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Deuteraphorura kozmani Parimuchová, Barjadze & Kováč sp. nov. urn:lsid:zoobank.org:act: BD48C182-8833-4537-9C44-024938DC47CD Fig. 5, Table 2 Etymology The species was named after the type locality, the Kozmani Cave in Georgia. Type material Holotype GEORGIA • ♂; Imereti, Kharagauli, Kozmani Cave; 42.10092528° N, 43.28852625° E; 14 Sept. 2021; Eter Maghradze leg.; hand collecting on detritus; IBE FS UPJS. Paratypes GEORGIA • 6 ♀♀, 2 ♂♂; same collection data as for holotype; IZISU • 8 ♀♀, 1 ♂; same collection data as for holotype; IBE FS UPJS. Description Body length 1.8–2.6 mm in females, 1.5–2.0 in males (average 2.0 mm; n = 18), shape cylindrical (Fig. 5a). Colour white to pale brownish in ethyl alcohol. Cuticular granulation fine and uniform, slightly dense around pseudocelli. Antennae almost as long as head, area antennalis relatively well marked. PAO with 14–16 compound vesicles. Ant. I with 8 chaetae in one row, Ant. II with 14–15 chaetae. AOIII with 5 papillae, 5 guard chaetae, 2 sensory rods almost as long as papillae, 2 rough sensory clubs and lateral ms (Fig. 5b). Lateral ms on Ant. IV placed basally at the level of second row of chaetae. Apical organite simple in unprotected cavity. Maxillary outer lobe simple with 1 basal chaeta and 2 sublobal hairs. Labium of AB-type, with 6 proximal chaetae. Basomedian field with 4 chaetae, basolateral field with 5 chaetae. Head ventrally with 5 postlabial chaetae. Pso formula dorsally as 33/133/4(3)4(5)3-45(6)3(4) (Fig. 5a); ventrally as 12/011/3222; head ventrally with 1 anterior, 1 postero-medial and 1 postero-lateral pso. Psx weakly visible. Subcoxae 1 of I–III pairs leg with 2,2,2 pso. Dorsal body chaetae only weakly differentiated into macro and mesochaetae. Th. I with 6–7 chaetae per half. ThII–AbdIII with 3 + 3 medial chaetae respectively. VT with 5–6 chaetae per half, basal chaetae mostly absent. Chaetae on Th. I–III sterna absent. Furca remnant with 2 + 2 thin chaetae in one row. MVO present only in fully adult males in form of 20–25 thickened, long and forked chaetae only on Abd. III sternum (Fig. 5c). Subcoxae 1 of legs I–III with 4, 4, 4 chaetae, subcoxae 2 with 3,14–17, 15–17 chaetae, trochanters with 8–10 chaetae each and femora with 14–15, 13–15, 13–15 chaetae, respectively. Tita I–III with 18, 19, 17 chaetae including 9 chaetae in distal whorl. Tita I with 6 + M chaetae in B row and 2 chaetae in C row, Tita II with 7 + M chaetae in B-row and 2 chaetae in C-row, Tita III with 6 + M chaetae in B row and 1 chaeta in C row. Claw without teeth. Empodium with basal lamella, tip of filament reaching two-thirds of the claw length (Fig. 5d). Ecology and distribution The species is known only from the type locality, occurring on guano and decaying organic material. It does not display any obvious troglomorphic adaptations. Remarks Both species belong to the species group of Deuteraphorura with 3 pso on hind margin of the head and possessing the pso on the first thoracic tergum. The vast majority of these species occupy caves in southern and central Europe. As morphological characters vary within both new species, reliable distinguishing from each other is possible only by ventral pseudocellar formula and shape of MVO in matured males. Deuteraphoruracolchisi sp. nov. has simple thickened chaetae in MVO, while modified chaetae in D.kozmani sp. nov. are longer and weakly forked at the tip. Similar to the new species, D. dashtenazensis Arbea, Yahyapour & Shayanmehr, 2020 has MVO only on Abd. III, but it differs in number of chaetae on this organ. Diagnostic morphological characters of both new species and other species of this group are listed in Table 2.

Deuteraphorura colchisi Parimuchová, Barjadze & Kováč sp. nov. urn:lsid:zoobank.org:act: AAC4848A-79D3-4A5C-8458-A8CB7BB2EA8F Fig. 4, Table 2 Deuteraphorura sp. – Zaragoza et al. 2021. Etymology The name is derived from ‘ Colchis ’ – the historical geographical, ethnical and political entity of Georgia which today is located in the west of the country. Type material Holotype GEORGIA • ♀; Imereti, Tskaltubo, Satsurblia Cave; 42.38805000° N, 42.60626700° E; 12 Mar. 2020; Eter Maghradze leg.; hand collecting on wood; IBE FS UPJS. Paratypes GEORGIA – Imereti, Tskaltubo • 3 ♀♀; Khomuli Cave; 42.31562° N, 42.63613° E; 11 Apr. 2020; Eter Maghradze leg.; hand collecting on wood, guano, water surface • 1 ♀, 3 ♂♂; Melouri Cave; 42.38752° N, 42.62819° E; 28 May 2019; Eter Maghradze leg.; pitfall traps with pork liver, hand collecting on guano and speleothems; IBE FS UPJS • 3 ♀♀; Prometheus Cave; 42.37716° N, 42.60086° E; 13 Feb. 2018; Eter Maghradze leg.; pitfall traps with pork liver, hand collecting on wood, guano and speleothems; IBE FS UPJS • 1 ♀; Datvi Cave; 42.37444° N, 42.59583° E; 27 Dec. 2019; Eter Maghradze leg.; pitfall traps with pork liver; IZISU • 1 ♀; same collection data as preceding; IBE FS UPJS • 2 ♀♀; Satsurblia Cave; same collection data as for holotype; IBE FS UPJS • 1 ♂; same collection data as preceding; IZISU • 1 ♀, 1 ♂; Sakadzhia Cave; 42.36756387° N, 42.59123348° E; 28 Dec. 2020; Eter Maghradze leg.; hand collecting on guano and detritus • 3 ♀♀, 3 ♂♂; Imereti, Khoni, Satevzia Cave; 42.43153377° N, 42.56590444° E; 10 Feb. 2020; Eter Maghradze leg.; hand collecting on guano and water surface; • 10 ♀♀, 2 ♂♂; same collection data as preceding; IZISU IBE FS UPJS • 1 ♀, 1 ♂; Imereti, Chiatura, Shvilobisa Cave; 42.3254° N, 43.26786° E; 8 Oct. 2021; Eter Magradze, Shalva Barjadze, Lado Shavadze, Mariam Gogshelidze leg.; hand collecting on guano, wood and detritus; IBE FS UPJS • 3 ♀♀; Samegrelo, Martvili, Inchkhuri Cave; 42.45678637°N, 42.40425674°E; 18 Jul. 2020; 10 Jul. 2021; Eter Maghradze, Shalva Barjadze, Lado Shavadze, Mariam Gogshelidze leg.; hand collecting guano, wood, water surface and walls; IZISU • 5 ♀♀; same collection data as preceding; IBE FS UPJS • 1 ♀; Motena Cave; 42.47657295° N, 42.39126228° E; 10 Jul. 2021; Shalva Barjadze, Lado Shavadze, Mariam Gogshelidze leg.; hand collecting on walls; IBE FS UPJS. Description Body length 1.3–2.3 mm in females, 1.85–2.1 in males (average 1.78 mm; n = 46), shape cylindrical (Fig. 4a). Colour white to pale brownish in ethyl alcohol. Cuticular granulation fine and uniform, slightly dense around pseudocelli. Antennae almost as long as head, area antennalis relatively well marked. PAO with 10–14 compound vesicles (Fig. 4c). Ant. I with 8–9 chaetae in one row, Ant. II with 14–15 chaetae. AOIII with 5 papillae, 5 guard chaetae, 2 sensory rods almost as long as papillae, 2 rough sensory clubs and lateral ms (as in Fig. 5b). Lateral ms on Ant. IV placed basally at the level of second row of chaetae. Apical organite simple in unprotected cavity. Maxillary outer lobe simple with 1 basal chaeta and 2 sublobal hairs. Labium of AB-type, with 6 proximal chaetae. Basomedian field with 4 chaetae, basolateral field with 5 chaetae. Head ventrally with 4 postlabial chaetae. Pso formula dorsally as 33/133/3(4)3(4)4(3)5(6)3-4 (Fig. 4a) (2 pso on Th. I sometimes appear); ventrally as 12/011/3212 (Fig. 4b for abdominal ventral pso formula); head ventrally with 1 anterior 1 postero-medial and 1 postero-lateral pso. Psx weakly visible. Subcoxae 1 of I–III pairs leg with 2,2,2 pso. Dorsal body chaetae only weakly differentiated into macro and mesochaetae. Th. I with 7 chaetae per half. ThII–AbdIII with 3 + 3 medial chaetae respectively. VT with 5–7 chaetae per half, basal chaetae absent. Chaetae on Th. I–III sterna absent. Furca remnant with 2 + 2 thin chaetae in one row. MVO present only in fully adult males in form of 10–20 thickened, short and bent spine-like chaetae only on Abd. III sternum (Fig. 4d). Subcoxae 1 of legs I–III 4,4,4 chaetae, subcoxae 2 with 3, 13–14, 14–16 chaetae, trochanters with 8–9 chaetae each and femora with 15, 13–15, 13–15 chaetae, respectively. Tita I–III with 18, 19, 17 chaetae including 9 chaetae in distal whorl. Tita I with 6 + M chaetae in B row and 2 chaetae in C row, Tita II with 7 + M chaetae in B-row and 2 chaetae in C-row, Tita III with 6 + M chaetae in B row and 1 chaeta in C row. Claw without teeth. Empodium with basal lamella, tip of filament reaching two-thirds of the claw length (as in Fig. 5d). Ecology and distribution The species is known only from caves in western Georgia where it inhabits warm caves at low altitudes. By its morphology, it does not display any obvious troglomorphic adaptations. Remarks See remarks for D.kozmani sp. nov.

Parimuchová, Andrea, Barjadze, Shalva, Maghradze, Eter, Kováč, Ľubomír (2023): Two new species of the genus Deuteraphorura Absolon, 1901 (Hexapoda, Collembola, Onychiuridae) from Georgian caves with remarks on the subterranean biodiversity of the Caucasus Mountains. European Journal of Taxonomy 879: 64-82, DOI: https://doi.org/10.5852/ejt.2023.879.2161, URL: http://dx.doi.org/10.5852/ejt.2023.879.2161

