Pheggomisetes serbicus, Vrbica & Petrović & Pantelić & Krmpot & Rabasović & Pavlović & Jovanić & Guéorguiev & Goranov & Vesović & Antić & Marković & Petković & Stanisavljević & Ćurčić, 2018

Vrbica, Maja, Petrović, Andjeljko, Pantelić, Dejan, Krmpot, Aleksandar J., Rabasović, Mihailo D., Pavlović, Danica, Jovanić, Svetlana, Guéorguiev, Borislav, Goranov, Stoyan, Vesović, Nikola, Antić, Dragan, Marković, Ðorđe, Petković, Matija, Stanisavljević, Ljubiša & Ćurčić, Srećko, 2018, The genus Pheggomisetes Knirsch, 1923 (Coleoptera: Carabidae: Trechinae) in Serbia: taxonomy, morphology and molecular phylogeny, Zoological Journal of the Linnean Society 183, pp. 347-371 : 351-356

PHEGGOMISETES SERBICUS ĆURČIĆ, VRBICA & B. GUÉORGUIEV SP. NOV. ( FIGS 1 AND 2A–H) Material examined: Holotype male labelled as follows: ‘Southeast Serbia, Stara Planina Mts., Pež Dupka Cave, 43°13′17.77″N 22°47′17.08″E, village of Dojkinci, 869 m a.s.l., near Pirot, 11.VII–10.X.2013, from pitfall traps, leg. D. Antić & M. Petković’ (white label, printed)/‘ Holotypus Pheggomisetes serbicus sp. nov. Ćurčić, Vrbica & Guéorguiev det. 2016’ (red label, printed) ( IZFB). Paratypes: six males and eight females, same data as for holotype ( IZFB); three males and four females labelled as follows: ‘Southeast Serbia, Stara Planina Mts., Pež Dupka Cave, village of Dojkinci, 869 m a.s.l., near Pirot, 11.VII.2013, leg. D. Antić & M. Petković’ ( IZFB). All paratypes are labelled with white printed locality labels and with red printed labels ‘ Paratypus Pheggomisetes serbicus sp. nov. Ćurčić, Vrbica & Guéorguiev det. 2016’. Description: TL R 5.55–6.675 mm ( M 6.11 mm) (HT 6.30 mm). Head oval, HL /HW R 1.18–1.34 ( M 1.26) (HT 1.27), widest somewhat before its middle, scarcely wider than pronotum ( Fig. 1). Frontal furrows long, slightly exceeding mid head level, deeply impressed anteriorly and sigmoidally curved. Neck narrow, HW/ NW R 1.97–2.75 ( M 2.50) (HT 2.19). Antennae long, around the same length as TL in males, while shorter than TL in females. Figure 1. Pheggomisetes serbicus sp. nov. from the Pež Dupka Cave, village of Dojkinci (near Pirot), Stara Planina Mts., Southeast Serbia. Holotype male, habitus (dorsal view). Scale = 5.0 mm. Pronotum widest somewhat after the anterior third, almost as long as wide ( Fig. 1). Anterior pronotal margin concave, shorter than pronotal base. Lateral pronotal margins rounded anteriorly and slightly concave posteriorly. Pronotal base somewhat concave in the middle. Fore pronotal angles obtuse, rounded. Hind pronotal angles acute, almost right. Elytra relatively long, oval, convex, with the lateral sides rounded anteriorly, widest slightly after the mid level, EL/EW R 1.61–1.83 ( M 1.725) (HT 1.81). Elytral base slightly narrower than pronotum ( Fig. 1). Humeral angles obtuse, rounded and quite elevated. Elytral apex rounded. Legs and claws long and thin ( Fig. 1). Median lobe of the aedeagus in lateral view curved, with a rounded somewhat elevated apex ( Fig. 2A, E). Basal bulb small, rounded. Parameres with three apical setae each. Median lobe in dorsal view straight, with a rounded apex, narrowing towards basal bulb ( Fig. 1B). Gutter-shaped copulatory piece covered with numerous spines ( Fig. 2B, F), wide at its basal three fifths and markedly narrowed at its apical two fifths. Male abdominal sternite IX (urite) subtriangular, slightly elongate, slightly longer than aedeagus ( Fig. 2C, G). Apophysis narrow, constricted proximally. Both gonocoxites IX and gonosubcoxites IX as presented in Figure 2D, H. Gonocoxites IX of moderate length, slightly curved, apically rounded, basally completely jointed