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Research Article
Collembola of the genus Protaphorura Absolon, 1901 (Onychiuridae) in the Eastern Palearctic: morphology, distribution, identification key
expand article infoIgor J. Kaprus', Wanda Weiner§, Grzegorz Pasnik|
‡ State Museum of Natural History, Ukrainian National Academy of Sciences, , Ukraine
§ Institue of Systematics and Evolution of Animals, Polish Academy of Sciences, Krakow, Poland
| Insitute of Systematics and Evolution of Animals, Polish Academy of Sciences, Krakow, Poland
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Abstract

Seven new species, Protaphorura jernika sp. n., Protaphorura abscondita sp. n., Protaphorura tuvinica sp. n., Protaphorura vasilinae sp. n., Protaphorura sayanica sp. n., Protaphorura oligopseudocellata sp. n. and Protaphorura nikolai sp. n. from different habitats of the southern Siberia and Far East of Russia, are described. Protaphorura ombrophila (Stach, 1960) is redescribed based on the type specimens. These species differ one from other and from all known species by dorsal and ventral pseudocellar formulae, number of pseudocelli on subcoxae 1 of legs I–III, parapseudocellar formula, chaetotaxy of body, structure of claw, size of postantennal organ and body length. Geographical distribution of all known Protaphorura species of Eastern Palearctic was analysed and an identification key to 50 species was provided.

Keywords

Protaphorurini, taxonomy, chaetotaxy, new species, redescription, Siberia, Far East

Introduction

The genus Protaphorura Absolon, 1901, widespread throughout Holarctic, is the most diverse taxon with almost 140 species described to date (Bellinger et al. 2016, Parimuchová and Kováč 2016), forty three of which is known from the Eastern Palearctic (Martynova 1976, Pomorski and Kaprus’ 2007, Kaprus’ and Pomorski 2008, Kaprus’ et al. 2014, Gulgenova and Potapov 2013, Sun, Wu and Gao 2013, Sun, Zhang and Wu 2013, Babenko and Kaprus’ 2014, Sun, Chang and Wu 2015 etc.). The boundaries of the Eastern Palearctic region we determined conventionally from the Ural Mountains and Caspian Sea to Japan Islands and Bering Strait. Siberia, which occupies most of the Eastern Palearctic, continues to be one of the poorly studied geographical regions. The results of this study allow to discover seven new species of Protaphorura. Additionally, Protaphorura ombrophila (Stach, 1960) is redescribed from Afghanistan, using the type material deposited in the Institute of Systematics and Evolution of Animals, Polish Academy of Sciences in Kraków (Poland). The present paper aims to provide a critical evaluation of all known Protaphorura species of the Eastern Palearctic.

Material and methods

Material of Protaphorura species was collected by the soil samples method. Samples were extracted using Berlese–Tullgren funnels. Specimens of new species were collected by Dr. Sophya Stebaeva (Severtsov Institute of Ecology and Evolution, Russian Academy of Sciences, Moscow) in southern Siberia from 1972 to 1994, Dr. Elena Sleptsova (North eastern Federal University in Yakutsk, Russia) in the north eastern Altai in 2002 and Dr. Nikolay Ryabinin (Institute of Water and Ecological Problems, Far Eastern Branch of Russian Academy of Sciences, Khabarovsk) in the Far East of Russia in 2011. Specimens were mounted in Faure’s medium, after clearing in lactophenol, and were studied using Olympus and Leica microscopes. Material is housed in the State Museum of Natural History, Ukrainian National Academy of Sciences, L’viv, Ukraine (SNHM), Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Kraków, Poland (ISEA) and Moscow Pedagogical State University, Russia (MPSU).

The studied type materials of Protaphorura ombrophila (Stach, 1960) are deposited in the Institute of Systematics and Evolution of Animals, Polish Academy of Sciences (Kraków).

Morphological terms. Labial types are named after Fjellberg (1999). Labium areas and chaetal nomenclature follow Massoud (1967) and D’Haese (2003). Tibiotarsal formula is presented after Deharveng (1983). Chaetae on furcal area are notated after Weiner (1996). Chaetae on anal valves are named following Yoshii (1996). Chaetae formula on thoracic tergum I is notated after Gisin (1952).

Abbreviations used in descriptions:

Abd. abdominal segments

Ant. antennal segments

AIIIO sensory organ of Ant. III

AS anal spines

pso pseudocellus

ms s-microchaeta

MVO male ventral organ

PAO postantennal organ

psp pseudopore

psx parapseudocellus

Th. thoracic segments

VT ventral tube

1m single psx or psp in medial position.

Species descriptions

Protaphorura abscondita sp. n.

Figs 1–9, 58

Type material

Holotype (female): Russia, Siberia, Krasnoyarsk Territory, Achinsk Province, 7 km from Nazarovo, steppe meadow, soil, ca 400 m alt., 57°02'N, 90°39'E, 14.VII.1987, leg. S.K. Stebaeva (SNHM). Paratypes: 8 males and 10 females, same data as holotype (SNHM – 7 paratypes: 1 male and 6 females, ISEA – 6 paratypes: 5 males and 1 female, MSPU – 5 paratypes: 2 males and 3 females).

Diagnosis

PAO with 20–23 simple vesicles. Pso formula dorsally 32/033/33343, ventrally 1/000/0000, subcoxae 1 of I–III legs with 1,1,1 pso respectively. Submedial pso a and b on Abd. terga I–II located close together. Psx formula on Abd. sterna: 111100. Th. tergum I with 12–15+12–15 chaetae, chaeta m present. Chaetae s' present on Abd. terga I–III. Manubrial field with 12 chaetae in 3 rows. Claw without lateral denticles.

Description

Holotype (female) length 1.2 mm, length of paratypes: 0.9–1.1 mm (males) and 1.0–1.3 mm (females). Shape of body typical of the genus: cylindrical with strong AS on distinct papillae (Fig. 1). Colour in alcohol yellowish-white. Granulation more or less uniform, distinct. Usually 10–11 grains around each pso.

Figures 1–9.

Protaphorura abscondita: 1 habitus and dorsal chaetotaxy 2 dorsal side of Ant. IV 3 ventral side of Ant. IV 4 right part of head ventrally 5 AIIIO 6 tibiotarsal chaetotaxy and claw of leg III 7 PAO and anterior cephalic pseudocelli 8 chaetotaxy of Abd. sternum IV 9 chaetotaxy of Abd. terga IVVI.

Antennae approximately as long as head, their base well marked. Ant. I with 10–11 chaetae, Ant. II with 17–18 chaetae. AIIIO consisting of 5 guard chaetae, 5 papillae, 2 smooth sensory rods, 2 straight and granulated sensory clubs, ventro-lateral microsensillum present (Fig. 5). Ant. IV with subapical organite in unprotected cavity without clear cuticular papilla. Microsensillum on Ant. IV in usual position above second proximal row of chaetae (Fig. 2). Ventrally Ant. IV with numerous chaetae (ca. 58–65) (Fig. 3). Ant. IV with 9–11 well-differentiated sensilla (Fig. 2, 3).

PAO of small length with 20–23 simple vesicles (Fig. 7). Labral formula of chaetae: 4/342. Maxillary outer lobe with simple palp, basal chaeta and with two sublobal hairs. Labial palp of type A. Labium with 6 proximal, 4 basomedian (E, F, G, and f), and 6 basolateral chaetae (a, b, c, d, e, e’). Papillae A-E with 1, 4, 0, 3, 3 guard chaetae respectively.

Pso formula dorsally 32/033/33343, ventrally 1/000/0000. Subcoxae 1 of I–III legs with one pso and one psx each. Submedial pso a and b on Abd. terga I–II located close together, i.e. closer than on Abd. tergum III, both set posteriorly to macrochaeta p5. Psx present on Abd. sterna I–IV (psx formula 0/000/111100). Psp formula dorsally 0/011/1111, ventrally 0/111/01m1m1m, coxae with 1 psp each.

Dorsal chaetotaxy rather symmetrical, as in Fig. 1, 4 and 9. Dorsal chaetae poorly differentiated into macrochaetae and microchaetae. Sensory chaetae s distinct on body. On head p2 chaetae on the same level as p1 and p3. Chaetae p6 on head located anterior to pso b. Th. tergum I with 12–15+12–15 chaetae, chaeta m present (chaetotaxy type i2–3m). Both Th. terga II and III with lateral microsensilla and with 5+5 or 6+6 axial microchaetae. Chaetae s' present on Abd. terga I–III, on Abd. tergum V present or absent. On Abd. tergum IV in axial area between M2 and P2 macrochaetae located 7– 8 chaetae, medial chaeta m0 present (rarely absent) (Fig. 1). Abd. tergum V usually with 1–2 unpaired microchaeta m0 and p0 (sometimes m0 absent) (Fig. 1). Abd. tergum VI with 1–2 medial chaetae a0 and m0 (often a0 absent). Relative position of prespinal microchaetae usually of subparallel type (Fig. 9). M/s ratio on Abd. tergum V as 10.5–11.4/9.0–9.5, (AS = 10). AS 1.2–1.3 times longer than inner edge of claw and 2.9–3.0 times longer than their basal diameter.