Plutomurus kharagauliensis Barjadze, Kováč & Parimuchová sp. nov. Figs 3–15 Type locality. GEORGIA, Imereti region, Kharagauli municipality, close to the village Amashuketi, Zemo Imereti Plateau karst massif, Kozmani Cave, 644 m asl. Type material. Holotype: Female on slide, dark zone, hand collecting, 27.xi.2021, leg. Sh. Barjadze, E. Maghradze, M. Gogshelidze, code GEOKOZ20211127-01 (IZISU); Paratypes (n=13): eight specimens on slides with the same data as holotype, code GEOKOZ20211127-02, 03, 04, 05, 08, 09, 10, 11 (IZISU); one specimen on slide, twilight zone, 14.ix.2021, leg. E. Maghradze, GEOKOZ20210914-14 (IZISU); two specimens on slides, dark zone, 27.xi.2021, leg. Sh. Barjadze, E. Maghradze, M. Gogshelidze, code GEOKOZ20211127-06, 07 (UPJ); two specimens on slides, twilight zone, 14.ix.2021, leg. E. Maghradze, GEOKOZ20210914-12, 13 (UPJ). Description. Body length 2.44–4.02 mm (Fig. 3). Colour. Body grey colored. Scale distribution. Scales are present dorsally on Ant. I–II, head, Th. and Abd. segments, all leg segments, both sides of ventral tube and ventral side of furca. Head. Ratio of antennal length: head length as 2.67–3.10. Ant. III–IV annulated. Eye patch typical for genus, with six well developed eyes (Fig. 12). Head dorsally with one unpaired (A 0), and seven paired Mc: two anterior (A 2, A 3), two interocular (S 2, S 4) and three postocular (Pa 3, Pa 4, Pa 5) distributed as in Fig. 12. Posterior margin of head with a row of mesochaetae. Prelabral and labral chaetae smooth: prelabral chaetae four (2+2) (Fig. 4); labrum with 554 papillate chaetae as typical for genus; distal margin of labrum with four elongate, thin–walled, flexible papillae. OML trifurcate, basal chaetae shorter than apical process; OML with four chaeta-like processes. Body. Dorsal bothriotrichal formula as 2,1/0,0,1,2,0 (Figs 13–14). Dorsal Mc formula: 4,2/3,3,4,6,3–4 (Figs 13–14). Thoracic macrochaetotaxy as in Fig. 13. Th. II with two anterior (m 3, m 4) and two posterior (p 3, p 5) Mc; Th. III with two posterior (p 3, p 4) Mc. Abdominal macrochaetotaxy as in Figs 14–15: Abd. I–II each with three long posterior Mc (m 2, m 3, m 4); Abd. III with four long Mc: two anterior (m 2, m 6) and two posterior (p 1, p 6) and one short Mc (p 7) as typical for genus; Abd. IV with six posterior Mc: two long (C 6, E 3) and four short Mc (A 6, B 6, D 3, T 7) (Fig. 15); Abd. V with 2–3 posterior (p 2, p 3 or p 2, p 3, p 4) long Mc and one long Mc laterally (Fig. 14). Mc length: Mc on Th. II (m 3, m 4, p 3, p 5) and Th. III (p 3, p 4)—120–167 µm; Mc on Abd. I–II (m 2, m 3, m 4) and Abd. III (m 2, m 6, p 1, p 6)—178–220 µm; Mc on Abd. IV-E 3 238 µm, C 6: 168 µm (found only in a single specimen), A 6: 93–135 µm, B 6: 110–140 µm, D 3: 100–155 µm and in T 7 Mc length is unknown (missing in all individuals). Diameter of Mc sockets on Abd IV-E 3 and C 6: 8–10 µm, 4 short Mc (A 6, B 6, D 3, T 7) subequalor in diameter.A 6 slightly longer (5.5–7.5 µm) than in remaining 3 short Mc: B 6: 5.5–6.5 µm; D 3: 5.0–6.5 µm and T 7: 5.5–6.0 µm. Legs. Hind legs with well–developed trochanteral (17–36 chaetae) and femoral (20–35 chaetae) organs (Fig. 9), with several elongated chaetae; tibiotarsal spine formula as 002, posterior side of tibiotarsus III with two outstanding inner, Spine-like chaetae, (arrows in Fig. 10). Tenent hair capitate (Figs 5, 10). Ratio of claw III: Emp. III: tenent hair III as 1.66–3.14: 0.91–1.71: 1. Inner edge of claw on all legs with one characteristically minute proximal unpaired tooth (as in Fig. 5) and 2–4 distal unpaired teeth, 3–5 in total (b, c, d in Fig. 5). Claw with long pseudonychia (Figs 5, 10). Ratio of pseudonychium III: claw III inner edge as 0.38–0.62: 1. Empodium lanceolate, tapered, with 2 internal lamellae bearing 0–6 teeth (Fig. 5). Ventral tube. Hirsute, in one specimen anterior basal, posterior basal and apicolateral sides with 36, about 24 and 35 smooth chaetae respectively (Fig. 8). Tenaculum. Corpus with one smooth chaeta; rami with 4 + 4 teeth (Fig. 6). Furca. Ratio manubrium: dens: mucro as 5.54–9.71: 8.11–11.43: 1. Outer margin of basal segment of dens with 2–6 apically acuminate macrochaetae (sockets with arrows in Fig. 11). Inner edge of dens with well differentiated spines (spinal sockets shown in Fig. 11); spines on basal segment of dens forming 2–3 short and poorly organized rows with 1–3 large spines apically; spines on distal segment of dens forming a single row extending maximum 1 / 3 or 1 / 2 of length of distal segment of dens; proximal edge of distal segment of dens always with small spines intercalated between long spines, terminal one always largest long spine in row (dental sockets in Fig. 11). Dental formula variable, as 4–11 I–III /1–7 I 1–4 I(II) 0–2 I 0–2 I 0–2 I (Arabic numbers represent small spines; Roman numerals in bold and Italics represent large spines, on basal/distal segment of dens). Mucro with two basal, 0 intermediate and two distal teeth (202 formula) (Fig. 7). Variability. All constant and variable characters are given in Tables 1–2. Discussion. The new species is most similar to P. birsteini Djanashvili & Barjadze, 2011 by the combination of the following characters: 1) number of prelabral chaetae (2+2); 2) shape of tenent hair (capitate); 3) formula of tibiotarsal spine-like chaetae (002); 4) number of eyes—6 and 5) number of Mc on Th. II (Djanashvili & Barjadze 2011; Barjadze et al. 2020a). Both species are cave dwellers. The new species differs from P. birsteini by the number of Mc on the head and abdominal terga: 1) three postocular Mc on the head dorsum present in the new species, while 2 Mc in P. birsteini and 2) Abd. IV with six Mc in the new species, while P. birsteini has three (Barjadze et al. 2020a). Plutomurus birsteini was described from eleven caves located in five different karst massifs of western Georgia and molecular investigations (COI) show that P. birsteini is a complex of genetically distinct cryptic species (Barjadze, unpublished data). Etymology. The specific name originates from the type locality––Kharagauli district in which the Kozmani Cave is situated. Ecology. It is a troglophilous species, considering the presence of body pigment and 6+6 well-developed eyes. It occupies both twilight and dark zones of the cave. Distribution. This species is known only from a single cave. This cave is the remotest one from Zemo Imereti plateau karst massif and it is isolated from other known caves in the same karst massif (Tatashidze et al. 2009). The new species is the first known taxon from Kozmani Cave., Published as part of Barjadze, Shalva, Parimuchová, Andrea, Raschmanová, Natália, Maghradze, Eter & Kováč, Ľubomír, 2022, Two new species of Plutomurus Yosii (Collembola: Tomoceridae) from the Caucasus and central Europe, pp. 252-266 in Zootaxa 5169 (3) on pages 254-258, DOI: 10.11646/zootaxa.5169.3.2, http://zenodo.org/record/6952146, {"references":["Djanashvili, R. & Barjadze, S. (2011) A new species of the genus Plutomurus Yosii, 1956 (Collembola, Tomoceridae) from Georgian caves. Journal of Cave and Karst Studies, 73, 28 - 30. https: // doi. org / 10.4311 / jcks 2010 lsc 0147","Tatashidze, Z. K., Tsikarishvili, K. D. & Jishkariani, J. M. (2009) The Cadastre of the Karst Caves of Georgia. Petiti Publishing House, Tbilisi, 670 pp. [in Georgian]"]}