with massive gonosubcoxites IX ( Fig. 2D, H). Chaetotaxy. Frons with six to seven (HT – 7) setae on each side. Pronotum with normal chaetotaxy (two pairs of setae). Five to seven setae on third interstria (HT – 7) on each elytron ( Fig. 1). Elytral umbilicate series: First three humeral setae close to marginal gutter, fourth being somewhat farther from the gutter, distance between umbilicate pores 2 and 3 shortest, while between pores 3 and 4 longest; median series at around the middle of the elytra, two setae being somewhat distanced from marginal gutter, distance between pores 5 and 6 about as long as distance between pores 2 and 3; apical series: setae 7 and 8 being somewhat distanced from marginal gutter, distance between pores 7 and 8 longer than distance between pores 3 and 4 ( Fig. 1). Differential diagnosis: The new species is compared here with the other known Pheggomisetes species ( Casale & Laneyrie, 1982; Moravec et al., 2003). A comparison of the new species with P. ninae is not provided here since the latter taxon is regarded as a subspecies of P. globiceps (see below). The new species differs from P. globiceps in having a smaller value of TL M (6.11 vs. ≥ 6.295 mm), a smaller value of HL M (1.39 vs. ≥ 1.40 mm), smaller values of AL M (5.91, males 6.01, females 5.70 vs. ≥ 6.525 mm, males ≥ 6.56 mm, females ≥ 6.30 mm), smaller values of AL / TL M (0.97, males 0.99, females 0.92 vs. ≥ 1.01, males ≥ 1.05, females ≥ 0.97), a greater value of HL / AL M (0.235 vs. ≤ 0.225), a greater value of FL M (0.75 vs. ≤ 0.68 mm), a greater value of FL/ HL M (0.54 vs. ≤ 0.48), different shape of the humeral angles (more rounded, quite elevated vs. more obtuse, less elevated), a greater value of EL/ TL M (0.55 vs. ≤ 0.54) and different shape of the copulatory piece in dorsal aspect (more markedly narrowed apically vs. gradually narrowed apically) ( Tables 2 and 3; Figs 1–13; Supporting Information, Table S1) ( Jeannel, 1928; Guéorguiev, 1964; this study). Even though the differences obtained between the mean values of certain measurements ( HL) and ratios ( HL / AL, EL/ TL) are very small (Supporting Information, Table S1), the distributions of ranges in the two species show statistically significant differences ( Table 3). Figure 2. Pheggomisetes serbicus sp. nov. from the Pež Dupka Cave, village of Dojkinci (near Pirot), Stara Planina Mts., Southeast Serbia. Bright-field (A–D) and TPEF (E–H) microscopy images. A, E, holotype male, aedeagus (lateral view). B, F, holotype male, aedeagus (dorsal view). C, G, holotype male, abdominal sternite IX (urite). D, H, paratype female, gonocoxites IX and gonosubcoxites IX. Scales = 0.1 mm. The new species differs from P. radevi in having a smaller value of TL R (5.55–6.675 vs. 7–8 mm), a greater value of HL /HW M (1.26 vs. 1.00), different shape of the head (widest slightly before its mid part, posteriorly somewhat convex vs. widest around its mid part, posteriorly very convex), a smaller value of HW/ NW M (2.50 vs. c. 3.00), different shape of the pronotum (weakly narrowed basally, strongly rounded anteriorly, well sinuate in back vs. strongly narrowed basally, weakly rounded anteriorly, strongly sinuate in back), a different value of PaW/PbW (pronotal apex between tips of the anterior angles narrower than pronotal base between tips of the posterior angles vs. pronotal apex between tips of the anterior angles wider than pronotal base between tips of the posterior angles), different shape of the hind pronotal angles (almost right, not prominent vs. acute, protruding backwards and outwards), different form of the elytra (less elongate, with more prominent