhovka Distr., vic. Raukhovka. 0 1.11.2008. A.V. Gontarenko (CS); RUSSIA: 1 ♂, 2 ♀: Volgograd Area, Elton Lake. 01- 10.08.2006. K. Makarov & A. Matalin (MSPU); 1 ♂: Rostov Area, Alexandrovka vill., Leninskiy Forest. 17.03.1977. E. Khachikov (CK); TURKEY: Niğde, Ulukişla, 5 km SW Kilan, Güney Mountains, 37°26'42''N, 34°25'55''E. 13.04.2018. S. Anlaş (CSA); IRAN: 1 ♀: Kopet-Dag, Khugdi env. 21.04.2005. lag3 [without collector`s name] (CS); KAZAKHSTAN: 2 ♂♂, 1 ♀: West Tian Shan, Chimkent Reg., Aksu-Dzhabagly State Nature Reserve, Unken-Kaindi, 70°35'N 42°23'E. 11.06.1999. 2200 m a.s.l., in bear dung. J. Cooter leg. (CR); 6 ♂♂, 6 ♀♀: same Natural Reserve, Kalbaidjailau, 70°42'N 42°25'E. 16.06.1999. 2500 m a.s.l., grubbing—snow melt. J. Cooter leg. (CR); 9 ♂, 6 ♀: same Natural Reserve, Ulken Kaindi Pass, 70°39'N 42°22'30''E. 14.06.1999. Ca 3000 m a.s.l., at snow melt. J. Cooter leg. (CR); 1 ♂: Karatau Mts. (plateau), 36 km W Karatau Mt., N 43°10'09'' E 69°56'32''. 1130 m a.s.l., 0 6.05.2015. A.S. Prosvirov (CS); KYRGYZSTAN: 2 ♂♂, 4 ♀♀: Kyrgyzskiy Alatau, Ala Artcha. 1650 m. 29.10.2000 (CSC); TAJIKISTAN: 1 ♀: Varzob District, Khodja obi Garm. 2000 m a.s.l. 21- 24.03.2015. A. Barševskis (CS); 1 ♀: Kondar Pass, Kvak. 2200 m a.s.l., near snow field. 10.05.1988. V.G. Shilenkov (CS); PAKISTAN: 1 ♂, 1 ♀: Swat, Malam Jabba. 2500-2600 m a.s.l. 18.05.1983. Besuchet & Löbl (MHNG); WITHOUT EXACT LOCALITY: 1 specimen [without apical segments of abdomen]: ‘34’ (ZMM; collection of A.L. Tichomirova).

Redescription. Measurements (n=50): HW: 0.55–0.64, HL: 0.30–0.41; AL: 0.77; OL: 0.12–0.17; PL: 0.47–0.57; PW: 0.80–1.02; ESL: 1.14–1.25; EW: 1.05–1.25; AW: 1.05–1.17; MTbL: 0.57, MTrL: 0.32 (MTrL 1–4: 0.20; MTrL 5: 0.12); AedL: 0.31–0.52; TL: 2.10–3.40 (holotype of Lathrimaeum mesasiaticum: 2.70; holotypes of Lathrimaeum cachemiricum and Deliphrosoma ivanovi: 2.80; holotype of Deliphrosoma kirgizicum: 3.40).

FIGURES 14–27. Body parts of Anthobium fusculum group (14, 16, 18, 20, 22, 24, 26— A. fusculum, 15, 17, 19, 21, 23, 25, 27— A. monticola): 14–15—ventral aspect of head, 16–17—ventral aspect of prothorax, 18–19—scutellum (dorsal view), 20–21—ventral aspect of elytron, 22–23—ventral aspect of mesothorax, 24–25—ventral aspect of metathorax (without metanepisternum), 26–27—metendosternite (ventral view). Scale bar: 0.1 mm.

