Lethrus (Lethrus) raymondi Reitter, 1890

Král, David & Hillert, Oliver, 2013, Three new Lethrus species close to L. raymondi (Coleoptera: Geotrupidae) from the Balkan Peninsula, Acta Entomologica Musei Nationalis Pragae 53 (1), pp. 219-244 : 230-233

Lethrus (Lethrus) raymondi Reitter, 1890

( Figs 7–9 , 19–21 , 31–33 , 37–38 , 41, 45, 49)

Lethrus Raymondi Reitter, 1890: 293 (description, key).

Lethrus raymondi: REITTER (1891) : 227 (female diagnosis); MIKŠIĆ (1955): 232 (distribution); MIKŠIĆ (1957): 177 (distribution); MIKŠIĆ (1959): 65, 117 (distribution); MIKŠIĆ (1965): 242 (key); MIKŠIĆ (1970): 15 (list, distribution); ANGELOV (1965): 101 (distribution).

Lethrus Reimondi (incorrect subsequent spelling): NEDJALKOV (1905): 422 (distribution).

Lethrus (Autolethrus) Raymondi: SÉMENOW (1892) :237 (classification, list); SÉMENOW (1894):484 (classification, list); REITTER (1893): 35 (revision, key); BOUCOMONT (1912): 38 (catalogue); WINKLER (1929): 1042 (catalogue).

Lethrus (Autolethrus) raymondi : SEMENOV- TIAN- SHANSKIJ (1935): 1398 (distribution, zoogeography); SÉMENOVTIAN- SHANSKIJ & MEDVEDEV (1936): 37, 74, 86; Figs 33 , 101, 164, 212, 268, 312 (description, key, classification, distribution); MIKŠIĆ (1958): 133, 135 (diagnostic charackters, key, distribution); BARAUD (1992): 82 (diagnostic charackters, key, distribution); BUNALSKI (2001): 167 (list).

Lethrus (Lethrus) raymondi: KRÁL et al. (2001b) : 257; Figs 2, 5, 8 (diagnostic characters, distribution); NIKOLAJEV (2003): 136, 151; Figs 96: 5, 99: 1, 110: 1 (description, key, classification, distribution, type designation); GUÉORGUIEV & BUNALSKI (2004): 268 (distribution); KRÁL & NIKOLAJEV (2006): 293 (catalogue).

Type locality.‘ Türkei (Salonicki)’ [= Greece, Thessaloníki].