Chaetotaxy of ventral side of head as in Fig. 4. Perilabial area with 4+4 a-chaetae (Fig. 4). Postlabial chaetae 5+5 along ventral groove. Th. sterna I–III with 1+1, 2+2, 2+2 chaetae respectively. VT with ca. 7–9+7–9 chaetae, and 2 chaetae at base. Chaetotaxy of Abd. sternum IV as in Fig. 8. Furcal rudiment: cuticular fold (located near the middle of sternum) with 2+2 dental microchaetae in 2 rows. Chaetotaxy of manubrial field rather stable: 4 chaetae present in ma-row, 4 chaetae in mm-row and 4 chaetae in mp-row (Fig. 8). MVO absent. Each lateral anal valves with a0, 2a1 and 1-2a2; upper anal valve with chaetae a0, 2a2, 2b1, 2b2, c0, 2c1 and 2c2 (as in P. jernika, Fig. 58).

Subcoxae 1 of I, II and III legs with 5, 7, 6 chaetae, subcoxae 2 with 1, 5, 5, coxae with 3, 10, 14, trochanters with 11, 11, 10, femora with 17 each, tibiotarsi with four rows of chaetae (distal whorl (A+T)+B+C): 11+8+3, 11+8+3, 11+8+4 chaetae respectively. Claw with very small (rarely without) denticle in 1/2 of inner edge of claw (Fig. 6). Empodial appendage of same length as inner edge of claw, without basal lamella (Fig. 6).

Etymology

The name of the new species refers to the Latin absconditus (hidden, concealed).

Discussion

P. abscondita sp. n. is characterized by a unique formula of dorsal pso: 2+2 posterior cephalic pso, 3+3 pso on Th. terga II and III and Abd. tergum V. Among seven known species with 3+3 pso on Th. terga II and III, the new species is most similar to the siberian P. tundricola (Martynova, 1976), P. submersa Kaprus’ & Pomorski, 2008 and P. merita Kaprus’ & Pomorski, 2008 due to number of pso on Abd. terga. P. abscondita sp. n. differs from all these species by the 9-11 well differentiated sensilla on Ant. IV. Additionally, it differs from P. merita by the absence of cauliflower like papilla on the tip of antenna and 1+1 ventral pso in posterolateral position on head. From P. submersa, the new species differs by having 3 pso on the base of antennae (4(5) pso in P. submersa) and from P. tundricola by relative position of prespinal microchaetae on Abd.6 (distinctly convergent type in P. tundricola and subparallel type in P. abscondita).

Protaphorura jernika sp. n.

Figs 10–17, 58

Type material

Holotype (female): Russia, N-E Altai, Turochak Region, Altyn-Tu Mt. Ridge, Archa Mt, mountain shrub tundra (=jernik tundra) with Betula rotundifolia, moss, 1700–1800 m alt., 51°31'N, 87°27'E, 9.VIII.2002, leg. E.V. Sleptsova (ISEA). Paratypes: 2 males, same data as holotype (SNHM).

Diagnosis

PAO with 39–44 simple vesicles. Pso formula dorsally 32/033/33342, ventrally 2/000/0001, subcoxae 1 of I–III legs with 1,1,1 pso respectively. Submedial pso a and b on Abd. terga I–II located far from each other. Psx formula on Abd. sterna: 111000. Th. tergum I with 12–15+12–15 chaetae, chaeta m present. Chaetae s' absent on Abd. terga I–III and V. Manubrial field with 16–17 chaetae in 4 rows. Claw without lateral denticles.

Description

Holotype (female) length 1.8 mm, length of paratypes: 1.4 mm (males). Shape of body typical for the genus: cylindrical with strong AS on distinct papillae (Fig. 10). Colour in alcohol yellowish-white. Granulation distinct, usually slightly coarser on head, Abd. tergum VI and around pso. Usually 9–11 grains around each pso.

Figures 10–17.

Protaphorura jernika: 10 habitus and dorsal chaetotaxy 11 left part of head ventrally 12 dorsal side of Ant. III–IV 13 chaetotaxy of Th. terga III aand Abd. tergum I 14 ventral side of Ant. IV 15 PAO and anterior cephalic pseudocelli 16 tibiotarsal chaetotaxy and claw of leg III 17 chaetotaxy of Abd. sternum IV.

Antennae slightly shorter than head, their base well marked. Ant. I with 10 chaetae, Ant. II with 18 chaetae. AIIIO consisting of 5 guard chaetae, 5 papillae, 2 smooth sensory rods, 2 straight and granulated sensory clubs, ventro-lateral microsensillum present (Fig. 12). Ant. IV with subapical organite in unprotected cavity without clear cuticular papilla. Microsensillum on Ant. IV in usual position on the level of second proximal row of chaetae. Ant. IV ventrally with very numerous chaetae (ca. 70–75) (Fig. 14). Sensilla indistinct on Ant. IV.

PAO of middle length with 39–44 simple vesicles (Fig. 15). Labral formula of chaetae: 4/342. Maxillary outer lobe with simple palp, basal chaeta and with two sublobal hairs. Labial palp of type A. Labium with 6 proximal, 4 basomedian (E, F, G, and f), and 6 basolateral chaetae (a, b, c, d, e, e’). Papillae A-E with 1, 4, 0, 3, 3 guard chaetae respectively.

Pso formula dorsally 32/033/33342, ventrally 2/000/0001 (Figs 10, 11, 13, 17). Subcoxae 1 of I–III legs with one pso and one psx each. Submedial pso a and b on Abd. terga I–II located far apart, i.e. on similar distance as on Abd. tergum III (Fig. 13). Psx present on Abd. sterna I–III (psx formula 0/000/111000). Psp formula dorsally 0/011/1111, ventrally 0/111/01m1m1m, coxae with 1 psp each.

Dorsal chaetotaxy, slightly asymmetrical and rather plurichaetotic, as in Figs 10 and 13. Dorsal chaetae rather well differentiated into macrochaetae and microchaetae. Sensory chaetae s indistinct on body. On head p1 chaetae are displaced forward in relation to p2–p4 (Fig. 10). Chaetae p6 on head located between pso a and b. Th. tergum I with 12–15+12–15 chaetae, chaeta m present (chaetotaxy type i2–3m). Both Th. terga II and III with lateral microsensilla and with 5+5 or 6+6 axial microchaetae. Chaetae s' absent on Abd. terga I–III and V. On Abd. tergum IV in axial area between M2 and P2 macrochaetae located 8–12 chaetae, medial chaeta m0 present (rarely absent) (Fig. 10). Abd. tergum V usually with 1–2 unpaired microchaeta m0 and p0 (sometimes m0 absent) (Fig. 10). Abd. tergum VI with 1–2 medial chaetae a0 and m0 (rarely a0 absent). Relative position of prespinal microchaetae usually of parallel type (Fig. 10). M/s ratio on Abd. tergum V as 13.6–17.6/5.6–6.9 (AS = 10). AS 1.1 times longer then inner edge of claw and 2.6 times longer then their basal diameter.

Chaetotaxy of ventral side of head as in Fig. 11. Perilabial area with 4+4 a-chaetae (Fig. 11). Postlabial chaetae 5-6+5-6 along ventral groove. Th. sterna I–III with 0+0, 1+1, 1+1 chaetae respectively. VT with ca. 8–9+8–10 chaetae and 1+2 chaetae at base. Furcal rudiment: cuticular fold (located on the anterior edge of sternum) with 2+2 dental microchaetae in 2 rows. Chaetotaxy of manubrial field: 4 chaetae present in ma-row, 4 chaetae in mm’-row, 4 chaetae in mm-row and 4–5 chaetae in mp-row (Fig. 17). MVO absent. Each lateral anal valves with a0, 2a1 and 2a2; upper anal valve with chaetae a0, 2a2, 2b1, 2b2, c0, 2c1 and 2c2 (Fig. 58).

Subcoxae 1 of I, II and III legs with 5–7, 6–8, 5–6 chaetae, subcoxae 2 with 1, 5, 5, coxae with 3, 8, 14, trochanters with 11, 11, 10, femora with 19 each, tibiotarsi with four rows of chaetae (distal whorl (A+T)+B+C): 11+8+3, 11+8+3, 11+8+4 chaetae respectively. Claw with strong denticle in 1/2 of inner edge of claw (Fig. 16). Empodial appendage of same length as inner edge of claw, without basal lamella (Fig. 16).

Etymology

The name of the new species refers to the Russian “jernik” (= shrub tundra or tundra with dwarf birch).

Discussion

Protaphorura jernika sp. n. belongs to the group of Protaphorura species with pseudocelli on subcoxa 1 of all legs and 2+2 pso ventrally on head. By the presence of 1+1 pso on Abd. sternum IV, the new species is similar to the P. vasilinae sp. n. Both species differ only in the formula of dorsal pso and ventral psx on Abd. sterna: the former has 32/033/33342 pso and 111000 psx whereas the latter 32/022/33332 pso and 110001mpsx (see also diagnosis of P. vasilinae sp. n.). P. jernika sp. n. differs from other two Eastern Palearctic representatives of this group, P. merita Kaprus’ & Pomorski, 2008 and P. buryatica Gulgenova & Potapov, 2013 by dorsal pso formula (32/033/33342 in the new species vs 32(3)/012/33342 in buryatica and 43/02(3)2(3)/3335(4,6)3(4) in merita), by the presence of 1+1 pso on abd. sternum IV in the new species and lack in the both other, by the number of vesicles in PAO (39-44 in the new species, 12-13 in buryatica and 16-22 in buryatica).