Plutomurus ruseki Barjadze, Kováč & Parimuchová sp. nov. Figs 16–28 Plutomurus sp. Barjadze et al. (2020b): 72 P. carpaticus Kováč & Krchová (2007): 85 P. carpaticus Kováč et al. (1999): 163 and 166 (Tab. 2) P. carpaticus Kováč et al. (2002): 163 (Tab. 3) P. carpaticus Kováč et al. (2007): 49 (Tab. 1) P. carpaticus Kováč et al. (2016): Appendix 1 P. carpaticus Lukáň et al. (2004): 170 P. carpaticus Papáč et al. (2020): 87 (Tab. 1) P. carpaticus Raschmanová et al. (2008): 463 and 469 (Tab. 3) P. carpaticus Raschmanová et al. (2013): 474 (Tab. 2) P. carpaticus Raschmanová et al. (2015): 806 (Fig. 2 and Appendix) P. carpaticus Raschmanová et al. (2018a): 253 and Appendix P. carpaticus Raschmanová et al. (2018b): 393 (Tab. 1) Type locality. SLOVAKIA, Muránska planina Plateau, Bobačka Cave, 680 m asl. Type material. Holotype: Female on slide, transition zone (20 m from small entrance), hand collecting, rotten wood, 05.x.2000, leg. Ľ. Kováč, Nr. 173-00 (UPJ). Type material is ordered alphabetically after geomorphological regions: Paratypes (n=42): Čierna hora Mts., Veľká ružínska jaskyňa Cave, 614 m asl, ca. 90 m from cave entrance, deep zone, 2 ex. on slides, pitfall trap, 10.viii.-14.x.1996, leg. Ľ. Kováč, Nr. 1327-96 (1 ex. in UPJŠ and 1 ex. in IZISU); Liptovská kotlina Basin, Liskovská jaskyňa Cave, 500 m asl, Guánová sieň, deep zone, 2 ex. on slides, extracted from bait (wood shavings), 16.iv.-14.ix.2004, leg. Ľ. Kováč, Nr. 481-04 (UPJŠ); ibidem: 50 m from cave entrance, transition zone, 1 ex. on slide, hand collecting from rotten wood, 14.ix.2004, leg. A. Mock, Nr. 491-04 (UPJŠ); Low Tatras Mts., Demänovská jaskyňa slobody Cave, 870 m asl, Sieň speleoterapie, transition zone (ca. 50 m from entrance), 1 ex. on slide, hand collecting on rotten wood, 27.ix.2000, P. Ľuptáčik, Nr. 127-00 (UPJŠ); Suchá Cave, 903 m asl, Predsieň, transition zone, 1 ex. on slide, hand collecting, rotten wood, 12.v.2011, leg. Ľ. Kováč and P. Ľuptáčik, Nr. 275-11 (UPJŠ); Slovak Karst, Jasovská Cave, 257 m asl, Komín, transition zone (50 m from show cave exit), 1 ex. on slide, hand collecting, dark cave sediment, 10.ix.1995, leg. Ľ. Kováč, Nr. 692-95 (UPJŠ); Silická ľadnica Ice Cave, 466 m asl, microdepression on a moderate scree slope in front of cave entrance, dense herbal cover, ass. Cardamino-Chrysosplenietum alternifolii, Lunario-Aceretum, well-developed soil profile, 2 ex. on slides, extracted from soil samples, 18.v.2005, leg. Ľ. Kováč, Nr. 110-05, 113-05 (UPJŠ); ibidem: 1 ex. on slide, extracted from soil samples, 11.v.2021, leg. N. Raschmanová, M. Marcin, (UPJŠ); ibidem: pioneer soil and vegetation on rocky debris, ass. Cardamino-Chrysosplenietum with sparse mosses and liverworts and dense growth of Chrysosplenium alternifolium, 1 ex. on slide, extracted from soil samples, 18.v.2017, leg. N. Raschmanová, Nr. 73-17 (UPJŠ); Stará brzotínska jaskyňa Cave, 258 m asl, entrance zone (~ 20 m from entrance), 2 ex. on slides, pitfall trap, 15.v.-08.vii.1998, leg. Ľ. Kováč, Nr. 296-98 (1 ex. in UPJŠ and 1 ex. in IZISU); ibidem: Veľká sieň, transition zone, 1 ex. on slide, pitfall trap, 12.iv.-06.ix.2010, leg. Ľ. Kováč, Nr. 426-10 (IZISU); ibidem: entrance zone (~ 10 m from entrance), 2 ex. on slides, hand collecting, cave wall/sediment, 2.vii.2021, leg. N. Raschmanová, J. Hankoščák, A. Parimuchová (UPJŠ); Zádiel Valley, 48°37.76’N, 20°49.81’E, 400 m asl, site on forested scree slope (E exposure, slope 35°) at the gorge bottom near the bank of Blatnica Creek (~10 meters), maple-lime wood (assoc. Aceri-Tilietum), mosses and sparse herbal cover, 1 ex. on slide, subterranean pitfall traps, depth of 65 cm, 19.x.2018 - 15.iv.2019, leg. N. Jureková, P. Ľuptáčik, A. Mock, Nr. 241-19 (UPJŠ); Spišsko-gemerský kras Karst, Slovak Paradise, Psie diery Cave, 942 m asl, dark zone, 1 ex. on slide, pitfall trap, 06.ii.1997, leg. V. Košel, Nr. 6/97 (UPJŠ); Vojenská jaskyňa Cave, 946 m asl, dark zone, 4 ex. on slides, pitfall trap, 28.i.-06.ii.1997, leg. V. Košel, Nr. 5/97 (UPJŠ); Tatry Mts., Belianske Tatry, Ľadová pivnica Cave, 1433 m asl, entrance zone, scree slope with coniferous forest soil and mosses, 2 ex. on slides, extracted from soil samples, 2.x.2018, leg. N. Raschmanová, Ľ. Kováč, M. Baňas, Nr. 305-18, 308-18 (UPJŠ); Západné Tatry Mts., Zadný úplaz Abyss, 1779 m asl, twilight zone, 3 ex. on slides, pitfall trap, 29.vi.-01.x.2010, leg. Ľ. Kováč, Nr. 715-10, 717-10 (UPJŠ); ibidem: twilight zone, 2 ex. on slides, pitfall trap, 29.vi.-01.x.2010, leg. A. Mock, Nr. 709-10 (UPJŠ); Nová Kresanica Abyss, twilight zone, small hall before Ženská priepasť, 1 ex. on slide, pitfall trap, 29.vi.-01.x.2010, leg. A. Mock, Nr. 721-10 (UPJŠ); Veľká Fatra Mts., Harmanecká Cave, 821 m asl, Izbica, entrance zone, 1 ex. on slide, pitfall trap, 07.v.-22.x.2002, leg. Ľ. Kováč, Nr. 187-02 (UPJŠ); ibidem: 1 ex. on slide, hand collecting, rotten wood, leg. P. Ľuptáčik, Nr. 202-02 (UPJŠ); Mažarná Cave, 830 m asl, entrance zone (15 m from entrance), 1 ex. on slide, hand collecting, surface of the sinter pool, 30.viii.1999, leg. Ľ. Kováč, Nr. 321-99 (IZISU); Volovské vrchy Mts., Jaskyňa nad kameňolomom Cave, 435 m asl, entrance zone, 2 ex. on slides, hand collecting, cave sediment and rotten wood, 21.vi.2021, leg. N. Raschmanová, Ľ. Kováč, M. Marcin, A. Parimuchová (UPJŠ); Kolónia II Cave, 807 m asl, dark zone, 3 ex. on slides, hand collecting, 02.iii.2011, leg. A. Mock, M. Rendoš, Nr. 18-11 (2 ex. in UPJŠ and 1 ex. in IZISU); Šarkania diera Cave, 690 m asl, dark zone, 2 ex. on slides, hand collecting on rotten wood, 29.iv.2005, leg. Z. Višňovská, Nr. 175-06 (UPJŠ). Additional material: Low Tatras Mts., pod Sinou, Acereto -Carpinetum, rendzina soil, 1 specimen on slide, 30.vi. 1960, leg. J. Rusek, Nr. 347 (ISB)–originally designated as paratype of Plutomurus carpaticus by Rusek & Weiner 1978;Vihorlat Mts., Okna creek, Fagetum carpaticum typicum, under stones, 1 specimen on slide, 23.vi.1959, leg. J. Rusek, Nr. 208 (ISB)—originally designated as paratype of P. carpaticus Rusek & Weiner 1978. Description. Body length 1.86–3.86 mm. Colour. Body grey colored. Scale distribution. Scales dorsally on Ant. I–II, head, Th. and Abd. segments, all leg segments, both sides of ventral tube and ventral side of furca. Head. Ratio of antenna as 1.50–3.23. Ant. III–IV completely annulated. Eye patch with six well developed eyes (Fig. 26). Head dorsally with one unpaired (A 0), and six paired Mc: two anterior (A 2, A 3), two interocular (S 2, S 4) and two postocular (Pa 3, Pa 5) distributed as in Fig. 26. Posterior margin of head with a row of mesochaetae. Prelabral and labral chaetae smooth: prelabral chaetae four (2+2) (Fig. 16); labrum with 554 papillate chaetae as typical for genus; distal margin of labrum with four elongate, thin–walled, flexible papillae. OML trifurcate, basal chaetae shorter than apical process; OML with four chaeta-like processes. Body. Dorsal bothriotrichal formula as 2,1/0,0,1,2,0 (Figs 27–28). Dorsal Mc formula: 5,2/3,3,4,3,2 or 3 (Figs 27–28). Thorax macrochaetotaxy as in Fig. 27. Th. II with three anterior (m 3, m 4, m 5) and two posterior (p 3, p 5) Mc; Th. III with two posterior (p 3, p 4) Mc. Abdominal macrochaetotaxy as in Fig. 28: Abd. I–II each with three long posterior Mc (m 2, m 3, m 4),; Abd. III with four long Mc: two anterior (m2, m6) and two posterior (p1, p6) and one short Mc (p7) as typical for genus; Abd. IV with three posterior Mc: two short (C6, T7) Mc and one very long and thick (E3) Mc, rarely one mesochaeta on A6 position; Abd. V with 2–3 long posterior Mc (p2, p3 or p2, p3 p4) and one long lateral Mc (Fig. 28). Legs. Hind legs with well–developed trochanteral (6–25 chaetae) and femoral (11–26 chaetae) organs (Fig. 17) with several elongated chaetae; tibiotarsal spine formula as 002, posterior side of tibiotarsus III with two outstanding inner Spine-like chaetae (arrows in Fig. 24). Tenent hair capitate (Fig. 18A). Ratio of claw III: Emp. III: tenent hair III as 1.21–2.23: 0.75–1.40: 1. Inner edge of claw on all legs with 2–4 teeth in total (b–c in Fig. 18A): one characteristically minute proximal unpaired tooth (a in Fig. 18A), one large unpaired tooth next to minute proximal unpaired tooth and 0–2 distal unpaired teeth, in total. Claw with long pseudonychia (Fig. 18A). Ratio of pseudonychium III: claw III inner edge as 0.32–0.54: 1. Unguiculus lanceolate, tapered, with 2 internal lamellae bearing 0–4 teeth (Fig. 18A and B). Ventral tube. anterior basal, posterior basal and apicolateral sides with 10, about 11 and 12 smooth chaetae, respectively (Fig. 22). Tenaculum. Corpus with one or rarely two smooth chaetae; rami with 4 + 4 teeth (Figs 19, 20). Furca. Ratio manubrium: dens: mucro as 2.56–5.93: 5.26–8.48: 1. Outer margin of basal segment of dens with 2–3 apically acuminate macrochaetae (Fig. 23d,e; sockets with arrows in Fig. 25). Inner edge of dens with well differentiated spines (Figs 23 a-c, 25); spines on basal segment forming 2–3 short and poorly organized rows with one or rarely two large spines apically; spines on distal segment forming a single row extending not more than half of distal segment’s length; basal edge of distal segment always with small spines intercalated between long spines, terminal one always largest long spine in row (Figs 23 a-c, 25). Dental formula is variable as 3–11 I-II /1–6 I(II) 1–4 I 0–4 I 0–3 I 0–2 I (Arabic numbers represent small spines; Roman numerals in bold and Italics represent large spines, on basal/distal segments of dens). Mucro with two basal, 0 intermediate (rarely one) and two distal teeth (formula 202 or 212) (Fig. 21). Variability. All constant and variable characters are given in Tables 1–2. Discussion. The following 6 species of Plutomurus have been reported from Europe (Barjadze et al. 2020b): P. baschkiricus (Skorikow, 1900), P. carpaticus Rusek & Weiner, 1978, P. jeleznovodski Kniss & Thibaud, 1999, P. kelasuricus Martynova, 1969, P. sorosi Kniss & Thibaud, 1999 and P. unidentatus (Börner, 1901). The new species is similar to P. baschkiricus (Skorikow, 1900), P. carpaticus Rusek & Weiner, 1978 and P. unidentatus (Börner, 1901) by the combination of the following characters: 1) number of prelabral chaetae (2+2); 2) shape of tenent hair (capitate); 3) formula of tibiotarsal spine-like chaetae (002) and 4) number of eyes (6) (Jordana et al. 2012; Barjadze et al. 2020b). The new species differs from P. baschkiricus by having well differentiated spines on dens, while P. baschkiricus has only narrow spines (Skorikow, 1900; Martynova 1969). The new species clearly differs from P. unidentatus by the number of Mc on the abdominal dorsum: 1) Abd. I with three Mc in the new species, while P. unidentatus has two on Abd. I and 2) Abd. IV with three Mc in the new species, while P. unidentatus has five on Abd. IV (Barjadze et al. 2020b). The new species is most similar to P. carpaticus, differing in the 1) number of postocular head Mc: two Mc in the new species, one in P. carpaticus and 2) number of Mc on Abd. II: three Mc in the new species, two in P. carpaticus (Barjadze et al. 2020b). Besides, based on mitochondrial COI gene, the pairwise genetic distance between P. carpaticus from Pieniny and P. ruseki sp. nov. is much greater than intraspecific distance in P. ruseki from two distant localities (Marcin et.al. unpublished data), thus confirming the new species status. P. ruseki sp. nov. also differs from P. carpaticus in habitat preference and geographic distribution. The new species prefers entrance and twilight zone of the caves (troglophilic species), while P. carpaticus is a surface form, found only in forest soil and mosses (Rusek & Weiner 1978 and our unpublished data), thus it may be considered as a trogloxene species. Moreover, P. carpaticus is geographically restricted to the surface habitats of Slovakian and Polish parts of the Pieniny Mountains, while the new species is known from subterranean habitats of several karst regions of Slovakia, and Western Carpathians. It is worth to mention that another possibly undescribed Plutomurus species was found on scree slope in the entrance zone of Silická ľadnica Ice Cave, which co-occurs with P. ruseki. It has head dorsum macrochaetotaxy as in P. ruseki and thoracic and abdominal dorsum macrochaetotaxy as in P. carpaticus. But further evaluation of this taxon will have to await the examination of individuals. Etymology. The species is named in the honour of the late prof. Dr. Josef Rusek (Biology Centre, v.v.i., Institute of Soil Biology, České Budějovice, Czech Republic), an outstanding Collembola taxonomist and ecologist, and Ph.D. supervisor and a great colleague of the last author. Ecology. It is a troglophilous species, considering the presence of pigmented body and 6+6 eyes. It lives mostly in entrance and twilight zone of caves, in several cases it was found to occupy even dark cave zone. It was also found to occupy forest soils and mosses. Distribution. This species is distributed in different karst regions of the Western Carpathians, Slovakia, with exception of the Pieniny Mountains. A Plutomurus specimen with 3 Mc on the Abd. I and II sampled in Perlynna Cave (Eastern Carpathians, Ugolka Karst Massif) in Ukraine, probably belong to this species (Robert Vargovitsh, personal communication)., Published as part of Barjadze, Shalva, Parimuchová, Andrea, Raschmanová, Natália, Maghradze, Eter & Kováč, Ľubomír, 2022, Two new species of Plutomurus Yosii (Collembola: Tomoceridae) from the Caucasus and central Europe, pp. 252-266 in Zootaxa 5169 (3) on pages 259-264, DOI: 10.11646/zootaxa.5169.3.2, http://zenodo.org/record/6952146, {"references":["Barjadze Sh., Kovac L. & Parimuchova, A. (2020 b) Contribution to the genus Plutomurus Yosii (Collembola, Tomoceridae): notes on the morphology of Plutomurus carpaticus Rusek & Weiner and P. unidentatus (Borner). Zootaxa, 4816 (3), 67 - 80. https: // doi. org / 10.11646 / zootaxa. 4816.1.3","Kovac, L. & Krchova, P. (2007) Collembolan communities (Hexapoda, Collembola) of the Krizova Cave, Medvedia Cave and Priepastova Cave in the Humenec Hill, Cierna hora Mts .. Slovensky kras (Acta carsologica Slovaca), 45, 79 - 92.","Kovac, L., Kosel, V. & Miklisova, D. (1999) Collembola (Hexapoda) of the associated with forest sites and caves. In: Tajovsky. K. & Pizl, V. (Eds.), Soil Zoology in Central Europe. ISB AS CR, Ceske Budejovice, pp. 16 l - 167.","Kovac, L., Hudec, I., Luptacik, P., Mock, A., Kosel, V. & Fenda, P. (2002) Spolocenstva kavernikolnych clankonozcov (Arthropoda) Demanovskych jaskyn. Abstract book of 3 rd conference Vyskum vyuzivanie a ochrana jaskyn, 2002, 155 - 164.","Kovac, L, Parimuchova, A. & Miklisova, D. (2016) Distributional patterns of cave Collembola (Hexapoda) in association with habitat conditions, geography and subterranean refugia in the Western Carpathians. Biological Journal of the Linnean Society, 119 (3), 571 - 592. https: // doi. org / 10.1111 / bij. 12555","Lukan, M., Rajecova, K., Kovac, L., Luptacik, P. & Mock, A. (2004) Predbezne vysledky prieskumu spolocenstiev terestrickych clankonozcov (Arthropoda) Jasovskej jaskyne. Abstract book of 4 th conference Vyskum vyuzivanie a ochrana jaskyn, 2004, 169 - 173.","Papac, V., Kosel, V., Kovac, L., Luptacik, P., Mock, A., Parimuchova, A., Raschmanova, N. & Visnovska, Z. (2020) Invertebrates of Dobsinska Ice Cave and Stratena cave system, Slovak Paradise, Slovakia. Slovensky kras (Acta carsologica Slovaca), 58 (1), 69 - 96.","Raschmanova, N., Kovac, L. & Miklisova, D. (2008) The effect of mesoclimate on Collembola diversity in the Zadiel Valley, Slovak Karst (Slovakia). European Journal of soil biology, 44, 463 - 472. https: // doi. org / 10.1016 / j. ejsobi. 2008.07.005","Raschmanova, N., Miklisova, D. & Kovac, L. (2013) Soil Collembola communities along a steep microclimatic gradient in the collapse doline of the Silicka ladnica Cave, Slovak Karst (Slovakia). Biologia, 68 (3), 470 - 478. https: // doi. org / 10.2478 / s 11756 - 013 - 0172 - 8","Raschmanova, N., Miklisova, D., Kovac, L. & Sustr, V. (2015) Community composition and cold tolerance of soil Collembola in a collapse karst doline with strong microclimate inversion. Biologia, 70 (6), 802 - 811. https: // doi. org / 10.1515 / biolog- 2015 - 0095","Raschmanova, N., Miklisova, D. & Kovac, L. (2018 a) A unique small-scale microclimatic gradient in a temperate karst harbours exceptionally high diversity of soil Collembola. International Journal of Speleology, 47 (2), 247 - 262. https: // doi. org / 10.5038 / 1827 - 806 X. 47.2.2194","Raschmanova, N., Sustr, V., Kovac, L., Parimuchova, A. & Devetter, M. (2018 b) Testing the climatic variability hypothesis in edaphic and subterranean Collembola (Hexapoda). Journal of Thermal Biology, 78, 391 - 400. https: // doi. org / 10.1016 / j. jtherbio. 2018.11.004","Rusek, J. & Weiner, W. M. (1978) Plutomurus carpaticus sp. n. (Collembola: Tomoceridae) from the Carpathian Mountains. Bulletin de l'Academie Polonaise des Sciences, Serie des Sciences Biologiques, 25 (11), 741 - 747.","Skorikow, A. S. (1900) One new Tomocerus species (Collembola) from the Eastern Russia. Annuaire du Musee Zoologique de l'Academie Imperiale des Sciences de Saint Petersbourg, 4, 473 - 480 [in German]","Kniss, V. & Thibaud, J. (1999) Genus Plutomurus in Russia and Georgia (Collembola, Tomoceridae). Revue francaise d'Entomologie, New Series, 21, 57 - 64. [in French]","Martynova, E. (1969) Springtails of the family Tomoceridae (Collembola) from the fauna of the USSR. Entomological Review, 43, 299 - 314. [in Russian]","Borner, C. (1901) About some new Collembolans from the caves of the area of Letmathe in Westphalia. Zoologische Anzeiger, 24, 333 - 345. [in German]","Jordana, R., Baquero, E., Reboleira, S. & Sendra, R. (2012) Reviews of the genera Schaefferia Absolon, 1900, Deuteraphorura Absolon, 1901, Plutomurus Yosii, 1956 and the Anurida Laboulbene, 1865 species group without eyes, with the description of four new species of cave springtails (Collembola) from Krubera-Voronya cave, Arabika Massif, Abkhazia. Terrestrial Arthropod Reviews, 5, 35 - 85. https: // doi. org / 10.1163 / 187498312 X 622430"]}

