Biologia 65/6: 1040—1048, 2010 Section Zoology DOI: 10.2478/s11756-010-0121-8

A new species of Pseudomaseus (Coleoptera: Carabidae: ) from Greece

Borislav Guéorguiev1 &VladimirSkoupý2

1National Museum of Natural History Sofia, 1 Blvd. Tzar Osvoboditel, 1000 Sofia, Bulgaria; e-mail: [email protected] 2Žilina, Spojovací 149,CZ-27 01 Kamenné Žehrovice, Czech Republic; e-mail: [email protected]

Abstract: A new species of carabid , Pterostichus hejkali sp. n. is described from Northwest Greece (type locality: Epirus District, Ioannina Prefecture, Ioannina Lake environs). Diagnostic characters include distinctive shape of the median lobe of aedeagus, of right paramere and of sexual modification on last visible sternum in males as well as shape of pronotum and size of body. Members of the new species are most similar to P. minor. A key to the identification of the Balkan taxa of Pseudomaseus is presented. Key words: Coleoptera; Carabidae; Pterostichus; new species; Greece.

Introduction to describe a new species of Pseudomaseus and point out the structural differences between it and related At present, the subgenus Pseudomaseus Chaudoir, 1838 species. (genus Pterostichus Bonelli, 1810) includes 21 species; two of them live in the Nearctic, and the other nine- teen species in the Palaearctic (Bousquet & Pilon 1983; Material and methods Solodovnikov 2001; Lorenz 2005; Angus et al. 2009); Pterostichus tsukubasanus Kasahara, 1988 also belongs Thirty three specimens of six species (P. anthracinus, P. to Pseudomaseus (Seiji Morita, pers. commun.), not to gracilis, P. hejkali, P. minor, P. nigrita, P. oenotrius)from Melanius Bonelli, 1810 as was given by Bousquet (2003: the Czech Republic, Italy, Slovenia, Bulgaria, Greece, and Turkey were examined. Male specimens were boiled in wa- 499) and Lorenz (2005: 273). The Nearctic species ter and their aedeagi were extracted and put in 10% KOH were reviewed by Bousquet (Bousquet & Pilon 1983; to become lighter. The genital figures were made based on Bousquet 1999) and this author also investigated seven preparations embedded in glycerine with the aid of a draw- Palaearctic species in ordertomakecompletecharac- ing tube mounted on a Zeiss transmitted-light microscope. teristics of the subgenus. However, such study is lacking After that, the aedeagus and parameres were embedded in in the Palaearctic Region. In 2001, Solodovnikov (2001) Euparal either on the same board with specimen from which proposed a key for the majority of known Palaearctic they were extracted or on a separate transparent label (three species and described three new taxa from the East- males of new species) beneath the specimen from which they ern Palaearctic, but recently two of them were syn- were removed. The measurements and other drawings were onymyzed (Angus et al. 2009). made with the aid of an Olympus SZX10 stereobinocular. Measurements: body length from apex of longer mandible The last catalogue of the Palaearctic Carabidae in- to elytral apex (BL); body width as maximum distance dicates four species of Pseudomaseus from the Balkan across body, namely the maximum distance across the elytra Peninsula (Bousquet 2003) – (BW); maximum transverse distance across head, including (Illiger, 1798) with ssp. anthracinus and ssp. biimpres- eyes (HW); maximum width of pronotum (PW); length of sus (K¨uster, 1853), P. gracilis (Dejean, 1828) with ssp. pronotum, measured from apical margin to basal margin gracilis, P. minor (Gyllenhal, 1827) with ssp. minor and along midline (PL); maximum width of elytra (EW); length ssp. turcicus (Brandmayr & Drioli, 1978), and P. nigrita of elytra, from basal edge to apex of longer elytron (EL). (Paykull, 1790). In fact, P. oenotrius Ravizza, 1975 Internal side of paramere marks this side of paramere which has also to be included in this list as it was recorded directly fastens the distal part of the median lobe of aedea- gus; conversely the external side of paramere is opposite to from Slovenia, Croatia, and Bosnia Herzegovina (Rav- internal side. Values for sizes and ratios among specimens izza 1975: 95; Brandmayr & Drioli 1978: 106), and we of the new species are shown in Table 1. do not know any subsequent source to disregard these The studied material could be found in the following records. In 2009, we studied various Pterostichini from collections: NMNHS – National Museum of Natural History, Greece and found a series, which did not conform with Sofia, Bulgaria; NMPC – National Museum, Prague, Czech any of the known species. The purpose of this paper is Republic; cRK – private collection Rudolf Kmeco; cRL –

c 2010 Institute of Zoology, Slovak Academy of Sciences A new species of Pseudomaseus 1041

Table 1. Data for measurements and ratios among type specimens of Pterostichus hejkali.

