REVIEW OF THE PALAEARCTIC PROTAPHORURA ABSOLON, 1901 SPECIES OF OCTOPUNCTATA GROUP (COLLEMBOLA: ONYCHIURIDAE)

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1 A N N A L E S Z O O L O G I C I (Warszawa), 2008, 58(4): REVIEW OF THE PALAEARCTIC PROTAPHORURA ABSOLON, 1901 SPECIES OF OCTOPUNCTATA GROUP (COLLEMBOLA: ONYCHIURIDAE) IGHOR KAPRUS 1 and ROMUALD J. POMORSKI 2 1 State Natural History Museum of NAS of Ukraine, Teatral na Str. 18, UA-79008, L viv, Ukraine; e-male: i-kaprus@mail.ru 2 Zoological Institute, Wrocław University, Sienkiewicza 21, PL Wrocław, Poland; onychus@biol.uni.wroc.pl Abstract. Protaphorura macfadyeni (Gisin, 1953) Protaphorura mongolica (Martynova, 1975) Protaphorura nutak (Yosii, 1972) and Protaphorura tetragrammata (Gisin, 1964) are redescribed based on the type specimens and new materials. Protaphorura rectopunctata Buşmakiu, 1996 and Protaphorura serbica (Loksa et Bagojevic, 1976) is a junior synonyms of Protaphorura sakatoi (Yosii, 1966). Protaphorura elenae sp. nov., Protaphorura licheniphila sp. nov., Protaphorura merita sp. nov., Protaphorura nazarovensis sp. nov. and Protaphorura submersa sp. nov. five new, closely related species from Siberia (Russia) are described. An key to all Palaearctic Protaphorura species with 4 and more pseudocelli at antennal base is provided. Key words. Collembola, Onychiuridae, Protaphorura, taxonomy, Palaearctic. INTRODUCTION The genus Protaphorura Absolon, 1901 is the most diverse within the tribe Onychiurini containing more than 120 species (Bellinger et al ). Species identification within this genus is mainly based on the pseudocellar formula the number and localisation of pseudocelli on different parts of body. Usage of this feature in identification keys allows distinguishing groups of species characterised by the same or similar number of pseudocelli. These groups are probably not natural, but have a great practical value. Group of species with four and more pseudocelli at antennal base is one of the richest among the genus Protaphorura. This species of octopunctatus group was erected by Pomorski and Kaprus (2007). In the same paper we redescribed the most known Protaphorura quadriocellata and Protaphorura octopunctata and described two new species, an identification key to known Palaearctic taxa of this group was also given. The material on several other species belonging to the same Protaphorura group, which can not be classify to any known species, is at our disposal. A considerable part of this material comes from Siberia thanks to the kindness of Russian collembologists Dr. Sophya Stebaeva, Anatoly Babenko and Mikhail Potapov. A minor part was collected by us, during our investigations of the Carpathians. The present paper is an attempt to give a critical review of all known Palearctic Protaphorura species with four and more pseudocelli at antennal base. It includes redescriptions of several species, descriptions of five new species and a identification key to all species of the group. PL ISSN Fundacja Natura optima dux doi: / X396602

2 668 I. KAPRUS and R. J. POMORSKI MATERIAL AND METHODS The group, mentioned above, contains many species which have been described long ago and needed modern redescriptions. That is why we tried to examine available type material of all species classified to this group. These species are listed below. ** the type material of the species has been studied, * we have investigated specimens presumable identical to the type material, the type material does not exist,? we have no information about the type material,! the species is well described or redescribed. Protaphorura asensitiva (Stach, 1954) Type locality: Slovenia, Julian Alps, Triglav Mt.;? Protaphorura caledonica (Bagnall, 1935) Type locality: Great Britain, Scotland, Cramond;? Protaphorura daviesi (Bagnall, 1935) Type locality: Great Britain, Scotland, N. Wales;? Protaphorura decempunctata (Kos, 1939) Type locality: Slovenia, Julian Alps, Triglav Mt.;? Protaphorura discrepans (Bagnall, 1948) Type locality: Switzerland, Beatus cave;! Protaphorura eichhorni (Gisin, 1954) Type locality: Luxembourg, Strassen (Kleepesch), beech forest; ** Protaphorura flavorufula (Martynova, 1976) Type locality: Russia, Jakutia, harbor Ambartchik near Medvezhka river; Protaphorura ianstachi Yosii, 1972 Type locality: Russia, Caucasus, ** Protaphorura macfadyeni (Gisin, 1953) Type locality: Jan Mayen Island, seashore; ** Protaphorura mongolica (Martynova, 1975) Type locality: Mongolia, Tüvshüülekh sum, Arkhangai aimag, in lichens; * Protaphorura nutak Yosii, 1972 Type locality: Japan, Hokkaido Island, summit of Poroshiri Mt.;! Protaphorura octopunctata (Tullberg, 1876) Type locality: Russia, North-East of Asia, delta of the Yenisei river near Dudino (at present Dudinka near Norilsk); ** Protaphorura pseudocellata (Naglitsch, 1962) Type locality: Germany;! Protaphorura quadriocellata (Gisin, 1947) Type locality: Switzerland, Langenthal, beech forest; ** Protaphorura rectopunctata Buşmakiu, 1996 Type locality: Moldova, satul Durlesti, raionul Ialoveni; ** Protaphorura sakatoi (Yosii, 1966) Type locality: Afghanistan, Karezmyl near Kabul;! Protaphorura saltuaria Pomorski et Kaprus, 2007 Type locality: Ukraine, East Carpathians, Chornoghora Mt., Vorochta, Piceetum forest; * Protaphorura serbica (Loksa et Bagojevic, 1976) Type locality: Croatia, Dubrovnik;? Protaphorura suboctopunctata (Khanislamova, 1986) Type locality: Russia, Bashkiria, Baltachevsk region, Bystry Tanyp river, floodplain forest; ** Protaphorura tetragrammata (Gisin, 1964) Type locality: Bosnia and Herzegovina, Igman, Malo Polje, Piceetum forest;! Protaphorura tolae Pomorski et Kaprus, 2007 Type locality: Russia, Jakutia, Suntar Khayata Mt., upper currents of Kyubyume river;! Protaphorura valsainensis (Acón, 1981) Type locality: Spain, Madrid, Sierra de Guaddarama. The studied material are deposited in the following institutions: IEEM Institute of Ecology and Evolution Russian Academy of Sciences, Moscow; MC Martynova s Collection in Moscow State Pedagogical University, Moscow; MNHG Museum of the Natural History, Geneve; SNHMU State Natural History Museum of Ukrainian National Academy of Sciences, L viv; ZIMAS Zoological Institute of Moldavian Academy of Sciences, Kishinev: ZIWU Department of Biodiversity and Evolutionary Taxonomy, Zoological Institute, Wrocław University, Poland. In our work we used the morphological nomenclature and standard of description after Pomorski (1998). Abbreviations: pso pseudocellar formula, psx parapseudocellar formula, chaetotaxy type on thoracic tergum I (i3m for example) (see Fig. 1), s and s chaetae on abdominal terga I III and V (see Fig. 1), p 0 chaeta and ratio M/s chaetae on abdominal tergum V (see Fig. 1). In our possession is also the material of several species, which we can not to classify it to any known species, also belonging to the group mentioned above. SPECIES DESCRIPTION Protaphorura elenae sp. nov. (Figs 1 5) Type material. Holotype: female, in detritus, Siberia, Russia, mouth of Kolyma River (left bank), ca. 60 km to the north from Pokhodsk [69 32 N, E], Sphagnum bog with Carex. lugens and Ledum sp., 19.VII.1994, leg. A. Babenko; paratypes: 4 males and 5 females, the same place as holotype.