Type material examined (8 specimens). GREECE: CENTRAL MACEDONIA PROVINCE: LECTOTYPE (♂) ( ZMAS) (designated by NIKOLAJEV 2003: 151), ‘ Salonichi // L. Raymondi / Reitt. 64. [hw, blue ink] // Type [p, red label] // L. Raymondi Rttr. / type [hw, Indian ink] / k. [kolekciya = collection of] V. Yakovleva [= V.Yakovlev; p, original in Cyrillics] // Syntypus [p, red label] // Lectotypus / Lethrus raymondi Reitter / design. G. Nikolajev / 30.08.2002 [p, red label]’. PARALECTOTYPES: ♀ ( ZMAS), ‘Salonichi // L. Raymondi / Reitt. 64. [hw, blue ink] // Lethrus / Raymondi m. / Salonichi [probably Reitter’s hw, Indian ink] // Lethr. Raymondi / ♀. Typ. Rttr. [hw, Indian ink] / A. Semenow det. [p] // Syntypus [p, red label]’; ♂ ( ZMAS), ‘ L. Raymondi / Reitt. 64. [hw, blue ink] // Lethr. Raymondi / ♂. Typ. Rttr. III. 98 [hw, Indian ink] / A. Semenow det. [p] // Syntypus [p, red label]’; ♂ ( ZMAS), ‘ L. Raymondi / Reitt. 64. [hw, blue ink] // Lethr. Raymondi / ♂. Typ. Rttr. III. 98 [hw, Indian ink] / A. Semenow det. [p] // L. Raymondi Rttr. / [hw, Indian ink] / k. [kolekciya = collection of] V. Yakovleva [= V. Yakovlev; p, original in Cyrillics] // Syntypus [p, red label]’; ♂ ( HNHM), ‘Salonicki / Emge [probably Reitter’s hw, Indian ink] // Holotypus [red p label] 1890 / Lethrus (Autolethrus) / Raymondi / Reitter [not Reitter’s hw, Indian ink, label with red margin] // Lectotypus [p] Lethrus / raymondi Reitter / 10.04.1973 Nikolajev [Nikolajev’s hw, red label]’, [not published by Nikolajev]; ♂ ( MNHN), ‘ L. Raymondi / m. Salonicki [probably Reitter’s hw, Indian ink] // Typus [red p label with black margin] // Museum Paris / 1938 / Coll.A. Boucomont [p] // Lethrus ♂ / raymondi / Reitter / Lectotypus / Det. R. Pittino 1993 [Pittino’s hw, red label]’, [not published by Pittino]; ♀ ( MNHN), ‘ L. Raymondi / Reitt. ♀ Salonicki [not Reitter’s hw but probably of Boucomont, Indian ink] // Typus [red p label with black margin] // Museum Paris / 1938 / Coll. A. Boucomont [p] // Lethrus ♀ / raymondi / Reitt. / Paralectotypus / Det. R. Pittino 1993 [Pittino’s hw, red label]’, [not published by Pittino]; ♂ ( ZSMC), ‘ Raymondi, Rttr. / Saloniky [hw] // Kiesenwetter [p] // Lethrus / raymondi / Reitter / Syntypus / Det. O. Hillert 2003 [hw]’.