Western Palaearctic Plutomurus The following 16 Plutomurus species are known from the Western Palaearctic: P. abchasicus, P. baschkiricus, P. birsteini, P. carpaticus, P. eristoi, P. jeleznovodski, P. jordanai, P. kelasuricus, P. kharagauliensis sp. nov., P. ortobalaganensis, P. pichkhaiai, P. revazi, P. ruseki sp. nov., P. shurubumuensis, P. sorosi and P. unidentatus (Barjadze et al. 2016; 2018; 2020a, b). Here we provide an identification key to Western Palaearctic species of Plutomurus. Key to species of Plutomurus Yosii from the Western Palaearctic 1 Prelabral chaetae 2+2................................................................................. 2 - Prelabral chaetae 3+3................................................................................ 10 2 Spine-like chaetae on tibiotarsus I, II, and III as 001......................................................... 3 - Spine-like chaetae on tibiotarsus I, II, and III as 002......................................................... 4 3 Eye patch with 4–5 eyes; tenaculum with 1 chaeta........................... P. sorosi (Russia, North Caucasus: cave) - Eye patch with 6 eyes; tenaculum with 3–4 chaetae.................... P. jeleznovodski (Russia, North Caucasus: cave) 4 Dens with very narrow spines........................................... P. baschkiricus (Russia, Mt Urals: caves) - Dens with normal spines............................................................................... 5 5 Tenent hair acuminate........................................................... P. abchasicus (Georgia: soil) - Tenent hair capitate................................................................................... 6 6 Th. II with 4 Mc (medial & posterior)..................................................................... 7 - Th. II with 5 Mc (medial & posterior)..................................................................... 8 7 Head with 2 postocular Mc; Abd. IV with 3 Mc....................................... P. birsteini (Georgia: caves) - Head with 3 postocular Mc; Abd. IV with 6 Mc........................... P. kharagauliensis sp. nov. (Georgia: cave) 8 Abd. I and IV with 2 and 5 Mc respectively.................................................................................... P. unidentatus (Belgium, Bulgaria, France, Germany, Italy, Luxembourg and Slovenia: soil & caves) - Abd. I and IV with 3 and 3 Mc respectively................................................................ 9 9. Head with 1 postocular Mc; Abd. II with 2 Mc... P. carpaticus (Poland, Czech Republic, Slovakia, Ukraine, Romania: soil) - Head with 2 postocular Mc; Abd. II with 3 Mc............................. P. ruseki sp. nov. (Slovakia: soil & caves) 10 Tenent hair capitate............................................................... P. revazi (Georgia: caves) - Tenent hair acuminate................................................................................ 11 11 Spine-like chaetae on tibiotarsus I, II, and III as 002........................................................ 12 - Spine-like chaetae on tibiotarsus I, II, and III as 001........................................................ 14 12 Eyes absent; Th. II with 6 (anterior, medial & posterior) Mc...................... P. ortobalaganensis (Georgia: caves) - Eyes present; Th. II with 4 (medial & posterior) Mc........................................................ 13 13 Abd. IV with 1–2 Mc: large lateral and sometimes a second smaller medial Mc......... P. shurubumuensis (Georgia: cave) - Abd. IV with 3 Mc: 1 large lateral and 2 smaller medial Mc........................... P. pichkhaiai (Georgia: caves) 14 Eyes present and distinct; empodium with a large tooth.............. P. kelasuricus (North Caucasus and Georgia: caves) - Eyes absent or indistinct; empodium with small teeth....................................................... 15 15 Th. II with 5 Mc; one chaeta–like process on the OML is much shorter than others............ P. jordanai (Georgia: cave) - Th. II without Mc; all chaeta-like process on the OML normally developed.................... P. eristoi (Georgia: cave) * Specimens determined as P. carpaticus from Czech Republic, Ukraine and Romania needs to be reinvestigated., Published as part of Barjadze, Shalva, Parimuchová, Andrea, Raschmanová, Natália, Maghradze, Eter & Kováč, Ľubomír, 2022, Two new species of Plutomurus Yosii (Collembola: Tomoceridae) from the Caucasus and central Europe, pp. 252-266 in Zootaxa 5169 (3) on pages 264-265, DOI: 10.11646/zootaxa.5169.3.2, http://zenodo.org/record/6952146, {"references":["Barjadze, Sh., Baquero, E., Soto-Adames, F., Giordano, R. & Jordana, R. (2016) New diagnosis for species of Plutomurus Yosii (Collembola, Tomoceridae), with descriptions of two new species from Georgian caves. Zootaxa, 4126 (1), 77 - 96. https: // doi. org / 10.11646 / zootaxa. 4126.1.3","Barjadze, Sh., Jordana, R., Baquero, E., Giordano, R. & Soto-Adames, F. (2018) Two new species of Plutomurus Yosii (Collembola, Tomoceridae) from the Caucasus. Zootaxa, 4526 (1), 29 - 40. https: // doi. org / 10.11646 / zootaxa. 4526.1.2."]}

Key to species of Plutomurus Yosii from Europe 1 Prelabral chaetae 3+3; tenent hair acuminate....................................... P. kelasuricus Martynova, 1969 * - Prelabral chaetae 2+2; tenent hair clavate.................................................................. 2 2 Tibiotarsi with 001 spine-like chaetae..................................................................... 3 - Tibiotarsi with 002 spine-like chaetae..................................................................... 4 3 Eye patch with 4–5 eyes; tenaculum with 1 chaeta.................................. P. sorosi Kniss & Thibaud, 1999 - Eye patch with 6 eyes; tenaculum with 3–4 chaetae........................... P. jeleznovodski Kniss & Thibaud, 1999 4 Dens with very narrow spines.................................................. P. baschkiricus (Skorikow, 1900) - Dens with normal spines............................................................................... 5 5 Teeth on empodium present; Abd. I and IV with 2 and 5 Mc respectively................... P. unidentatus (Börner, 1901) - Teeth on empodium absent; Abd. I and IV with 3 and 3 Mc respectively.............. P. carpaticus Rusek & Weiner, 1978, Published as part of Barjadze, Shalva, Kováč, Ľubomír & Parimuchová, Andrea, 2020, Contribution to the genus Plutomurus Yosii (Collembola, Tomoceridae): notes on the morphology of Plutomurus carpaticus Rusek & Weiner and P. unidentatus (Börner), pp. 67-80 in Zootaxa 4816 (3) on pages 78-79, DOI: 10.11646/zootaxa.4816.1.3, http://zenodo.org/record/3954115, {"references":["Martynova, E. (1969) Springtails of the family Tomoceridae (Collembola) from the fauna of the USSR. Entomological Review, 43, 299 - 314. [in Russian]","Kniss, V. & Thibaud, J. (1999) Genus Plutomurus in Russia and Georgia (Collembola, Tomoceridae). Revue francaise d'Entomologie, New Series, 21, 57 - 64. [in French]","Skorikow, A. S. (1900) One new Tomocerus species (Collembola) from the Eastern Russia. Annuaire du Musee Zoologique de l'Academie Imperiale des Sciences de Saint Petersbourg, 4, 473 - 480 [in German]","Borner, C. (1901) About some new Collembolans from the caves of the area of Letmathe in Westphalia. Zoologische Anzeiger, 24, 333 - 345. [in German]","Rusek, J. & Weiner, W. M. (1978) Plutomurus carpaticus sp. n. (Collembola: Tomoceridae) from the Carpathian Mountains. Bulletin de l'Academie Polonaise des Sciences, Serie des Sciences Biologiques, 25 (11), 741 - 747."]}

Barjadze, Shalva, Kováč, Ľubomír, Parimuchová, Andrea (2020): Contribution to the genus Plutomurus Yosii (Collembola, Tomoceridae): notes on the morphology of Plutomurus carpaticus Rusek & Weiner and P. unidentatus (Börner). Zootaxa 4816 (3): 67-80, DOI: https://doi.org/10.11646/zootaxa.4816.1.3

