Review of the subgenus Lyrothorax of the genus Pterostichus (Coleoptera: Carabidae) in the Western Palaearctic, with descriptions of two new species Author Muilwijk, Jan 0000-0002-0230-226X uilwijk@gmail.com Author Kataev, Boris M. 0000-0001-9890-2595 harpal@zin.ru Author Wrase, David W. 0000-0003-1779-2684 carterus@gmx.de text Zootaxa 2024 2024-06-12 5468 2 255 274 http://dx.doi.org/10.11646/zootaxa.5468.2.2 journal article 298438 10.11646/zootaxa.5468.2.2 7a281190-31de-47e2-8ba9-85a1c6924faf 1175-5326 11615558 0FF33E52-AEAB-4207-935C-4E11F551A96D Pterostichus ( Lyrothorax ) caspius ( Ménétriés, 1832 ) ( Figures 1 , 2a – 11a , 12 , 15a ) Feronia ( Omaseus ) caspia Ménétriés, 1832: 120 : “ Lenkoran ”. Pterostichus ( Lyrothorax ) caspius : Heyden et al. , 1906: 88 : “Casp.” (part.). Platysma ( Lyrothorax ) caspium : Jakobson, 1907: 344 : “Talysh, N. Persien , Transkaspien” (part.). Pterostichus ( Lyrothorax ) caspius : Csiki, 1930 ; pars 112: 645: “Transcaspia, Lenkoran, N. Persien ” (part.). Pterostichus ( Lyrothorax ) caspius : Kryzhanovskij, 1983 : “SE Azerbaijan and N Iran ”. Pterostichus ( Lyrothorax ) caspius : Kryzhanovsky et al. , 1995: 104: “Talysh Mts.”. Pterostichus ( Lyrothorax ) caspius : Bousquet, 2017: 719 : Azerbaijan , Czech Republic (introduced), Iran (part.). Type material examined. ( Fig. 1 ) Lectotype : male (present designation), labelled “Daghest.” [printed] and “Caspica Menet. / Mar. casp.” [a former bottom label written by Ménétriés] and “ LECTOTYPE Feronia caspia Ménétriés, 1832 , Muilwijk, Kataev & Wrase des. 2024” ( ZIN ). Paralectotypes : 1 female , labelled “ Daghest .” [printed] ( ZIN ) ; 1 female , labelled “ Lenkoran .” [printed] ( ZIN ); both paralectotypes are also supplied with the label “ PARALECTOTYPE Feronia caspia Ménétriés, 1832 , Muilwijk, Kataev & Wrase des. 2024” (see Remarks below) . Additional material studied . Azerbaijan . “ Ad mare Casp , Brandt leg.” ( 1 male , MFNB ); Allar , Yardymli env, 23.5.2007 , 2000 m , K. Orszulik leg. ( 1 male , CKO ) [aedeagus examined]; Talysh , Yardymly District , 2 km S of Allar village , 38° 47’ N / 48° 14’ 10” E , H= 1800 m , 25.VII.1984 , I. Belousov leg. [vic. of Allar] ( 2 males , 2 females , CIB ) [aedeagus examined in 2 males]; Talysh , Yardymly District , 6 km NE of Allar village , 38° 49’ 23” N / 48° 17’ 28” E , H= 1300–1400 m , 26.VII.1984 , I. Belousov leg. [ 10 km NE] ( 1 male , 1 female , CIB ) [aedeagus examined in 1 male]; Astara district : Talysh , Palikesh vill., 1700 m , 26. VI.2003 , I. Pljushtch leg. ( 2 males , CAP , CDW ) [aedeagus examined]; Astara Istisu, W Astara, 100 m , 2.–6.VI.1996 , W. Schawaller leg. ( 2 males , 1 female , CDW , SMNS ); Astara , Istisu , 27.IV.1984 , Kh. Aliev leg. ( 4 males , 1 female , CIB ) [aedeagus examined in 1 male]; Astara Distr. , Shiyakeran vill. env., 9.VII.1988 , B. Kosov leg. ( 2 males , 1 female , MPU ) [aedeagus examined in both males]; Astara Distr. , upp. Istisuchay Riv. , Siov Vill. env., h~ 1000 m , 38°26’28”N 48°39’01”E , 18–19.IV.2004 , K. Makarov leg. ( 1 male , MPU ) [aedeagus examined]; “ Caspi. -M.- Gebiet , Hamarat, Leder leg.” ( 1 male , 1 female , MFNB ); “ Caspi.-Gebiet , Lirik Leder (Reitter)” ( 1 female , ZIN ); “ Caspi.Gebiet , Hamarat , Leder ( Reitter )” ( 2 females , ZIN ); “ Caucasus ” ( 1 male , 1 female , RMNH )” [aedeagus examined]; “ Lenkoran , Reitter” ( 3 males , 1 female , SMNS ); “Lenkoran” X.1983 , S. Golovatch leg. (1 female, SMNS); “ Lenkoran , v. Bodemeyer leg.” ( 3 males , 6 females , MNP ); Lenkoran , Apo below Bilasar , 350 m , 8.