3 REVIEW OF THE PALAEARCTIC PROTAPHORURA ABSOLON, 1901 SPECIES Figures 1 5. P. elenae sp. nov.: (1) habitus and dorsal chaetotaxy; (2) postantennal sense organ and anterior cephalic pseudocelli; (3) antennal III sense organ; (4) chaetotaxy of abdominal terga V and VI; (5) tibiotarsal chaetotaxy and claw of leg III.

4 670 I. KAPRUS and R. J. POMORSKI Other material examined. 3 males, Siberia, Russia, mouth of Kolyma River (right bank), Vysokaya Mt., [69 21 N, E], moss association, 19.VII.1994, leg. A. Babenko. The type material is preserved in the following collections: SNHMU holotype end 3 paratypes (2 males and 1 female); ZIWU 2 paratypes (1 unreproductive male and 1 female); IEEM 4 paratypes (1 male and 3 females) and other material. Etymology. The species is named in honour of an eminent Russian collembologist, Elena Martynova. Diagnosis. P. elenae sp. nov. shares the same number of dorsal pseudocelli (taking variability into account) with P. merita sp. nov., but the tips of its antennae are typical for Protaphorura species, claws with teeth and 1+1 pseudocelli on ventral side of a head. Contrary, P. merita sp. nov. is characterized by the presence of cauliflower like papilla on antennal tip, head with 2+2 ventral pseudocelli and toothless claws. Description. Color in alcohol yellowish white. Length without antennae: males mm, females mm. Body shape cylindrical, with small anal spines set on distinct papillae. Antennae approximately as long as head. Granulation more or less uniform, distinct. Base of antenna well marked. Antennal segment IV with subapical organite. Microsensillum on antennal segment IV in latero-external position, ca. 1 / 3 length from the base. Antennal segment III with microsensillum slightly below antennal III sense organ. Antennal III sense organ built of 5 guard chaetae, 2 sensory rods, 2 straight and granulated sensory clubs and 5 papillae (Fig. 3). Postantennal sense organ consists of simple vesicles (Fig. 2). Pseudocellar formula dorsally variable: 4(5,6)4/03 (2)3(2)/4(3)4(3)4(3,5)5(6)4(3); ventrally: 1/000/00000, all subcoxae1 with one pseudocellus. Formula of parapseudocelli ventrally: 1/000/100001? m ; on subcoxa1 parapseudocelli invisible. Position of pseudocelli is presented in Fig. 1. Dorsal chaetotaxy, usually with some asymmetry, well differentiated into macro- and microchaetae as in Fig. 1. Sensilla weakly marked. Thoracic terga II and III with microsensilla laterally. Thoracic tergum I with chaetae (chaetotaxy type i3 4m). Abdominal terga I III with chaetae s. Abdominal tergum V with s and p 0 chaetae (Fig. 4). Ratio M/s on abdominal tergum V = Abdominal tergum VI with one medial chaeta. Straight lines, passing through bases of short chaetae situated above anal spines, parallel (Fig. 1). Between legs on pro-, meso- and metathorax 1+1, 2+2 and 2+2 chaetae respectively. Ventral tube with 9+9 (seldom ) chaetae and 2+2 chaetae at the base. Furca reduced to cuticular pocket with 2+2 setulae 1+1 setulae located on a cuticular fold, remaining 1+1 setulae set located distinctly below of the fold. Claws with small denticle (Fig. 5). Empodial appendage without basal lamella, as long as inner edge of the claw. Tibiotarsi with distal whorl, composed of 11 chaetae. Anal spines distinctly shorter than inner edge of the claw ratio Male ventral organ is absent. Biology. P. elenae has been collected in moist green mosses and Sphagnum bog in the tundra zone. Bisexual. Remarks. Probably, P. elenae is the same as Protaphorura octopunctata sensu Martynova (1976). Listed as Protaphorura sp. A in Babenko and Fjellberg (2006). Protaphorura licheniphila sp. nov. (Figs 6 10) Type material. Holotype adult female and 12 paratypes (4 adult males and 8 adult females): Russia, Middle Siberia, Putorana Plateau, Dynkengda Mts., Sobachje Lake (Yt Kyuel ) [69 06 N, E], Laricetum forest, in soil and litter, 70 m alt., 20.VII.1997, leg. A. Babenko. The type material is preserved in the following collections: ZIWU holotype and 5 paratypes (2 males and 3 females); SNHMU 5 paratypes (2 males and 3 females); IEEM 2 females. Etymology. The name of species descends from the Latin word lichenes lichens and Greek word philos friend. Diagnosis. The same number of pseudocelli on body tergites (except of abdominal tergum V), the presence of pseudocelli on all subcoxae 1 and the absence of s chaetae on abdominal terga I III and V allow suggesting a close relationship between P. licheniphila sp. nov. and P. tetragrammata (Gisin, 1964) (see diagnosis of P. tetragrammata). These species distinctly differ in parapseudocelar formula, in ratio of M/s chaetae and in the number of pseudocelli on abdominal tergum V. Description. Color in alcohol white. Length without antennae: males mm, females mm. Body shape cylindrical, with anal spines set on distinct papillae. Antennae approximately as long as head. Granulation more or less uniform, fine, grains around pseudocellus. Base of antenna well marked. Antennal segment IV with subapical organite. Microsensillum on antennal segment IV in latero-external position, ca. 1 / 3 length from the base. Antennal segment III with microsensillum slightly below antennal III sense organ. Antennal III sense organ built of 5 guard chaetae, 2 sensory rods, 2 straight and granulated sensory clubs and 5 relatively short papillae (Fig. 7).

5 REVIEW OF THE PALAEARCTIC PROTAPHORURA ABSOLON, 1901 SPECIES Figures P. licheniphila sp. nov: (6) habitus and dorsal chaetotaxy; (7) antennal III sense organ; (8) postantennal sense organ and anterior cephalic pseudocelli; (9) tibiotarsal chaetotaxy and claw of leg III; (10) chaetotaxy of abdominal terga V and VI.