Specimen BL BW PW/HW PW/PL EW/PW EL/PL EL/EW

Holotype 8.9 mm 3.0 mm 1.41 1.24 1.20 2.42 1.63

Paratype 1 8.7 mm 2.9 mm 1.39 1.26 1.21 2.58 1.70

Paratype 2 9.3 mm 3.0 mm 1.33 1.21 1.22 2.45 1.65

Paratypes ¾ 1 8.9 mm 3.0 mm 1.34 1.26 1.23 2.55 1.65

Paratypes ¾ 2 8.6 mm 2.8 mm 1.32 1.21 1.22 2.52 1.71

Paratypes ¾ 3 8.8 mm 3.0 mm 1.34 1.26 1.23 2.58 1.67

Paratypes ¾ 4 8.8 mm 3.0 mm 1.38 1.25 1.20 2.47 1.65

Average 8.86 mm 2.96 mm 1.36 1.24 1.22 2.51 1.67

Figs 1–6. Pronotum, dorsal view: Pterostichus (Pseudomaseus) hejkali, female paratype (1); P.(Pseudomaseus) minor minor,Czech Republic, Dyje River, female (2). Last abdominal sternum of males, ventral and lateral view: Pterostichus (Pseudomaseus) hejkali, paratype (3, 4); P.(Pseudomaseus) minor minor, Bulgaria, Pancharevo Reservoir (5, 6). Scales 1 mm. private collection Roman Lohaj; cVS – private collection Head as long as wide, dorsal surface with dou- Vladimir Skoupý. ble punctuation (a very fine punctuation spread al- most throughout dorsum and a coarser punctuation over and behind posterior supraorbital setiferous punc- Pterostichus (Pseudomaseus) hejkali sp. n. tures); frontal furrows well impressed, arched, diver- (Table 1, Figs 1, 3, 4, 7–9, 15–17, 23–25, 31–33) gent posteriorly. Eyes relatively big, prominent, hemi- spherical; temporal slopes laterally as long as or longer Description. An elongate, convex and shiny Pseudo- of 1/3 length of eyes. Labrum subrectangular, with maseus-species with length of body from 8.6 to 9.3 mm 3+3 marginal setiferous punctures anteriorly. Clypeus (average 8.86 mm), maximal width from 2.8 to 3.0 mm subtrapezium-shaped, hardly concave anteriorly, con- (average 2.96 mm), black coloured torso, black to dark vex posteriorly, posterior margin ± reduced. Antennae brown appendages (palpomeres, tibiae, and tarsomeres relatively long, pubescent from proximal part of arti- dark brown to red-brown), and without microsculpture cle 4, terminal two articles reaching base of pronotum. of tegument (present only on labrum). Mentum tooth bifid. Gula smooth; ventral lobes of head 1042 B. Guéorguiev &V.Skoupý

Figs 7–14. Median lobe of aedeagus, left lateral view: Pterostichus (Pseudomaseus) hejkali, holotype (7); Pterostichus (Pseudomaseus) hejkali, paratype (8); P.(Pseudomaseus) hejkali, paratype (9); P.(Pseudomaseus) minor minor, Czech Republic, Dyje River (10); P. (Pseudomaseus) minor minor, Bulgaria, Pancharevo Reservoir (11); P.(Pseudomaseus) minor turcicus, Turkey, Salmanka¸s Pass (12); P.(Pseudomaseus) minor ssp., Greece, Prespa Lake (13); P.(Pseudomaseus) oenotrius, Italy, Lago di Doberdo Lake (14). Scale 0.5 mm.

