Unexpected diversity of Lacon Laporte, 1838 (Coleoptera: Elateridae: Agrypninae) in the Levant: revised species concepts, new species, and an identification key Author Kundrata, Robin Author Mertlik, Josef Author Németh, Tamás text Zootaxa 2019 2019-10-03 4679 3 401 449 journal article 25321 10.11646/zootaxa.4679.3.1 c6d3d19c-627e-425f-8824-193a7213994a 1175-5326 3471061 4F71C33E-8B87-4D9E-ACAD-CED563A7033B Lacon drusus ( Marseul, 1870 ) ( Figs 100–118 , 190, 191 , 194 ) Adelocera drusa Marseul, 1870: 380 . Adelocera bruleriei Desbrochers des Loges, 1875: 37 ; synonymized by Hayek (1973: 63) . Lacon drusus : Hayek (1973: 63) [as drusa ]. Type material . Lectotype of L. drusa , male, „ Adelocera drusa , Kar-Eliz [ Lebanon : Kab Elias] [further data unreadable]; Type, Drusa Mars. type; Museum Paris, coll. de Marseul 1890, Liban, Lectotype Adelocera drusa Marseul. C.M.F. von Hayek det. 1973“ ( MNHN ). Lectotype of L. bruleriei , female, „ Brulerii [sic!] ( ♂ ?) m., Type, probable ♀ , CMFH 1968, Lectotype Adelocera bruleriei Desbr., C.M.F. von Hayek 1972 “ ( MNHN ). Type locality . Lebanon : Kab Elias . Other material examined . Israel , 2 males , „Jaffa, Collect Godart, Muséum Paris, Coll. A. Argod, 1931 [one with additional label: graeca Cand ]“ ( MNHN ); 1 male , „Israel, N. Golan, 1350 m , Mt. Hermon, 10.V.1996 , D. Gianasso [leg.], Lacon drusa (Mars.) det. Platia 2009“ ( PCGP ); 1 male , „ ♂ , Bruleriei Desbr. , Naplouse, Desbrochers“ ( SDEI ); 1 female , „Israel, Hula reserve, 1.VII.1993 , V. Chikatunov [leg.], Lacon graecus (Cand.) det. Preiss 2000, Lacon drusum (Baudi) det. Preiss, 2004“ ( SMNH ); 1 female , „Israel, N. Oren, 25.02.[19]97, V. Chikatunov leg., Lacon graecus (Candeze) det. Preiss 2000, Lacon drusum (Baudi [sic!]) det. Preiss 2004” ( SMNH ); Lebanon , 1 male , „Liban, Peyron, Drusa Mars , Museum Paris, Coll. L. Bedel, 1922, Graeca Cand. “ ( MNHN ); 1 male , „Liban — Jbail, Laklouk, m. 1400, 8/ 10.V.2000 , G. Sama leg., Lacon graecus (Cd.) det. G. Platia 2000 “ ( PCGP ); 1 female , „Liban, Museum Paris, Coll. Abeille de Perrin 1919, ♀ , brulerei [sic!] Desbr., C. M. F. von Hayek det. 196 [sic!]” ( MNHN ); 1 female , “Liban central, 329, Museum Paris, Collection Léon Fairmaire 1906, Adelocera drusa “ ( MNHN ); 1 female , „Lebanon, Northern gov., Tannourine env., 2 km N Harissa, Tannourine Cedars Nat. Reserve, swept & hand-collected, 34°12‘34“N , 35°55‘45“E , 1750 m , 1.V.2017 , leg. A. Kotán, P. Nemes & T. Németh (No. 30)“ ( HNHM ); 1 male , 4 females , „Lebanon, Northern gov., Tannourine env., 2 km N Harissa, Tannourine Cedars Nat. Reserve, swept & singled, 34°12‘34“N , 35°55‘45“E , 1750 m , 26.VI.2016 , leg. A. Kotán, P. Nemes & T. Németh (No. 24)“ ( HNHM ); 1 male , 1 female , „Lebanon, Northern gov., Ehden, S to Horsh Ehden Natural Reserve, from beneath stones, 34°18‘33“N , 35°59‘14“E , 1895 m , 4.V.2017 , leg. M. Boustani, A. Kotán, P. Nemes, T. Németh & C. Tanios (No. 36)“ ( HNHM ); 1 female , „Lebanon, Northern gov., Tannourine env., 2 km N Harissa, Tannourine Cedars Nat. Reserve, swept & singled, 34°12‘34“N , 35°55‘45“E , 1750 