6 672 I. KAPRUS and R. J. POMORSKI Postantennal sense organ consists of simple vesicles (Fig. 8). Pseudocellar formula dorsally: 43/022/33342; ventrally: 1/000/00000, all subcoxa1 with one pseudocellus (Fig. 6). Formula of parapseudocelli ventrally: 1/000/ m, all subcoxa 1 with parapseudocellus ventrally. Dorsal chaetotaxy, usually with some asymmetry, well differentiated into macro-, meso- and microchaetae as in Fig. 6. Sensilla weakly marked. Thoracic terga II and III with microsensilla laterally. Thoracic tergum I with chaetae (chaetotaxy type i3m). Abdominal terga I III and V without chaetae s. Abdominal tergum V with chaeta p 0 present (Figs 6, 10). Ratio M/s on abdominal tergum V = Abdominal tergum VI with one medial chaeta. Straight lines, passing through bases of short chaetae situated above anal spines parallel (Fig. 10). There are 1+1, 2+2 and 2+2 chaetae between legs on pro-, meso- and metathorax respectively. Ventral tube with chaetae and 2+2 chaetae at the base. Furca reduced to shallow cuticular pocket with 1+1 setulae located distinctly on a cuticular fold. Claws always with small denticle (Fig. 9). Empodial appendage without basal lamella, as long as inner edge of the claw. Distal whorl of chaetae on tibiotarsi with 11 chaetae. Anal spines shorter than inner edge of the claw ratio Male ventral organ is absent. Biology. P. licheniphila was found in northern taiga in periodically drying lichen cushions, covering rocks. It was also numerous in one of the studied tundra regions (north bank of Taimyr Lake, foothills of Byrranga Mts.) characterized by the presence of many rocky outcrops. In all other tundra areas was rather rare. Listed as Protaphorura sp. B in Babenko and Fjellberg (2006). Bisexual. Protaphorura macfadyeni (Gisin, 1953) (Figs 11 15) Onychiurus macfadyeni Gisin, 1953: (230). Type locality. Jan Mayen Island, seashore. Material examined. Holotype dried specimen (details invisible), S9 Jan Mayen Island. 2 females (Pa1: S9) Jan Mayen Island. 1 female, 1 unreproductive male (Pa1: B11) Jan Mayen Island. 3 females, 1 unreproductive male (Jn10: North-Deutschland, litoral, 1953, leg. Strenzke (MHNG). Redescription. Color in alcohol white. Length without antennae: unreproductive males 1.3 mm, females mm. Body shape cylindrical, with strong anal spines set on distinct papillae. Antennae approximately as long as head. Granulation more or less uniform, distinct. Base of antenna well marked. Antennal segment IV with subapical organite. Microsensillum on antennal segment IV in latero-external position, ca. 1 / 3 length from the base. Antennal segment III with microsensillum slightly below antennal III sense organ. Antennal III sense organ built of 5 guard chaetae, 2 sensory rods, 2 straight and granulated sensory clubs and 5 papillae (Fig. 12). Postantennal sense organ consists of simple vesicles (Fig. 13). Pseudocellar formula dorsally variable: 43/023/4(3) 4(3)4(3)53; ventrally: 1/000/00000, all subcoxae1 with one pseudocellus. Formula of parapseudocelli ventrally: 1/000/ m, each subcoxae1 with one parapseudocellus. Position of pseudocelli dorsally is presented in Fig. 11. Dorsal chaetotaxy of adults poorly differentiated into macrochaetae and microchaetae, usually asymmetrical in submedial part of terga, as in Fig. 11. Sensilla weakly marked. Thoracic terga II and III with microsensilla laterally. Thoracic tergum I with chaetae. Abdominal terga I III and V without chaetae s. Abdominal tergum V with chaeta p 0 present (Fig. 15). Ratio M/s on abdominal tergum V = 2.1. Abdominal tergum VI with one medial chaeta. Straight lines, passing through bases of short chaetae situated above anal spines, parallel or subparallel (Fig. 15). Between legs on pro-, meso- and metathorax 0+0, 2+2 and 3+3 chaetae respectively. Ventral tube with 2+2 (3+3) chaetae at the base. Furca reduced to cuticular pocket with 2+2 setulae. Claws always without teeth. Empodial appendage without basal lamella, a little shorter than inner edge of the claw (Fig. 14). Distal whorl of tibiotarsi composed of 11 chaetae. Male ventral organ is absent. Remarks. P. macfadyeni belongs to the group of Protaphorura species with 2+2 and 3+3 pseudocelli on thoracic terga II and III respectively and with one pseudocellus on all subcoxae1. Within this group it is characterised by variable number pseudocelli on abdominal terga I III. Due to dorsal pseudocellar formula, ventral parapseudocellar formula and some chaetotactic characters P. macfadyeni is related to described below P. nazaroviensis sp. nov. (differences see also diagnosis of this species). Biology. P. macfadyeni lives in marine littoral wrack and beach meadows. Bisexual. Protaphorura merita sp. nov. (Figs 16 23) Type material. Holotype reproductive male, Russia, S. Tuva, ca km W of Samagaltay, southern foothills of East Tannu-Ola Mt. Range, between

7 REVIEW OF THE PALAEARCTIC PROTAPHORURA ABSOLON, 1901 SPECIES Figures P. macfadyeni (Gisin, 1953): (11) habitus and dorsal chaetotaxy; (12) antennal III sense organ; (13) postantennal sense organ and anterior cephalic pseudocelli; (14) tibiotarsal chaetotaxy and claw of leg III; (15) chaetotaxy of abdominal terga IV, V and VI.

8 674 I. KAPRUS and R. J. POMORSKI Figures P. merita sp. nov.: (16) habitus and dorsal chaetotaxy; (17) chaetotaxy and localization of parapseudocelli on ventral side of body.

9 REVIEW OF THE PALAEARCTIC PROTAPHORURA ABSOLON, 1901 SPECIES Figures P. merita sp. nov.: (18) antennomere IV, antennal III sense organ; (19) tip of antenna with cauliflower like papilla; (20) sensory clubs and sensory rods of antennal III sense organ; (21) postantennal ense organ and anterior cephalic pseudocelli; (22) distal part of leg III; (23) chaetotaxy of abdominal terga V and VI.