(outside gular suturae) with several transversal wrin- stria 3 in anterior 1/3 elytron, second one on stria 2 kles and punctures. just after middle of elytron, and last one on stria 2 Pronotum subcordate (Fig. 1), widest in middle. in posterior 1/4 elytron); intervals flat, hardly convex. Disc convex, smooth (except basal part); midline fine, Ratio EW/PW 1.20–1.23 (average 1.22); ratio EL/PL almost reaching both anterior and posterior margin. 2.42–2.58 (average 2.51); ratio EL/EW 1.63–1.70 (av- Basal part punctured, with two impressions from each erage 1.67). Wings fully developed in most specimens; side of midline, punctuation spread on and in area be- only one possesses wings shorter than length of elytra. tween impressions (outside limited from convex fold), Proepisternae, frontal parts of mesoepisternae, and central 1/8–1/9 of pronotum smooth, impunctate. Lat- lateral parts of metathorax coarsely punctured, epipleu- eral margin of pronotum convex forward, backward sub- rae, central parts of prothorax and metathorax smooth, sequently convex, straight and hardly concave before other parts of thorax (including metaepisternae) finely posterior angles; lateral border fine, equal in width from punctured. Abdominal sterna with ± distinct longitudi- end to end; anterior margin slightly concave; posterior nal wrinkles in base and laterally (first sternum entirely margin straight in middle, oblique towards obtuse pos- wrinkled). Last visible sternum in males marginally terior angles. Ratio PW/HW 1.32–1.41 (average 1.36); with 2 setiferous punctures, medially with quite ele- ratio PW/PL 1.21–1.26 (average 1.24) (Table 1). vated, wide and massive longitudinal keel, prolonged Elytra subparallel, convex dorsally, widest just af- medially on apical 2/3 to 3/4 of length of sternum ter middle; shoulders round, with small denticles (vis- and terminally clearly bent to left (Figs 3, 4); last ible at dorsal-apical view). Lateral margin more con- visible sternum in females entirely smooth medially, stricted in apical fifth, slightly concave just before apex. marginally with 4 setiferous punctures. Legs relatively Basal margin slightly wider than basal margin of prono- long and slender. tum, basal border wide; elytral striae fine, impunctate, Median lobe of aedeagus in lateral view (Figs 7–9, each elytron with 3 setiferous punctures (first one on 15–17) bent in middle, basal part (basal bulb + medial A new species of Pseudomaseus 1043

Figs 15–22. Median lobe of aedeagus, right lateral view: Pterostichus (Pseudomaseus) hejkali, holotype (15); Pterostichus (Pseudo- maseus) hejkali, paratype (16); P.(Pseudomaseus) hejkali, paratype (17); P.(Pseudomaseus) minor minor, Czech Republic, Dyje River (18); P.(Pseudomaseus) minor minor, Bulgaria, Pancharevo Reservoir (19); P.(Pseudomaseus) minor turcicus, Turkey, Salmanka¸s Pass (20); P.(Pseudomaseus) minor ssp., Greece, Prespa Lake (21); P.(Pseudomaseus) oenotrius, Italy, Lago di Doberdo Lake (22). Scale 0.5 mm. stalk) somewhat shorter than apical part, latter with Etymology. It is our pleasure to name this species pronounced apex; endophallus with a distinct spinu- after Jiří Hejkal, an excellent Czech specialist on Cara- late area (at left lateral-apical part of median lobe). bidae and genus Amara Bonelli, 1810. Right paramere (Figs 23–25, 31–33) sickle-like, distal part with blunt end, medial part almost straight, prox- Diagnosis and recognition. Adults of this species imal part robust, ± obliquely bent towards end. can be distinguished from those of all other Palaearctic Pseudomaseus species by the following combination of Types. Holotype: male, labelled “W – GREECE: Ioan- character states: black coloured specimens with size of nina : Ioannina lake env. 17.IV.2003 K. Adamík & I. body between 8.6 mm and 9.3 mm (Table 1), pronotum Boščík leg.” [typeset white label] / “collection Skoupý” [typeset white label] / “Holotype Pterostichus hejkali sp. (Fig. 1) with lateral border straight or hardly concave

n. Guéorguiev & Skoupý det. 2009” [typeset red label] before obtuse posterior angles and base to posterior an- ¾¾ (NMPC). Paratypes: 2 ,3 , labelled as holotype gles oblique, very elevated and massive longitudinal keel