m , 18.VI.2016 , leg. A. Kotán, P. Nemes & T. Németh (No. 6)“ ( HNHM ); 1 female , „Lebanon, Northern gov., Tannourine env., 2 km N Harissa, Tannourine Cedars Nat. Reserve, swept & singled, 34°12‘34“N , 35°55‘45“E , 1750 m , 16.VI.2016 , leg. A. Kotán, P. Nemes & T. Németh (No. 1)“ ( HNHM ); 1 female , „Lebanon, Northern gov., Bcharre env., 1 km E Ariz, Horsh Arz el-Rab, ancient Cedrus forest, swept & hand collected, 34°14.494‘N , 36°2.884‘E , 1900 m , 9.V.2017 , leg. A. Kotán, P. Nemes, T. Németh & C. Tanios (No. 45)“ ( HNHM ); 1 male , „Lebanon, Northern gov., Tannourine env., 2 km N Harissa, Tannourine Cedars Nat. Reserve, swept & singled, 34°12‘34“N , 35°55‘45“E , 1750 m , 19.VI.2016 , leg. A. Kotán, P. Nemes & T. Németh (No. 10)“ ( HNHM ); 1 female , „Lebanon, Northern gov., Tanou-rine env., 2 km N Harissa, Tanourine [sic!] Cedars Natural Reserve, 34°12’34”N , 35°55’45”E , 1750 m , from beneath stones, 23.V.2015 , leg. András Márkus & Tamás Németh” ( HNHM ); 1 male , “Libanon: Dj. Sannin (b. Faqra), 1900–2100 m , 17/ 31. V.1999 , Heinz leg., CCW, Lacon graecus (Cand.) det. Platia 2013“ ( PCGP ); 1 male , „Lebanon-Bcharré des Cedres, 1960 m , 12.VI.1999 , P. Rapuzzi, Lacon graeca (Cd.) det. Platia 000“ ( PCGP ); 1 male , „Libano c. Byblos, Qartaba, 6.VI. [19]93, P. Rapuzzi, Lacon graecus (Cd.) det. Platia 000“ ( PCGP ); 1 male , „Libano — Chouf, Djebel Barouk m. 1700, Passo W. Kefraya, 4/ 7.VI. [19]99, G. Sama leg., Lacon graecus (Cd.) det. Platia 000“ ( PCGP ); 1 ex. , „Lebanon, Akkar gov., Fnaidek, Quercus forest, hand collected, 34°28‘19“N , 36°12‘28“E , 3.VIII.2018 , leg. P. Nemes, T. Németh & C. Tannios (No. 84)“ ( PCRK ); Syria , 1 male , 2 females , „Syria, muh. Dimashq , 4 km NE Bloudan, N 33°45.955, E 36°10.033, 2240 m, 30.IV.2005 , leg. N. Rahmé, A. Márkus, A. Kotán & A. Podlussány, Lacon drusa (Mars.) det. Platia 2010 “ ( HNHM ); 2 males , „BLD, Museum Paris, Coll. Abeille de Perrin 1919 [one with additional label: Adelocera graeca Cand. ]” ( MNHN ); 2 females , “Syria, muh. Dimashq , 4 km NE Bloudan, N 33°45.955, E 36°10.033, 2240 m, 28.IV.2005 , leg. N. Rahmé, A. Márkus, A. Kotán & A. Podlussány, Lacon drusa (Mars.) det. Platia 2010 “ ( HNHM ); 2 males , „Syria, pr. Damascus , Jabal Lubnan ash Sharqi, 1700 m , Ayn an Nusur, 23.V.2004 , N. Rahmé, L. Nádai & K. Székely [leg.], Lacon drusa (Mars.) det. Platia 2010 “ ( HNHM ); 1 male , „Syria — W. Damascus , Bludan, m. 1700–2000, 26/5/ 07–04/06/07 , leg. L. Saltini, Lacon drusa (Mars.) det. Platia 2009“ ( PCGP ); 4 males , „Syria mer. occ., Bludan mts., Anti Lebanon, 17.–23.V.1995 , leg. Kabátek [one with additional labels: Lacon drusa (Mars.) det. C. Wurst 1997, Lacon drusum (Mars.) det. R. Preiss 2003 , Lacon drusa (Mars.) det. Platia 2010 “]” ( PCJM ); 1 male , 1 female , “Syria SW, Blúdán, NW Damascus , 21.–22.V.1998 , lgt. E. & P. Hajdaj [male with additional data: Lacon drusa (Mars.) det. Platia 2010 ]” ( PCJM ); 1 female , “Syria SW, Blúdán, 21.–22.V.1998 , lgt. E. & P. Hajdaj” ( PCJM ); 5 males , “Sy — Mts. Anti-Lebanon, 5 km NW of Blúdán, 4.VI.1998 , Z. Košťál + V. Bíža lgt.” ( PCJM ); 2 males , “Syr. Antilebanon mts., Bludan 1900–2000 m , 22.5.2002 , Skoupý leg.