10 676 I. KAPRUS and R. J. POMORSKI Shevelig-Khem River and Aryskannyg-Khem River [50 40 N, E], in dry steppe petrophyte vegetation with Caragana bungei, Potentilla acaulis and Artemisia frigida, soil, 1100 m alt., 15.VII.1993, leg. S. Stebaeva; paratypes 7 males and 15 females, the same place as holotype. Other material examined. 3 males and 4 females, Russia, S. Tuva, 20 km W of settl. Erzin, basalt cliff Onchalaan [50 10 N, E], soil under Orostachys spinosa, 1300 m alt, 6.VII.2000, leg. S. Stebaeva. The type material is preserved in the following collections: ZIWU holotype and paratype (female); SNHMU 13 paratypes (4 males and 9 females); IEEM 5 paratypes (1 male and 4 females) and other material; MC 3 paratypes (2 males and 1 female). Etymology. The species name derived from the Latin word meritus meritorious, just, fair and is dedicated to Sophya Stebaeva, eminent Russian collembologist. Diagnosis. Among the group of examined Protaphorura species P. merita sp. nov. is characterised by presence of cauliflower-like papilla on tip of antenna, presence of 2+2 pseudocelli on ventral side of head and relatively small number vesicles in postantennal sense organ, but relationship of the new species is difficult to determine. According to pseudocellar formula it can be related with described above P. elenae sp. n. Description. Color in alcohol white. Length without antennae: males mm, females mm. Body shape cylindrical with strong anal spines set on distinct papillae. Antennae approximately as long as head. Granulation uniform and distinct with small areas of stronger granules located on frontal part of head, around of posterior cephalic pseudocelli, thoracic dorsal pseudocelli and near anal spines on end of abdomen. Base of antenna well marked. Antennal segment IV with a subapical organite and microsensillum in latero-external position, ca. 1 / 3 length from the base. Tip of antenna with cauliflower like papilla (Fig. 19). Antennal segment III with microsensillum slightly below antennal III sense organ. Antennal III sense organ built of 5 guard chaetae, 2 sensory rods, 2 straight and granulated sensory clubs and 5 papillae (Figs 18, 20). Postantennal sense organ consists of simple vesicles (Fig. 21). Pseudocellar formula dorsally variable: 43/02(3)3 (2)/3335(4,6)3(4); ventrally: 2/000/00000, all subcoxae1 with one pseudocellus. Formula of parapseudocelli ventrally: 0/000/ m, all subcoxae1 with parapseudocellus ventrally. Position of pso and psx is presented in Figs 16, 17. Dorsal chaetotaxy, usually with some asymmetry, well differentiated into macro-, meso- and microchaetae as in Fig. 16. Sensilla poorly marked. Thoracic terga II and III with microsensilla laterally. Thoracic tergum I with chaetae (chaetotaxy type i2 3m). Abdominal terga I III and V without chaetae s. Abdominal tergum V with p 0 chaeta present (Fig. 23). Ratio M/s on abdominal tergum V = Abdominal tergum VI with one medial chaeta. Straight lines, passing through bases of short chaetae situated above anal spines parallel or subparallel (Fig. 23). Between legs on pro-, meso- and metathorax 1+1, 2+2 and 2+2 chaetae respectively. Ventral tube with chaetae and 2+2 chaetae at the base. Ventral abdominal chaetotaxy as in Fig. 17. Furca reduced to cuticular pocket with 2+2 setulae 1+1 setulae located on cuticular fold, remaining 1+1 setulae located distinctly below of the fold. Claws always without denticle (Fig. 22). Empodial appendage without basal lamella, a little shorter than inner edge of the claw (ratio c. 0.95). Distal whorl of chaetae on tibiotarsi with 11 chaetae. Anal spines as long as claw or a little shorter (ratio ). Male ventral organ is absent. Biology. Protaphorura merita inhabits xerophytic habitats of Siberian steppes. Bisexual. Protaphorura mongolica (Martynova, 1975) comb. nov. (Figs 24 27) Onychiurus quadriocellatus mongolicus Martynova, 1975: (10). Type locality. Mongolia, Tüvshüülekh sum, Arkhangai aimag, in lichens. Type material. Holotype reproductive male, Mongolia, Tüvshüülekh sum, Arkhangai aimag, in lichens Hypogymnia bitteri on Larix sibirica, 1650 m alt., 28.VII.1970, leg. G. Medvedev (MC). Redescription. Color on slide yellowish white. Length without antennae 1.2 mm. Body shape cylindrical, with small anal spines set on distinct papillae. Antennae approximately as long as head. Granulation more or less uniform, distinct. Antennal segment IV with a subapical organite. Microsensillum on antennal segment IV in latero-external position, ca. 1 / 3 length from the base. Antennal III sense organ not seen. Postantennal sense organ consists of simple vesicles (Fig. 25). Pseudocellar (pso) dorsally: 43/022/ (Fig. 24), ventrally: 1/000/ Ventral parapseudocelli invisible (?). Subcoxae1 of I III pairs of legs with 1, 0, 0 pseudocelli and 1, 1, 1 parapseudocelli respectively. Dorsal chaetotaxy, more or less symmetrical, well differentiated into macro- and microchaetae as in Fig. 24. Sensilla weakly marked. Thoracic terga II and III

11 REVIEW OF THE PALAEARCTIC PROTAPHORURA ABSOLON, 1901 SPECIES Figures P. mongolica (Martynova, 1975): (24) habitus and dorsal chaetotaxy; (25) postantennal sense organ and anterior cephalic pseudocelli; (26) tibiotarsal chaetotaxy and claw of leg III; (27) chaetotaxy of abdominal terga IV, V and VI.