Protaphorura cykini sp. nov. Parimuchová & Kováč (Figs. 2–16) Diagnosis. PAO with 65–71 simple vesicles. Pso formula dorsally: (2)32/022/33343, ventrally: 1/000/0 0 0 0 0. Subcoxae 1 with 1,1,1 pso each. Antennae longer than head (Fig. 2). Subapical organite on Ant. IV with two papillae (Fig. 3; 12). VT with 12–15 distal and 2–3 basal chaetae on each side. Type material. Holotype (female) and 12 paratypes (4 females, 8 males): Russia, Siberia, Irkutsk region, Pribaikalsky National Park, Primorsky Range, Okhotnichya Cave, collected by hand on mouldy excrements of pika (Ochotona sp.) (Fig.1a, site 1), 12.vi. 2016, leg. O.I. Kadebskaya and A. V. Osintsev. Holotype and paratypes herein designated are deposited in IBE FS UPJS, Košice, Slovakia. Other material. Russia, Siberia Irkutsk region, Pribaikalsky National Park, Primorsky Range, Okhotnichya Cave, 3 females and 1 male collected by hand on excrements of pika (Ochotona sp.), (Fig.1a site 2) 4.x.2014, leg. O.I. Kadebskaya and P. Holúbek. Other material from type locality deposited in IBE FS UPJS, Košice, Slovakia. Description. Body length 4.3–4.7 mm in males, 5.2–5.6 mm in females, body shape typical of genus (Fig. 1c), with anal spines on distinct papillae. Colour white in ethyl alcohol. Granulation fine and uniform, coarser around pseudocelli (Fig. 9). Antennae slightly longer than head (head/ antenna ratio 1:1.1), area antennalis well marked (Fig. 2). Ant. I with 14–16 chaetae. AOIII with 5 guard chaetae, 5 papillae, 2 smooth sensory rods shorter than papillae and 2 morel-like sensory clubs of equal size. Microsensillum in ventro-lateral position, at level of last guard chaeta. Subapical organite of Ant. IV placed in cavity protected by two granulated papillae, one slightly forward the organite and the second behind it (Fig. 3; 12). Ms on Ant. IV in ventro-lateral position, in 1/2 of segment length. PAO with 65–71 simple vesicles (Fig. 14). Labial palp of type A with 8 proximal chaetae. Basomedian field of labium with 4+4 chaetae, 9–12 postlabial chaetae with left/right asymmetry (Fig. 6). Maxillary outer lobe with simple palp, 1 basal and 2 sublobal chaetae. Pso formula dorsally: (2)32/022/33343, ventrally: 1/000/00000. Subcoxae 1 I–III with 1 pso each, psx absent. Ventral psx formula: 0/000/?111(2)00. Dorsal chaetotaxy more-less symmetrical with macrochaetae well differentiated. Th. I with 18–22 chaetae on each side. Chaetae s on Abd. I–III short, s´missing (Fig. 4). Abd. IV as in Fig. 7. Abd. V always with chaeta s between pso a–b, ratio of chaetae M/ s= 7/5 (Fig. 8). Straight lines passing through the bases of K and K’ prespinal chaetae slightly convergent (Fig. 11). Furca remnant with distinct, arched cuticular fold with 2+2 chaetae in two rows (1+1 placed on fold, 1+1 posterior to fold) (Fig. 16), arrangement of manubrial chaetae as in Fig. 10. Male ventral organ absent even in males with genital plate well developed (Fig. 15). VT with 12–15 distal and 2–3 basal chaetae on each side. Legs. Tita of all legs with 11 chaetae in distal verticil (A+T), 7 chaetae and chaeta M in verticil B and 7 chaetae in verticil C. Two proximal chaetae Y (placed above verticil C) present (Fig. 5). Claw without lateral and inner teeth (Fig. 13). Ratio length/ width of claw ~2.2. Etymology. The species is named after R. A. Cykin, an important personality in speleology of the Siberia. Biology. Specimens of P. cykini sp. nov. were discovered in Okhotnichya Cave, aggregated on mouldy pika´s excrements (Ochotona sp.), approximately 80 m from the entrance. The cave is mesotrophic with organic material (needles, leaf litter) accumulated near the entrance (Fig. 1b), and deer´s and elk´s bones and excrements of small mammals (Ochotona sp. and Microtus sp.) dispersed across the whole cave system. Fungi colonizing organic remnants and excrements probably serve as important food source for collembolans occupying the cave (Fig. 1d). P. cykini sp. nov. displays troglomorphic traits, i.e. elongated antennae and claws and larger body compared to other congeners. It lives permanently in cold thermal conditions; in June 2016 the cave air temperature measured at collecting sites ranged from +1.1 to +1.9 °C and the floor ice was present on the bottom of the cave passages. Along with the new species, the collembolan Oligaphorura schoetti was collected in Okhotnichya Cave in the same period. Distribution. Known only from the type locality: Okhotnichya Cave, Primorski Range, Irkutsk Region, Siberia, Russia. Discussion. The new species possess a unique combination of diagnostic features (Table 1). With the body length from 4.3 to 5.6 mm, long PAO consisting of 65–71 simple vesicles and subapical organite of Ant. IV surrounded by two cuticular papillae, P. cykini sp.nov. is a well-defined species. Subapical organite protected by the papillae is characteristic of P. tschernovi from the Western Taimyr, Siberia, P. borealis from eastern Europe (Babenko & Kaprus´2014), and P. nikolai from the Primorsky Krai, Far East (Kaprus´ et al. 2016). In contrast to P. cykini, these three species have body length P. borealis and P. nikolai differ from the new species also in number of pso on subcoxae 1 of all legs (Table 1). Presence of 60 or more simple vesicles in PAO was documented mostly in large species such as P. macrodentata, P. borealis, P. armata multituberculata and also in P. aksuensis, the taxonomical status of the last two species being unclear. P. janosik is similar to new species in lacking psx on ventral side of head but differ in number of chaetae on Th. I tergum and in lack of cuticular papillae in subapical organite on Ant. IV. Only few Palaearctic Protaphorura species have been described from caves, namely P. stalagmitorum (Absolon, 1900); P. armata multituberculata (Stach, 1934); P. teres (Yosii, 1956); P. ombrophila (Stach, 1960); P. triparallata (Gisin, 1960); P. dallaii (Nosek & Paoletti, 1981); P. brevispinata (Yosii, 1966); P. septempapillata (Palissa, 1986); P. janosik Weiner, 1990; P. aconae Arbea & Jordana, 1994 and P. zlatiborensis Lučić, Ćurčić, Pavković-Lučić & Tomić, 2008. However, the first six species are considered as species dubia due to insufficient morphology data in their original descriptions (Parimuchová & Kováč 2016). Length PAO Species Dorsal pso Ventral pso Psx SOp Th. I S-cox1 Distribution (mm) vesicles aksuensis (Martynova, 1972) 1.6 33/022/33333?? 90??? Kyrgyzstan armata multituberculata (Stach, 1934) 4.2 32/022/33343?? 54–62??? Westphalia, Germany borealis (Martynova, 1973, in Martynova 2.5–2.9 32/022/33333 0/- 1/000/ 111101m 40–65 + 14–18 0 0 0 pso + 111 psx Eastern Europe to Alaska al. 1973) sensu Babenko & Kaprus´ 2014) janosik Weiner, 1990 * 2.9–4.3 33/022–3/33343 1/- 0/000/0(1)0(1)0(1)000 36–58 - 14–17 111 pso + 0 0 0 psx Western and Eastern Carpathians macrodentata (Hammer, 1953) 2.4–3.4 32/022/33342 1/000/0001 1/000/ 111001m 50–60 - 25–30 111pso + 111 psx Canada, Chukotka sensu Babenko & Fjellberg (2016) Peninsula nikolai Kaprus´, Weiner & Paśnik 2016 1.5–1.7 33/022/33342 1/- 1/000/100000 29–36 + 11–12 100pso + 0 0 0 psx Primorsky Krai, Far East sayanica Kaprus´, Weiner &Paśnik 2016 2.7–2.9 32/022/33343 1/- 1/000/ 111101m 41–48 - 18–21 111pso + 111 psx southern Siberia tschernovi (Martynova, 1976) 1.8–2.3 32(3)/022/33343(2) 1/- 1/000/ 111101m 30–42 + 15–18 111pso + 111 psx Western Taimyr, Siberia sensu Babenko &Kaprus´(2014) cykini sp.nov. 4.3–5.6 (2)32/022/33343 1/- 0/000/?111(2)00 65–71 + 18–22 111 pso + 0 0 0 psx Primorski range, Siberia based on recent data (Parimuchová et al, 2017) of material from the type locality Totally 22 troglobiotic collembolan species have been registered in southern karst regions of Russia (Turbanov et al. 2016a, b, c), most of them distributionally limited to Crimea and Caucasus Mts that undoubtedly represent diversity hotspots of the cave fauna. The both regions are centered along the latitudinal band 42–45° N, corresponding by geographic position to the mid-latitude biodiversity ridge in Europe defined by Culver et al. (2006). Considering the number of biospeleologically surveyed caves in Russia, diversity of the cave-adapted Collembola in this country is still underestimated. Although, we cannot expect much addition by the further explorations in Siberian caves, since biodiversity of this vast region was repeatedly diminished during the cold Pleistocene glacial periods. The recent cave adapted fauna of Siberia probably evolved from the ancestors that occupied the soil or superficial subterranean habitats during the warmer Pleistocene periods. Accordingly, we consider the new species a glacial relict, supported by the morphological traits that do not show marked troglomorphisms. Molecular data. Obtained sequences are part of the COI gene at the 5´end corresponding to the position from 1564 to 2131 of the Drosophila yakuba mitochondrion (KF824900.1). We performed distance analysis by the Neighbor-joining distance tree (Fig. 17) based on the barcode sequences of the new species, P. borinensis Parimuchová & Kováč, 2016 and the species available in GenBank. Results strongly support the species differentiation. All the corresponding clusters have the highest bootstrap support, showing thus clear separation between the congeners (Hogg & Hebert 2004). Clustering of species analysed follows neither their geographic distribution nor morphological characteristics (Table 2). For example, interspecific K2P distances suggest that oligopseudocellar P. genheensis is the closest relative of P. cykini sp.nov. with the smallest genetic distance of 0.157 between the species involved in the analysis. P. janosik and P. borinensis, both from the Western-Carpathian caves, represent genetically well-separated species in spite of the low morphological differentiation (Parimuchová & Kováč 2016). interspecific intraspecific n 1 2 3 4 5 6 1 P. janosik 0.017 0.019 0.015 0.015 0.017 0.018 0.017 0.004 5 2 P. cykini sp.nov. 0.180 0.017 0.016 0.019 0.017 0.018 0.002 0.001 6 3 P. aurantiaca 0.184 0.192 0.018 0.019 0.018 0.017 0.011 0.003 2 4 P. genheensis 0.148 0.157 0.170 0.016 0.017 0.018 0.014 0.003 5 5 P. changbaiensis 0.142 0.201 0.193 0.170 0.017 0.019 0.001 0.001 6 6 P. maoerensis 0.166 0.180 0.192 0.171 0.178 0.017 0.008 0.003 3 7 P. borinensis 0.194 0.193 0.170 0.183 0.194 0.169 - - 1

Protaphorura borinensis sp. nov. (Fig. 1–13) Diagnosis. PAO with 36–43 simple vesicles. Pso formula dorsally: 33 /023/ 33343 (4), ventrally: 1 /000/00000. Subcoxae I with 1,1, 1 pso each. Coarse granulation on head with up to 8 granules between p 1 –p 1 chaetae. Abd. I– II with s and s' chaetae, s' always missing on Abd. III. Abd. V with chaeta s closest to pso c (Fig. 8). Verticil C of each Tita with 3 chaetae. On Abd. I pso a and b placed between macrochaetae p 2 and p 5. VT with 11–15 distal and 2–3 basal chaetae on each side. Type material. Holotype female and 14 paratypes (13 females, 1 male: Slovakia, Little Carpathians, Sedmička Cave, Veľký dóm Hall, collected by pitfall traps, 19.vi.– 28.xi. 2014, leg. A. Mock, P. Ľuptáčik, A. Parimuchová. Other material: Slovakia, Little Carpathians, Sedmička Cave, 6 females and 4 males, collected on walls and surface of water puddles on the cave sediment, 19.vi. 2014, leg. A. Parimuchová. Description. Body length from 2.7 mm (males) to 3.3 mm (females), body shape typical for the genus, with anal spines on distinct papillae (Fig. 1). Colour white alive and in ethyl alcohol. Granulation distinct and coarse especially on head, with up to 8 granules between p 1 chaetae on hind margin of head (Fig. 9). Antennae equally long as head, area antennalis well marked. Ant. I with 11–14 chaetae (mostly 12). AOIII with 5 guard chaetae, 5 papillae, 2 smooth sensory rods shorter than papillae and 2 morel-like sensory clubs of equal size. Microsensillum in ventro-lateral position, at level of last guard chaeta (Fig. 6 a, b). Subapical organite of Ant. IV placed in unprotected cavity. Ms on Ant. IV in ventro-lateral position, in 1 / 3 of segment length measured from base. PAO with 36–43 simple vesicles (Fig. 3). Labial palp of type A with mostly 7 (6–8) proximal chaetae (Fig. 10). Basomedian field of labium with 4 + 4 chaetae, 4 + 4 postlabial chaetae in symmetrical arrangement. Maxillary outer lobe with simple palp, 1 basal and 2 sublobal chaetae (Fig. 2). Pso formula dorsally: 33 /023/ 33343 (4), ventrally: 1 / 000/00000. Subcoxae I–III with 1 pso each. Ventral psx formula: 1 /000/ 11110, femur and subcoxae I with 1 psx each, femoral psx sometimes open and visible as larger area of ungranulated cuticle (Fig. 13). 1 pseudoporus on each subcoxa II, pseudopora on Th. I–III and Abd. II present but hardly visible. Dorsal chaetotaxy plurichaetotic, on Th. I variable, chaetotaxy type from i 3 m to 2 i 3 (2)m with modifications, mostly with 19–25 chaetae per half side of tergum (Fig. 10). Chaetae s and s' on Abd. I and Abd. II present, s' seldom missing. On Abd. I–II pso a and b located between macrochaetae p 2 and p 5 (Fig. 7). Abd. III without s', only pso a between macrochaetae. Abd. V always with chaeta s closest to pso c, and mostly also with chaeta s' and s" (Fig. 8); ratio of chaetae M/ s= 19 / 8. Straight lines passing through the bases of K and K’ prespinal chaetae distinctly convergent; chaeta K x sometimes present (Fig. 12). Furca remnant with distinct, arched cuticular fold with 2 + 2 chaetae in two rows (1 + 1 placed on fold, 1 + 1 posterior to fold), 4 chaetae in ma row, 4–5 chaetae in mm row and 5–7 chaetae in mp row (Fig. 5). Male ventral organ absent. VT with 11–15 distal and 2–3 basal chaetae on each side. Legs. Tita of all legs with 11 chaetae in distal verticil (A+T), 7 chaetae and chaeta M in verticil B and 3 (seldom 4) chaetae in verticil C. Proximal chaeta Y (placed above verticil C) present. Claw with 2 lateral teeth and 1 small inner tooth in the middle of claw length (Fig. 4). Ratio length/ width of claw ~2.0. Etymology. The species is named after the Borinský kras Karst, a small karst area near Bratislava, where the type locality (Sedmička Cave) is situated. Biology. Specimens of P. borinensis sp. nov. were discovered in Sedmička Cave, Borinský kras Karst, in the Little Carpathians that is the westernmost mountain range of the Western Carpathians. The cave appears to be oligotrophic without active underground stream, bat guano or other deposited organic materials. Specimens were distributed in hind parts of the cave distant from the entrance, collected on walls and surface of the water pools. The new species shows obvious affinity to subterranean environment, up to date it is known only from type locality in spite of intensive sampling in nearby caves of the same karst area. P. borinensis sp. nov. displays troglomorphic traits, i.e. slightly elongated antennae and claws. Distribution. Known only from type locality, Sedmička Cave, Little Carpathians, Slovakia. Discussion. The new species is similar to the Holarctic P. ar m a t a in dorsal and ventral pso formula (33 /023/ 33343, 1 /000/00000) and distinctly convergent straight lines passing through the bases of prespinal chaetae. It differs from P. armata by larger body, higher number of vesicles in PAO and plurichaetotic Th. I tergum (Table 1). The new species is the most similar to troglobiotic P. janosik Weiner, 1990, species endemic to the Western Carpathians, in number of vesicles in PAO (~ 40) and strong plurichaetosis on Th. I tergum especially. However, macrochaeta p 3, typical for P.janosik, is absent in the new species. Furthemore, P. borinensis sp. nov. differs from P. janosik by 11–15 distal chaetae on VT (18–22 in P. j a no s i k). Granulation, especially on head and Th. I, is more distinct in P. borinensis sp. nov. than in P. janosik; only 8 granules are present between p 1 chaetae in the new species compared to 12 in P. janosik (specimens from the type locality and several other populations from the Western Carpathians). Distinguishing between juveniles/subadults of P. janosik and P. borinensis sp. nov. can be problematic since subadult specimens of P. janosik often has indistinct macrochaeta p 3 and lower number of granules between p 1 chaetae. Anyway, cuticular fold of furca remnant in the new species is arched and well visible while it is very small and shallow in P. janosik Weiner, 1990. Specimens of the new species and specimens of five different populations of P. janosik (incl. the population from the type locality) were sequenced by COI locus. Maximum likelihood tree placed P. janosik and P. borinensis population in two different branches with strong statistical support showing a clear separation of analysed specimens of P. janosik and P. borinensis sp. nov. (Parimuchová et al., unpubl.). Morphology and molecular variability of P. janosik populations from the Western Carpathian caves will be shown in the next paper. Thus, we assume P. borinensis, despite being morphologically similar to P. janosik, as a new species. According to Pomorski (1990) the presence of chaeta s' on Abd. I affiliates the new species into common group together with P. s-vontoernei (Gisin, 1957) and P. pseudovanderdrifti (Gisin, 1957). The same author, when evaluating variability of abdomin s' is missing. If s' is present, then both pso a and b are demarcated by macrochaetae. In the new species, there is a new combination: both pso a and b are set between macrochaetae and chaeta s' is absent. The new species is also similar to P. octopunctata (Tullberg, 1876) in body length, length of antennae (equal to head), and variable tergal chaetotaxy with tendency to plurichaetosis. Nevertheless, there are specific characters separating the new species from others of “ octopunctata ” group characterized by area antennalis with 4 pso and parallel straight lines passing through the bases of prespinal chaetae. According to number of vesicles in PAO (~ 36), body length (~ 3 mm), 3 antero-dorsal and 3 postero-dorsal pso on head and presence of 2–3 pso on Th. II–III, we propose to assign the new species to “ multivesiculata ” group together with Protaphorura janosik, P. septempapillata and P. armata multituberculata.