Body elongate, subconvex. Body and antennae brown to dark brown or black, usually with lateral portions of pronotum and elytra yellow-brown to reddish-brown; mouthparts and legs yellow to reddish-yellow; basal portion of antennomere 1 and tarsi (in some specimens) yellow. Body shiny, without microsculpture on elytra; head between supra-antennal elevation and anterior margin of eye with distinct diagonal cellular-shaped meshes, median part of head without or with transverse microsculpture beween punctures, more coarse in posterior portion between ocelli, infraorbital ridges of head with dense, small and coarse microsculpture (some specimens with very fine microsculpture on middle portion of head except of posterior part of infraorbital ridges with distinct transverse meshes between punctures); middle part of neck and abdominal tergites with distinct isodiametric microsculpture; pronotum with indistinct to distinct transverse microsculpture stronger in middle of anterior third (some specimens without microsculpture here); scutellum with indistinct to distinct transverse microsculpture or without it. Head with small or moderately large and deep punctation, markedly denser on posterior portion of infraorbital ridges; middle part of neck with dense smooth punctation; pronotum with irregular punctation as that on head, denser in middle, with interspaces between punctures as long as about diameter of two-four nearest punctures, sometimes sparser in mediobasal and lateral portions; scutellum with several small smooth to deep punctures; punctation of elytra as that in middle part of pronotum, but distinctly denser and deeper and/or smaller on parascutellar portion and along suture in some specimens, with interspaces between punctures in middle as long as diameter of one or two nearest punctures, on each elytron not forming longitudinal rows of punctures and more or less irregularly scattered, or with very indistinct three to four vague and tangled longitudinal rows along suture; abdominal tergites without or with sparse and very indistinct small punctures. Body glabrous; clypeus with several long tactile setae; anterior part of frons with several irregular small setae; middle part of pronotum and each elytron with irregular, very sparse, small and erect setae; abdominal tergites with regular sparse and small setae. Habitus as in Fig. 1.

Head 1.5–1.8 times as wide as long; middle portion of head slightly flattened or irregularly elevated, with indistinct to distinct semicircular impression between eyes at posterior third, usually with distinct and deep elongate grooves in front of ocelli, reaching middle or anterior third of eye; postocular ridges smooth and indistinct in some specimens; anterior portion between antennal insertion and anterior margin of eye with moderately deep semicircular notch. Ocelli large, situated at level or slightly behind of level of postocular ridges; distance between ocelli slightly longer than distance between ocellus and posterior margin of eye. Labrum with widely rounded membranous latero-apical portions, moderately deep apical emargination and long thin basolateral projections (Fig. 6). Mandibles short, with narrow basal and elongate apical portions, with indistinct small tooth in middle on right mandible (Fig. 8). Labium and labial palpomeres moderately narrow, submentum small, elongate (Fig. 12). Apical segment of maxillary palp slightly more than twice as long as preceding segment (Fig. 11). Gular sutures with widely rounded apical parts, narrowly diverging posteriad; shortest distance between sutures located at level posterior third of eyes (Fig. 14). Antenna moderately short, slightly exceeding shoulders of elytra when reclined; length × width of antennomeres (holotype of L. mesasiaticum): 1: 0.11 × 0.07; 2: 0.08 × 0.06; 3: 0.08 × 0.04; 4: 0.07 × 0.04; 5–7: 0.06 × 0.04; 8–9: 0.05 × 0.05; 10: 0.05 × 0.06; 11: 0.10 × 0.06.

Pronotum 1.7 times as wide as long, 1.4–1.5 times as wide as head, widest at or about middle, evenly or slightly more narrowed posteriad than anteriad; apical margin distinctly shorter than posterior margin, with straight or slightly rounded apical margin; anterior angles widely rounded, not or slightly protruded anteriad; posterior angles widely rounded; lateral edges of pronotum with irregular and small, flattened crenulation; pronotum with very wide elevated middle portion, with wide and deep longitudinal impression (indistinct in some), with pair of indistinct to moderately deep semioval impression on mediobasal third; lateral portions slightly explanate, each with deep oval pit in front of middle. Prothorax (Fig. 16) with elongate intercoxal process and moderately wide slightly protruded mesosternal processes. Mesothorax (Fig. 22) transverse, narrow in middle, with thin and acute intercoxal process reaching posterior margin of mesocoxae. Scutellum (Fig. 18) with widely rounded apex. Metathorax (Fig. 24) narrow, with wide and deep intercoxal cavities and narrow intercoxal process. Metendosternite (Fig. 26) with strongly reduced paired medioapical protrusions.

Elytra about as long as wide, long, markedly more than twice as long as pronotum, more or less subparallelsided and indistincly widened apicad from middle, reaching apical margin of tergite IV or V, with straight or slightly rounded apical margins, truncated at suture; shoulders of elytra widely rounded, not protruded apicad (Fig. 20); lateral portions of elytra narrow, slightly explanate, latero-apical margins with small, irregular and smooth crenulation; each elytron with indistinct transverse elevations between punctures in parascutellar portions or with distinct transverse and diagonal elevations, significantly coarser in mediobasal portions in some.

Legs moderately long and slender, tibiae covered by elongate setae, distinctly stronger on lateral margin, with a few strong setae on apical margins near apex; first four protarsomeres of both sexes dilated; metatarsus less than half as long as metatibia.