Protaphorura nikolai sp. n.

Figs 18–25, 58

Type material

Holotype (male): Russia, Primorsky Krai, Khasansky district, Barabash village, mixed forest with Quercus, Acer and Juglans, in soil and leave litter, 9.VII.2011, leg. N.A. Ryabinin (SNHM). Paratypes: 6 males and 6 females, same data as holotype (SNHM – 9 paratypes: 5 male and 4 females, ISEA – 3 paratypes: 1 male and 2 females).

Diagnosis

PAO with 29–36 simple vesicles. Pso formula dorsally 33/022/33342, ventrally 1/000/0000, subcoxae 1 of I–III legs with 1,0,0 pso respectively. Submedial pso a and b on Abd. terga I–II located close together. Psx formula on Abd. sterna: 100000. Th. tergum I with 11–12+11–12 chaetae, chaeta m present. Chaetae s' absent on Abd. terga I–III and V. Manubrial field with 14–15 chaetae in 3 rows. Claw without lateral denticles.

Description

Holotype (male) length 1.5 mm, length of paratypes: 1.45–1.55 mm (males) and 1.58–1.72 mm (females). Shape of body typical for the genus: cylindrical with strong AS on distinct papillae (Fig. 18). Colour in alcohol yellowish-white. Granulation more or less uniform, distinct. Usually 12–14 grains around each pso.

Figures 18–25.

Protaphorura nikolai: 18 habitus and dorsal chaetotaxy 19 dorsal side of Ant. III–IV 20 tip of Ant. IV 21 PAO and anterior cephalic pseudocelli 22 right part of head ventrally 23 tibiotarsal chaetotaxy and claw of leg III 24 chaetotaxy of Abd. sternum IV 25 chaetotaxy of Abd. tergum VI.

Antennae approximately as long as head, their base well marked. Ant. I with 11–12 chaetae, Ant. II with 17–18 chaetae. AIIIO consisting of 5 guard chaetae, 5 papillae, 2 smooth sensory rods, 2 straight and granulated sensory clubs, ventro-lateral microsensillum present (Fig. 19). Ant. IV with subapical organite in cavity protected by cuticular papillae (Fig. 20). Microsensillum on Ant. IV situated on level or below of second proximal row of chaetae. Ventrally Ant. IV with numerous chaetae (ca. 68–72). Ant. IV without differentiated sensilla (Fig. 19).

PAO is relatively small with 29–36 simple vesicles (Fig. 21). Labral formula of chaetae: 4/342. Maxillary outer lobe with simple palp, basal chaeta and with two sublobal hairs. Labial palp of type A. Labium with 7 proximal, 4 basomedian (E, F, G, and f), and 6 basolateral chaetae (a, b, c, d, e, e’). Papillae A-E with 1, 4, 0, 3, 3 guard chaetae respectively.

Pso formula dorsally 33/022/33342, ventrally 1/000/0000 (Figs 18, 22, 24). Subcoxae1 of legs I, II and III with 1,0,0 pso respectively. Psx on subcoxae1 of legs I, II and III absent. Submedial pso a and b on Abd. terga I–II located close together, i.e. much closer than on Abd. tergum III, both set posteriorly to macrochaeta p5 (Fig. 18). Ventral psx formula 1/000/100000). Psp formula dorsally 0/011/1111, ventrally 0/111/01m1m1m, coxae with 1 psp each.

Dorsal chaetotaxy slightly asymmerical, chaetae well differentiated into macrochaetae, mesochaetae and microchaetae as in Fig. 18. Sensory chaetae s indistinct on body. On head p2 chaetae on same level as p1 and p3. Chaetae p6 on head located anterior to pso b (Fig. 18). Th. tergum I with 11–12+11–12 chaetae, chaeta m present (chaetotaxy type i2–3m). Both Th. terga II and III with lateral microsensilla and with 4+4 or 5+5 axial microchaetae. Chaetae s' absent on Abd. terga I–III and V. On Abd. tergum IV in axial area between M2 and P2 macrochaetae located 9–11 chaetae, medial chaeta m0 present or absent, p0 present or absent (Fig. 18). Abd. tergum V usually with 1 unpaired microchaeta p0 (m0 absent) (Fig. 18). Abd. tergum VI with 1 medial chaetae m0. Relative position of prespinal microchaetae of distinctly divergent type (Fig. 25). M/s ratio on Abd. tergum V as 33–40/20–22, (AS = 10). AS 0.6–0.7 times as long as inner edge of claw and 2.0 times longer than their basal diameter.

Chaetotaxy of ventral side of head as in Fig. 22. Perilabial area with 4+4 a-chaetae. Postlabial chaetae 5+5 along ventral groove. Th. sterna I–III with 1+1, 2+2, 2+2 chaetae respectively. VT with ca. 8–9+8–9 chaetae, and 2(1)+2(1) chaetae at base. Chaetotaxy of Abd. sternum IV as in Fig. 22. Furcal rudiment: cuticular fold (located near the middle of sternum) with 2+2 dental microchaetae in 2 rows. Chaetotaxy of manubrial field: 4 chaetae present in ma-row, 6-7 chaetae in mm-row and 4 chaetae in mp-row (Fig. 24). MVO absent. Each lateral anal valves with a0, 2a1 and 2a2; upper anal valve with chaetae a0, 2a2, 2b1, 2b2, c0, 2c1 and 2c2 (as in P. jernika, Fig. 58).

Subcoxae 1 of I, II and III legs with 5–6, 6–7 and 5–6 chaetae respectively, subcoxae 2 with 1, 5, 5, coxae with 3, 11, 13, trochanters with 11, 11, 10, femora with 21, 21, 18, tibiotarsi with four rows of chaetae (distal whorl (A+T)+B+C): 11+8+4, 11+8+4, 11+8+4–5 chaetae respectively. Claw with very strong denticle in the 1/2 of inner edge of claw (Fig. 23). Empodial appendage 0,9–1,0 times as long as inner edge of claw, without basal lamella (Fig. 23).

Etymology

The species is cordially dedicated to Russian oribatologist Dr. Nikolay Ryabinin, who collected the type material of new species in Primorsky Krai of Russia.

Discussion

Protaphorura nikolai sp. n. belongs to the group of Protaphorura species with 1,0,0 pseudocelli on subcoxa 1 of I, II and III legs and 1+1 pso on head ventrally. Among Asiatic species this group includes P. zori (Martynova, 1975 in Martynova & Chelnokov, 1975)(although Martynova did not mention subcoxal pso, the examined by us type has 1,0,0 pso on subcoxae), P. brevispinata (Yosii, 1966), P. changbaiensis Sun, Zhang & Wu, 2013, P. mongolica (Martynova, 1975 in Martynova & Chelnokov, 1975), P. sakatoi (Yosii, 1966) and P. maoerensis Sun, Wu & Gao, 2013. Within this group, it shares dorsal pso formula with P. zori but differs from the latter by the presence of inner denticle on claw, the absence of chaeta a0 on Abd. tergum VI (in P. zori inner denticle absent and chaeta a0 present) and by arrangement of prespinal chaetae (placed divergently in P. nikolai and convergently in P. zori). Perhaps there are other differences in the morphology of these two species, but P. zori is not well described and needs more detailed study.

Protaphorura oligopseudocellata sp. n.

Figs 26–32, 58

Type material

Holotype (female): Russia, Siberia, Western Sayan, Oiskii Mt. Range, vicinity of weather station Olenya Rechka, mountain tundra with Betula rotundifolia, Salix sp, Sphagnum sp., 1800 m alt., in moss and soil, 52°48'N, 93°13'E, 27.VI.1990, leg. S.K. Stebaeva (SNHM). Paratypes: 3 females and juvenile, same data as holotype (ISEA – 1 paratype, MSPU – 1 paratype and juvenile).

Diagnosis

PAO with 32–34 simple vesicles. Pso formula dorsally 32/011/22232, ventrally 1/000/0000, subcoxae 1 of I–III legs without pso. Psx formula on Abd. sterna: 111000. Th. tergum I with 23–25+23–25 chaetae, one, two or three chaetae m present. Chaetae s' present on Abd. terga I–III and absent or present on Abd. tergum V. Manubrial field with 12–13 chaetae in three rows. Claw with pair of lateral denticles.

Description

Holotype (female) length 2.2 mm, length of paratypes: 2.0–2.3 mm (females). Shape of body typical of the genus: cylindrical with strong AS on distinct papillae (Fig. 26). Colour in alcohol yellowish-white. Granulation more or less uniform, distinct. Usually 7–10 grains around each pso.

Figures 26–32.

Protaphorura oligopseudocellata: 26 habitus and dorsal chaetotaxy 27 PAO and anterior cephalic pseudocelli 28 dorsal side of Ant. IIIIV 29 and 30 distal part of leg III 31 right part of head ventrally 32 chaetotaxy of Abd. sternum IV.

Antennae as long as the head, their base well marked. Ant. I with 11–12 chaetae, Ant. II with 18 chaetae. AIIIO consisting of 5 guard chaetae, 5 papillae, 2 smooth sensory rods, 2 straight and granulated sensory clubs, ventro-lateral microsensillum present (Fig. 28). Ant. IV with subapical organite in unprotected cavity without clear cuticular papilla. Microsensillum on Ant. IV in usual position above second proximal row of chaetae. Ventrally Ant. IV with numerous chaetae (ca. 74–78). Sensilla indistinct on antennal segment IV (Fig. 28).