Deuteraphorura sp. 1 D. cf. closanica Kováč et al. 2014 D. cf. closanica Kováč et al. in press. The specimens collected in Pružinská Dúpna jaskyňa Cave, Strážovské vrchy Mts, north-western Slovakia, are similar to Romanian obligate cave species D. closanica (Gruia, 1965) by dorsal pseudocellar formula (33/133/ 33353). Only few adult specimens are available (see material below) and the number of ventral pso is not clearly seen. This form probably represents a new troglobiotic species, however, more specimens are necessary for detailed morphology study. Material examined. Slovakia: Strážovské vrchy Mts, Pružinská Dúpna jaskyňa Cave, “ Hlavná chodba” passage, 2 specimens (1 female and 1 juvenile), on rotten wood, 17 Sep. 2012, leg. A. Parimuchová; ibid., terminal part of cave, 3 specimens (1 female and 3 juveniles), on surface of water puddle, 17 Sep. 2012, leg. A. Parimuchová; ibid. part behind the disphotic zone, 1 juvenile, pitfall trap, 27Apr.–17 Sep. 2012, leg. Ľ. Kováč.

Deuteraphorura kratochvili (Nosek, 1963) Figs 1–10, 14–15, Tab. 1 Onychiurus (Onychiurus) kratochvili Nosek, 1963: 77 Deuteraphorura cf. hussoni Kováč 1999: 68 Deuteraphorura cf. hussoni Kováč & Košel 1998: 67 Deuteraphorura cf. fimetaria Kováč 1998: 97 Deuteraphorura cf. fimetaria Kováč 1999: 68 Deuteraphorura cf. fimetaria Kováč 2000: 73 Deuteraphorura cf. kratochvili Kováč et al. 2004: 61 Deuteraphorura cf. kratochvili Kováč 2005: 91 Deuteraphorura cf. kratochvili Kováč et al. 2010: 143 (in fig.) Deuteraphorura cf. kratochvili Kováč et al. 2014: 124 Deuteraphorura cf. kratochvili Kováč et al. in press Deuteraphorura cf. kratochvili Barciová et al. 2010: 271 Deuteraphorura cf. kratochvili Košel 2012: 88 Deuteraphorura sp. Kováč et al. 2003: 35 Deuteraphorura sp.1 Kováč et al. 1999: 161 Deuteraphorura sp. 2 Kováč et al. 1999: 161 Diagnosis. Dorsal pso formula: 33/033/33354, ventral pso formula: 2-3/00(1)0(1)/2212, subcoxae 1 of legs I–III with 2, 2, 2 pso. Subcoxae 2 of legs I–III with 0, 3, 3 chaetae. Tita of legs I–III with 2 chaetae in C-verticil. Neotype, male. Slovakia: Low Tatras, Demänovská Cave System, Demänovská jaskyňa mieru Cave, entrance part, 1 male (Nr. 790-11), on surface of water in a container, 6.ix.2011, leg. A. Parimuchová and P. Ľuptáčik. Neotype herein designated, deposited in IBE FS UPJS, Košice.[Holotype male and allotype female, type locality: Low Tatras, Demänovská Cave System, Demänovská Ice Cave, in depth of 60 m, 15.viii.1961, leg. J. Nosek; type specimens were lost] Other material from type locality. Slovakia: Low Tatras, Demänovská Cave System, Demänovská Ice Cave, 5 females on permanent slides, 12.v.2000, leg. P. Ľuptáčik and Ľ. Kováč; Demänovská jaskyňa mieru Cave, “ Koncový sifón”, 2 specimens (1 male and 1 female), surface of water pools, 12.v.2011, leg. Ľ. Kováč; “ Chodba trosiek”, 2 females, on baits, 27.ix.2000, leg. P. Ľuptáčik; “at the exit”, 1 female, on baits, 27.ix.2000, leg. P. Ľuptáčik; Demänovská Slobody Cave, “ Za Veľkým dómom”, 2 females, on surface of water pools, 11.v.2000, leg. P. Ľuptáčik; Pustá jaskyňa Cave, “ Zrútená sieň” Hall, 1 male, on rotten wood, 9.ix.2015, leg. Ľ. Kováč; ibid., 1 male, on surface of a water pool, 9.ix.2015, leg. Ľ. Kováč; Suchá jaskyňa Cave, “ Vysoký dóm” Hall, 1 male, on surface of water pools, 7.ix.2011, leg. A. Parimuchová. Other material from type locality deposited in IBE FS UPJS, Košice. Other material. Slovakia: Low Tatras Mts, Jaskyňa mŕtvych netopierov Cave, entrance of “ Biela chodba” passage, 1 female, on mouldy guano, 8.ix. 2011, leg. A. Parimuchová. Kozie chrbty Mts, Važecká jaskyňa Cave, “Kamenný dóm” Hall, 1 female, on surface of a water pool, 23.vi.2015, leg. A. Parimuchová; ibid.,”Zrútená sieň” Hall, 3 females, on surface of water in a bucket, 23.vi.2015, leg. A. Parimuchová; Zápoľná jaskyňa Cave, “Sintrové jazierka”, 1 female and 1 male, on surface of sinter pools, 22.vi.2015 and 7.xi.2015, leg. P. Ľuptáčik, Ľ. Kováč, V. Papáč and A. Parimuchová. Pieniny Mts, Aksamitka Cave, internal parts of the cave, 2 females, on surface of cave pools, 26.v.1998, leg. A. Mock. Slovak Paradise, Duča Cave, “Veľká sieň” Hall, 2 females and 1 male, on bait, 7.xii.1998, leg. V. Košel; ibid: 1 female and 1 male, on rotten wood, 11.vii.2012, leg. A.Parimuchová; Stratenská jaskyňa Cave, “SNP” Hall, pitfall trap, 8.viii.1986 – 30.vii.1987, leg.V. Košel. Muránska planina Plateau, Bobačka Cave, “Čapík”, 1 female, on bat guano, 5.x.2000, leg. P. Ľuptáčik, and Ľ. Kováč; “Veľryba”, 1 female, pitfall trap and 1 female, bait extraction, 5.x.–9.xi.2000, leg Ľ. Kováč; ibid. 1 female, on surface of a water pool, 9.xi.2000, leg. Ľ. Kováč; Dielik Cave, middle part of entrance passage, 7 females and 3 males on stony debris with wood, 5.i.2010, leg V. Papáč; Zlatnica Cave, “Hall of SNP”, 5 females and 2 males on rotten wood, 21.v.2009, leg V. Papáč. Revúcka vrchovina Mts, Ochtinská aragonitová jaskyňa Cave, “Mliečna cesta”, 2 females, on rotten wood, 15.x.2003, leg. Lukáň; “Veľká sieň” Hall, 1 male, on surface of water pool, 15.x. 2003, leg.Ľ. Kováč; “Spojovacia chodba” Passage, 1 male, on rotten wood, 15.x.2003, leg. P. Ľuptáčik. Slovak and Aggtelek Karst, Nová Brzotínska jaskyňa Cave, 60 m from the entrance, 3 females, extracted from bait, 15.v.‒8.vii.1998, leg. Ľ. Kováč; Stará Brzotínska jaskyňa Cave, hind part of the cave, 3 males and 1 female, on surface of water pools, 6.v.2009, leg. Ľ. Kováč and P. Ľuptáčik; Marciho jaskyňa Cave, “Zadná sieň” Hall, 3 females and 2 males, on rotten wood, 22.x.1995; Ardovská jaskyňa Cave, “Zrútený dóm” Hall, 3 females and 1 male, on bat guano, 30.x.1997, leg. Ľ. Kováč; Jasovská jaskyňa Cave, hind part of the cave, 1 female, on thick layer of organic material, 10.x.1995, leg. Ľ. Kováč; ibid: “Husitská sieň” Hall, 1 female, extracted from a bait, 21.i.–3.iii.1999; Krásnohorská jaskyňa Cave, “Ábonyiho dóm” Hall, 2 females and 1 male, on surface of water pools, 12.ix.2003, K. Rajecová. Hungary: Slovak and Aggtelek karst, Vass-Imre barlang Cave (Hungary), passage to discovery entrance, 1 female, on marten excrements, 5.ix. 2013, leg. A. Mock. Other material deposited in IBE FS UPJS, Košice. Redescription. Body length 1.5–2.2 mm, shape cylindrical (Figs. 1, 2). Colour white in ethyl alcohol. Cuticular granulation rather fine and uniform. Antennae almost as long as head, area antennalis well marked. PAO with 14–17 compound vesicles. Ant. I with 8 chaetae in one row, Ant. II with 14 chaetae. AOIII with 5 papillae, 5 guard chaetae, 2 sensory rods shorter than papillae, 2 smooth sensory clubs and lateral ms. Apical organite simple in unprotected cavity. Lateral ms on Ant. IV placed basally, equally long as sensory rod of AOIII (Fig. 5). Labium of AB-type, with 6 proximal chaetae. Basomedian field with 4 chaetae. Maxillary outer lobe simple with 1 basal chaeta and 2 sublobal hairs, maxillary palp in apical third slightly serrated (Fig. 7). Labral formula as 4/142 (Fig. 9). Pso formula dorsally: 33/033/33354 (Fig. 2); head dorsally with 3 anterior pso arranged in triangle and 3 posterior; lateral pso poorly visible, located at hind margin of head, more distant from 2 axial pso. Pso formula ventrally: 2-3/000/2212 (Fig. 4); head ventrally with 1 anterior and 1 postero-lateral pso (Fig.3) or 1 anterior, 1 postero-central and 1 postero-lateral pso (Fig.10). Ventral psx formula on thorax and abdomen I–IV: /011/1010, psx on head, anal valve, subcoxae and femora not seen. Subcoxae 1 of I–III pairs leg with 2, 2, 2 pso. Th. I with 7–9 chaetae per half. Th. II–III with 4+4 axial chaetae and lateral ms. VT mostly with 6 chaetae per half; 4 in proximal and 2 in distal row, basal chaetae absent, chaetae on Th. I–III sterna absent. MVO present in form of thickened leaf-like chaetae: 4 on Abd. II sternum and 6–8 on Abd. III sternum (Figs. 8, 14). Furca remnant with 2+2 thin chaetae in one row, ma row with 2+2 chaetae in level of dental microchaetae, mm row with 3+3 chaetae and mp row with 2+2 chaetae, external ones as macrochaetae. Subcoxae 1 of legs I– III with 3, 4, 4 chaetae, subcoxae 2 with 0, 3, 3, coxae with 3, 10, 13, trochanters with 9 chaetae each and femora with 15, 15, 14 chaetae, respectively. Tita I with 6+M chaetae in B row (B1 absent) and 2 chaetae in C row, Tita II with 7+M chaetae in B-row and 2 chaetae in C-row, Tita III with 6+M chaetae in B row and 2 chaetae in C row. Claw slightly elongated without teeth, length/width ratio 2.4–2.8. Empodium with basal lamella, tip of filament reaching two-thirds of the claw length (Fig. 6). Variability. We investigated material of Deuteraphorura also from caves in southern karst areas of the Western Carpathians in Slovakia, which were listed in recent literature as D. cf. kratochvili (see references above). Populations from the Slovak and Aggtelek Karst differ slightly from those from type locality: 3 pso on ventral side of the head (instead of 2 pso), psx on Th. II–III sterna are often replaced by pso (Fig.10), 0, 4, 3 chaetae (instead of 0, 3, 3) are frequently registered on subcoxae 2, and only 1chaeta (instead of 2 chaetae) is often present in C row of Tita III. Based on examination of variability of these characters in numerous specimens from different caves and karst regions, we consider the observed differences as intraspecific variability. In specimens of the same population modified chaetae in MVO may differ in shape, from distinct leaf-like structures (Fig. 14) to simple thickened chaetae (Fig. 15), probably regarding developmental stage. Remarks. We have not studied the type material of D. kratochvili since its deposition is unknown (J. Hollier, Natural History Museum Geneva, pers. comm.), thus considered to be lost. The specification of the type specimens is following: Slovakia, Low Tatras, Demänovská Valley, Demänovská Ice Cave, 2 specimens (1 male and 1 female), 15.vii.1961, leg. J. Nosek. Any form similar to D. kratochvili has not been registered in the type locality. Thus, we consider freshly collected specimens used in this redescription to be identical to Nosek´s species. Compared with the original description (Nosek, 1963), we specified dorsal pso formula as 33/033/ 33354 (originally 32/033/33344), posterodorsal pso on head and lateral pso on Abd. IV are located more laterally from others. We also specified ventral pso formula as 2-3/000/ 2212 in contrast to 1/111/2220 (orig. description) that was incorrect due to misinterpretation of hardly visible ventral cuticular structures (psx, pseudopores) on thoracic sterna. Discussion. Totally 32 species of the genus Deuteraphorura lacking pseudocelli on Th. I have been described worldwide to date (Weiner & Fiera 2014). Of this species-group only D. rendsinae (Haybach, 1961) and D. doftana Weiner & Fiera, 2014 are characterized by the same dorsal pseudocellar formula (33/033/33354) as in D. kratochvili (Nosek, 1963). These species are distinguished by ventral pso numbers: 2/000/ 1112 in D. rendsinae, 3/ 011/ 3212 in D. doftana and 2-3/000/ 2212 in D. kratochvili as well as by number of vesicles in PAO: 15–17 in D.kratochvili, 11 in D. doftana and 8–10 in D. rendsinae (Tab. 1). Distribution. Distribution range of D. kratochvili covers several karst regions of the Western Carpathians in their central (Strážovské vrchy Mts, Veľká Fatra Mts, Považský Inovec Mts, Low Tatras, Kozie chrbty Mts, Slovak Paradise, Muránska planina Plateau) and southern parts (Revúcka vrchovina Mts, Slovak and Aggtelek karst). It also occupies Aksamitka Cave in the Pieniny Mts in the north of the Western Carpathians along with other troglobiotic collembolan species‒‒ Neelus koseli Kováč & Papáč, 2010.