(NMNHS; cVS); 1 ¾, labelled: “W – GREECE: Ioannina on last visible sternum in males (Figs 3, 4), median lobe : Ioannina lake env. 17.IV.2003 Karel Adamík leg.” [typeset of aedeagus in lateral aspect (Figs 7–9, 15–17) with api- white label] / “Pterost.((Pseudomas.) spec. D.W. Wrase cal part somewhat longer than basal part, longer apex, det. 06” [handwritten white label] / “collection Skoupý” and a noticeable spinulate area of endophallus (situ- [typeset white label] / “Paratype Pterostichus hejkali sp. n. ated in left lateral-apical part of median lobe), right Guéorguiev & Skoupý det. 2009” [typeset red label] (cVS). paramere with relatively long medial part and blunt Type Locality. Northwest Greece, Epirus District, Ioan- nina Prefecture, Ioannina Lake environs. Ioannina Lake rep- distal end (Figs 23–25, 31–33). resents a natural endorheic basin (a closed drainage basin The new species is distinct from its closest relative, that retains water and allow no outflow). The lake is situ- P. minor by the following five characters. The pronotum ated at ca. 470–480 m a.s.l. of P. hejkali (Fig. 1) has lateral borders less constricted 1044 B. Guéorguiev &V.Skoupý

Figs 23–30. Right paramere, internal side: Pterostichus (Pseudomaseus) hejkali, holotype (23); Pterostichus (Pseudomaseus) hejkali, paratype (24); P.(Pseudomaseus) hejkali, paratypes (25); P.(Pseudomaseus) minor minor, Czech Republic, Dyje River (26); P. (Pseudomaseus) minor minor, Bulgaria, Pancharevo Reservoir (27); P.(Pseudomaseus) minor turcicus, Turkey, Salmanka¸s Pass (28); P.(Pseudomaseus) minor ssp., Greece, Prespa Lake (29); P.(Pseudomaseus) oenotrius, Italy, Lago di Doberdo Lake (30). Scale 0.2 mm. to anterior margin, backward straight to hardly concave right paramere smaller, with shorter medial part and before obtuse posterior angles, base (towards posterior less robust proximal part (Figs 26–29, 34–37). Finally, angles) oblique, and basal punctuation not spread on P. hejkali has larger size of body (8.6–9.3 mm) while P. central 1/8–1/9 of pronotum and within midline (sepa- minor is relatively smaller species (6.7–8.5 mm). rate punctures may present there as an exception). Con- P. hejkali is also closely related to P. oenotrius. versely, the pronotum of P. minor (Fig. 2) has lateral The former is distinct from the latter in the, median borders more constricted to anterior margin, backward lobe of aedeagus as a whole more robust with relatively more distinctly concave before right to slightly obtuse longer apical part (in relation to basal part) and en- posterior angles, base (towards posterior angles) less dophallus with distinct spinulate area (Figs 7–9, 15– oblique, and basal punctuation spread on central 1/8– 17), right paramere robust with wide blunt distal end, 1/9 of pronotum and within midline as a rule. The sex- straighter medial part and more robust proximal part ual modification on the last visible male sternum in P. (Figs 23–25, 31–33), as well as last visible sternum in hejkali (Figs 3, 4) is quite elevated, wide (at least two the males with protruding (easily visible in lateral view) times higher and wider than that in its congener), and sexual modification (Figs 3, 4); P. oenotrius has median massive longitudinal keel, prolonged medially on apical lobe of aedeagus as a whole more slender with relatively 2/3 to 3/4 of length of sternum and terminally clearly shorter apical part (Figs 14, 22) and endophallus with- bent to left; P. minor has keel with similar situation and out spinulate area (see Brandmayr & Drioli 1978: 103, prolongation, but slightly elevated, narrow, slender and 107, Figs I b and III a), right paramere slender with terminally almost right (Figs 5, 6). The median lobe of narrow round distal end, oblique medial part and less aedeagus in lateral aspect in P. hejkali larger, with rel- robust proximal end (Figs 30–38), as well as last visible atively longer apical part, longer apex, and visible spin- sternum in the males with indistinct sexual modifica- ulate area of endophallus in left lateral position (Figs tion. 7–9, 15–17); its congener has smaller median lobe, with relatively shorter apical part, shorter apex, and hidden Phylogenetic relationships. Based on characters of position of the spinulate area of endophallus (Figs 10– the internal morphology, P. hejkali is most closely re- 13, 18–21). Furthermore, the right paramere of the new lated to P. minor. Males of this species share uniquely species larger, with longer medial part and more ro- with those of the latter species a ± elevated longitudi- bust proximal part (Figs 23–25, 31–33); P. minor has nal keel on last visible sternum in males, an endophallus A new species of Pseudomaseus 1045