PAO relatively small, consisting of 32-34 simple vesicles (Fig. 27). Labral formula of chaetae: 4/342. Maxillary outer lobe with simple palp, basal chaeta and with two sublobal hairs. Labial palp of type A. Labium with 7 proximal, 4 basomedian (E, F, G, and f), and 6 basolateral chaetae (a, b, c, d, e, e’). Papillae A-E with 1, 4, 0, 3, 3 guard chaetae respectively.

Pso formula dorsally 32/011/22232, ventrally 1/000/0000 (Figs 26, 31, 32). Subcoxae 1 of I–III legs without pso and with one psx each. Psx formula 1/000/111000. Th. terga II and III with 1+1, 1+1 psx in lateral position (Fig. 26). Psp formula dorsally 0/011/1111, ventrally 0/111/01m1m1m, coxae with 1 psp.

Dorsal chaetotaxy plurichaetotic, usually with some asymmetry, all dorsal chaetae rather short (except macrochaetae), well differentiated into macro- meso- and microchaetae, as in Fig. 26. Sensory chaetae s indistinct on body. On head p2 chaetae are displaced forward in relation to p1 and p3. Chaetae p6 on head located anterior to pseudocelli b (Fig. 26). Th. tergum I with 23–25+23–25 chaetae, 1–3 chaetae m and 1–2 chaetae i present (chaetotaxy type i(1–2)3–4m(1–3)). Both Th. terga II and III with lateral microsensilla and with 6+6 or 7+7 axial microchaetae. Chaetae s' present on Abd. terga I–III and absent or present on Abd. tergum V (Fig. 26). On Abd. tergum IV in axial area between M2 and P2 macrochaetae located 23–24 chaetae, medial chaetae p0 and m0 present (sometimes these chaetae absent). Abd. tergum V with one unpaired microchaeta p0 (Fig. 26). Abd. tergum VI with medial chaetae m0. Relative position of prespinal microchaetae of convergent type (Fig. 26). M/s ratio on Abd. tergum V as 23.5–23.9/15 (AS = 10). AS 0.7–0.8 times as long as inner edge of claw and 2.3 times longer than their basal diameter.

Chaetotaxy of ventral side of head as in Fig. 31. Perilabial area with 4–5+4–5 a-chaetae. Postlabial chaetae 4-5+4-5 along ventral groove. Thoracic sterna I–III with 1+1, 2–3+2–3, 2–3+2–3 chaetae respectively. VT with ca. 10+10 chaetae, and 2–3 chaetae at base. Chaetotaxy of Abd. sternum IV as in Fig. 32. Furcal rudiment: cuticular fold (located near the middle of sternum) with 2+2 dental microchaetae in 2 rows. Chaetotaxy of manubrial field: 4–5 chaetae present in ma-row, 4 chaetae in mm-row, 4 chaetae in mp-row (in adult specimens) (Fig. 32). Each lateral anal valves with a0, 2a1 and 2a2; upper anal valve with chaetae a0, 2a2, 2b1, 2b2, c0, 2c1 and 2c2 (as in P. jernika, Fig. 58).

Subcoxae 1 of I, II and III legs with 6–8, 7–8, 7–9 chaetae, subcoxae 2 with 1, 5, 5, coxae with 4, 10, 15, trochanters with 13, 15, 15, femora with 21, 23, 22–23, tibiotarsi with four rows of chaetae (distal whorl (A+T)+B+C): 11+8+3–4, 11+8+5–6, 11+8+5 chaetae respectively. Claw with strong denticle in 1/2 of inner edge of claw and pair of lateral denticles (Figs 29, 30). Empodial appendage 0.9 times as long as inner edge of claw, without basal lamella (Fig. 29).

Etymology

The name of the new species refers to the Latin oligo (a few) and pseudocellus (false ocellus) ‒ characteristic structure in Onychiuroidea.

Discussion

Protaphorura oligopseudocellata sp. n. is characterized by the reduced number of pso on body dorsally – 32/011/22232. Only four species with 1+1 pso on Th. tergum III is currently known: P. januarii (Weiner, 1977), P. stiriaca (Stach, 1946), P. pseudostyriaca (Loksa, 1964) and P. pseudarmata (Folsom, 1917). The first three species are described from Europe and the last one from North America. Among these species P. oligopseudocellata sp. n. is probably the most similar to P. januarii and P. stiriaca due to the absence of pso on subcoxa 1 of all legs and some similarity of dorsal pso formulae. The new species can be easily distinguished from these species by the number of pso on Abd. terga I–V (22232 in P. oligopseudocellata sp. n., 23232 in P. januarii and 33232 in P. stiriaca), the plurichaetotic chaetotaxy and by the presence of strong lateral denticles on claws and 1+1 pso on head ventrally (lateral denticles and pso absent in P. januarii and P. stiriaca).

Protaphorura ombrophila (Stach, 1960)

Figs 33–36, 59

Onychiurus ombrophilus Stach, 1960: 509 – 514, pl. LXV

Type material

Lectotype (female) (by present designation): Afghanistan, ”Tchehel Sotoun” Cave near Jalrayz, W Kabul, with the original label: “Tchehel Sotoun-Höhle (nahe Djalrez), 20.III.1959”, leg. Dr. K. Lindberg. Paralectotypes: 1 male and 8 females, same data as lectotype.

Redescription

Lectotype (female) length 1.9 mm, length of paralectotypes: 1.8 mm (male) and 1.8–2.2 mm (females). Shape of body typical of the genus: cylindrical with strong AS on distinct papillae. Colour in alcohol white. Granulation more or less uniform, distinct. Usually 11–13 grains around each pso.

Antennae slightly shorter than head, their base well marked. Ant. I with 10 chaetae, Ant. II with 16–18 chaetae. AIIIO consisting of 5 guard chaetae, 5 papillae, 2 smooth sensory rods, 2 straight and granulated sensory clubs, ventro-lateral microsensillum present. Ant. IV with subapical organite in unprotected cavity without clear cuticular papilla. Microsensillum on Ant. IV in usual position above second proximal row of chaetae. Sensilla indistinct on Ant. IV.

PAO of middle length, consisting of 24–38 simple vesicles. Labral formula of chaetae: 4/342. Maxillary outer lobe with simple palp, basal chaeta and with two sublobal hairs. Labial palp of type A. Chaetotaxy of labium invisible.

Pso formula dorsally 32/022(3)/33(2)3(2)43, ventrally 2/000/0001 (Figs 33–36). Subcoxae 1 of I–III legs without pso. Submedial pseudocelli a and b on Abd. terga I–II located far apart, i.e. on similar distance as on Abd. tergum III (Fig. 33). Psx formula 0/000/11?00?.

Figures 33–36.

Protaphorura ombrophila: 33 position of pso and p-chaetae in midsection of Abd. tergum I 34 position of p-chaetae on posterior margin of head 35 chaetotaxy of Abd. terga V–VI 36 chaetotaxy of Abd. sternum IV.

Dorsal chaetotaxy rather symmetrical. Dorsal chaetae well differentiated into macrochaetae and microchaetae. On head p2 chaetae are displaced forward in relation to p1 and p3 (Fig. 34). Chaetae p6 on head located between pseudocelli a and b. Th. tergum I with 8–10+8–10 chaetae, chaeta m absent (chaetotaxy type i2-). Both Th. terga II and III with lateral microsensilla. Chaetae s' absent on Abd. terga I–III and V (Fig. 35). On Abd. tergum IV in axial area between M2 and P2 macrochaetae located 6–7 chaetae, medial chaeta m0 present. Abd. tergum V usually with 1 unpaired microchaeta m0 (p0 absent) (Fig. 35). Abd. tergum VI with 1 medial chaetae m0. Relative position of prespinal microchaetae usually divergent or parallel type (Fig. 35). M/s ratio on Abd. tergum V as 18.2/8.8 (AS = 10). AS 0.8–0.9 times as long as inner edge of claw and 2.8-3.4 times longer then their basal diameter.

Perilabial area with 4+4 a-chaetae. Th. sterna I–III without chaetae. VT with ca. 8–9+8–9 chaetae, and 1 chaetae at base. Furcal rudiment: cuticular fold (located on the anterior edge of the sternum) with 2+2 dental microchaetae in 2 rows. Chaetotaxy of manubrial field: 4 chaetae present in ma-row, 2 chaetae in mm’ -row, 4 chaetae in mm-row and 5 chaetae in mp-row (Fig. 36). MVO absent. Each lateral anal valves with a0 and 2a1 (a2 absent); upper anal valve with chaetae a0, 2a2, 2b1, 2b2, c0, 2c1 and 2c2 (as in P. vasilinae, Fig. 59).

Subcoxae 1 of I, II and III legs with 5, 6, 5–6 chaetae, tibiotarsi with four rows of chaetae (distal whorl (A+T)+B+C): 11+8+3, 11+8+3, 11+8+4 chaetae respectively. Claw with very small denticle in 1/2 of inner edge of claw. Empodial appendage 0.7–0.8 times as long as inner edge of claw, without basal lamella.