shoulders vs. more elongate, with less prominent shoulders), different shape of the median lobe (less bent vs. more bent) and different shape of the basal bulb (relatively small, rounded vs. mediumsized, relatively elongate) ( Jeannel, 1928; Guéorguiev, 1964; this study). The new species differs from P. buresi in having a smaller value of TL R (5.55–6.675 vs. 7.20–9.00 mm), different shape of the head (less elongate, posteriorly more convex, abruptly narrowing towards the neck vs. more elongate, posteriorly less convex, gradually narrowing towards the neck), a different position of maximum head width (slightly in front of the middle vs. anteriorly), a greater value of HW/NW M (2.50 vs. c. 2.00), different form of the lateral pronotal margins (a little rounded anteriorly, slightly sinuate posteriorly vs. somewhat arcuate), different shape of the hind pronotal angles (almost right, not prominent vs. acute, protruding backwards and outwards), different shape of the elytra (less elongate, with more pronounced shoulders vs. more elongate, with less pronounced shoulders), different shape of the median lobe (less elongate, less bent vs. more elongate, more bent) and different shape of the basal bulb (relatively small, rounded vs. relatively massive, elongate) ( Jeannel, 1928; Guéorguiev, 1964; this study). Table 2. Results of ANOVA for each variable between P. serbicus sp. nov. and P. globiceps (exact significance level P ≤ 0.05, marked in bold) Variable F P -value HL 4.760 0.033 HW 0.273 0.603 FL 11.519 0.001 HL/HW 1.346 0.251 HL/AL 54.485 < 0.001 HL/PL 0.559 0.458 HL/TL 22.596 < 0.001 HW/NW 1.875 0.176 HW/PW 0.040 0.843 AL/TL 17.269 < 0.001 PW 1.092 0.300 PL/PW 3.788 0.056 PaW 5.207 0.026 PbW 0.287 0.594 PaW/PW 3.582 0.063 PbW/PW 0.047 0.829 PaW/PbW 3.744 0.058 EW 11.428 0.001 EWP 0.673 0.415 BW/PW 0.643 0.426 Among the three known species of the genus, P. globiceps is the smallest one and the only species that has right hind angles of the pronotum ( Jeannel, 1928; Guéorguiev, 1964). It has a pronotum that is weakly narrowed basally, with lateral margins moderately rounded anteriorly and weakly sinuate posteriorly, in addition to a relatively narrow elytral base ( Jeannel, 1928; Guéorguiev, 1964). Thus, it is quite clear that P. serbicus sp. nov. shares the same character states and is closely related to it. In addition, P. buresi differs from all other congeners in having a rather thick neck. No significant differences within the genus are evident in the male genitalia (especially in regard to shape of the median lobe), even at the species level ( Jeannel, 1928; Guéorguiev, 1964; this article). Variability: The number of setae on both frons (six to seven on each side) and elytra can vary (five to seven on each side). Etymology: The new species is named after Serbia, its terra typica. Distribution: The type locality is the Pež Dupka Cave in the village of Dojkinci (near Pirot) in the Stara Planina Mountains of Southeast Serbia. The new species inhabits a few caves in the villages of Dojkinci and Bela in the Stara Planina Mountains of Southeast Serbia; P. buresi and P. radevi live in caves near the town of Vratsa and villages of Chiren, Eliseyna, Chelopek, Druzhevo and Milanovo in the West Stara Planina Mountains of Western Bulgaria; and P. globiceps inhabits numerous caves in the West Stara Planina Mountains and Pre-Balkan region of Western Bulgaria, as well as a few caves in the villages of PetrlaŠ and Činiglavci in the Stara Planina Mountains of Southeast Serbia ( Guéorguiev & Guéorguiev, 1995; this study). Table 3. Results of Mann–Whitney U test between P. serbicus sp. nov. ( n = 24) and P. globiceps ( n = 44) (exact significance level P ≤ 0.05, marked in bold) Variable U P -value TL 116.5 < 0.001 HWP 447.5 0.305 AL 1.5 < 0.001 FL/HL 250.0 < 0.001 HW/EW 320.0 0.014 PL 345.5 0.026 PWP 233.5 < 0.001 PL/TL 224.5 < 0.001 PW/EW 245.0 0.001 EL 431.0 0.325 EL/EW 351.5 0.042 EL/TL 206.0 <0.001 PHEGGOMISETES SERBICUS BELENSIS ĆURČIĆ, VRBICA & B. GUÉORGUIEV SUBSP. NOV. ( FIGS 4 AND 5A–H) Material examined: Holotype male labelled as follows: ‘Southeast Serbia, Stara Planina Mts., Suva Dupka Cave, 43°14′40.9″N 22°44′16.5″E, village of Bela, 801 m a.s.l., near Pirot, 25. V.2014, leg. S. Ćurčić, D. Antić & I. Petrović’ (white label, printed)/‘ Holotypus Pheggomisetes serbicus belensis subsp. nov. Ćurčić, Vrbica & Guéorguiev det. 2016’ (red label, printed) ( IZFB). Paratypes: three males and two females, same data as for holotype ( IZFB); four males and eight females labelled as follows: ‘Southeast Serbia, Stara Planina Mts., Suva Dupka Cave, village of Bela, 801 m a.s.l., near Pirot, 25.V- 05.VII.2014, from pitfall traps, leg. D. Antić’ ( IZFB); one female labelled as follows: ‘Southeast Serbia, Stara Planina Mts., a cave in the vicinity of the Suva Dupka Cave, 43°14′30.84″N 22°44′9.40″E, village of Bela, 793 m a.s.l., near Pirot, 25.V-05.VII.2014, from pitfall traps, leg. D. Antić’ ( IZFB). All paratypes are labelled with white printed locality labels and with red printed labels ‘ Paratypus Pheggomisetes serbicus belensis subsp. nov. Ćurčić, Vrbica & Guéorguiev det. 2016’. Description: TL R 5.70–6.60 mm ( M 6.045 mm) (HT 5.925 mm). HL /HW R 1.23–1.33 ( M 1.27) (HT 1.24) ( Fig. 4). Frontal furrows reaching mid head level. HW/ NW R 2.09–2.50 ( M 2.24) (HT 2.25). Anterior pronotal margin clearly (in males) or slightly (in females) concave, shorter than pronotal base ( Fig. 3). Lateral pronotal margins very slightly concave posteriorly. Elytra with lateral sides almost straight in the anterior half, EL/EW R 1.62–1.92 ( M 1.77) (HT 1.71) ( Fig. 4). Median lobe of the aedeagus in lateral view slightly convex dorsally at around the level of two fifths, having an almost straight apex ( Fig. 5A, E). Median lobe Figure 3. Shape of the shoulders in the Pheggomisetes subspecies analysed. A, P. serbicus serbicus subsp. nov. B, P. globiceps ciniglavcensis subsp. nov. C, P. globiceps ilandjievi. D, P. serbicus belensis subsp. nov. E, P. globiceps ninae comb. & stat. nov. F, P. globiceps globiceps. Scales = 0.5 mm. Elytral umbilicate series: First three humeral setae close to marginal gutter, fourth being somewhat farther from the gutter, distance between umbilicate pores 2 and 3 shortest, distance between pores 1 and 2 approximately the same as between pores 3 and 4; median series at around the middle of the elytra, two setae being somewhat distanced from the gutter, distance between pores 5 and 6 somewhat shorter than distance between pores 2 and 3; apical series: setae 7 and 8 being somewhat distanced from marginal gutter, distance between pores 7 and 8 longer than distance between pores 3 and 4 ( Fig. 4). Figure 4. Pheggomisetes serbicus belensis subsp. nov. from the Suva Dupka Cave, village of Bela (near Pirot), Stara Planina Mts., Southeast Serbia. Holotype male, habitus (dorsal view). Scale = 5.0 mm. in dorsal view as presented in Figure 5B, F and inner sac as presented in Figure 5A, B, E, F. Male abdominal sternite IX (urite) as presented in Figure 5C, G, subtriangular, slightly elongate, slightly longer than aedeagus. Apophysis narrow, constricted proximally. Both gonocoxites IX and gonosubcoxites IX as presented in Figure 5D, H. Chaetotaxy. Frons with six to eight (HT – 6) setae on each side. Pronotum