Additional material examined (815 specimens). GREECE: CENTRAL MACEDONIA PROVINCE: 1 ♂ 2♀♀ ( MNHN), 1 ♂ ( RPCM), Saripazar, 1910, R. Michel, Museum Paris , Mission du Vardar ; 2 ♂♂ ( BMNH), Kalamaria , iv.1918, J. Waterson; 2 ♂♂ 1 ♀ ( MNHN), Salonique env., Région du Mont Prophéte Élie, 786 m, v.1918, Dr. A. Berton; 4 ♂♂ 4 ♀♀ ( ZSMC), Langadas , 2.v.1942, F. Kühlhorn leg. ; 1♂ 1♀ ( ZSMC), Kolchikon nr. Loutra Langadas , E of Saloniki, 7.v.1942, P. Babiy leg.; 66 spec. ( RPCM), 7 km W Petroto 20 km N of Thessaloniki , 300 m, 3.iv.1990, L. & R. Pittino leg.; 13 spec. ( RPCM), same data but 22.iv.1990; 103 spec. ( RPCM), 5 ♂♂ 6 ♀♀ ( MNHN), same data but 20.iii.1991; 11 spec. ( RPCM), same data but 20.iv.1993; 5 ♂♂ 6 ♀♀ ( MNHN), 7 km W Petroto , 200 m, 20.iii.1991, Pittino leg. ; 27 ♂♂ 30 ♀♀ ( DKCP), Néa Filadélfia env., 40°46′31″N 022°49′24″E, 54 m, 19.iv.2005, David Král leg. ; 41 ♂♂ 102 ♀♀ ( JSCP), same data but Jan Schneider leg. ; 37 ♂♂ 42 ♀♀ ( OHCB), same data but Oliver Hillert leg. ; 2 ♂♂ 2 ♀♀ ( DKCP), Néa Filadélfia env., 40°46′31″N 22°49′24″E, 54 m, 12.iv.2006 ; 15 ♂♂ 10 ♀♀ ( RCCP), same data but Radek Červenka leg. ; 4 ♂♂ 4 ♀♀ ( SJCP), same data but Radek Dunda leg. ; 7 ♂♂ 3 ♀♀ ( OHCB), same data but Oliver Hillert leg. ; 7 ♂♂ 11 ♀♀ ( NMPC), ca. 3 km S Nea Filadelfia, 40°46.5′N 22°49.3′E, ca. 85 m, steppe, field, river, 27.–28.iv.2007, Jiří Hájek leg.; 10 ♂♂ 15 ♀♀ ( RCCP), same data but Radek Červenka leg. ; 2 ♂♂ 5♀♀ ( TRCP), same data but Tomáš Růžička leg. ; 8 ♂♂ 13 ♀♀ ( JSCP), same data but Jan Schneider leg. ; 1 ♂ ( MHNG) , 1 ♂ ( SMFD) , 1 ♂ ( ZMHB), Kortatsch , A. Schatzmayr; 1 ♂ ( SOFM), Solun , col. Nedelkow ; 1 ♂ ( DEIC), Salonick , G. Müller; 1 ♂ ( DEIC), Salonicki, Dieck ; 1 ♂ ( OHCB), Salonicki, Gr., Emge ; 3 ♂♂ 1 ♀ ( DEIC), 1 ♂ 1 ♀ ( MNHN), 1 ♂ ( OHCB), Salonik ; 1 ♂ 1 ♀ ( DEIC), Salonik, Bruck ; 1 ♂ ( NHMB), 1 ♂ 1 ♀ ( SMFD), Saloniki, Scriba ; 2 ♂♂ 1 ♀ ( HNHM), 1 ♂ 1 ♀ ( MNHN), Saloniki ; 1 ♂ ( HNHM), Saloniki ; 1 ♂ ( HNHM), 1 ♂ 1 ♀ ( MNHN), Salonique, Collection Le Moult ; 1 ♂ ( MNHN), Salonique, Museum Paris , Mission du Vardar ; 1 ♂ ( MNHN), Salonique , P. Donkier ; 1 ♂ ( MNHN), Saloniki ; 2 ♂♂ 1 ♀ ( ZMHB), Saloniki ; 2 ♂♂ 1 ♀ ( SMTD), Salonicki, Kirsch ; 2 ♂♂ ( SMTD), Salonicki ; 1 ♀ ( SMTD), Salonik, Kiesenwetter ; 1 ♂ ( NLHD), 1 ♂ ( SMTD), Saloniki, Macedon ; 1 ♀ ( SMTD), Salonicki, Müller ; 6 ♂♂ 1 ♀ ( ZSMC), Saloniky, Kiesenwetter ; 9 ♂♂ 2 ♀♀ ( ZSMC), Saloniky ; 1 ♂ 1 ♀ ( MNHN), Griechenland, Reitter ; 2 ♂♂ ( NHMW), Graecia ; 1 ♂ 1 ♀ ( ZSMC), same data . MACEDONIA: 2 ♂♂ ( ZSMC), Veles, 3.iv.[19]18 ; 1 ♂ ( ZSMC), Veles, 20.iii.[19]18 ; 2 ♂♂ ( SMFD), Veles, 12.iv.[19]18; 8 ♂♂ 4 ♀♀ ( ZSMC), same data; 14 ♂♂ 6 ♀♀ ( ZSMC), Veles, 15.iv.[19]18; 4 ♂♂ 1 ♀ ( ZMHB), Veles, P. Schulze S.G.; 10 ♂♂ 3 ♀♀, ( ZSMC), Stip, 7.iv.[19]18; 3 ♂♂ ( DEIC), Štip, 15.v.[19]37, W. Liebmann; 1 ♂ ( OHCB), Stip; 5 ♂♂ 1 ♀ ( ZSMC), Usküb, 7.iv.[19]18, Macedon. Exp. [19]17–[19]18; 1 ♂ ( ZSMC), Usküb, 2.vii.[19]18, Macedon. Exp. [19]17–[19]18; 6 ♂♂ 2 ♀♀ ( ZSMC), Usküb, 12.iv.[19]18, Macedon. Exp. [19]17–[19]18; 3 ♂♂ ( ZSMC), Usküb, Macedon. Exp. [19]17–[19]18; 1 ♂ 1 ♀ ( SMFD), Uskub ; 1 ♂ ( ZMHB), Usküb ; 1 ♂ ( NHMB), Üsküb ; 1 ♂ 1 ♀ ( SMFD), same data; 2 ♀♀ ( ZSMC), Kalučkovo, Macedon. Exp. [19]17–[19]18; 1 ♂ ( ZSMC), Plavuša Planina, Macedon. Exp. [19]17–[19]18; 1 ♂ ( ZSMC), Plavuša pl.; 1 ♂ ( SMFD), Wodno , 7.iv.[19]28; 3 ♂♂ 2 ♀♀ ( VKCS), Nogaevci / Ubogo , 41.62484899 N 21.95277169E, 216 m a.s.l., 12–13.v.2006, Václav Křivan leg. ; 7 ♂♂ 5 ♀♀ ( VKCS), Penuš , 10 km SE Štip, 21.v.2006, 41°42′02″N 22°06′53″E, 270 m a.s.l., Václav Křivan leg. ; 2 ♂♂ ( DEIC), 1 ♂ ( MNHN), Keretschkoi , A. Schatzmayr; 1 ♂ ( ZFMK), Macedonien, Doiran lake 1 ♂ 3 ♀♀ ( NMPC), Skoplje, Petrbok [lgt.]. Not located or imprecisely localized: 1 ♂ 1 ♀ ( HNHM), Balkán , coll. E. Friv. ; 2 ♂♂ ( DEIC), Macedonia ; 1 ♂ ( ZMHB), Macedonien ; 1 ♂ ( MNHN), Turcia ; 1 ♂ ( SMFD), Türk .