Deuteraphorura schoenviszkyi (Loksa, 1967) Figs 11–13, Tab. 1 Onychiurus (Onychiurus) schoenviszkyi Loksa 1967: 292 Onychiurus schöenviszkyi sp. nov. Bajomi 1968: 121 Onychiurus schoenviszkyi Bajomi 1969: 243 Onychiurus schoeviszki Traser 1999: 49 Orthonychiurus schoenviszkyi Dányi & Traser 2007: 23 Orthonychiurus schoenviszkyi Dányi 2011: 423 Diagnosis. Dorsal pso formula: 33/033/33343, ventral: 2/000/0111, subcoxae 1 of legs I–III with 1 pso each. Subcoxae 2 of legs I–III with 0, 3, 3 chaetae, respectively. C-verticil of Tita I –III with 2, 1, 1 chaetae, respectively. Neotype, female. Slovakia: Slovak and Aggtelek Karst, Gombasecká jaskyňa Cave (approximately 15 km west from the type locality, in the same karst area), “ Mramorová sieň” Hall, 1 female (Nr. 920-09), collected on rotten wood, 1.x.2009, leg P. Ľuptáčik. Neotype herein designated, deposited in IBE FS UPJS, Košice [Type material not specified in the original description and lost; type locality: Hungary, Slovak and Aggtelek Karst, Kifli Shaft Cave]. Other material from type karst area. Slovakia: Slovak and Aggtelek Karst, Gombasecká jaskyňa Cave, “ Blatistá chodba” passage, 2 females, on surface of water pool, 22.x.1999, leg. Ľ. Kováč; ibid., “ Mramorová sieň” Hall, 1 female, on rotten wood, 3.v.2008, leg P. Ľuptáčik; Majkova jaskyňa Cave, “ Západná chodba” passage, 2 females, on bait, 15.v.1998, leg. Ľ. Kováč; Stará Brzotínska Cave, chamber 50 m from entrance, 1 male, on bait, 9.vii.1998, leg. Ľ. Kováč. Examined material deposited in IBE FS UPJS, Košice. Redescription. Body length 1.3–1.5 mm, shape cylindrical. Colour white in ethyl alcohol. Citicular granulation rather fine and uniform. Antennae almost as long as head, area antennalis well marked. PAO with 13– 14 compound vesicles. Ant. I with 8 chaetae in one row, Ant. II with 14 chaetae. AOIII with 5 papillae, 5 guard chaetae, 2 sensory rods shorter than papillae, 2 smooth sensory clubs and lateral ms. Apical organite simple. Lateral ms on Ant IV placed basally, equally long as sensory rod of AOIII. Labium of AB-type, with 6 proximal chaetae. Basomedian field with 4 chaetae. Maxillary outer lobe simple with 1 basal chaeta and 2 sublobal hairs. Maxillary palp in apical third slightly serrated. Labral formula as 4/142 (as in Fig. 9). Dorsal pso formula: 33/033/33343; head dorsally with 3 anterior pso arranged in triangle and 3 posterior; lateral pso hardly visible, located at hind margin of head, more distant from 2 axial pso. Ventral pso formula: 2/000/ 0111 (Fig. 13); head ventrally with 2 pso: anterior and postero-lateral pso. Subcoxae 1 of legs I–III with 1, 1, 1 pso respectively. Psx on sterna and legs not seen. Th. I with 7–9 chaetae per half, Th. II–III with 4+4 axial chaetae and 1 lateral ms. VT with 6–7 chaetae per half: 4 chaetae in proximal and 2–3 chaetae in distal row, basal chaetae absent (Fig. 12). MVO present in form of thickened chaetae: 4 on Abd. II sternum and 6 on Abd. III sternum (shape as in Fig. 15). Furca remnant with 2+2 thin chaetae in one row, ma row with 2+2 chaetae in level of dental microchaetae, mm row with 3+3 chaetae and mp row with 2+2 chaetae, external ones as macrochaetae. Subcoxae 1 of legs I–III with 3, 4, 4 chaetae, subcoxae 2 with 0, 3, 3 chaetae, coxae with 3, 10, 12–13 chaetae, trochanter with 9 chaetae each and femora with 15, 14–15, 14 chaetae, respectively. Tita I with 6+M chaetae in B row (B1 absent) and 2 chaetae in C row, Tita II with 7+M chaetae in B-row and 1 chaeta in C-row and Tita III with 6+M chaetae in B row and 1 chaeta in C row. Claw elongated without teeth, length/width ratio 2.4–2.9. Empodium with small basal lamella, tip of filament reaching two-thirds of the claw length (Fig. 11). Remarks. As in Deuteraphorura kratochvili, the type material of D. schoenviszkyi is inaccessible to study due to unknown place of its deposition (L. Dányi, Hungarian Natural History Museum in Budapest, pers. comm.), thus considered to be lost. Specification of the material is following: Hungary, Slovak and Aggtelek Karst, Kifli Shaft Cave, 4.viii.1965, leg. I. Loksa. The specimens of recently collected D. schoenviszkyi in caves of the Slovak and Aggtelek Karst differ from the original description (Loksa 1967) by higher number of pso on Th. II (3 vs. 2), ventral pso on head (2 vs. 1) and papillae in AOIII (5 vs. 4). We consider this form with only 1 pso on subcoxae and Abd. II –IV sterna identical to Loksa´s species in spite of the outlined differences that are most probably the result of inaccurate original description. Distribution. The species represent troglobiotic and endemic form with distribution range restricted to the Slovak and Aggtelek Karst situated in southern Slovakia and northeastern Hungary. Discussion. D. schoenviszkyi is the only representative of the species-group lacking pso on Th.I that has reduced number of pso on abdominal sterna (0,1,1,1) and subcoxae 1 of legs I-III (1,1,1). It is rather similar to D. kratochvili but clearly differs by dorsal (33/033/33343 vs 33/033/33354) and ventral (2/000/0111 vs 2-3/00(1)0(1)/ 2212 pseudocellar formula and the number of pseudocelli on subcoxae 1 (1,1,1 vs 2,2,2) (Tab. 1).

Protaphorura Absolon, 1901 Diagnosis of the genus. After Weiner (1996) and Pomorski (1998) the genus Protaphorura is characterized by following characters: hind margin of head with pso, PAO with numerous simple vesicles oriented perpendicularly to long axis of the organ in its central part, d0 on the head dorsally absent, ms on Ant. IV in the middle of posterior half of segment, Th. I without pso, Th. II–III with lateral ms, Abd. VI with anal spines on distinct papillae or with small spines without papillae, furca reduced to cuticular fold with 2 + 2 or 1 + 1 chaetae, distal verticil of Tita with 11 chaetae and male ventral organ present or absent. AOIII mostly with 5 papillae, some recently described species have only 4 papillae (Gulgenova & Potapov 2013). Labium is of A-type, i.e. with blunt tipped terminal sensillum on papilla A (Fjellberg 1998 b) but this character seems to be variable within the genus. For instance, P. janosik from Slovakia (caves) has all papillae with acuminate terminal sensilla.

Parimuchová, Andrea, Kováč, Ľubomír (2016): A new cave species of the genus Protaphorura Absolon, 1901 (Collembola, Onychiuridae) from the Western Carpathians (Slovakia) with critical comments to the Palaearctic representatives of the genus. Zootaxa 4098 (2): 254-272, DOI: http://doi.org/10.11646/zootaxa.4098.2.3

Parimuchová, Andrea, Kováč, Ľubomír (2016): Redescription of two troglobiotic species of Deuteraphorura Absolon, 1901 (Collembola, Onychiuridae) from the Western Carpathians. Zootaxa 4168 (2): 327-340, DOI: http://doi.org/10.11646/zootaxa.4168.2.6