Abdomen about as wide as elytra or slightly narrower, with pair of rounded tomentose wing-folding spots in middle of tergite V.

Male. Apical margin of abdominal tergite VIII and sternite VIII slightly emarginate (Fig. 2 8–29). Aedeagus wide and suboval (Fig. 38–43; Shavrin 2013: Fig. 1–6), with widely rounded apex; parameres wide, slightly exceeding apex of median lobe, with four short apical setae; internal sac variable, consisting of numerous acute thorns, spiralled in basal part in some specimens. Aedeagus laterally see in Hlaváč et al. (2016: 2).

Female. Apical margin of abdominal tergite VIII vaguely sinuate (Fig. 30). Apical margin of abdominal sternite VIII rounded (Fig. 31). Genital segment as in Fig. 32.

Comparative notes. Anthobium fusculum differs from the remaining species of the fusculum group by the dark brown to black coloration of the body, by the shape of the suboval aedeagus, wide parameres, and by the structure of the internal sac.

Distribution. Anthobium fusculum is widely distributed in the western Palaearctic Region (Fig. 44), from the middle (Austria, Czech Rebublic, France, Germany, Italy, Switzerland), eastern (Belarus, Poland, Russia, Ukraine) and northern Europe (Denmark, Finland, Norway, Sweden) to Caucasus (Armenia), Middle East (Turkey, Iran) and Central Asia (Kasakhstan, Kyrgyzstan, Tajikistan, Uzbekistan, Pakistan and India (Kashmir)).

The record of the species from Spain (Smetana 2004; Schülke & Smetana 2015) is most likely based on the result of old misidentification (see also Gamarra & Outerelo 2008).

Bionomics. According to published data, A. fusculum can be regarded as hygrophilous species; it was noted as psammophilous and phytodetriticolous (Uhlig & Zerche 1981; Koch 1989; Vogel 2013), stenotopic ripicolous alpine (Schatz 2007) and riparian species (Andersen & Hanssen 2005). The species is recorded from intermediate elevations beginning from about 100 m a.s.l. to high elevations about 3000 m a.s.l. It inhabits riparian and forest habitats, where it can be found under stones, mosses, litter and debris. Some specimens were observed under bark of trees, occasionally taken from bear dung, mushrooms, etc. Many populations in the Alps and some specimens from Aksu-Dzhabagly State Nature Reserve (Kazakhstan) and Kondar Pass (Tajikistan) were collected near the snow. The beetles have been collected from March to midwinter in the European part of its distribution where it was noted as winter active species (Schatz 2007).

Remarks. Lathrimaeum fusculum was originally described from unspecified number of types from “Mark Brandenburg ”, northern Germany. More than 150 years from the description, the species has been many times recorded from Europe, included in the regional European keys (see above) and it was redescribed several times (Fairmaire & Laboulbène 1849, Luze 1905, Zanetti 1987, etc.). Zerche (1991) noted that he studied specimens of A. fusculum from Uzbekistan (Chimgan) that in habitus were somewhat different from European specimens and remotely looked like some species of the genus Deliphrosoma Reitter, 1909. The external variability of the species was a long-standing base for descriptions of new taxa, which were later synonymised with A. fusculum: Assing (2005) studied holotypes of L. melanochromum Iablokoff-Khnzorian, 1961 (described from Armenia) and Olophrum kondarense Iablokoff-Khnzorian, 1964 (described from Tajikistan), and synonymised them with A. fusculum; Shavrin (2013) studied holotype of L. kashmiricum Coiffait, 1982 (described from Kashmir, India), and, together with Deliphrosoma ivanovi Shavrin, 2012 and D. kirgizicum Shavrin, 2012, which were described from the Central Asia, synonymised them with A. fusculum. Another new synonymy is presented here. Lathrimaeum mesasiaticum Kirschenblat, 1961 was described from the holotype and two “cotypes” (all females) from “Stalinabad” (= Dushanbe; Tajikistan), “Pendzhiksit” (=Panjakent, Tajikistan) and “Golodnaya Step`” (Uzbekistan). The first author compared this species with L. fusculum and noted that L. mesasiaticum “...easily differs [from L. fusculum] by more long elytra, more gentle and denser punctation of pronotum and many other characters [translated from Russian]”. The first author studied these types in ZIN and found no morhological differences between these and other specimens of A. fusculum. The photograph of habitus (Fig. 7) of A. fusculum in Boháč (2016) shows Olophrum fuscum (Gravenhorst, 1806).

The species is reported from Pakistan for the first time.