Diagnostic characters of males. Total body length 20–29 mm. Dorsal surface black, except of moderately shiny pronotum almost alutaceous; ventral surface black with weak blue tinge, moderately shiny.

Head ( Figs 7 , 19 , 31 , 41 ). Clypeus transverse, trapezoidal with anterior angles round. Eye canthus exceeding eyes, projecting anterolaterad, almost rectangular, lateral margins divergent posteriad, anterolateral angle round, oblique keel above eyes absent. Pleurostomal process evenly arcuate, hardly exceeding ventrolateral mandibular outline. Punctation of frons double, consisting of coarse, transversally rugose, regularly and densely distributed punctures, intermixed with fine, unevenly irregularly distributed ones; coarse punctures separated by approximately less than their diameter, punctation becoming distinctly sparser posteriad and on occiput; clypeus and eye canthus distinctly rugose.

Mandibles in maximally developed (hyperthelic) male with well developed ventral mandibular processes symmetrical, external outline almost semicircular, pointed subapically, in dorsal aspect ( Figs 31 , 41 ) with maximum width approximately at middle of mandibular length.

Ventral mandibular processes ( Figs 7 , 20 , 41 ) symmetrical, distinctly longer than horizontal length of mandible; base thickened, not exceeding lateral mandibular outline in dorsal aspect, with strongly concave external outline in basal half in frontal aspect; longitudinal keel on base laterally present, in dorsal aspect straight and distinctly subparallel to lateral mandibular outline, so broad as maximum width of mandibular outline basally; in lateral aspect very weakly arcuate, distinctly subparallel to lateral mandibular outline, divergent gradually basad approximately from middle of its length; anterior subbasal tooth absent; posterior subbasal tooth absent (lateral aspect); inferiobasal tooth present, round; both processes bent inward aproximately middle of mandibular length in frontal view; anterior subapical tooth present, distinctly visible in lateral aspect, distinctly extended in frontal aspect; apical emargination deep; apical tooth sharp, strongly extended apically.

Pronotum transverse, distinctly broader than base of elytra, broadest just behind middle; margin entirely bordered, slightly crenulate in anterior parts. Anterior angles not projecting anterolaterad, with evenly arcuate outline; lateral margin approximately straight to round posterior angle; posterior margin straight. Punctation of dorsal surface simple, consisting of deep, sparsely and irregularly distributed punctures; punctures separated by approximately two to four times their diameters discally, surface near lateral margins considerably shagreened and alutaceous.