Key to Palaearctic species of the genus Protaphorura Totally 85 species are included, the key is primarily constructed on papers of Pomorski (1998), Pomorski & Kaprus´(2007); Kaprus´& Pomorski (2008); Gulgenova & Potapov (2013), Babenko & Kaprus´(2014) and Kaprus´ et al. (2014), and on original descriptions of other Palaearctic species not provided in this literature sources. 1 Th. II with 0 or 1 pso.................................................................................. 2 - Th. II with 2 or 3 pso................................................................................. 17 2 Base of antennae with 3 pso............................................................................ 3 - Base of antennae with 4 pso........................................................................... 16 3 Subcoxae I of leg 1, 2, 3 with 1, 0, 0 pso respectively........................................................ 4 - Subcoxae I of all legs with 1 pso......................................................................... 5 - Subcoxae I of all legs without pso...................................................................... 10 4 Abd. III with 2 pso, dorsal pso: 32 /012/ 33232....................... P. changbainensis Sun, Zhang & Wu, 2013; China - Abd. III with 1 pso, dorsal pso: 32 /012/ 33132............................. P. brevispinata (Yosii, 1966); Korea, caves 5 Hind margin of head dorsally with 2 pso.................................................................. 6 - Hind margin of head dorsally with 3 pso.................................................................. 7 6 Ventral pso on head absent, dorsal pso: 32 /012/ 23132.............................. P. b o r z ic a (Gruia, 1999) Romania - Ventral side of head with 2 pso, dorsal pso: 32 /012/ 33342...... P. buryatica Gulgenova & Potapov, 2013; Russia (Buryatia) 7 Ventral pso on head present............................................................................. 8 - Ventral pso on head absent, PAO with 20–22 vesicles, dorsal pso: 33 /012/ 33233... P. fistania Thibaud & Peja, 1994; Albania 8 PAO with 28–32 vesicles, mvo absent, dorsal pso: 33 /012/ 33342............................................... 9 - PAO with 20–26 vesicles, mvo present on Abd. II–III, dorsal pso: 33 /012/ 33333.................................... ..................................................... P. christiani Pomorski, Leithner & Bruckner, 2003; Austria 9 Abd. II–IV without ventral psx................... P. taimyrica (Martynova, 1976) sensu Sun, Chang & Wu, 2015; China - Abd. II–IV with 1 ventral psx on each side.............................. P. genheensis Sun, Chang & Wu, 2015; China 10 AOIII with 5 papillae................................................................................ 11 - AOIII with 4 papillae................................................................................ 15 11 Head without ventral pso.............................................................................. 12 - Head with ventral pso................................................................................ 14 12 Th. II without dorsal pso: 32 /001/ 33232........... P. stiriaca (Stach, 1946) sensu Kaprus´, Paśnik & Weiner, 2014; Austria - Th. II with 1 dorsal pso............................................................................... 13 13 Th. III with 2 pso, dorsal pso: 33 /012/ 33332, mvo present on anal valves........ P. minima Sun, Zhang & Wu, 2013; China - Th. III with 1 pso, dorsal pso: 32 (3)/011/ 23232, mvo present on Abd. II–IV..................................................................................................... P. januarii (Weiner, 1977); Poland (Pieniny Mts.) 14 Head with 1 ventral pso, dorsal pso: 33 /01- 22 / 3334 (3) 2, mvo absent............... P. jacutica (Martynova, 1976); Russia - Head with 2 ventral pso, dorsal pso: 33 /01(2) 2 / 331 (2) 32, mvo present on Abd. II–III..................................................................................... P. bakhchisaraica Kaprus´, Paśnik & Weiner, 2014; Ukraine 15 Head with ventral pso, dorsal pso: 33 /012/ 33342.............. P. dzherga Gulgenova & Potapov, 2013; Russia (Buryatia) - Head without ventral pso, dorsal pso: 33 /012/ 33332........... P. dorzhievi Gulgenova & Potapov, 2013; Russia (Buryatia) 16 AOIII with 5 papillae, head with ventral pso, PAO with 36–40 vesicles, dorsal pso: 43 /012/ 332-343............................................................................... P. tolae Pomorski & Kaprus´, 2007; Russia (Yakutia) - AOIII with 4 papillae, head without ventral pso, PAO with 22–28 vesicles, dorsal pso: 43 /012/ 33353......................................................................... P. uniparis Gulgenova & Potapov, 2013; Russia (Buryatia) 17 Base of antennae with more than 3 pso................................................................... 18 - Base of antennae with 3 pso........................................................................... 39 18 Subcoxae 1 of legs without pso, pso dorsally: 4-63 - 4 /022/ 3334 - 53 - 5.............................................. ................................ P. octopunctata (Tullberg, 1876), sensu Pomorski & Kaprus´, 2007; Russia (Beringia) - Subcoxae 1 of legs 1,2, 3 with 1, 0, 0 pso................................................................. 19 - Subcoxae 1 of legs 1,2, 3 with 1, 1, 1 pso................................................................. 21 19 Abd. V with 2 pso, claw with strong (large) denticle, pso dorsally: 43 /022/ 33342.................................... ....................................... P. mongolica (Martynova, 1975) sensu Kaprus´& Pomorski 2008; Mongolia - Abd. V with 3 pso................................................................................... 20 20 Abd. IV with 5 pso, dorsal pso: 43 /022/ 33353............................ P. maoerensis Sun, Wu & Gao, 2013; China - Abd. IV with 4 pso, dorsal pso: 43 /022/ 33343..................... P. s a k a to i (Yosii, 1966); Kaprus´& Pomorski (2008); Poland, Ukraine, Moldova, Russia, Kyrgyzstan, Kazakhstan, Afghanistan 21 Th. II–III with variable number of pseudocelli (2–3, 2 – 3 respectively), head ventrally with 2 + 2 pseudocelli or abdominal ter- gum V with chaetae s' mvo absent....................................................................... 38 - Th. II–III with stable number of pseudocelli (2 or 3 symmetricaly), Abd. V without chaetae s'; if Abd. V with chaetae s', then PAO is small (18–26 simple vesicles), mvo present (Abd. II–IIIor Abd. II–IV; or VT) or absent...................... 22 22 Th. II–III with 3 + 3 and 3 + 3 pseudocelli respectively........................................................ 23 - Th. II–III with 2 + 2 and 3 + 3 pseudocelli respectively........................................................ 25 - Th. II–III with 2 + 2 and 2 + 2 pseudocelli respectively........................................................ 29 23 PAO with 32–36 vesicles, claws without denticle, pso dorsally 43 /033/ 33353...... P. daviesi (Bagnall, 1935); Great Britain - PAO with 18–26 vesicles, claws always with denticle....................................................... 24 24 Abd. I–III without chaeta s ´, straight lines passing through bases of short prespinal chaetae distinctly convergent, pso dorsally 4 (3) 4 /033/ 3335 (4) 3........................................................ P. pseudocellata (Naglitsch, 1962); Great Britain, Czech Republic, Germany, Poland, Ukraine, Moldova - Abd. I–III with chaetae s', straight lines passing through bases of short prespinal chaetae parallel, pso dorsally 4 (5) 3 (4,5)/033/ 4 (3) 4 (3) 4 (3) 5 (6) 3 (4).............................. P. s u bm er s a Kaprus´& Pomorski, 2008; Russia (Southern Siberia) 25 Head with 4 + 4 posterior pseudocelli, PAO with 34–40 vesicles................................................ 26 - Head with 3 + 3 posterior pseudocelli, PAO with 18–30 vesicles............................................... 27 26 Base of antennae and Abd. V with 5 pso, pso dorsally: 54 /023/ 454 (5-7) 5 (6) 5............................................................................................... P. decempunctata (Kos, 1939); Austria (Alps), Slovenia - Base of antennae and Abd. V with less than 5 pso, pso dorsally: 44 /023/ 34353............................................................................................... P. caledonica (Bagnall, 1935); Great Britain, Scotland 27 Abd. V dorsally with 2 pso, pso dorsally: 43 /023/ 33342, PAO with 18–22 vesicles, claws with denticle............................................................... P. nazarovensis Kaprus´& Pomorski, 2008; Russia (Southern Siberia) - Abd. V dorsally with 3 pso, claws without denticle......................................................... 28 28 Pso dorsally: 43 /023/ 33353, PAO with 16–22 vesicles, ms on Ant. VI in 1 / 4 from base................................................................................................. P. jiamusiensis Sun, Wu & Gao, 2013; China - Pso dorsally: 43 /023/ 4 (3) 4 (3) 4 (3) 53, PAO with 26–30 vesicles, ms on Ant. VI in 1 / 3 from base............................................................................ P. macfadyeni (Gisin, 1953) sensu Kaprus´& Pomorski 2008; Denmark, Finland, Germany, Iceland, Norway (Jan Mayen, Svalbard) Sweden, Japan (Hokkaido) 29 Abd. I–IV with 1 + 1 pseudocelli respectively, AOIII with 4 papillae, PAO with 22–27 vesicles, claws without denticle.............................................................. P. discrepans (Bagnall, 1948); Great Britain, Switzerland - Abd. I–IV with more than 1 + 1 pseudocelli................................................................ 30 30 Abd. IV without anterolateral pseudocelli (Abd. I–V with 3 + 3 pseudocelli respectively)............................ 31 - Abd. IV with anterolateral pseudocelli (Abd. I–V with variable number of pseudocelli)............................ 33 31 Head with 4 + 4 posterior pseudocelli, claws without denticle... P. suboctopunctata (Khanislamova, 1986); Russia (Bashkiria) - Head with 3 + 3 posterior pseudocelli, claws with denticle.................................................... 32 32 Abd. II–IV sterna with 1 + 1 parapseudocelli, abdominal tergum VI usually with 2 medial chaetae, VT with 13-19 + 13-19 chaetae............................................. P. quadriocellata (Gisin, 1947) sensu Pomorski & Kaprus´, 2007; Spain, Great Britain, Austria, Switzerland, Germany, Denmark, Norway, Slovakia, Afghanistan - Abd. II–IV sterna without parapseudocelli, Abd. VI with 1 medial chaeta, VT with 9 + 9 chaetae............................................................. P. saltuaria Pomorski & Kaprus´, 2007; Ukraine, Poland (Eastern Carpathians) 33 Sensory rods of antennal III sense organ are higher than accompanying papillae, the first one situated at external surface of first internal papilla, second one between third and fourth papillae (fig. 33); pso dorsally: 43 /022/ 33353 (4).................................................. P. nutak (Yosii, 1972) sensu Kaprus´& Pomorski 2008; Japan, China, Russia (Far East) - Sensory rods of AntIIIO shorter than accompanying papillae, typically arranged and set close together behind papillae in the middle part of the organ.............................................................................. 34 34 Abd. V with 5 pso, claw without inner tooth, pso dorsally: 43 /022/ 46655............ P. asensitiva (Stach, 1954); Slovenia - Abd. V with 2 pso, straight lines passing through bases of short prespinal chaetae parallel, claw with small tooth........ 35 - Abd. V with 3 or 4 pso, straight lines passing through bases of short prespinal chaetae convergent.................... 36 35 Head ventrally with 2 + 2 pso, mvo present on Abd. II–III, pso dosally: 42 /022/ 33332................................................................................... P. ianstachi (Yosii, 1972); sensu Kaprus´ et al. (2014); Georgia - Head ventrally with 1 + 1 pso, mvo absent, pso dosally: 43 /022/ 33342............................................................................................ P. licheniphila Kaprus´& Pomorski, 2008; Russia (Middle Siberia) 36 PAO with P. valsainensis (Acón, 1981); Spain - PAO with> 30 vesicles............................................................................... 37 37 AOIII with one morel-like and one sponge-like sensory clubs, macrochaetae retused and slightly knobbed, mvo absent, head ventrally with 1 pso, dorsal pso: 43 /022/ 33343........... P. tetragrammata (Gisin, 1961) sensu Kaprus´& Pomorski 2008; Bosnia and Herzegovina, Poland, Ukraine - AOIII with two morel-like sensory clubs, macrochaetae typical, mvo present on Abd. III–IV, head ventrally with 1 or 2 pso, dorsal pso: 44 (3)/022/ 3335 (4,6) 3................. P. eichhorni (Gisin 1954) sensu Weiner & Stomp (1995); Luxembourg 38 Head ventrally with 2 + 2 pso, Ant. IV with cauliflower-like papillae, PAO with 16–22 vesicles, dorsal pso: 43 /02(3) 3 (2)/ 3335 (4,6) 3 (4)................................................. P. merita Kaprus´& Pomorski 2008; Russia (Tuva) - Head ventrally with 1 + 1 pso, Ant. IV without cauliflower-like papillae, PAO with 36–40 vesicles, dorsal pso: 4 (5,6) 4 / 03(2) 3 (2)/ 4 (3) 4 (3) 4 (3,5) 5 (6) 4 (3)............................... P. elenae Kaprus´& Pomorski, 2008; Russia (Siberia) 39 Subcoxae 1 of leg 1,2, 3 with 1,0,0 pso respectively, furca rudiment with 1 + 1 chaeta, mvo present on VT; dorsal pso: 33 /022/ 33342............................................................ P. stogovi Pomorski, 1993; Russia (Karelia) - Sucoxae 1 of all legs without pso........................................................................ 40 - Sucoxae 1 of all legs with 1 pso each..................................................................... 47 40 Head ventrally without pso, PAO with 40–65 vesicles, dorsal pso: 32 /022/ 33333................................................ P. borealis (Martynova, 1973) sensu Babenko & Kaprus´, 2014; Russia, Siberia (also Novaya Zemlya, Taimyr) - Head ventrally with 2 pso, Abd. V without s', mvo present.................................................... 41 - Head ventrally with 1 pso............................................................................. 43 41 Abd. III with 2 pso, short chaetae above anal spines convergent or nearly so; ms on Ant. IV in 1 / 3 of segment length..... 42 - Abd. III with 3 pso, short chaetae above anal spines parallel, ms on Ant. IV in 1 / 2 of segment length; mvo present on Abd. II– III; dorsal pso: 32 /022/ 33332............................................ P. kopetdagi Pomorski, 1994; Turkmenia 42 Hind margin of head with 2 pso, claw with small tooth; mvo present on Abd. II–IV; dorsal pso: 32 /022/ 33232.......................................................... P. ajudagi Pomorski, Skarżyński & Kaprus´1998; Ukraine (Crimea) - Hind margin of head with 3 pso, claw with large tooth; mvo present on Abd. II–III; dorsal pso: 33 /022/ 3324 (3) 3............................................................ P. s a l s a Kaprus´, Paśnik & Weiner, 2014; Russia (South Siberia) 43 Abd. III with 2 pso, dorsal pso: 32 /022/ 33232.............................. P. microcellata (Dunger, 1978); Mongolia - Abd. III with 3 pso.................................................................................. 44 44 Abd. IV with 3 pso, short chaetae above anal spines convergent, Abd. V with or without s'......................... 45 - Abd. IV with more than 3 pso, short chaetae above anal spines parallel, Abd. V without s'.......................... 46 45 Dorsal pso: 33 /022/ 33333, psx on subcoxae and Abd. III–IV absent........................................................................... P. islandica (Bödvarsson, 1959) sensu Fjellberg 1998 a; Norway, Denmark, Iceland, Poland - Dorsal pso: 33 /022/ 33332, psx on subcoxae and Abd.III–IV present........................................................ P. bicampata (Gisin, 1956) sensu Sun, Zhang & Wu (2013); Fjellberg (1998 a); Denmark, Norway, Iceland, China 46 Abd. IV with 4 pso; Th. II with 2 pso; PAO with 28–35 vesicles; dorsal pso: 33 /022/ 33343 P. aurantiaca (Ridley, 1880) sensu Pomorski, 1998; Europe except arctic regions - Abd. IV with 5 pso; Th. III with 3 pso PAO with ≤ 30 vesicles, dorsal pso: 34 /023/ 33353...................................................................... P. pseudovanderdrifti (Gisin, 1957) sensu Pomorski (1998); Western Europe 47 Head ventrally without pso............................................................................ 48 - Head ventrally with 1 + 1 pso........................................................................... 49 48 AOIII with 4 papillae, Abd. IV with 4 pso, PAO with ~ 30 vesicles, dorsal pso: 33 /022/ 23343.......................... ............................................................... P. edentata (Kos, 1939); Slovenia (Julian Alps) - AOIII with 5 papillae, Abd. IV with 2 pso, PAO with ~ 40 vesicles, macrochaetae long, claws distinctly elongated; dorsal pso: 32 /022/ 22223........................... P. zlatiborensis Lucić, Čurcić, Pavković-Lučić & Tomić, 2008; Serbia, caves 49 Straight lines passing through bases of short chaetae above anal spines parallel................................... 50 - Straight lines passing through bases of short chaetae above anal spines convergent................................ 57 50 Abd. I–III with chaeta s', dorsal pso: 32 /022/ 3334 (3) 2...................................................................................... P. pjasinae (Martynova, 1976) sensu