12 678 I. KAPRUS and R. J. POMORSKI with microsensilla laterally. Thoracic tergum I with 11 chaetae (chaetotaxy type i2m). Abdominal terga I III and V without chaetae s. Abdominal tergum V with p 0 chaeta present (Fig. 27). Ratio M/s on abdominal tergum V = 1.7. Abdominal tergum VI with one medial chaeta. Straight lines, passing through bases of short chaetae situated above anal spines, parallel (Fig. 27). Furca reduced to cuticular pocket with 2+2 setulae 1+1 setulae located on a cuticular fold, remaining 1+1 setulae located distinctly below of the fold. Claws with denticle (Fig. 26). Empodial appendage without basal lamella, shorter than inner edge of the claw (ratio 0.8). Distal whorl of tibiotarsi with 11 chaetae. Male ventral organ absent. Remarks. P. mongolica was described by Martynova (1975) on the base of one specimen from Mongolia as a subspecies of P. quadriocellata (Gisin, 1947). Examination of holotype let us to ascertain that this species isn t connected with P. quadriocellata, but he belongs to the group of Protaphorura species with 1,0,0 pseudocelli on subcoxae1 of all legs. It is closely related to P. sakatoi (Yosii, 1966) from which differs in presence of 2+2 pseudocelli on abdominal tergum V and in presence of strong inner denticle on claws (P. sakatoi has 3+3 pseudocelli on abdominal tergum V and no denticle, or rarely small denticle on claws). The species needs redescription on the base on new specimens. Protaphorura nazarovensis sp. nov. (Figs 28 31) Type material. Holotype 1 male on slide, Russia, Krasnoyarsk Territory, vicinity Nazarovo, undisturbed steppe meadow, near Melilotus albus Desr., soil, 20.VII.1988, leg. S. Stebaeva; paratypes 11 males and 24 females on slides, the same place as holotype. The type material is preserved in the following collections: SNHMU holotype and 20 paratypes (5 males and 15 females); ZIWU 7 paratypes (2 males and 5 females); IEEM 4 paratypes (2 males and 2 females) and other material; MC 4 paratypes (2 males and 2 females). Etymology. The species is dedicated to Nazarovo town (Siberia), near which it has been collected. Diagnosis. P. nazarovensis sp. nov. is probably the most related to P. macfadyeni (Gisin, 1953) due to the presence of one pso on subcoxae1 of all legs and the same number of dorsal pso on thoracic terga II III. These two species can be easily distinguished by psedocellar formulas on abdominal terga, numbers of vesicles in postantennal sens organ, and by the presence/ absence of denticle on claws (see also redescription of P. macfadyeni). Description. Color in alcohol white. Length without antennae: males mm, females mm. Body shape cylindrical, with relatively small anal spines set on distinct papillae. Antennae approximately as long as head. Granulation more or less uniform, distinct. Base of antenna well marked. Antennal segment IV with subapical organite. Microsensillum on antennal segment IV in latero-external position, ca. 1 / 3 length from the base. Antennal segment III with microsensillum slightly below antennal III sense organ. Antennal III sense organ built of 5 guard chaetae, 2 sensory rods, 2 straight and granulated sensory clubs and 5 papillae (Fig. 31). Postantennal sense organ consists of simple vesicles (Fig. 30). Pseudocellar formula dorsally: 43/023/33342; ventrally: 1/000/00000, all subcoxa1 with one pseudocellus (Fig. 28). Formula of parapseudocelli ventrally: 1/000/ m, all subcoxae1 with parapseudocellus ventrally. Dorsal chaetotaxy, usually with some asymmetry, well differentiated into macro- and microchaetae as in Fig. 28. Sensilla weakly marked. Thoracic terga II and III with microsensilla laterally. Thoracic tergum I with chaetae (chaetotaxy type i2m). Abdominal terga I III and V without chaetae s. Abdominal tergum V with p 0 chaeta present (Fig. 28). Ratio M/s on abdominal tergum V = Abdominal tergum VI with one medial chaeta. Straight lines, passing through bases of short chaetae situated above anal spines parallel. Pro-, meso- and metathorax with 1+1, and 2+2 ventral chaetae respectively. Ventral tube with chaetae and 2+2 chaetae at the base. Furca reduced to shallow cuticular pocket with 2+2 setulae 1+1 setulae located on a cuticular fold, remaining 1+1 setulae set located distinctly below the fold. Claws always with small denticle (Fig. 29). Empodial appendage without basal lamella, as long as inner edge of the claw. Distal whorl of chaetae on tibiotarsi with 11 chaetae. Anal spines as long as of inner edge of the claw. Male ventral organ is absent. Biology. The species has been collected on dry xerophytic meadow. Bisexual. Protaphorura nutak (Yosii, 1972) comb. nov. (Figs 32 36) Onychiurus (Protaphorura) longisensillatus nutak Yosii, 1972: (86). Type locality. Japan, Hokkaido Island, summit of Poroshiri Mt.

13 REVIEW OF THE PALAEARCTIC PROTAPHORURA ABSOLON, 1901 SPECIES Figures P. nazarovensis sp. nov.: (28) habitus and dorsal chaetotaxy; (29) distal part of leg III; (30) postantennal sense organ and anterior cephalic pseudocelli; 31) antennal III sense organ.

14 680 I. KAPRUS and R. J. POMORSKI Figures P. nutak (Yosii, 1972): (32) habitus and dorsal chaetotaxy; (33) antennal III sense organ; (34) postantennal sense organ and anterior cephalic pseudocelli; (35) remnant of furca; (36) tibiotarsal chaetotaxy and claw of leg III.

15 REVIEW OF THE PALAEARCTIC PROTAPHORURA ABSOLON, 1901 SPECIES 681 Material examined. Many adult specimens of both sexes: Russia, Primorskyi Kray, Khasan Region, Kedrovaya Pad Natural Reserve, detritus and soil, on stream, 29.IX.2004, leg. R. J. Pomorski; many adult specimens of both sexes: Russia, Primorskyi Kray, Shkotovo Region, Anisimovka, Livadiysky Range 2, [43 09 N, E], gravel and roots of plants in dry river bed (exhaustor), 12.IX.2004, leg. R. J. Pomorski. Redescription. Color in alcohol yellowish white. Length without antennae: males mm, females ) mm. Body shape cylindrical, with relatively strong anal spines set on distinct papillae. Antennae approximately as long as head. Granulation more or less uniform, distinct. Base of antenna well marked. Antennal segment IV with subapical organite. Microsensillum on antennal segment IV in latero-external position, ca. 1 / 3 length from the base. Antennal segment III with microsensillum slightly below antennal III sense organ. Antennal III sense organ built of 5 guard chaetae, 2 straight and granulated sensory clubs and 5 papillae, 2 very long sensory rods. Sensory rods of antennal III sense organ higher than accompanying papillae, one of the rods is located at external surface of first (internal) papilla, the second one set between third and fourth papillae (Fig. 33). Postantennal sense organ consists of simple vesicles (Fig. 34). Pseudocellar formula dorsally: 43/022/33353(4); ventrally: 1/000/ Formula of parapseudocelli (psx) ventrally: 1/000/ m. Each subcoxae1 with one pseudocellus and one parapseudocellus. Position of dorsal pso is presented in Fig. 32. Dorsal chaetotaxy, usually with some asymmetry, well differentiated into macro-, meso- and microchaetae as in Fig. 32. Sensilla weakly marked. Thoracic terga II and III with microsensilla laterally. Thoracic tergum I with chaetae (chaetotaxy type i3 4m). Abdominal terga I III and V without chaetae s. Abdominal tergum V with p 0 chaeta present (Fig. 32). Ratio M/s on abdominal tergum V = 1.4. Abdominal tergum VI with one medial chaeta. Straight lines, passing through bases of short chaetae situated above anal spines, parallel. Between legs on pro-, meso- and metathorax 1+1, 2+2 and 2+2 chaetae respectively. Ventral tube with chaetae and 2+2 chaetae at the base. Furca reduced to cuticular pocket with 2+2 setulae 1+1 setulae located on a cuticular fold, remaining 1+1 setulae located distinctly below of the fold (Fig. 35). Inner side of cuticular pocket with remnant of linea ventralis (Fig. 35). Claws with strong denticle (Fig. 36). Empodial appendage without basal lamella, as long as inner edge of the claw. Distal whorl of tibiotarsi with 11 chaetae. Anal spines shorter than inner edge of the claw (ratio 0.85). Male ventral organ absent. Remarks. The species was described from Hokkaido Island in Japan (Yosii 1972) as a subspecies Onychiurus (Protaphorura) longisensillatus nutak Yosii, 1972 and was separated from the nomina species by presence of 4+4 pseudocelli at the base of antenna (versus 3+3). We have examined numerous specimens from Russian Far East, identical with the original description, and we have ascertained that this character is very constant and therefore we come to conclusion, that they for sure represents separate species. P. nutak together with Protaphorura longisensillata (Yosii, 1969) and Protaphorura diplosensillata (Dunger, 1978) from Mongolia form a group of Protaphorura species marked by peculiar structure of the antennal III sense organ. Sensory rods are higher than accompanying papillae and their localisation is exceptional within the genus and even in all Onychiurinae tribe. The first one rod is located at external surface of base of first internal papilla, the second one is located between of third and fourth papillae. P. nutak have the same pseudocellar formula like P. valsainensis (Acon, 1981) from Spain, but details of structure of antennal III sense organ shows that it is similarity only. Biology. P. nutak appears to be one of the commonest onychiurid species collected in humid habitats along streams and rivers in the Russian Far East (Primorskyi Kray). In the Japan it inhabits high alpine belt of Poroshiri Mt. (Hidaka Mountains in the Hokkaido island). Bisexual. Protaphorura sakatoi (Yosii, 1966) Onychiurus sakatoi Yosii, 1966: (335). Protaphorura serbica (Loksa et Bagojevic, 1976) syn. nov. Protaphorura rectopunctata Buşmakiu 1995 syn. nov. Type locality. Afghanistan, Karezmyl near Kabul. Type material. Paralectotypes of Protaphorura sakatoi 2 females: Afghanistan, Karezmyl near Kabul, 16.VI.1960 (MNHG ); paratypes of Protaphorura rectopunctata Buşmakiu females: Moldova, village Durleşhti, soil under grapes culture, 4.IX.1992, leg. G. Buşmakiu (ZIMAS). Other material examined. 20 females, 12 males, Ukraine, Chornomorskyi Biosphere Reservation, Solenoozerna part, Kcherson district, steppe vegetation, soil, 1.V.2006, leg. I. J. Kaprus ; 2 females, 1 male, Ukraine, Striletskyi Step Reservation, Lugansk district, steppe with Elytrigia sp., soil, 19.V.1998, leg.