–9.VI.1996 , leg. W. Schawaller ( 1 male , CDW ) [aedeagus examined]; “Aus Circ Lenkoran Transcauc. Or.” ( 1 male , ZIN ); “ Lenkoran Leder ( Reitter )” ( 1 male , 5 females , ZIN ) [aedeagus examined]; “ Lenkoran Caucasus” ( 2 males , ZIN ); Lenkoran Distr. , Alekseevka–Dashdatuk , 6.VIII.1988 , V. Belov leg. ( 1 male , MPU ) [aedeagus examined]; Lenkoran Distr. , Avrora Vill. , Hirkan forest , 28.IV.1966 , Zagulyaev , Pastukhov leg. ( 1 male , ZIN ); same data but 20.IV.1966 ( 1 female , ZIN ); same data but Pastukhov leg. ( 1 male , ZIN ) [aedeagus examined]; same data but 1.V.1966 ( 1 male , ZIN ); Lenkoran Distr. , Avrora vill. env., 1.IV.1980 , S. Alekseev leg. ( 1 male , MPU ) [aedeagus examined]; Lenkoran Distr. , Avrora vill. env., oak forest, 26–27.I.1982 , K. Makarov leg. ( 1 male , 1 female , MPU ) [aedeagus examined]; Bileser , Avrora , Hirkan forest , 26–5–2007 , Orszulik leg. ( 2 males , 1 female , CKO ); Talysh , Avrora , 13.IV.1979 , M. Danilevsky leg. ( 1 male , MPU ) [aedeagus examined]; Lenkoran Distr. , Dashdatuk vill. env., oak forest, 28.I.1988 , V. Dushenkov leg. ( 1 female , MPU ); Talysh , Lenkoran , 28.IV.1969 , V. Dolin leg. ( 1 male , MNP ) [aedeagus examined]; “ Schach-Agač Talysch III–1904 ” ( 1 male , ZIN ); “ Schach-Agatsch ” ( 1 female , ZIN ); “ Talysch Gebirge , Transcaucas , Leder leg.” ( 1 male , MFNB ); “ Talyschgebg. Transcaucas. Leder , Reitter ” ( 2 males , 3 females , ZIN ); “ Talysch , Hauser leg.” ( 3 males , 4 females , MNP ); “ Talysch , Kaukas. ” ( 1 male , NHMUK ); “ Talysch , Carl Rost leg.” ( 1 male , MNP ) [aedeagus examined]; Talysh , Alazapin , climbing the Mt. Angelovyband , beech forest, 3.VII.1929 , K. Arnoldi leg. ( 1 male , 1 female , ZIN ) [aedeagus examined]; Talysh , Alekseevka , 28.V.1956 , V. Kurnakov leg. ( 1 male , ZIN ); Talysh , Alekseevka , 28.V.1936 , A. Bogachev leg. ( 3 males , 1 female , ZIN ) [aedeagus examined in 2 males]; Talysh , Bilyasar , 6.VI.1936 , K. Arnoldi leg. ( 1 male , ZIN ); same data but 9.VI.1936 ( 1 female , ZIN ); Talysh , Hirkan Nature Reserve , 21.VIII.1983 , I. Belousov leg. ( 1 male , ZIN ); Talysh , Lerik Distr. , Hircan Nature Reserve , E Lake Khanbulan & Ranger station, 38°39’36” N / 48°46’35” E , H= 100 m , 31.VIII.1983 , I. Belousov leg. ( 13 males , 10 females , CIB ) [aedeagus examined in 4 males]; Talysh , 30 km of road Lenkoran–Lerik, 38°45’32”N , 48°35’41”E , 29.VII.1984 ,. I. Belousov leg. ( 1 male , 2 females , CIB ); Talysh , road Lenkoran–Lerik, W Piran , forest, 38°43’58”N 48°37’43”E , 30.VII.1984 , I. Belousov leg. ( 2 males , CIB ); Talysh , Lerik –“Chertov most”, 38°44’35”N 48°24’10”E , H= 1250 m , 11.VII.1983 , I. Belousov leg. ( 1 male , 1 female , CIB ); Talysh , Lenkoran , Lerik, V.1969 , V. Dolin leg. ( 1 female , MNP ); Talysh , Astara District , WNW of Sym village , 38°29’18”N , 48°37’41”E , H= 700 m , 15.X.2017 , S. Alexeenko leg. ( 2 males , 3 females , CIB ); Talysh , Diabar Hollow , vic. Orand village , 38°45’20”N 48°21’30”E , H= 1500 m , 9.VII.1983 , I. Belousov leg. ( 1 male , 2 females , CIB ); Talysh , Diabar Hollow , vic. Amburdere village , 38°45’22”N 48°21’30”E , H= 1400 m , 10.VII.1983 , I. Belousov leg. ( 1 male , CIB ); Talysh , Polikesh , 20.vi.1994 , A.G. Koval leg. ( 1 male , CKO ) [aedeagus examined]; Talysh , W Polikesh (= Palikesh ) env., 38°30’55”N , 48°38’06”E , H= 620 m , 6.V.2017 , S. Alexeenko leg. ( 5 males , 7 females , CIB ); Talysh , vic. of Polikesh Vill. , ca 1200 m , 12–13.VI. 1994 , A. Koval leg. ( 2 males , 1 female , CAK ); same data but ca 1500 m , 14.VI.1994 , A. Koval leg. ( 1 male , 1 female , CAK ); Talysh , same data but ca 1000–1400 m , pitfall traps , 13–20.VI.1994 , A. Koval leg. ( 1 male , CAK ); W Bilyasary , road Alazapin–Kagoy , 38°36’53”N 48°36’49”E , 18–19.VII.1984 , I. Belousov leg. ( 2 males , 2 females , CIB ) . Iran . Ardebil . E. Ardebil, 12.VII.2005 , Atamehr leg. ( 1 male , CJM ) [aedeagus examined]. Gilan . Hasthpar , Talysch , 25.VII.1976 , Bohac leg.( 1 male , 5 females , MNP ); Charasu-dagh ,östl. Heroabad 2000–2300 m , 4.VIII.1968 , W. Heinz leg. ( 1 female , CWH ); Asålem (pass), NW Bandar-e Anzali , 3737N 4848E, 1630 m , 29–30.VI.2000 , P. Kabátek leg. ( 1 male , CDW ) [aedeagus examined]; Asålem Pass S., 5.vii 2003 , 1600 m , I. Rapuzzi leg. ( 1 female , CJM ); 25 km SW di Asålem , m 850, D. Gianasso leg. ( 1 male , CGI ); A 11, 22 km WSW Kala , Chay 1200 m , 2001, Müller leg. ( 1 female : SMNS ); Chalkaser , 15.VIII.1972 , W. Heinz leg. ( 1 male , CWH ); Charasu-dagh SW , Assalem 12–1800 m , ( Waldzone ), 3.VIII.1972 , Gilan (Iran), W. Heinz leg. ( 1 male , CWH ) [aedeagus examined]; Dare-dasht , 1000 m , E Rudbarak, 4.VII.1968 , W. Heinz leg. ( 2 males , CWH ) [aedeagus examined]; Sardab-rud , Rudbarak , 1700 m , 20.VIII.1998 , ( 2 males , CWH ) [aedeagus examined]; SE of Tutkabon , 10.6. 2014 , 900 m , N 36°50′18′′ E 49°39’47” F. Černý leg. ( 1 male , CPV ) [aedeagus examined]; Tutkabon near Rudbar, N 36°83’ E 49°66’, 960 m 16.6.2010 , W. Grosser leg. ( 1 female : CZM ); Ghaleroudkhan , 1.sep.2011 , 27 03 33 N 49 14 15E , Moslem Salari leg. ( 1 male , CRF ) [aedeagus examined]; Masuleh , 1.VIII.1967 , südl. Talysch 1000–1800 m , W. Heinz leg. ( 2 females , CWH ); Masuleh , 9.VIII.1968 , 1400–2000 m , W. Heinz leg. ( 2 females , CWH ); Masuleh , 13.IV.1996 , 1400 m , W. Heinz leg. ( 3 males , 1 female , CWH ) [aedeagus examined in all males]; Rostamabad (near Rudbar ) 12 km NW, 3655N 4924E, 1351 m , 30.–31.V.2003 . P. Kabátek leg. ( 1 male , 1 female , CPW ) [aedeagus examined]; Rostamabad 1650 m 1.6. 2010 M. Marik leg. ( 1 female , CVS ); “ Nord Persien , Elburs geb., v. Bodemeyer” ( 1 male , MNP ) [aedeagus examined]; Elbourz occ., Sud Siahkal , 22.VI.2003 , I. Rapuzzi leg. ( 1 female , CJM ) . Material for further examination. (see intraspecific variation below). Iran . Gilan , “ Gassan kiade [near the mouth of Sefid-Rud River ], Persia, 14.VI. 915 Iljin (in Russian), Platysma caspius Men. V. Lutshnik d.” ( 1 male , ZIN ) [aedeagus examined]; Gilan , Rasht Distr. , near Dizkoh vill., 36°59’19”N 49°34’56”E 5–6.V.2016 , D. Kasatkin leg. ( 1 male , MPU ) [aedeagus examined]; Māzandarān , Elburs Mts. Vak Kash env. Galugah–Yaneh Sar road, 36°40’N , 53°46’E , 670 m , 28.V.2007 21, O. Šauša leg.” ( 1 female . CPB ); Mazandaran prov., 15 km S Alamdeh, 6. 6. 