Figs 31–38. Right paramere, external side: Pterostichus (Pseudomaseus) hejkali, holotype (31); Pterostichus (Pseudomaseus) hejkali, paratype (32); P.(Pseudomaseus) hejkali, paratype (33); P.(Pseudomaseus) minor minor, Czech Republic, Dyje River (34); P. (Pseudomaseus) minor minor, Bulgaria, Pancharevo Reservoir (35); P.(Pseudomaseus) minor turcicus, Turkey, Salmanka¸s Pass (36); P.(Pseudomaseus) minor ssp., Greece, Prespa Lake (37); P.(Pseudomaseus) oenotrius, Italy, Lago di Doberdo Lake (38). Scale 0.2 mm. of aedeagus with a distinct spinulate area (cf. Brand- ments and their activities are long time existing). mayr & Drioli 1978), and a right paramere with blunt or The wings of the new form are well developed in truncate distal end. Next, the new species is closest to most specimens (macroptery), and only one female pos- P. oenotrius. Both taxa share similar shape of median sesses wings shorter than length of elytra (brachyptery). lobe of aedeagus and right parameres, but endophallus A dimorphic wing status of P. hejkali is here supposed, of latter species without spinulate area (cf. Brandmayr but further studies should give answer to this conjec- & Drioli op. cit.), and its right paramere with a round ture. (rather than a blunt) distal end. Remarks. Oertzen (1886) and Apfelbeck (1904: 261) Variations. It is interesting to note that one of male cited several localities of P. minor from Central Greece paratypes shows three striking anomalies in characters (“Veluchi”, “Akarnania”, Missolonghi”, and “Vrahori of primary importance. The median lobe of aedeagus See”), situated south and southeast of the Ioannina (Figs 9, 17) has shortened basal part (in relation to the Lake. As all these finds lie in the area of the Pindus apical one) while the same parts in other two male types mountain range, some of them or all may refer to P. are relatively longer (Figs 7, 8, 15, 16). There is also an hejkali. Further investigations, including genetic and unusual differing shape of the right paramere (Figs 25, molecular research methods, are needed to corroborate 33) with prominent angular juts and proximal excision the status of the new species as well as to specify the while the parameres of the other two types (Figs 23, 24, geographical distribution of P. hejkali, P. minor and 31, 32) seems very similar each other with well devel- P. oenotrius in the western and southern parts of the oped proximal parts. Finally, the pronotum of the male Balkan Peninsula where the populations of three species in question has one lateral setiferous punctures on left meet each other. lateral margin and two closely situated lateral setifer- ous punctures on right lateral margin; all other types Pterostichus (Pseudomaseus) minor minor of P. hejkali have single lateral setiferous puncture on (Gyllenhal, 1827) each side of pronotum. Similar aberrations have high (Figs 2, 5, 6, 10, 11, 18, 19, 26, 27, 34, 35) frequency of occurrence in genus Pterostichus (Gandhi & Herms 2008). Most likely such abnormalities are re- Material examined. Czech Republic: 1 , labelled “Mor.

sponse of disturbances and changes in the environment mer. 23.7.1995 Pavlov Dyje Skoupý leg.” (cVS); 1 ¾,la- (in the area around the Ioannina Lake urban settle- belled “Boh. c. 11–8–89 Kalspot 5850 C Skoupý leg.” (cVS). 1046 B. Guéorguiev &V.Skoupý

Bulgaria: 1 , “Bulgaria Pancharevo, 600 m, 17.10.1993, (GO) 6.12.90 Lago di Doberdo Zanella leg.” (cVS). SLOVE- ¾ B. Petrov” (NMNHS); 1 , labelled “Bulgaria, Arkutino NIA: 1 ¾, labelled “Cerknisko jezzero Cerknica, 4-VIII-94

25.5.1995 B. Gueorguiev” (NMNHS); 1 ¾, labelled “Bul- Slovenia leg. J. Muilwijk” (cVS). garia, Urvich, 600 m, 27.7.1995, B. Gueorguiev” (NMNHS). Taxonomic notes. See part “Discussion”.

Pterostichus (Pseudomaseus) minor turcicus (Brandmayr et Drioli, 1978) Discussion (Figs 12, 20, 28, 36)

The intraspecific relationships within the Palaearctic ¾ Material examined.Turkey:1 ,1 ,labelled“Tr.bor. representatives of Pseudomaseus seem very complex or. 22.5.2000 Salmanka¸sge¸c. 1800 m, Skoupý leg.” (cVS); and still not well settled. Previous works demonstrated