16 682 I. KAPRUS and R. J. POMORSKI O. Starostenko; 4 females, 1 male, Ukraine, Chomutovskyi Step Reservation, Donetsk district, steppe vegetation, soil, 6.VI.1993, leg. I. Bondarenko; 3 females, Ukraine, Dnipropetrovsk district, Kapitanovskyi Bajrak, Samara river, Quercus-Fraxinus-Tilia forest, litter, 25.VI.1985, leg. I. Vtorov; 13 females, 2 males, Ukraine, Medobory Reservation, Ternopil district, meadow steppe vegetation, soil, 24.V.1993, leg. I. J. Kaprus ; 3 females, Ukraine, Khmelnytsk district, Ustya village, Dnister river, meadow steppe vegetation on limestone, soil, 26.V.2005, leg. S. G. Bakajeva; numerous specimens, Ukraine, Crimea Peninsula, Nikitskyi pass, 1400 m alt., mountain meadow between stones, 12.IX.1997, leg. R. J. Pomorski, D. Skarzynski, I. Kaprus ; 4 females, 1 male, 2 juv.: Ukraine, Crimea Peninsula, Morskoje Village, [44 o 49 N 34 o 47 E], plant detritus on the stream, 10.IX.1997, leg. R.J. Pomorski; 3 females, 1 juv.: Kyrgyz Republic, Tian Shan Mts., Kyrgyskyi Mt. Range, Ala Archa National Park, 3200 alt., 19.IV.2001, soil and detritus near tree, leg. A. Pomorska; 6 females, 2 males: N-E Kazakhstan, Pavlodar district, settl. Matogul, steppe with Stipa sp., [53 43 N, E] 7.IX.1977, leg. S. Stebaeva; 2 females: Russia, Central Caucasus, valley of Baksan River, Elbrus village, ca m alt., [43 15 N, E], birch forest, moss on the stone, 23.IX.1999, leg. A. Babenko; 2 reproductive males, 2 females: Russia, Krasnodarskyi Territory, Adygea, 2000 m alt., birch forest, [43 50 N E, 1.IV.2004, leg. V. Bulavintsev; 3 females, 1 male: Russia, W. Siberia, S-W of Novosibirsk district, 17 km W of Karasuk, southern bank of Krotovaya Lyaga lake, soil, steppe, [53 43 N, E], 23.VI.1998, leg. S. Stebaeva; 2 unreproductive males, 1 female, 2 juv.: Russia, N Altai, settl. Cherga, [51 35 N, E], stone steppe, Sedum community, 6.VII.1988, leg. S. Stebaeva. Remarks. Detailed comparison of the type specimens of P. sakatoi and P. rectopunctata with rich material of P. serbica revealed lack of distinct differences in morphology among these three forms. The only difference is the larger variability of dorsal pseudocellar formula found in P. rectopunctata, shown by numerous asymmetric aberrations within a single population, and therefore we are considering hereby P. serbica and P. rectopunctata as junior synonyms of Protaphorura sakatoi. Examination of listed above materials shows a little broader range of variability of some morphological characters, not taken into consideration in the first redescription of P. serbica (Pomorski, 1998). Thus, ventral parapseudocellar formula of P. sakatoi in Europe (Pomorski, 1998) is as following: 1/000/1101(0)1. Asian populations of this species more often has parapseudocelli on abdominal sternum IV and a small denticle on claws, which is completely absent in European specimens. Biology. P. sakatoi lives only in xerophytic habitats. Probably it is connected with steppe plant communities of Europe and Asia. Protaphorura submersa sp. nov. (Figs 37 41) Type material. Holotype 1 female, Russia, Tuva, near settl. Erzin [50 16 N, E], 1100 m alt., floodplain forest with Populus laurifolia soil and litter, 1.VIII.1995, leg. S. Stebaeva; paratypes 12 females and 8 males, the same place as holotype. Other material examined. 3 females and 1 male, Russia, S-E Tuva, Sangilen Plateau, 8 km to settl. Moren [50 24 N, E], ca m alt., meadow steppe with Artemisia glauca, soil, 2.VII.2000, leg. S. Stebaeva; 7 females and 13 males, Russia, S-E Tuva, 8 km to setll. Moren, Larix sibirica forest, soil and litter, 1200 m alt., 8.VII.2000, leg. S. Stebaeva; 3 females, Russia, Tuva, East Tannu-Ola Mt. Range,??. 30 km NW of Khol -Oozhu, mountain Larix sibirica forest nearby Kara-Khol lake, 1700 m alt., [50 55 N, E], 16.VII.1993, leg. S. Stebaeva; 5 females and 2 males, Russia, S-E Tuva, ca. 30 km NW of Erzyn, Sangilen Plateau, Khorumnug-Taiga Mt. Range, right bank of Bayan-Kol river, Larix sibirica forest with Rhododendron dahurica, soil and litter, 1200 m alt., 6.VII. 1995, leg. S. Stebaeva; 3 females, Russia, W. Sayany Mts., Oiskiy Mt. Range, vicinity meteostation Olenya Rechka, mountain tundra, moss Sphagnum and Carex sp., ca m alt., [52 48 N, E], 27.VI. 1990, leg. S. Stebaeva. The type material is preserved in the following collections: SNHMU holotype and 8 paratypes (3 males and 5 females) end other material; ZIWU 3 males and 5 females; IEEM 2 males, 2 females and other material; MC other material. Etymology. The species name derived from the Latin word submergo I am sinking, I am immersing. Diagnosis. Because of the presence of pseudocelli on subcoxa of all legs and 3+3 pseudocelli on thoracic terga II-III P. submersa sp. n. is the most similar to P. pseudocellata. The new species distinctly differs from the latter having s chaetae on abdominal terga I III and different arrangement of microchaetae above anal spines. Straight lines, passing through bases of these chaetae are parallel in P. submersa sp. n. and convergent in P. pseudocellata. Description. Color in alcohol white. Length without antennae: males mm, females mm. Body shape cylindrical, with relatively small anal spines set on distinct papillae. Antennae approximately as long as head. Granulation more or less uniform, distinct. Base of antenna well marked. Antennal segment IV with a subapical organite. Microsensillum on antennal segment IV in latero-external