“ ( PCJM ); 1 male , „Syria, Bludan env. (NW Damascus ), 1900 m , 21.5.1998 , St. Kadlec leg.“ ( PCJM ); 1 male , „Syria, Bludan, 6.5.2008 , Skoupý leg.“ ( PCJM ); 1 female , „Syria m. occ., Anti-Lebanon mts., Bludan env., alp. (NW Damascus ), 2000 m , 3.v.2000 , S. Benedikt leg.“ ( PCJM ); 3 females , „Syria, Bludan, 6.5.2008 , Štěpánek leg.“ ( 1x PCJM , PCPS ); 2 females , „Syria m. occ., Hermon mts., alp, Burqush env., 1800–2200 m a.s.l., 5.v.2000 , S. Benedikt leg.“ ( PCJM ); 1 male , „Antiliban [Anti-Lebanon Mountains]” ( MNHN ); 1 male , “Antiliban [Anti-Lebanon Mountains], Museum Paris, Coll. Abeille de Perrin 1919” ( MNHN ); 1 male , “ Adelocera graeca [further text unreadable]; Museum Paris, Coll. de Marseul 1890, Graeca Cand. “ ( MNHN ); 1 male , „[yellow label with unreadable data], Janson coll. 1903–130., Says des Drucks Chevr.; Adelocera drusa Chv. [further text unreadable], Adelocera sp. nov. ( A. drusae , Chev. in mus.), ( A. Drusorum ?) [+ further long text]; Adelocera drusa Marseul., C.M.F. von Hayek 1974 , Paris Mus. Camp [further text unreadable], Adelocera bruleriei DesB, camp [further text unreadable], Coll du B. Paris, C.M.F. von Hayek” ( MNHN ); 1 male , „Syria, Slenfah, 27.4.2008 , 1 ♂ , Stěpánek [sic!] leg.” ( PCJM ); 1 male , “Sy — Mts. Anti-Lebanon, 5 km NW of Blúdán, 4.VI.1998 , Z. Košťál + V. Bíža lgt., Lacon drusa Marseul V. Dušánek det. 2002“ ( PCVD ); 3 males , „SYR. Antilebanon Mts., Blúdán env. 2000 m , 22. 5. 2002 Košťál leg“ ( PCVD ); 1 male , „SYR. Antilebanon Mts., Blúdan env. 2000 m , 22.5.2002 Košťál & Voříšek leg“ ( PCVD ); 1 fe- male, „Syria Occ. Bludan, 40 km W of Damascus 1700–2300m , Antilebano, leg. A. Wrzecionko, 6.5.2008 , Lacon drusa (Marseul) J. Mertlik det. 2011“ ( PCVD ); 4 males , „SY-Antilibanon Mts., NW of Blúdán env., 4.VI.1998 , Z. Košťál lgt.“ ( PCVD ); 1 female , „Syria, Museum Paris, Coll. Abeille de Perrin 1919, Adelocera drusa Mars. ♀ =? graeca ♀ , brulerei [sic!] Desbr., C. M. F. von Hayek det. 1968” ( MNHN ). Comparative remarks . This species is easily recognizable by its typically matt, pitch black body coloration and dorsum with black pubescence, convex pronotum with rather small punctures on disk, moderately long antennae in males, reaching last third of pronotal length (in females shorter, reaching only the second third of pronotal length), elytral surface rugose, forming distinct cell-like structures, especially in basal half, elytral punctures distinctly of two sizes, with smaller punctures of about half size of larger puncture, and male genitalia wih median lobe shorter than parameres, and parameres with robust apical lobes ( Figs 100–106 , 112, 115–117 , 191 ). The only other species with pitch black body coloration is L. safitensis sp. nov. , but it is easily distinguishable from L. drusus by the elytra less than twice as long as pronotum and distinctly wider median antennomeres ( Figs 126, 128 ). Lacon graecus ( Candèze, 1857 ) , which was often misidentified with L. drusus , differs in the