Remarks

Protaphorura ombrophila has been described by Stach (1960) from Afghanistan about 55 years ago, when many important diagnostic characters remained unknown. Latter Yosii (1966), during his research on some Collembola of Afghanistan, India and Ceylon, discovered three females of the species in Afghanistan and wrote: “They (i.e. P. ombrophila) coincide fairly well with the detailed description of Stach. However, the posterior margin of head has 3+3, 3+2 and 2+2 pseudocelli. In other respects no difference is to be found”. Parimuchová and Kováč in their recent publication (2016) devoted to the critical analysis of Palearctic species of the genus Protaphorura and assigned this species to the group “species dubia”. Here we present first redescription of P. ombrophila based on characters currently used in taxonomy of Protaphorura. See also the discussion in P. tuvinica sp. n.

Protaphorura sayanica sp. n.

Figs 37–44, 58

Type material

Holotype (male): Russia, Siberia, Western Sayan, Oiskii Mt. Range, vicinity of weather station Olenya Rechka, mountain tundra, 1800 m alt., in moss and soil, 52°48'N, 93°13'E, 10.VII.1990, leg. S.K. Stebaeva (SNHM). Paratypes: 2 females and 2 juveniles, same data as holotype (SNHM – 1 paratype female, ISEA – 1 paratype female and 2 juveniles); 2 females: Russia, Krasnoyarsk Territory, Khakasia, Kuznetskii Alatau Mt. Range, ca 5 km NW of settl. Kommunar, mountain tundra with Dryas oxyodontha, 1500 m alt., 54°20'N, 89°17'E, 24.VII.1990, leg. S.K. Stebaeva (ISEA); 2 males, female and 2 juveniles: Russia, Kuznetskii Alatau Mt. Range, Kemerovo Prov., 10 km NW of Mezhdurechensk, mixed taiga with rich herbaceous cover, under Abies sibirica, soil, 500-600 m alt., 53°45'N, 88°00'E, 1.VII.1982, leg. S.K. Stebaeva (SNHM); male: Russia, Salair Range, 130 km SE of Novosibirsk, 11 km N of Mirnyi, chern forest, 500 m alt., soil, 54°38'N, 84°45'E, 7.VI.1972, leg. S.K. Stebaeva (MPSU); female subadult: Russia, West Siberia, 25 km S of Novosibirsk, Akademgorodok, glade in birch forest, soil, 400 m alt., 54°49'N, 83°08'E, 7.X.1994, leg. S.K. Stebaeva (SNHM).

Diagnosis

PAO with 41–48 simple vesicles. Pso formula dorsally 32/022/33343, ventrally 1/000/0000, subcoxae 1 of I–III legs with 1,1,1 pso respectively. Submedial pso a and b on Abd. terga I–II located far apart. Psx formula on Abd. sterna: 111101m. Th. tergum I with 18–21+18–21 chaetae, one or two chaetae m present. Chaetae s' present on Abd. terga I–III and V. Manubrial field with 14 chaetae in three rows. Claw with pair of lateral denticles.

Description

Holotype (male) length 2.7 mm, length of paratypes: 2.7–2.9 mm (females). Other specimens length: 2.62.7 mm males and 2.8 mm female. Shape of body typical of the genus: cylindrical with strong AS on distinct papillae (Fig. 37). Colour in alcohol yellowish-white. Granulation more or less uniform, distinct. Usually 7–9 grains around each pso.

Figures 37–44.

Protaphorura sayanica: 37 habitus and dorsal chaetotaxy 38 right part of head ventrally 39 tip of Ant. IV 40 dorsal side of Ant. IIIIV 41 PAO and anterior cephalic pseudocelli 42 and 43 distal part of leg III 44 chaetotaxy of Abd. sternum IV.

Antennae as long as the head, their base well marked. Ant. I with 11 chaetae, Ant. II with 16–19 chaetae. AIIIO consisting of 5 guard chaetae, 5 papillae, 2 smooth sensory rods, 2 straight and granulated sensory clubs, ventro-lateral microsensillum present (Fig. 40). Ant. IV with subapical organite in unprotected cavity without clear cuticular papilla (Fig. 39). Microsensillum on Ant. IV in usual position above second proximal row of chaetae. Ventrally Ant. IV with numerous chaetae (ca. 65–70) (Fig. 40). Sensilla indistinct on Ant. IV (Fig. 40).

PAO large, consisting of 41–48 simple vesicles (Fig. 41). Labral formula of chaetae: 4/342. Maxillary outer lobe with simple palp, basal chaeta and with two sublobal hairs. Labial palp of type A. Labium with 7 proximal, 4 basomedian (E, F, G, and f), and 6 basolateral chaetae (a, b, c, d, e, e’). Papillae A-E with 1, 4, 0, 3, 3 guard chaetae respectively.

Pso formula dorsally 32/022/33343, ventrally 1/000/0000 (Figs 37, 38, 44). Subcoxae 1 of I–III legs with one pso and one psx each. Submedial pso a and b on Abd. terga I–II located far apart, i.e. on similar distance as on Abd. tergum III (Fig. 37). Psx formula 1/000/111101m. Psp formula dorsally 0/011/1111, ventrally 0/111/01m1m1m, coxae with 1 psp each.

Dorsal chaetotaxy, usually slightly asymmetrical, all dorsal chaetae rather long, well differentiated into macro- meso- and microchaetae, as in Fig. 37. Sensory chaetae s indistinct on body. On head p2 chaetae on the same level as p1 and p3. Chaetae p6 located anterior to pso b on head (Fig. 37). Th. tergum I with 18–21+18–21 chaetae, 1–2 chaetae m and 1–2 chaetae i present (chaetotaxy type i(1–2)2–4m(1–2)). Both Th. terga II and III with lateral microsensilla and with 5+5 or 6+6 axial microchaetae. Chaetae s' present on Abd. terga I–III and V. On Abd. tergum IV in axial area between M2 and P2 macrochaetae located 15–18 chaetae, medial chaeta p0 present (sometimes p0 absent). Abd. tergum V with one unpaired microchaeta p0 (Fig. 37). Abd. tergum VI with medial chaetae m0. Relative position of prespinal microchaetae of convergent type (Fig. 37). M/s ratio on abdominal tergum V as 18.9–26.6/15.7–20.6 (AS = 10). AS 0.8–1.1 times as long as inner edge of claw and 2.9 times longer than their basal diameter.

Chaetotaxy of ventral side of head as in Fig. 38. Perilabial area with 5(4)+5(4) a-chaetae. Postlabial chaetae 4-5+4-5 along ventral groove. Th. sterna I–III with 1–2+1–2, 2–3+2–3, 2–3+2–3 chaetae respectively. VT with ca. 11–12+11–12 chaetae, and 2–3 chaetae at base. Furcal rudiment: cuticular fold (located near middle of sternum) with 2+2 dental microchaetae in 2 rows. Chaetotaxy of manubrial field: 4 chaetae present in ma-row, 6 chaetae in mm-row, 4 chaetae in mp-row (in adult specimens) (Fig. 44). MVO absent. Each lateral anal valves with a0, 2a1 and 2a2; upper anal valve with chaetae a0, 2a2, 2b1, 2b2, c0, 2c1 and 2c2 (as in P. jernika, Fig. 58).

Subcoxae 1 of I, II and III legs with 7–9, 8–9, 7–8 chaetae, subcoxae 2 with 1, 5, 5, coxae with 4, 10, 12-15, trochanters with 11, 13, 13, femora with 20–21, 20–23, 20–23, tibiotarsi with four rows of chaetae (distal whorl (A+T)+B+C): 11+8+3, 11+8+4–5, 11+8+4–5 chaetae respectively. Claw with strong denticle in 1/2 of inner edge of claw and pair of lateral denticles (Figs 42, 43). Empodial appendage as long as the claw, without basal lamella. (Fig. 43).

Etymology

The name of the new species refers to the Sayan Mountains in Southern Siberia, an area where the type specimens were collected.

Discussion

Protaphorura sayanica sp. n. is probably the most similar to such Asiatic Protaphorura species as P. pjasinae (Martynova, 1976), P. microtica (Dunger, 1978) and P. subarctica (Martynova, 1976) due to the presence of the same number of pso on subcoxae 1 of all legs, ventral and dorsal side of head, Th. terga I–II and Abd. terga I–IV. However, P. sayanica sp. n. may easily be distinguished from these species by the number of pso on Abd. tergum V (3+3 pso in the new species and 2+2 pso all other species presented above) and presence of pair of lateral denticles on claw (absent in other four species).

Protaphorura tuvinica sp. n.

Figs 45–50, 59

Type material

Holotype (male): Russia, S-W Tuva, ca 30 km SW of Mugur-Aksy, upper reaches of Mugur River, Mongun-Taiga Mts, mountain tundra, moss under Betula rotundifolia, 2700 m alt., 50°22'N, 90°05'E, 23.VII.1993, leg. S.K. Stebaeva (SNHM). Paratypes: 10 males, 3 females and 7 juveniles, same data as holotype (SNHM – 6 paratypes: 5 males and 1 female, ISEA – 4 paratypes: 3 males and 1 female, MSPU – 4paratypes: 3 males and 1 female, and 7 juveniles).