17 REVIEW OF THE PALAEARCTIC PROTAPHORURA ABSOLON, 1901 SPECIES Figures P. submersa sp. nov.: (37) habitus and dorsal chaetotaxy; (38) antennal III sense organ; (39) postantennal sense organ and anterior cephalic pseudocelli; (40) tibiotarsal chaetotaxy and claw of leg III; (41) chaetotaxy of abdominal terga V and VI.

18 684 I. KAPRUS and R. J. POMORSKI position, c.1/3 length from the base. Antennal segment III with microsensillum slightly below antennal III sense organ. Antennal III sense organ built of 5 guard chaetae, 2 sensory rods, 2 straight and granulated sensory clubs and 5 papillae (Fig. 38). Postantennal sense organ consisting of simple vesicles (Fig. 39). Pseudocellar formula dorsally is highly variable: 4(5)3(4-5)/033/4(3)4(3)4(3)5(6)3(4); ventrally: 1/000/00 000, all subcoxae1 with one pseudocellus (Fig. 37). Formula of parapseudocelli (psx) ventrally: 1/000/ m, all subcoxae1 with parapseudocellus ventrally. Dorsal chaetotaxy, usually with some asymmetry, well differentiated into macro- meso-and microchaetae as in Figs 37, 41. Sensilla well marked. Thoracic terga II and III with microsensilla laterally. Thoracic tergum I with chaetae (chaetotaxy type i3m). Abdominal terga I III with chaetae s. Abdominal tergum V with or rarely without-s chaetae, p 0 chaeta present (Fig. 41). Ratio M/s on abdominal tergum V = Abdominal tergum VI with one medial chaeta. Straight lines, passing through bases of short chaetae situated above anal spines parallel (Fig. 41). Between legs on pro-, mesoand metathorax , 2+2 and 2+2 chaetae respectively. Ventral tube with chaetae and 2+2 chaetae at the base. Furca reduced to shallow cuticular pocket with 2+2 setulae 1+1 setulae located on a cuticular fold, remaining 1+1 setulae located distinctly below of the fold. Claws always with denticle (Fig. 40). Empodial appendage without basal lamella, as long as inner edge of the claw. Distal whorl of chaetae on tibiotarsi with 11 chaetae. Anal spines shorter than inner edge of the claw (ratio ). Male ventral organ is absent. Biology. The species lives in warm, wet, dry and sandy places in highlands and lowlands of Siberia. It has been collected in forests and meadow steppe. Bisexual. Protaphorura tetragrammata (Gisin, 1964) (Figs 42 48) Onychiurus tetragrammatus Gisin, 1964: (7) Type locality. Bosnia and Herzegovina, Igman, Malo Polje, Piceetum forest. Type material. Holotype reproductive male (slide No 10) Igman, Malo Polje, 1160 m alt. Piceetum excelsae; Bosnia and Herzegovina; 6.VII.1961; 2 paratypes (reproductive males); the same data as holotype (MNHG). Other material examined. 4 males and 11 females, Poland, Bieszczady Mts., Sękowiec village, Fagus-Abies forest, soil and litter, 1.X.1996, leg. I. J. Kaprus ; 1 female, Poland, Bieszczady Mts., Muczne village, Fagus-Abies forest, soil and litter, 30.IX.1996, leg. I. J. Kaprus ; 1 male, Poland, Bieszczady Mts., Pszczeliny village, Fagus-Abies forest, soil and litter, 30.IX. 1996, leg. I. J. Kaprus ; 2 males, Poland, Bieszczady Mts., Hylaty village, Fagus-Abies forest, soil and litter, 1.X.1996, leg. I. J. Kaprus ; 1 male and 3 females, Ukraine, Verchnio-Dnistrovs ki Beskydy Mts., Lybochora village, Turkivs kyi region, Fagus forest, soil and litter, 8.VIII.1993, leg. I. J. Kaprus (ZIW and SNHMU). Diagnosis. P. tetragrammata belongs to the group of Protaphorura species with pseudocelli on upper subcoxa of all legs, 2+2 pseudocelli on both II and III thoracic terga and with anterolateral pseudocelli on abdominal tergum IV. Among this group is characterised by presence distinctly differentiated macroand mesochaetae, which are apically retused and a little knobbed. P. tetragrammata is closely related with described above P. licheniphila sp. nov. (differences see diagnosis of P. licheniphila). Redescription. Color in alcohol white. Length without antennae: males mm ( mm according original description). Body shape cylindrical, with strong anal spines set on distinct papillae. Antennae approximately as long as head. Granulation more or less uniform, distinct. Base of antenna well marked. Antennal segment IV with a subapical organite. Microsensillum on antennal segment IV in latero-external position, at level of second row of posterior chaetae. Antennal segment III with microsensillum slightly below antennal III sense organ. Antennal III sense organ built of 5 guard chaetae, 2 sensory rods, 2 similarly sized sensory clubs: one is morel-like distinctly granulated, the other sponge-like and 5 papillae (Figs 45, 46). Postantennal sense organ consists of simple vesicles (30 40 simple vesicles in material from Ukraine and Poland) (Fig. 44). Pseudocellar formula dorsally variable: 43/022/ 33333(4)3 [4(3)3/022/33343 in material from Poland]; ventrally: 1/000/00000, all subcoxae1 with one pseudocellus and one parapseudocellus. Formula of parapseudocelli ventrally: 1/000/111100? m. Position of pso dorsally is presented in Fig. 47. Dorsal chaetotaxy very distinctly differentiated into apically retused and slightly knobbed macrochaetae and pointed mesochaetae (Fig. 48). Microchaetae very short, apically pointed. Generally chaetotaxy symmetrical, but some asymmetries in submedial part of terga and abdominal tergum VI are visible, as in Figs. 42, 47. Sensilla weakly marked. Thoracic terga II and III with microsensilla laterally. Thoracic tergum I with chaetae (chaetotaxy type i2 3m). Abdominal terga I III