2014 / N 36° 26’ 27”, E 51° 54’ 11”, / 534 m , Walter Grosser lgt. (1 female, CZM) ; Mazandaran occ. Gavar Deh env. 30.5.2001 lgt. Orszulik ( 1 female , CKO ) ; “ Masendran , Persien Em. Reitter ” ( 1 male , NHMUK ) ; “ Elburs Gebirge , Nord-Persien , v. Bodemeyer leg.” ( 5 males , 2 females , MNP , NME , RMNH ) ; “ Elburs Gebirge” ( 1 female , RMNH ) ; “ Alburs ” ( 1 male , 1 female , MFNB ) ; “ Elburs , Carl Rost leg.” ( 8 males , 6 females , MFNB , MNP ) . Specimens (probably) mislabelled. (see Remarks below). “UdSSR, Armenia , Talis , 1-6.V.1991 , J. Voldřich leg.,Az. Nagorno Karabach, Talys vill.” ( 1 male , CDW ) [aedeagus examined]; “UdSSR, Armenia , Talis, 1–6.V.1991 , J. Voldřich leg.” ( 1 female , CRF ); “[ Kazakhstan ] Pavlodar 31.V. / 928 Siberia [in Russian] // Platysma / caspium Men. / V. Lutshnik d.” ( 1 male , ZIN ) [aedeagus examined] . Redescription. Habitus as in Figs 1–2a , 3a . Body length: 10.5–14.9 mm (N=215). Pronotum transverse (PW/PL = 1.37–1.50), wider than head (PW/HW = 1.31–1.40); hind angles sharp. Inner and outer basolateral foveae more or less punctured, outer fovea faintly impressed, inner fovea longer than outer one; inner and outer foveae not separated; with a sharp keel between lateral bead and outer fovea; basal transverse impression absent; lateral gutter narrow, ending mostly before hind angles. Microsculpture consisting of fine, irregular meshes, somewhat suppressed on disc, with sparse micropores. FIGURE 2. Habitus in dorsal view and pronotum increased, (a) P. caspius ( Ménétriés, 1832 ) from Talysh, (b) P. fritzei sp. n. holotype, (c) P. vagus sp. n. , from different localities, c1 holotype, c2 from Bylnice, Czech Republic. Scale bar =10 mm. Elytra ovoid-elongate (EL/EW = 1.39–1.44). Basal margin mostly straight, continuing fluently in lateral margin. Inner intervals flat, outer slightly convex. Microsculpture consisting of transverse, irregular meshes. Ventral side. Metepisternum slightly longer than wide, clearly less densely punctured than metasternum ( Fig. 4a ). Ventrite 7 in females with a superficial apical impression, in some specimens bordered on outside by a short keel, or medially divided by a keel ( Fig. 5a ). Microsculpture in apical impression consisting of deep isodiametric meshes. Median lobe of aedeagus as in Figs 6a – 9a , robust, its length 2.4–3.0 mm. Middle part bent at an angle of about 90 degrees to basal part. Margin of the folder of the membranous surface of apical orifice (pft) short, longer than wide, apically rounded, in some specimens outwards directed, terminal lamella (tl) in lateral view as a rule oblique and with a tooth (tltl) ( Fig. 6a ); in dorsal view relatively short (WL-al: 2.38–2.60, in lectotype 2.49), with apical margin mostly rectilinear ( Fig 7a ); left lateral margin sinuated outwards; right lateral margin sinuated to the apex. Gonoporal band in ventral view small, triangle shaped, with long curved point. Right pigmented band in middle (slightly) curved dorsally ( Fig. 8a ). Apical part slightly wider than basal part. Inflated endophallus ( Fig. 9a ) elongated apically; apical lobe (al) large; ostium lobe (ol), preapical lob (pl), ventral lobe (vl) small. Parameres. Right paramere short, flat, with apex right-angled ( Fig. 10a ); left paramere square, with two strongly sclerotized apophyses. Female genitalia as in Fig. 11a . Gonocoxite 1 with a row of apical setae, gonocoxite 2 with one ensiform seta ventrolateral, three ensiform setae dorsolateral of which basal seta is small. FIGURE 3. Habitus in ventral view, (a) P. caspius ( Ménétriés, 1832 ) from Talysh Mts., Hyrkan, (b) P. fritzei sp. n. holotype, (c) P. vagus sp. n. holotype. Scale bar = 10 