Diagnosis

PAO with 37–45 simple vesicles. Pso formula dorsally 32/022/33332, ventrally 2/000/0001, subcoxae 1 of I–III legs without pso. Submedial pso a and b on Abd. terga I–II located far apart. Psx formula on Abd. sterna: 110–1001 m. Th. tergum I with 9–11+9–11 chaetae, chaeta m absent. Chaetae s' absent on abdominal terga I–III and V. Manubrial field with 19 chaetae in 4 rows. Claw without lateral denticles.

Description

Holotype (male) length 1.9 mm, length of paratypes: 1.7–1.8 mm (males) and 1.9–2.2 mm (females). Shape of body typical for the genus: cylindrical with strong AS on distinct papillae (Fig. 45). Colour in alcohol yellowish-white. Granulation more or less uniform, distinct. Usually 11–12 grains around each pso.

Figures 45–50.

Protaphorura tuvinica: 45 habitus and dorsal chaetotaxy 46 right part of head ventrally 47 dorsal side of Ant. IIIIV 48 PAO and anterior cephalic pseudocelli 49 tibiotarsal chaetotaxy and claw of leg III 50 chaetotaxy of Abd. sternum IV.

Antennae slightly shorter than head, their base well marked. Ant. I with 9–10 chaetae, Ant. II with 17 chaetae. AIIIO consisting of 5 guard chaetae, 5 papillae, 2 smooth sensory rods, 2 straight and granulated sensory clubs, ventro-lateral microsensillum present (Fig. 47). Ant. IV with subapical organite in unprotected cavity without clear cuticular papilla. Microsensillum on Ant. IV in usual position above second proximal row of chaetae. Ventrally Ant. IV with numerous chaetae (ca. 68–70). Sensilla indistinct on Ant. IV (Fig. 47).

PAO of middle length, consisting of 37–45 simple vesicles (Fig. 48). Labral formula of chaetae: 4/342. Maxillary outer lobe with simple palp, basal chaeta and with two sublobal hairs. Labial palp of type A. Labium with 7 proximal, 4 basomedian (E, F, G, and f), and 6 basolateral chaetae (a, b, c, d, e, e’). Papillae A-E with 1, 4, 0, 3, 3 guard chaetae respectively.

Pso formula dorsally 32/022/33332, ventrally 2/000/0001 (Figs 45, 46, 50). Subcoxae 1 of I–III legs without pso and with one psx each. Submedial pso a and b on Abd. terga I–II located rather far apart, i.e. on similar distance as on Abd. tergum III (Fig. 45). Psx formula 0/000/110(1)01. Psp formula dorsally 0/011/1111, ventrally 0/111/01m1m1m, coxae with 1 psp each.

Dorsal chaetotaxy rather symmetrical and plurichaetotic, chaetae well differentiated into macrochaetae and microchaetae (fig. 45). Sensory chaetae s indistinct on body. On head p2 chaetae displaced forward in relation to p1 and p3. Chaetae p6 on head located between pso a and b (Fig. 45). Th. tergum I with 9–11+9–11 chaetae, chaeta m absent (chaetotaxy type i2-). Both Th. terga II and III with lateral microsensilla and with 5+5 or 6+6 axial microchaetae. Chaetae s' absent on Abd. terga I–III and V. On Abd. tergum IV in axial area between M2 and P2 macrochaetae located 7–8 chaetae, medial chaeta m0 present (Fig. 45). Abd. tergum V usually with 1–2 unpaired microchaeta m0 and p0 (often m0 absent) (Fig. 45). Abd. tergum VI with medial chaetae m0. Relative position of prespinal microchaetae of parallel type (Fig. 45). M/s ratio on Abd. tergum V as 14.6–17.2/4.6–6.2 (AS = 10). AS 0.9–1.0 as long as inner edge of claw and 3.1 times longer than their basal diameter.

Chaetotaxy of ventral side of head as in Fig. 46. Perilabial area with 5+5 a-chaetae (Fig. 46). Postlabial chaetae 5-6+5-6 along ventral groove. Th. sterna I–III without chaetae. VT with ca. 8–9+8–9 chaetae and 2 chaetae at base. Furcal rudiment: cuticular fold (located on the anterior edge of the sternum) with 2+2 dental microchaetae in 2 rows. Chaetotaxy of manubrial field: 5 chaetae present in ma-row, 4 chaetae in mm’-row, 6 chaetae in mm-row and 4 chaetae in mp-row (Fig. 50). MVO absent. Each lateral anal valves with a0 and 2a1 (a2 absent); upper anal valve with chaetae a0, 2a2, 2b1, 2b2, c0, 2c1 and 2c2 (as in P. vasilinae, Fig. 59).

Subcoxae 1 of I, II and III legs with 5–6, 6, 5 chaetae, subcoxae 2 with 1, 5, 5, coxae with 3, 10, 13, trochanters with 11, 12, 10, femora with 20, 20, 19–20, tibiotarsi with four rows of chaetae (distal whorl (A+T)+B+C): 11+8+3, 11+8+3, 11+8+3-4 chaetae respectively. Claw with strong denticle in 1/2 of inner edge of claw (Fig. 49). Empodial appendage of the same length as inner edge of claw, without basal lamella. (Fig. 49).

Etymology

The name of the new species refers to the Tuva Republic (Russian Federation), the place where the type specimens were collected.

Discussion

Protaphorura tuvinica sp. n. belongs to the group of Protaphorura species without pseudocelli on subcoxa 1 of all legs and with 2+2 pso ventrally on head: P. ombrophila (Stach, 1960), P. kopetdagi Pomorski, 1994, P. salsa Kaprus’, Paśnik & Weiner, 2014, P. bakhchisaraica Kaprus’, Paśnik & Weiner, 2014 and P. ajudagi Pomorski, Skarżyński & Kaprus’, 1998. All these species inhabit the territory of southern Palearctic from Crimean Peninsula to central Asia and southern Siberia.

The new species has the pseudocellar formula the same as in kopetdagi (32/022/33332) when the other posses the different number of pseudocelli. The males of P. kopetdagi, P. salsa, P. bakhchisaraica and P. ajudagi are armed with the male ventral organ whereas the new species and P. ombrophila have males devoided of the organ. P. tuvinica differs also from the latter species by the number of pso on Abd. terga IV-V (3,2 in the new species and 4,2 in P. ombrophila).

Protaphorura vasilinae sp. n.

Figs 51–57, 59

Type material

Holotype (female): Russia, West Siberia, 25 km S of Novosibirsk, Akademgorodok, lawn, soil, 400 m alt., 54°49'N, 83°08'E, 2.X.1994, leg. S.K. Stebaeva (SNHM). Paratypes: 7 females and 6 juveniles, same data as holotype (SNHM – 3 paratype females and 3 juveniles, ISEA – 4 paratype females and 3 juveniles); 2 females and 3 juveniles: Russia, N-E Altai, Turochak Region, meadow, soil, 11.VI.2002, leg. E. Sleptsova (SNHM).

Diagnosis

PAO with 32–36 simple vesicles. Pso formula dorsally 32/022/33332, ventrally 2/000/0001, subcoxae 1 of I–III legs with 1,1,1 pso respectively. Submedial pso a and b on Abd. terga I–II located far apart. Psx formula on Abd. sterna: 110001m. Th. tergum I with 10–11+10– 11 chaetae, chaeta m absent. Chaetae s' absent on Abd. terga I–III and present on Abd. tergum V. Manubrial field with 25–28 chaetae in 6 rows. Claw without lateral denticles.

Description

Holotype (female) length 1.5 mm, length of paratypes: 1.4–1.7 mm (females). Shape of body typical of the genus: cylindrical with strong AS on distinct papillae (Fig. 51). Colour in alcohol yellowish-white. Granulation more or less uniform, distinct. Usually 10–12 grains around each pso.

Figures 51–57.

Protaphorura vasilinae: 51 habitus and dorsal chaetotaxy 52 AIIIO 53 ventral side of Ant. IV 54 PAO and anterior cephalic pseudocelli 55 tibiotarsal chaetotaxy and claw of leg III 56 right part of head ventrally 57 chaetotaxy of Abd. sternum IV.

Antennae slightly shorter than the head, their base well marked. Ant. I with 10 chaetae, Ant. II with 16 chaetae. AIIIO consisting of 5 guard chaetae, 5 papillae, 2 smooth sensory rods, 2 straight and granulated sensory clubs, ventro-lateral microsensillum present (Fig. 52). Ant. IV with subapical organite in unprotected cavity without clear cuticular papilla (Fig. 53). Microsensillum on antennal segment IV in usual position above second proximal row of chaetae. Ventrally Ant. IV with numerous chaetae (ca. 50–55). Ant. IV with 8–11 slightly differentiated sensilla (Fig. 53).

PAO of middle length, consisting of 32–36 simple vesicles (Fig. 54). Labral formula of chaetae: 4/342. Maxillary outer lobe with simple palp, basal chaeta and with two sublobal hairs. Labial palp of type A. Labium with 7 proximal, 4 basomedian (E, F, G, and f), and 6 basolateral chaetae (a, b, c, d, e, e’). Papillae A-E with 1, 4, 0, 3, 3 guard chaetae respectively.