19 REVIEW OF THE PALAEARCTIC PROTAPHORURA ABSOLON, 1901 SPECIES Figures P. tetragrammata (Gisin, 1964): (42) chaetotaxy of abdominal terga V and VI; (43) distal part of leg III; (44) postantennal sense organ and anterior cephalic pseudocelli; (45) antennal III sense organ; (46) sensory clubs and sensory rod at antennal III sense organ; (47) habitus and dorsal chaetotaxy; (48) macrochaeta.

20 686 I. KAPRUS and R. J. POMORSKI without chaetae s. Abdominal tergum V with p 0 chaeta present (Fig. 42). Ratio M/s on abdominal tergum V = Because of asymmetries, the number of medial chaetae on abdominal tergum VI difficult to recognize. Straight lines, passing through bases of short chaetae situated above anal spines, convergent (Fig. 42). Between legs on pro-, meso- and metathorax 1+1, 2+2 and 2+2 chaetae respectively. Ventral tube with chaetae and 2+2 chaetae at the base. Furca reduced to cuticular pocket with 2+2 setulae and 2+2 setulae below. Claws with strong denticle (Fig. 43). Empodial appendage without basal lamella, as long as inner edge of the claw. Tibiotarsi with distal whorl composed of 11 chaetae. Anal spines shorter than inner edge of the claw (ratio ). Male ventral organ is absent. Remarks. Available type material of P. tetragrammata contains only 3 reproductive males. All of them have some asymmetries in chaetotaxy of abdominal tergite VI (one paratype have only one anal spine developed teratologically), and therefore a confirmation of constant occurrence of two medial chaetae mentioned by Gisin (1964) is impossible. Besides, the second paratype have only 3+3 pseudocelli on abdominal tergum IV (anterolateral pso is absent). Despite of these teratology and asymmetries all other diagnostic characters of P. tetragrammata are well visible and we are able to ascertain, that specimens of Protaphorura collected in Polish and Ukrainian Carpathians belongs to this species. Biology. The species is connected with moist soil and litter of mountain forests. Bisexual. Key to the Palaearctic species of Protaphorura with 4 and more pseudocelli at antennal base (only for adults or subadults) 1. Subcoxae1 of I, II and III pair of legs without pseudocelli Subcoxae1 of I, II, and III pair of legs with 1, 0, 0 pseudocelli respectively Subcoxae1 of I, II, and III pair of legs with 1, 1, 1 pseudocelli respectively Pseudocellar formula dorsally: 4(5,6)3(4)/022/3335 (4)3(4,5), dorsomedial pseudocelli on thoracic tergum II and anterolateral pseudocelli on abdominal tergum IV present; North Asiatic part of Russia P. octopunctata (Tullberg, 1876). Pseudocellar formula dorsally: 43/012/33343, dorsomedial pseudocelli on thoracic tergum II and anterolateral pseudocelli on abdominal tergum IV absent; Russia (Yacutia) P. tolae Pomorski et Kaprus, Pseudocellar formula dorsally: 43/022/33342, claws always with strong inner denticle; Mongolia P. mongolica (Martynova, 1970). Pseudocellar formula dorsally: 43/022/33343, claws without or rarely with very small denticle (in asian populations); Central and South East Europe, Crimea Mts., Russia (Caucasus Mts., Southern Siberia), Afghanistan, Kazakhstan, Tadzhikistan P. sakatoi (Yosii, 1966) 4. Thoracic terga II and III with variable number of pseudocelli (2 3, 2 3 respectively), head ventrally with 2+2 pseudocelli or abdominal tergum V with chaetae s Thoracic terga II and III with stable number of pseudocelli, head ventrally with 1+1 pseudocelli, abdominal tergum V without chaetae s ; if abdominal tergum V with chaetae s, than postantennal sense organ is small (18 26 simple vesicles) Thoracic terga II and III with 3+3 and 3+3 pseudocelli respectively Thoracic terga II and III with 2+2 and 3+3 pseudocelli respectively Thoracic terga II and III with 2+2 and 2+2 pseudocelli respectively Postantennal sense organ consists of vesicles, claws without denticle, pseudocellar formula dorsally 43/033/33353; Great Britain P. daviesi (Bagnall, 1935). Postantennal sense organ consists of vesicles, claws always with denticle Abdominal terga I-III without chaeta s, straight lines passing through bases of short chaetae situated above anal spines distinctly convergent, pseudocellar formula dorsally 4(3)4/033/3335(4)3; Great Britain, Czech Republic, Germany, Poland, Ukraine P. pseudocellata (Naglitsch, 1962). Abdominal terga I III with chaetae s, straight lines passing through bases of short chaetae situated above anal spines parallel, pseudocellar formula dorsally 4(5)3(4,5)/033/4(3)4(3)4(3)5(6)3(4); Russia (Southern Siberia) P. submersa sp. nov. 8. Head with 4+4 posterior pseudocelli, postantennal sense organ consists of vesicles Head with 3+3 posterior pseudocelli, postantennal sense organ consists of vesicles Pseudocellar formula dorsally: 54/023/454(5-7)5 (6)5, ratio claw/anal spines as 2.0; Austria (Alps) P. decempunctata (Kos, 1939). Pseudocellar formula dorsally: 44/023/34353, ratio claw/anal spines as 1.7; Great Britain P. caledonica (Bagnall, 1937) 10. Pseudocellar formula dorsally: 43/023/4(3)4(3)4 (3)53, postantennal sense organ consists of vesicles, claws without denticle; Denmark, Finland, Germany, Iceland, Norway, Jan Mayen, Svalbard, Sweden P. macfadyeni (Gisin, 1953)