more shiny and paler body with dorsal pubescence black and white ( Figs 181, 182 ), antennae of about the same length in both males and females ( Figs 181, 182 ), less convex pronotum with larger punctures, paramere with a relatively longer and narrower apical lobe ( Fig. 183 ), and the large sclerite of bursa copulatrix with much longer spines ( Fig. 187 ). Diagnostic characters . Male ( Figs 100, 101, 104–106 , 107, 109–117 ). Body 7.0– 11.8 mm long and 2.1–3.3 mm wide ( lectotype : 8.2 mm long, 2.4 mm wide). Body dorsally matt, pitch black (except historical specimens which are lighter; Figs 100–103 ), with either antennomeres II–XI, apices of serrate antennomeres, or posterior angles of pronotum lighter (in many cases only one of these body parts lighter, in some cases more); tarsi reddish brown to dark brown; dorsum densely covered with black, decumbent, moderately long, thick setae; ventral parts more shiny, with dark brown to black setae (very occassionally with several single white setae, especially in older specimens). Antenna moderately long, surpassing 2/3 of pronotal length or almost reaching pronotal posterior angles. Antennomere III 1.20–1.25 times as long as antennomere IV; median antennomeres relatively short, strongly serrate, about 1.25–1.35 times as wide as long. Pronotum 1.15–1.20 times as long as wide, sides almost subparallel, posterior angles more or less divergent, disk sparsely covered with small, deep punctures, usually separated by about diameter of puncture. Scutellar shield 1.4–1.5 times as long as wide, anterior margin slightly concave. Elytra 2.15–2.35 times as long as wide, and 2.15–2.35 times as long as pronotum, with punctures forming distinct lines, distinctly of two sizes, with smaller punctures of about half size of larger punctures; surface between punctures forming distinct cell-like structures, especially in basal half. Aedeagus with median lobe shorter than parameres, slightly surpassing lateral subapical hook of paramere; paramere robust, elongate, subparallel-sided in more than basal half, then sinuate, apically with inner sides distinctly diverging, apex rounded; apical parameral lobe robust, with outer margin sinuate. Female (102, 103, 108, 118). Body 8.6–14.0 mm long and 2.5–4.2 mm wide. Antenna shorter, reaching or slightly surpassing half of pronotal length; median antennomeres slightly less serrate, about 1.20 times as wide as long. Pronotum more robust, shorter and wider, distinctly convex, about 1.10 times as long as wide; sides widely to equally rounded; posterior angles slightly divergent. Scutellar shield relatively slightly wider, 1.3–1.4 times as long as wide. Elytra 2.05–2.20 times as long as wide, and 2.05–2.15 times as long as pronotum. Sclerites of bursa copulatrix as in Fig. 118 . Immature stages unknown. Intraspecific variability . There are some differences in body proportions (relative length of antennae, pronotum and elytra, and tarsomeres) between the coastal population from Lebanon and the population from the Anti- Lebanon Mountains (mainly from Syria ). Both males and females in the