1 , labelled “Tr. bor. or. 27.5.2000 Yalniz¸cam env. Skoupý presence of sympatric sibling species, such as P. luctuo- leg.” (cVS); 2 ¾¾, labelled “Turkey, Bolu vil. Mengen env., 700–750 m, 18.–20.VI.2003 I. Smatana lgt.” (cRL). sus – P. tenuis and P. nigrita – P. rhaeticus (Bousquet & Pilon 1983; Koch 1984, 1986) or otherwise ‘semis- pecies’ (according to Brandmayr & Drioli 1978), such Pterostichus (Pseudomaseus)?minor (or P. as P. minor – P. oenotrius (Ravizza 1975; Brandmayr oenotrius) & Drioli 1978). The sibling species have no differences (Figs 13, 21, 29, 37) in the external morphology, but internally they are dis-

tinct each other in the structure of the internal sac of ¾ Material examined. Greece: 1 ,1 , labelled “N. Greece Prespa lake Lefkonas env. Wetlands at lake 7.6.2008 Rudolf the aedeagus and right paramere in the males as well Kmeco leg.” (cRK). in the structure of the sternum 8 in the females (Koch 1984). On the other hand, the ‘semispecies’, such like P. minor – P. oenotrius represent also closely related Taxonomic and distributional notes on Ptero- species, as probably their distribution is result of para- stichus minor s.l. patric speciation. Brandmayr & Drioli (1978: 116) have generalized that, in spite of the new diagnostic char- The study of males of the nominotypical subspecies acters they found in P. oenotrius, a restricted “belt from the Czech Republic and Bulgaria and males of of secondary intergradation” exists in Switzerland be- ssp. turcicus from Northeast Turkey revealed insignifi- tween this species and P. minor as in this area “only cant variations in the form of the median lobe of aedea- intermediate individuals are present”, while such hybrid gus and right paramere. Externally, the specimens from phenotypes are missing in other zone of contact. In fact, both subspecies are indiscernible each other, too. The natural cross-species-hybrids and hybrid zones between only distinct constant special feature we know to dis- species are known in ground- species (see Turin et criminate ssp. turcicus from ssp. minor is the size and al. 2003: 70, 305). Within these zones, populations are situation of the sexual modification on the last visi- characterizes by intermediate values of morphometric ble sternum in males (see the key to the species be- characters. Brandmayr & Drioli (op. cit) have found low). The females from Mengen come from a region of that shape of the median lobe of aedeagus interme- Northwest Turkey closely situated to Abant Lake, the diate in hybrids between P. minor and P. oenotrius. type locality of ssp. turcicus. Other three specimens Most hybrid zones in Europe originated during the come from high-mountain spots of Northeast Turkey, Pleistocene interglacials or postglacial, when popula- e.g., the Salmanka¸sPass(G¨um¨u¸shane Vilayet) and the tions which were separated in different refuges, met Yalniz¸cam Pass (Artvin Vilayet). The last two finds sig- each other during subsequent expansion (Turin et al. nificantly extend the known range of ssp. turcicus to the op. cit.). Although that a partial hybridization between east. P. minor and P. oenotrius has occurred, we consider Compared with the other males, the male from them distinct species (since the nomenclature of semis- North Greece (Lefkonas) has quite prominent apex of pecies is arbitrary, for species “in statu nascendi” we median lobe (similar to that of P. oenotrius)andspe- can use binominal names). There are at least four con- cific structure of right paramere (similar to that de- stant characters in the genital morphology allowing us picted for P. oenotrius by Ravizza 1975: 94, Fig. 1); to distinguish both taxa (Ravizza 1975; Brandmayr & the last visible sternum of this specimen was destroyed Drioli 1978; present work): 1/ endophallus of aedeagus before our examination. For the time being, we refer without spinulate area in P. oenotrius (vs. endophal- the species attribution of this finding as doubtful until lus with spinulate area in P. minor); 2/ median lobe of more material from Albania and Republic of Macedonia aedeagus in lateral aspect with prominent thin apex in is available for study. P. oenotrius (vs. median lobe of aedeagus in lateral view with no or scarcely prominent apex in P. minor); 3/ Pterostichus (Pseudomaseus) oenotrius Rav- right paramere distally with round end, proximally “less izza, 1975 closed” in P. oenotrius (vs. right paramere distally with

(Figs 14, 22, 30, 38) blunt or truncate end, proximally “more closed” in P. ¾ Material examined.Italy:1 ,1 , labelled “Padule di minor); 4/ sexual modification on last visible sternum

Fucecchio (FI) 2.I.77 Magrini” (NMNHS); 1 , labelled “It. (lateral aspect) in males not prominent in P. oenotrius A new species of Pseudomaseus 1047