Pso formula dorsally 32/022/33332, ventrally 2/000/0001 (Figs 51, 56, 57). Subcoxae 1 of I– III legs with one pso and one psx each. Submedial pso a and b on Abd. terga I–II located far apart, i.e. on similar distance as on Abd. tergum III (Fig. 51). Psx present on Abd. sterna I–II and VI (psx formula 0/000/110001m). Psp formula dorsally 0/011/1111, ventrally: 0/111/01m1m1m , coxae with 1 psp each.

Dorsal chaetotaxy rather symmetrical, as in Fig. 51. Dorsal chaetae well differentiated into macrochaetae and microchaetae. Sensory chaetae s indistinct on body. On head p2 chaetae are displaced forward in relation to p1 and p3. Chaetae p6 located between pseudocelli a and b on head. Th. tergum I with 10–11+10–11 chaetae, chaeta m absent (chaetotaxy type i2–3-). Both Th. terga II and III with lateral microsensilla and with 5+5 or 6+6 axial microchaetae. Chaetae s' absent on Abd. terga I–III and present on Abd. tergum V. On Abd. tergum IV in axial area between M2 and P2 macrochaetae located 7–8 chaetae, medial chaeta m0 present (Fig. 51). Abd. tergum V usually with 2 unpaired microchaeta m0 and p0 (sometimes m0 absent) (Fig. 51). Abd. tergum VI with medial chaetae m0. Relative position of prespinal microchaetae of subparallel type (Fig. 51). M/s ratio on abdominal tergum V as 14.9–16/5.6–5.2 (AS = 10). AS 1.1 times longer than inner edge of claw and 3.1 times longer than their basal diameter.

Chaetotaxy of ventral side of head as in Fig. 56. Perilabial area with 4+4 a-chaetae. Postlabial chaetae 4–5+4–5 along ventral groove. Th. sterna I–III without chaetae. VT with ca. 8–9+8–9 chaetae, and 2 chaetae at base. Chaetotaxy of Abd. sternum IV as in Fig. 57. Furcal rudiment: cuticular fold (located on the anterior edge of the sternum) with 2+2 dental microchaetae in 2 rows. Chaetotaxy of manubrial field: 4 chaetae present in ma-row, 4 chaetae in ma’-row, 4–5 chaetae in mm’’-row, 5–6 chaetae in mm’-row, 4 chaetae in mm-row and 4–5 chaetae in mp-row (in adult specimens) (Fig. 56). MVO absent. Each lateral anal valves with a0 and 2a1 (a2 absent); upper anal valve with chaetae a0, 2a2, 2b1, 2b2, c0, 2c1 and 2c2 (Fig. 59).

Figures 58–59.

Chaetotaxy of anal valves: 58 Protaphorura jernika 59 Protaphorura vasilinae.

Subcoxae 1 of I, II and III legs with 5, 6–7, 6 chaetae, subcoxae 2 with 1, 5, 4, coxae with 3, 10–11, 13, trochanters with 11, 11, 10, femora with 18, 18, 18, tibiotarsi with four rows of chaetae (distal whorl (A+T)+B+C): 11+8+3, 11+8+3, 11+8+4 chaetae respectively. Claw with strong denticle in the 1/2 of inner edge of claw (Fig. 55). Empodial appendage of the same length as inner edge of claw, without basal lamella. (Fig. 55).

Etymology

The species is cordially dedicated to Vasilina, a granddaughter of Dr. Sophya Stebaeva.

Discussion

The same number of pso on body ventrally (2+2 on head and 1+1 on Abd. sternum V), the presence of pseudocelli on subcoxae 1 of all legs, 2+2 posterior cephalic pso and 2+2 pso on Abd. tergum V allow suggesting a close similarity between P. vasilinae sp. n. and P. jernika sp. n. (see also discussion of P. jernika sp. n.). These species distinctly differ in the number of pso on Th. terga II–III and Abd. tergum IV (2+2,2+2 and 3+3 in P. vasilinae vs 3+3,3+3 and 4+4 in P. jernica respectively), in the formula of psx on Abd. sterna (110001m in P. vasilinae vs 111000 in P. jernica) and in the chaetotaxy of Th. tergum I (chaetotaxy type i2–3- in P. vasilinae vs i2–3m in P. jernica) and Abd. tergum V (s’ present in P. vasilinae vs s’ absent in P. jernica).

Key to Protaphorura species of the Eastern Palearctic

For the species with high variability in the pseudocellar formula we used in the key the most common type.