with normal chaetotaxy (two pairs of setae). Five to seven setae on third interstria (HT – 6–7) on each elytron ( Fig. 4). Differential diagnosis: The new subspecies is compared with the nominal subspecies, P. serbicus serbicus subsp. nov. The new subspecies clearly differs from P. serbicus serbicus subsp. nov. in having a smaller value of TL M (6.045 vs. 6.11 mm), a different value of FL/ HL M (frontal furrows reaching mid head level vs. frontal furrows somewhat exceeding mid head level), a smaller value of HW/NW M (2.24 vs. 2.50), a smaller value of FL M (0.68 vs. 0.75 mm), a greater value of AL M (5.99 vs. 5.91 mm), different shape of the anterior pronotal margin in females (less concave vs. more concave), different shape of the pronotal base in the middle (less concave vs. more concave), different shape of the lateral margins of the elytra anteriorly (more straight vs. rounded), a greater value of PaW/PbW M (0.78 vs. 0.74), a greater value of EL/EW M (1.77 vs. 1.725), a different position of certain humeral and median setae (distance between pores 2 and 3 shortest, distance between pores 1 and 2 approximately the same as between pores 3 and 4, distance between pores 5 and 6 somewhat shorter than distance between pores 2 and 3 vs. distance between pores 2 and 3 shortest, distance between pores 3 and 4 longest, distance between pores 5 and 6 about as long as distance between pores 2 and 3) belonging to the elytral umbilicate series, different shape of the median lobe apex in lateral aspect (almost straight vs. somewhat elevated) and different shape of the median lobe’s basal bulb in lateral aspect (more curved vs. less curved) (Supporting Information, Table S1) (this study). Variability: The number of setae on both frons (six to eight on each side) and elytra can vary (five to seven on each side). Etymology: The new subspecies is named after the village of Bela, in which the type locality is situated. Distribution: It inhabits two caves in the village of Bela (near Pirot) in the Stara Planina Mountains of Southeast Serbia – the Suva Dupka Cave and a cave in its vicinity.

Figures

Figure 1. Pheggomisetes serbicus sp. nov. from the Pež Dupka Cave, village of Dojkinci (near Pirot), Stara Planina Mts., Southeast Serbia. Holotype male, habitus (dorsal view). Scale = 5.0 mm.

Figure 2. Pheggomisetes serbicus sp. nov. from the Pež Dupka Cave, village of Dojkinci (near Pirot), Stara Planina Mts., Southeast Serbia. Bright-field (A–D) and TPEF (E–H) microscopy images. A, E, holotype male, aedeagus (lateral view). B, F, holotype male, aedeagus (dorsal view). C, G, holotype male, abdominal sternite IX (urite). D, H, paratype female, gonocoxites IX and gonosubcoxites IX. Scales = 0.1 mm.

Figure 3. Shape of the shoulders in the Pheggomisetes subspecies analysed. A, P. serbicus serbicus subsp. nov. B, P. globiceps ciniglavcensis subsp. nov. C, P. globiceps ilandjievi. D, P. serbicus belensis subsp. nov. E, P. globiceps ninae comb. & stat. nov. F, P. globiceps globiceps. Scales = 0.5 mm.

Figure 4. Pheggomisetes serbicus belensis subsp. nov. from the Suva Dupka Cave, village of Bela (near Pirot), Stara Planina Mts., Southeast Serbia. Holotype male, habitus (dorsal view). Scale = 5.0 mm.

Figure 5. Pheggomisetes serbicus belensis subsp. nov. from the Suva Dupka Cave, village of Bela (near Pirot), Stara Planina Mts., Southeast Serbia. Bright-field (A–D) and TPEF (E–H) microscopy images. A, E, holotype male, aedeagus (lateral view). B, F, holotype male, aedeagus (dorsal view). C, G, holotype male, abdominal sternite IX (urite). D, H, paratype female, gonocoxites IX and gonosubcoxites IX. Scales = 0.1 mm.