Figs 39–44. Left basal half of pronotum: Pterostichus (Pseudomaseus) nigrita, Bulgaria, Central Stara Planina, male (39); Pterostichus (Pseudomaseus) anthracinus anthracinus, Bulgaria, Kiten, male (40). Median lobe of aedeagus, left lateral view: Pterostichus (Pseudo- maseus) anthracinus anthracinus, Bulgaria, Kiten, male (41); Pterostichus (Pseudomaseus) gracilis gracilis, Bulgaria, Dragoman, male (42). Right paramere, internal view: Pterostichus (Pseudomaseus) anthracinus anthracinus, Bulgaria, Kiten, male (43); Pterostichus (Pseudomaseus) gracilis gracilis, Bulgaria, Dragoman, male (44). Scales 1 mm (Figs 39–42), 0.5 mm (Figs 43, 44).

(vs. sexual modification on last visible sternum in males like (Fig. 43). Size larger, 9.5–12 mm ...... prominent in P. minor)...... P. anthracinus (Illiger, 1798) We should remark that the Note: P. anthracinus biimpressus (K¨uster, 1853) dif- species complex, including P. hejkali, P. minor,and fers from the nominotypical subspecies in the presence P. oenotrius, is at least at this moment of our knowl- of ± deep excision just before tip (as tip forms acute, edge, a classic set of allopatric species or subspecies (a protruding denticle) of each elytron in the female sex. “rassenkreis” in the oldest sense). The semispecies con- – Basal impressions of pronotum shallower. Abdom- cept is probably the best to use for P. minor/P. oenos- inal sterna laterally without punctuation, ± wrin- tius, but P. hejkali could be a well-separated taxon, kled; last visible sternum of male without impres- with specific rank. sion, smooth or with ± protruding sexual modifica- tion. Right paramere never hook-like. Size smaller, Key to the Balkan species of Pseudomaseus less than 10 mm ...... 3 Chaudoir, 1838 3 Median lobe of aedeagus in lateral view with dis- tinct ventral projection before apex (Fig. 42); right 1 Posterior angles of pronotum (Fig. 39) with a small, paramere with pointed distal end (Fig. 44). Last vis- well visible, ± protruding denticle; sides towards pos- ible sternum of male without protruding sexual mod- terior angles convex .....P. nigrita (Paykull, 1790) ification. Larger species (8.0–10.0 mm) ...... – Posterior angles of pronotum (Figs 1, 2, 40) right, ...... P. gracilis (Dejean, 1828) without protruding denticle; sides of pronotum to- – Median lobe of aedeagus in lateral view never with wards posterior angles straight or slightly concave . 2 ventral projection before apex (Figs 7–22); right 2 Basal impressions of pronotum relatively deep, pit- paramere with blunt or round distal end (Figs 23–38). like. Abdominal sterna laterally with both distinct Last visible sternum of male with or without protrud- punctuation and longitudinal wrinkles; last visible ing sexual modification. Smaller species (6.7–9.3 mm) sternum of male with large flat impression. Median ...... 4 lobe of aedeagus as in Fig. 41; right paramere hook- 4 Last visible sternum of male laterally without pro- 1048 B. Guéorguiev &V.Skoupý