1 AIIIO with four papillae 2
AIIIO with five papillae 5
2 Th. terga II and III with 3+3 pso each (formula of dorsal pso: 33/033/33333) P. matsumotoi (Kinoshita, 1923) (Japan)
Th. terga II and III with 1+1 and 2+2 pso respectively 3
3 Ventrally on head 1+1 pso present in posterolateral position, formula of dorsal pso: 33/012/33342 P. dzherga Gulgenova & Potapov, 2013 (Russia: Transbaikal)
Ventral pso on head absent 4
4 Formula of dorsal pso: 33/012/33332, furcal area with two pairs of dental microchaetae P. dorzhievi Gulgenova & Potapov, 2013 (Russia: Transbaikal)
Formula of dorsal pso:43/012/33353, furcal area with one pair of dental microchaetae P. uniparis Gulgenova & Potapov, 2013 (Russia: Transbaikal)
5 AIIIO with two slender, long sensory rods: one inserted dorsal to the papillae, secod between papillae 6
AIIIO with two normal sensory rods inserted behind the papillae , between the sensory cluba 8
6 Antennal base with four pso (formula of dorsal pso: 43(4)/022/3(4)3(4)3(4)5(6)3(4)), PAO with 4246 vesicles P. nutak (Yosii, 1972) (Russia: Far East, Kunashir Island and Eastern Siberia, Japan)
Antennal base with three pso 7
7 Formula of dorsal pso: 33/022/33343, PAO with about 45 vesicles P. longisensillata (Yosii, 1969) (Japan)
Formula of dorsal pso: 32/022/33342, PAO with 3640 vesicles P. diplosensillata (Dunger, 1978)* (Mongolia)
8 Subcoxae1 of legs I, II and III without pso 9
Subcoxae1 of legs I, II and III with 1,0,0 pso respectively 21
Subcoxae1 of legs I, II and III with 1,1,1 pso respectively 27
9 Ventral pso on head absent 10
Ventrally on head 1+1 or 2+2 pso present 11
10 Formula of dorsal pso: 32/022/33333, PAO with 4065 vesicles, MVO absent P. borealis (Martynova, 1973 in Martynova, Gorodkov & Chelnokov, 1973) (Eastern Palearctic)
Formula of dorsal pso: 33/012/33332, PAO with 2126 vesicles, MVO in a form of two brush-shape chaetae on each anal valve P. minima Sun, Zhang & Wu, 2013 (North Eastern China)
11 Ventrally on head 2+2 pso present, Abd. sternum IV with 1+1 pso 12
Ventrally on head 1+1 pso present in anteromedial position, Abd. sternum IV without pso 15
12 Abd. tergum V with 3+3 pso 13
Abd. tergum V with 2+2 pso 14
13 Anterolateral pso on Abd. tergum IV present, formula of dorsal pso: 32(3)/022(3)/33(2)3(2)43, MVO absent P. ombrophila (Stach, 1960) (Afghanistan)
Anterolateral pso on Abd. tergum IV absent, formula of dorsal pso: 33/022/3324(3)3, MVO present on Abd. sterna IIIII with 2+2 and 1+1 modified chaetae respectively P. salsa Kaprus’, Paśnik & Weiner, 2014 (Russia: southern Siberia)
14 PAO with 3745 vesicles, formula of dorsal pso: 32/022/33332, MVO absent P. tuvinica sp.n. (Russia: southern Siberia)
PAO with 2636 vesicles, formula of dorsal pso: 32/022/33332, MVO present on Abd. sterna IIIII with 2+2 and 2+2 modified chaetae respectively P. kopetdagi Pomorski, 1994 (Turkmenistan: Kopetdag Mts.)
15 Antennal base with four or more pso 16
Antennal base with three pso 17
16 Dorsomedial pso on Th. tergum II and anterolateral pso on Abd. tergum IV present (formula of dorsal pso: 4(5,6)3(4)/022/3335(4)3(4,5)) P. octopunctata (Tullberg, 1876) (Russia: central Siberia)
Dorsomedial pso on Th. tergum II and anterolateral pso on Abd. tergum IV absent (formula of dorsal pso: 43/012/333(2)43) P. tolae Pomorski & Kaprus’, 2007 (Russia: eastern Siberia)
17 Posterior cephalic pso 2+2, claws with pair lateral denticles 18
Posterior cephalic pso 3+3, claws without lateral denticles 20
18 Th. tergum I in adult specimens with 11+11 chaetae, claws with strong lateral denticles, formula of dorsal pso: 32/022/33232 P. microcellata (Dunger, 1978) (Mongolia)
Th. tergum I in adult specimens with 1725+1725 chaetae 19
19 Th. tergum I with 1720+1720 chaetae, formula of dorsal pso: 32/022/33332, claws with very small lateral denticles P. cf. microcellata (Dunger, 1978) (Russia: central Siberia after Babenko & Kaprus’, 2014)
Th. tergum I with 2325+2325 chaetae, formula of dorsal pso: 32/011/22232, claws with strong lateral denticles oligopseudocellata sp. n. (Russia: southern Siberia)
20 Formula of dorsal pso: 33/022/33332, ventral psx formula: 01/000/111100, chaetae s’ present on Abd. terga IIII and V P. bicampata (Gisin, 1956) (Northern Europe, Eastern Palearctic)
Formula of dorsal pso: 33/01(2)2/3334(3)2, ventral psx formula: 01/000/100000, chaetae s’ absent on Abd. terga IIII and V P. jacutica (Martynova, 1976) (north eastern Europe, eastern Asia)
21 Antennal base with four pso 22
Antennal base with three pso 24
22 Abd. tergum IV with 5+5 pso (formula of dorsal pso: 43/022/33353), PAO with 4042 vesicles P. maoerensis Sun, Wu & Gao, 2013 (noth eastern China)
Abd. tergum IV with 4+4 pso 23
23 Formula of dorsal pso: 43/022/33342, claws always with strong inner denticle, PAO with 2627 vesicles P. mongolica (Martynova, 1975) (Mongolia)
Formula of dorsal pso: 43/022/33343, claws without or rarely with very small inner denticle (in Asian populations), PAO with 3035 vesicles P. sakatoi (Yosii, 1966) (central and south-eastern Europe, Russia: Caucasus Mts and southern Siberia, Afghanistan, Kazakhstan, Tajikistan)
24 Abd. sternum IV with 1+1 pso, formula of dorsal pso: 32/012/33132 P. brevispinata (southern Korea)
Abd. sternum IV without pso 25
25 Posterior cephalic pso 2+2 (formula of dorsal pso: 32/012/33232), psx formula on Abd. sterna IVI: 100000 P. changbaiensis Sun, Zhang & Wu, 2013 (north eastern China)
Posterior cephalic pso 3+3 (formula of dorsal pso: 33/022/33342) 26
26 Claws without inner denticle, chaeta a0 present on Abd. tergum VI, prespinal chaetae placed convergently P. zori (Martynova, 1975 in Martynova & Chelnokov, 1975) (Tajikistan: eastern Pamir)
Claws with strong inner denticle, chaeta a0 absent on Abd. tergum VI, prespinal chaetae placed divergently P. nikolai sp. n. (Russia: Far East)
27 Ventrally on head 2+2 pso present 28
Ventrally on head 1+1 pso present in anteromedial position 31
28 Abd. sternum IV without pso 29
Abd. sternum IV with 1+1 pso 30
29 Formula of dorsal pso: 32(3)/012/33342, claws with inner denticle, PAO with 1213 vesicles P. buryatica Gulgenova & Potapov, 2013 (Russia: Transbaikal)
Formula of dorsal pso: 43/02(3)2(3)/3335(4,6)3(4), claws without inner denticle, PAO with 1622 vesicles P. merita Kaprus’ & Pomorski, 2008 (Russia: southern Siberia)
30 Formula of dorsal pso: 32/033/33342, psx formula on Abd. sterna IVI: 111000 P. jernika sp. n. (Russia: southern Siberia)
Formula of dorsal pso: 32/022/33332, psx formula on Abd. sterna IVI: 110001m P. vasilinae sp. n. (Russia: southern Siberia)
31 Antennal base with four or more pso 32
Antennal base with three pso 36
32 Th. tergum III with 2+2 pso (formula of dorsal pso: 43/022/33342), psx formula 1/000/110001m P. licheniphila Kaprus’ & Pomorski, 2008 (Russia: central Siberia)
Th. tergum III with 3+3 pso 33
33 Abd. tergum V with 2+2 pso (formula of dorsal pso: 43/023/33342), PAO with 1822 vesicles P. nazarovensis Kaprus’ & Pomorski, 2008 (Russia: south Siberia)
Abd. tergum V with 3+3 or more pso 34
34 Abd.terga I–III and V without chaetae s’, formula of dorsal pso: 43/023/33353, PAO with 1622 vesicles, psx formula on Abd. sterna IVI: 111101m P. jiamusiensis Sun, Wu & Gao, 2013 (north eastern China)
Abd.terga I–III and V with chaetae s’ 35
35 PAO with 1826 vesicles, psx formula on Abd. sterna IVI: 111101m, formula of dorsal pso: 4(5)3(4,5)/033/4(3)4(3)4(3)5(6)3(4) P. submersa Kaprus’ & Pomorski, 2008 (Russia: southern Siberia)
PAO with 3640 vesicles, psx formula on Abd. sterna IVI: 100001?m , formula of 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: eastern Siberia)
36 Posterior cephalic pso 2+2 37
Posterior cephalic pso 3+3 42
37 Th. terga II and III with 3+3 pso each (formula of dorsal pso: 32/033/33343) P. abscondita sp. n. (Russia: southern Siberia)
Th. Terga II and III with 2+2 pso 38
38 Abd. terga IIII without chaetae s’ 39
Abd. terga IIII with chaetae s’ 40
39 Subapical organite on Ant. IV in cavity protected by cuticular papillae, PAO with 30–42 simple vesicles, most common formula of dorsal pso: 32/022/33343, but some specimens may have 3+3 posterior pso on head and 2+2 pso on Abd. tergum V P. tschernovi (Martynova, 1976) (Russia: western Taimyr, central Siberia)
Subapical organite on Ant. IV in unprotected cavity, PAO with 25–40 simple vesicles, formula of dorsal pso: 32/022/3333(4)2 P. subarctica (Martynova, 1976) (Northern Palearctic)
40 Abd. tergum V with chaetae s’ PAO with about 41–48 vesicles, formula of dorsal pso: 32/022/33343 and ventral pso: 1/000/0000 P. sayanica sp. n. (Russia: southern Siberia)
Abd. tergum V without chaetae s’ 41
41 Submedial pso a and b on Abd. terga I-II in nearby position and both these pso set medially to macrochaetae p5, formula of dorsal pso: 32/022/3334(3)2, PAO with 24‒40 simple vesicles P. pjasinae (Martynova, 1976) (northern Asia, western Siberia)
Submedial pso a and b on Abd. terga I-II set far apart and pso b set laterally to macrochaetae p5, formula of dorsal pso: 32/022/33342, PAO with 22 simple vesicles P. microtica (Dunger, 1978) (Mongolia)
42 Th. tergum II with 1+1 pso (formula of dorsal pso: 33/012/33342), psx formula on Abd. sterna IVI: 111101m, PAO with 2432 vesicles P. genheensis Sun, Chang & Wu, 2015 (north eastern China)
Th. tergum II with 2+2 or more pso 43
43 Abd. tergum IV with 3 +3 pso (formula of dorsal pso: 33/022/33333), claws without inner denticle P. fimata (Gisin, 1952) (Kyrgyzstan, Iran)
Abd. tergum IV with 4 +4 or more pso 44
44 Chaetae s’ present on Abd. terga IIII or V 45
Chaetae s’ absent on Abd. terga IIII or V 48
45 AS less than 0.5 length of claws III, formula of dorsal pso: 33/022/33343 P. ussurica (Martynova, 1981) (Russia: Far East)
AS 0.7–1.0 length of claws III 46
46 Relative position of prespinal microchaetae on Abd. tergumVI parallel type, formula of dorsal pso: 33(2)/022/33342(3), psx formula on Abd. sterna IVI: 110001m P. boedvarssoni Pomorski, 1993 (Russia: western and central Siberia)
Relative position of prespinal microchaetae on Abd. tergumVI distinctly convergent type 47
47 Formula of dorsal pso highly variable: 33(2)/03(2)3(2)/4(3,5)4(3,5)4(3,5,6)5(4,6)3(2,4), chaetae s on Abd. tergum V 1.01.1 times longer than AS P. tundricola (Martynova, 1976) (north eastern Europe, western and central Siberia)
Formula of dorsal pso: 33/022(3)/3334(5,6)2(3), chaetae s on Abd. tergum V 1.5 times longer than AS P. neriensis (Martynova, 1976) (Russia: eastern Siberia)
48 Th. tergum II with 3+3 pso (formula of dorsal pso: 3(4)3/033/33342), claws with hardly noticeable inner denticle P. kaszabi (Dunger, 1978) (Mongolia, north eastern China)
Th. tergum II with 2+2 or rarely 1+1 pso (formula of dorsal pso: 33/02(1)2(3)/33342(3), claws with clear inner denticle P. taimyrica (Martynova, 1976) (northern Asia)

Species insufficient described which are not included in the key

P. aksuensis (Martynova, 1972), formula of dorsal pso: 33/022/33333, (Kyrgyzstan)

P. tridentata (Stebaeva, 1982), formula of dorsal pso: 32/022/33342, (southern Siberia)

P. teres (Yosii, 1956), formula of dorsal pso: 32/022/33333, (Japan)

P. yagii (Miyoshi, 1923), formula of dorsal pso: 32/022/33232, (Japan)

Acknowledgements

We are greatly indebted to Russian zoologists Dr. Sophya Stebaeva, Dr. Anatoly Babenko (Severtsov Institute of Ecology and Evolution, Moscow), Dr. Elena Sleptsova (North eastern Federal University in Yakutsk) and Dr. Nikolay Ryabinin (Institute of Water and Ecological Problems, Far Eastern Branch of Russian Academy of Sciences, Khabarovsk) for their materials and information about biology and geographical distribution of Protaphorura species. The work was supported by the cooperation between Ukrainian, Polish and Russian Academies of Sciences.

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