coastal population are usually smaller than the specimens from the Anti-Lebanon Mountains (body length usually 7–9 mm in males and 9–11 mm in females vs 10–11 mm in males and 11–14 mm in females). Additionally, they have slightly shorter antennae, relatively shorter and wider tarsomeres (see Figs 113 and 114 for the most different ones), and relatively shorter elytra. Males have usually elytra 2.15–2.30 times as long as wide (vs 2.25–2.35 times in the specimens from the Anti-Lebanon Mountains), and 2.15–2.25 times as long as pronotum (vs 2.25–2.35 times). Females have usually elytra 2.05–2.15 times as long as wide (vs 2.15–2.20 times in the specimens from the Anti-Lebanon Mountains), and 2.05–2.10 times as long as pronotum (vs 2.10–2.15 times). The body surface, especially on pronotal disk, also slightly differ among specimens (even within a single population), and although the punctures on pronotum are usually separated by about their diameter, in some cases punctation is denser or sparser (i.e., surface is more or less matt), with punctures separated by about 0.5 or 1.5 times their diameter, respectively. The surface of elytra is also slightly variable, from less rugose to distinctly rugose. A single male specimen from the Syrian Coastal Mountain Range ( Syria : Latakia , Slenfah) represents another population ( Figs 105, 106 , 109, 110 ). This specimen is 11.0 mm long and 3.0 mm wide, with pronotum 1.20 times as long as wide, elytra 2.35 times as long as wide, and 2.35 times as long as pronotum. It has somewhat larger punctures on pronotal disk. With a large body size, body proportions and more elongate tarsi, it is more similar to the specimens from the Anti-Lebanon Mountains. More material from that part of Syria is needed to make more precise conclusions about this population. Distribution . Israel (new country record), Lebanon , Syria ( Fig. 194 ). For the habitat photograph see Fig. 190 . Biology . During the day, adult specimens of L. drusus in northern Lebanon were usually collected under stones, either inside dry cedar or mixed forests or in meadows without any trees or bushes. At night, they were actively crawling on stones (T. Németh, personal observation). Literature . Marseul (1870: 380) : original description of A. drusa ; Candèze (1874: 21) : revision [as Adelocera ]; Desbrochers des Loges (1875: 37) : original description of A. bruleriei ; Schenkling (1925: 10) : catalogue [under A. graeca ]; Hayek (1973: 63) : revision, Cate (2007: 101) : catalogue; Platia (2014: 74) : distributional note (as L. grae- cus , all specimens from Lebanon ); Németh (2019: 129) : catalogue. Remarks . Lacon drusus was often considered a similar species to L. graecus , which occurs in Greece and the Asia Minor (e.g., Candèze 1874 ). Schenkling (1925) even synonymized these species, but Hayek (1973) did not accept it and considered both species valid. Platia (2014) reported several specimens of L. graecus from Lebanon (cited also by Németh (2019)) , but all of them were here re-examined, compared with the type material, and clearly identified as L. drusus .