truding sexual modification. Endophallus of aedea- References gus without spinulate area; right paramere with round distal end (Figs 30, 38)...... Angus R.B., Galian J., Wrase D.W. & Chaladze G. 2009...... The western Palaearctic species of the P. oenotrius Ravizza, 1975 (Paykull) complex, with the description of a new species – Last visible sternum of male laterally with distinctly from Spain and a new subspecies of P. nigrita from Ana- protruding sexual modification (Figs 3–6). Endophal- tolia (Coleoptera, Carabidae). Nouv. Rev. Entomol. (N.S.) lus of aedeagus with spinulate area (Figs 7–9); right 25: 297–316. Apfelbeck V. 1904. Die K¨aferfauna der Balkanhalbinsel, mit paramere with blunt or truncate distal end (Figs 23– Ber¨ucksichtigung Klein-Asiens und der Insel Kreta. Erster 29, 31–37) ...... 5 Band: Familienreihe Caraboidea. Berlin: R. Friedl¨ander und 5 Sides of pronotum backward straight to hardly con- Sohn, IX + 422 pp. cave before obtuse posterior angles; punctuation of Bousquet Y. 1999. Supraspecific classification of the Nearctic Pterostichini (Coleoptera: Carabidae). Fabreries, Supplément inner basal impression not spread on central 1/8–1/9 9: 1–292. of pronotum and within midline (Fig. 1). Last ster- Bousquet Y. 2003. Tribe Pterostichini Bonelli, 1810, pp. 469–521. num in males with more elevated and wider longitu- In: L¨obl L. & Smetana A. (eds), Catalogue of Palaearctic dinal keel, clearly bent to left terminally (Figs 3, 4). Coleoptera. Volume 1. Archostemata – Myxophaga – Ade- phaga. Apollo Books, Stenstrup, Denmark. Median lobe of aedeagus as in Figs 7–9, 15–17; right Bousquet Y. & Pilon J.G. 1983. Redescription of Pterostichus paramere as in Figs 23–25, 31–33. Larger species (8.6– (Pseudomaseus) tenuis (Casey), a valid species (Coleopteran: 9.3 mm)...... P. hejkali sp. n. Carabidae). Coleopt. Bull. 37: 389–396. – Sides of pronotum backward more distinctly concave Brandmayr P. & Drioli G. 1978. Semispeciazione pleistocenica in Platysma (Melanius) minus (Gyll.). Mem. Soc. Entomol. before right to slightly obtuse posterior angles; punc- Ital. 57: 101–116. tuation of inner basal impression spread on central Gandhi K.J.K. & Herms D.A. 2008. Report on the largest known 1/8–1/9 of pronotum and within midline (Fig. 2). occurrence of morphological anomalies in ground beetles 62: Last sternum in males with less elevated and narrow (Coleoptera: Carabidae). Coleopt. Bull. 104–113. Koch D. 1984. Pterostichus nigrita, ein Komplex von Zwillings- longitudinal keel, almost straight terminally (Figs 5, arten. Entomol. Bl. 79: 141–152. 6). Median lobe of aedeagus as in Figs 10–13, 18– Koch D. 1986. Morphological-Physiological Studies on Ptero- 21; right paramere as in Figs 26–29, 34–37. Smaller stichus “nigrita” (Col., Carab.), a Complex of Sibling Species, ... pp. 267–279. In: Boer P.D. den, Luff M. L., Mossakowski D. species (6.7–8.5 mm) P. minor (Gyllenhal, 1827) & Weber F. (eds), Carabid Beetles. Their Adaptations and Note: P. minor turcicus (Brandmayr & Drioli, 1978) Dynamics, Fischer, Stuttgart. differs from the nominotypical subspecies in the size Lorenz W. 2005. Systematic list of extant ground beetles of the and situation of the sexual projection on the last vis- World. (Insecta, Coleoptera, Adephaga: Trachypachidae & Carabidae incl. Paussinae, Cicindelinae, Rhysodinae). Second ible sternum in the males. In the nominotypical race, edition. Tutzing: Lorenz Publishing, 530 pp. there is a long keel which extends longitudinally along Oertzen E.V. 1886. Verzeichnis der Coleopteren Griechenlands medial and apical 2/3 length of sternum; in ssp. tur- und Cretas (mit Bemerkungenuber ¨ ihre geographische Ver- cicus, there is a shorter projection which extends lon- breitung und Sammelberichten). Dtsch. Entomol. Z. 30: 189– 293. gitudinally along apical 1/3–1/4 length of sternum. Ravizza C. 1975. Pterostichus (Melanius) oenotrius n. sp. (Coleoptera, Carabidae). Boll. Soc. Entomol. Ital. 107: 92– 96. Acknowledgements Solodovnikov I. 2001. New and little-known species of ground- beetles of the subgenus Melanius Bon (sic) of the genus We thank Dr. S. Hristovski (Skopje, Republic of Macedo- Pterostichus Bon. (Coleoptera: Carabidae) from Palearctic nia) for data about P. minor from Republic of Macedonia, region. Baltic J. Coleopterol. 1: 19–29. Dr. S. Morita (Tokyo, Japan) for the information about the Turin H., Penev L. & Casale A. 2003. The Genus Carabus in Eu- systematic position of P. tsukubasanus, and an anonymous rope. A Syntesis. Co-published by Pensoft Publishers, Sofia- reviewer for making useful recommendations to an earlier Moscow & European Invertebrate Survey; Leiden XVI + 512 version of the manuscript. Mr. R. Kmeco (Litovel, Czech pp. Republic) and Mr. R. Lohaj (Košice, Slovakia) sent us ma- Received January 19, 2010 terial for study. Accepted July 30, 2010