mm. FIGURE 4. Metepisterna and metasternum, (a) P. caspius ( Ménétriés, 1832 ) from Gilan, South Siahkal, (b) P. fritzei sp. n. holotype, (c) P. vagus sp. n. holotype. c1 holotype, c2 from Asålem Pass. FIGURE 5. Ventrite 7 (a1, b1– b2, c1– c2) and sternite 8 (a3–c3), (a) P. caspius ( Ménétriés, 1832 ) a1, a3 from Talysh Mts., Hyrkan, (b) P. fritzei sp. n. from different localities, b1 from Asålem Pass, b2 from Nahár Khorán, b3 from Chalandar, ( c) P. vagus sp. n. , c1, c3 from Gorgan, Shaskola, c2 from Břeclav, Czech Republic . Scale bar = 1 mm . FIGURE 6. Median lobe in left lateral view, (a) P. caspius ( Ménétriés, 1832 ) lectotype, (b) P. fritzei sp. n. from Khelard, (c) P. vagus sp. n. from Ramsar Curti, preputial field tooth (pft), terminal lamella (tl), tooth on terminal lamella (tltl). Scale bar = 1 mm. Distribution. Azerbaijan and North Western Iran : provinces Ardebil and Gilan . The occurrence in Mazandaran needs further investigation ( Fig. 15a ). Pterostichus caspius has been collected together with P . fritzei sp. n. at the Asålem Pass ( Gilan ). Intraspecific variation. Terminal lamella of some males from Gilan is slightly longer (WL-al: 2.0–2.5) and/or has a (slightly) convex apex; lateral tooth (tltl) smaller or nearly disappeared ( Fig. 12 ). In habitus we could not find any differences from the P. caspius specimens from Azerbaijan . Additionally , some old specimens from the Western Elburz without precise locality data could only be identified as belonging to the P. caspius complex as we decided not to dissect genitalia due to a certain fragility of these specimens, so the endophallus was also not available for examination in these specimens. In future, other methods could also be available for their determination. Specimens from these localities are placed in ‘material for further examination’ and their status requires further study. From Masuleh ( Gilan ) three males were collected on 13.iv.1996 . One male has the characteristic median lobe of P. caspius , the other two males are slightly aberrant, however no differences in the habitus could be found. Remarks. According to the original description, this species is based on material collected in the forests of Lenkoran (former Russian Empire , now Azerbaijan ), seven versts (about 7.5 km ) from the sea (“les forêts de Lenkoran à 7 verstes de la mer”), without indicating the number of specimens ( Ménétriés, 1832 ). In the collection of ZIN, there are three specimens ( one male and two females ) that appear to be from the original type series with a former common bottom label written by Ménétriés “Caspica Menet. / Mar. casp.”, pinned now under the male. Although only one female is labelled “ Lenkoran .” and the other two specimens are labelled “Daghest.” (= Dag(h)estan), we identified all three as syntypes , since P. caspius does not occur in Dagestan and these printed labels were obviously added subsequently. Two specimens collected by J. Voldřich have a locality label stating: “ Armenia , Talis, 1-6.V.1991 ”.A handwritten label has been added to one of the two “Nagorno Karabach, Talys vill.”, this record is undoubtedly wrong, as there are no suitable living conditions for P. caspius in this area, it certainly refers to the Talysh region. The locality label, given by V. Lutshnik with a locality from Kazakhstan is very doubtful, because it is far outside the known distribution area.