Scutellar shield widely triangular, finely shagreened.

Elytra almost semicircular, apices not prominent, each apex forming independent arcus. Epipleuron strongly narrowed apicad, epipleural keel not reaching elytron apex. Whole surface alutaceous, finely transversally rugose; striae not indicated, entirely vanishing in rugosities.

Legs. Profemur not armed, protibia with seven to nine gradually diminishing external denticles proximad, and with row of tubercles on ventromedial edge.

Aedeagus as in Figs 37–38 .

Variability in males. Mandibular processes in medium developed ( Figs 8 , 20 , 32 ) and underdeveloped (hypothelic) ( Figs 9 , 21 , 33 ) males short, more or less straight with only indicated teeth or without them at all, simply round to almost acute apically.

Females differ from males as follows: external outline of mandibles almost straight, only in apical quarter rounded in dorsal aspect; ventral mandibular process absent; protibia broader, row of tubercles on ventromedial edge less pronounced.

Differential diagnosis. For differential diagnosis see character matrix ( Table 1 ).

Collecting circumstances. The material collected by the authors was taken on intensively grazed pasture on moderately steep slopes consisting of loess soil ( Fig. 49 ).

Distribution. Macedonia: basin of the river Vardar (= Axiós) approximately between Skopje and Gevgelija; Greece: left side of the Axiós (= Vardar) river basin ( Fig. 45 ). Precise published records available are as follows: Greece: Thessaloníki ( BARAUD 1992, NEDJALKOV 1905, REITTER 1893); Macedonia: Gevgelija, Skopje, Štip, Vodno ( BARAUD 1992, MIKŠIĆ 1955, 1959).

Records from the Thracian lowlands in Bulgaria – Chirpan, Khaskovo and Stara Zagora by ( MIKŠIĆ 1959) or Plovdiv ( ANGELOV 1965) concern almost certainly L. schaumii as well as records from ‘Roumélie’ ( BARAUD 1992), ‘Rumelien’ ( BOUCOMONT 1912, WINKLER 1929) or ‘Vostochnaya Rumeliya [in cyrillic]’ (SEMENOV- TIAN- SHANSKIJ & MEDVEDEV 1936). Records from ‘Kleinasien’, ‘Asia minor’ or ‘Malaya Aziya [in cyrillic]’ (cf. BOUCOMONT 1912; REITTER 1890, 1893; SEMENOV- TIAN- SHANSKIJ & MEDVEDEV 1936; WINKLER 1929) concern other species, probably L. rotundicollis .

Figures

Figs 1–12. Head, frontal aspect.Maximally developed males:1 – Lethrus (Lethrus) halkidikensis sp. nov. (holotype); 4 – L. (L.) perun sp. nov. (holotype); 7 – L. (L.) raymondi Reitter, 1890 (Greece: Néa Filadélfia – OHCB); 10 – L. (L.) strymonensis sp. nov.(holotype). Medium developed males: 2 – Lethrus (L.) halkidikensis sp. nov.(paratype, Greece: Tagarádes – OHCB); 5 – L.(L.) perun sp. nov.(paratype, Bulgaria:Khotovo – OHCB); 8 – L. (L.) raymondi (Greece: Néa Filadélfia – OHCB); 11 – L.(L.) strymonensis sp.nov.(paratype,Greece:Séres – OHCB).Underdeveloped males: 3 – Lethrus (L.) halkidikensis sp. nov. (paratype, Greece: Tagarádes – OHCB); 6 – L. (L.) perun sp. nov. (paratype, Bulgaria: Khotovo – OHCB); 9 – L. (L.) raymondi (Greece: Néa Filadélfia – OHCB); 12 – L. (L.) strymonensis sp. nov. (paratype, Greece: Séres – OHCB). Lettering: b – subapical tooth of ventral mandibular process, c – apical tooth of ventral mandibular process, d – inferiobasal tooth of ventral mandibular process, e – apical emargination of ventral mandibular process. Schematically, not to scale.

Figs 13–24. Head, lateral aspect. Maximally developed males: 13 – Lethrus (Lethrus) halkidikensis sp. nov. (holotype); 16 – L. (L.) perun sp. nov. (holotype); 19 – L. (L.) raymondi Reitter, 1890 (Greece: Néa Filadélfia – OHCB); 22 – L. (L.) strymonensis sp. nov. (holotype). Medium developed males: 14 – Lethrus (L.) halkidikensis sp. nov. (paratype, Greece: Tagarádes – OHCB); 17 – L. (L.) perun sp. nov. (paratype, Bulgaria: Khotovo – OHCB); 20 – L. (L.) raymondi (Greece:Néa Filadélfia – OHCB); 23 – L.(L.) strymonensis sp.nov. (paratype, Greece: Séres – OHCB). Underdeveloped males: 15 – Lethrus (L.) halkidikensis sp. nov. (paratype, Greece: Tagarádes – OHCB); 18 – L. (L.) perun sp. nov. (paratype, Bulgaria: Khotovo – OHCB); 21 – L. (L.) raymondi (Greece: Néa Filadélfia – OHCB); 24 – L. (L.) strymonensis sp. nov. (paratype, Greece: Séres – OHCB). Lettering: a – lateral longitudinal keel on base of ventral mandibular process, b – subapical tooth of ventral mandibular process, c – apical tooth of ventral mandibular process. Schematically, not to scale.

Figs 25–36. Head, dorsal aspect. Maximally developed males: 25 – Lethrus (Lethrus) halkidikensis sp. nov. (holotype); 28 – L. (L.) perun sp. nov. (holotype); 31 – L. (L.) raymondi Reitter, 1890 (Greece: Néa Filadélfia – OHCB); 34 – L. (L.) strymonensis sp. nov. (holotype). Medium developed males: 26 – Lethrus (L.) halkidikensis sp. nov. (paratype, Greece: Tagarádes – OHCB); 29 – L. (L.) perun sp. nov. (paratype, Bulgaria: Khotovo – OHCB); 32 – L. (L.) raymondi (Greece: Néa Filadélfia – OHCB); 35 – L. (L.) strymonensis sp. nov. (paratype, Greece: Séres – OHCB). Underdeveloped males: 27 – Lethrus (L.) halkidikensis sp. nov. (paratype, Greece: Tagarádes – OHCB); 30 – L. (L.) perun sp. nov. (paratype, Bulgaria: Khotovo – OHCB); 33 – L. (L.) raymondi (Greece: Néa Filadélfia – OHCB); 36 – L. (L.) strymonensis sp. nov. (paratype, Greece: Séres – OHCB). Lettering: a – lateral longitudinal keel on base of ventral mandibular process. Schematically, not to scale.

Figs 37–38.Lethrus (Lethrus) raymondi Reitter, 1890 (Greece:Néa Filadélfia – OHCB).37 – aedeagus in dorsal aspect; 38 – the same, but in lateral aspect. Schematically, not to scale.

Figs 39–42. Habitus in frontolateral aspect. 39 – Lethrus (Lethrus) halkidikiensis sp. nov. (holotype), 40 – L. (L.) perun sp. nov.(holotype), 41 – L. (L.) raymondi Reitter, 1890 (Greece:Néa Filadélfia – OHCB, body length: 28 mm), 42 – L. (L.) strymonensis sp. nov. (holotype).

Fig. 49. Collecting habitat of Lethrus (L.) raymondi Reitter, 1890 (Greece: Néa Filadélfia, April 2006; photo David Král).

Fig. 45. Sketch map of the Vardar / Axiós river basin (Macedonia, Greece) with marked distribution of Lethrus (L.) raymondi Reitter, 1890 empty dot represents the type locality.