Research Article |
Corresponding author: Grzegorz Pasnik ( pasnik@isez.pan.krakow.pl ) Academic editor: Louis Deharveng
© 2016 Igor J. Kaprus', Wanda Weiner, Grzegorz Pasnik.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Kaprus' I, Weiner W, Pasnik G (2016) Collembola of the genus Protaphorura Absolon, 1901 (Onychiuridae) in the Eastern Palearctic: morphology, distribution, identification key. ZooKeys 620: 119-150. https://doi.org/10.3897/zookeys.620.9372
|
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.
Protaphorurini , taxonomy, chaetotaxy, new species, redescription, Siberia, Far East
The genus Protaphorura Absolon, 1901, widespread throughout Holarctic, is the most diverse taxon with almost 140 species described to date (
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
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
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.
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 (
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.
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.
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 5AIIIO6 tibiotarsal chaetotaxy and claw of leg III 7PAO 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.
PAO of small length with 20–23 simple vesicles (Fig.
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.
Chaetotaxy of ventral side of head as in Fig.
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.
The name of the new species refers to the Latin absconditus (hidden, concealed).
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).
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 (
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.
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.
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 15PAO 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.
PAO of middle length with 39–44 simple vesicles (Fig.
Pso formula dorsally 32/033/33342, ventrally 2/000/0001 (Figs
Dorsal chaetotaxy, slightly asymmetrical and rather plurichaetotic, as in Figs
Chaetotaxy of ventral side of head as in Fig.
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.
The name of the new species refers to the Russian “jernik” (= shrub tundra or tundra with dwarf birch).
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).
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 (
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.
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.
Protaphorura nikolai: 18 habitus and dorsal chaetotaxy 19 dorsal side of Ant. III–IV 20 tip of Ant. IV 21PAO 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.
PAO is relatively small with 29–36 simple vesicles (Fig.
Pso formula dorsally 33/022/33342, ventrally 1/000/0000 (Figs
Dorsal chaetotaxy slightly asymmerical, chaetae well differentiated into macrochaetae, mesochaetae and microchaetae as in Fig.
Chaetotaxy of ventral side of head as in Fig.
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.
The species is cordially dedicated to Russian oribatologist Dr. Nikolay Ryabinin, who collected the type material of new species in Primorsky Krai of Russia.
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.
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 (
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.
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.
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.
PAO relatively small, consisting of 32-34 simple vesicles (Fig.
Pso formula dorsally 32/011/22232, ventrally 1/000/0000 (Figs
Dorsal chaetotaxy plurichaetotic, usually with some asymmetry, all dorsal chaetae rather short (except macrochaetae), well differentiated into macro- meso- and microchaetae, as in Fig.
Chaetotaxy of ventral side of head as in Fig.
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
The name of the new species refers to the Latin oligo (a few) and pseudocellus (false ocellus) ‒ characteristic structure in Onychiuroidea.
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).
Onychiurus ombrophilus Stach, 1960: 509 – 514, pl. LXV
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.
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
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.
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.
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.
Protaphorura ombrophila has been described by
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 (
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.
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.
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.
PAO large, consisting of 41–48 simple vesicles (Fig.
Pso formula dorsally 32/022/33343, ventrally 1/000/0000 (Figs
Dorsal chaetotaxy, usually slightly asymmetrical, all dorsal chaetae rather long, well differentiated into macro- meso- and microchaetae, as in Fig.
Chaetotaxy of ventral side of head as in Fig.
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
The name of the new species refers to the Sayan Mountains in Southern Siberia, an area where the type specimens were collected.
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).
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 (
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.
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.
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.
PAO of middle length, consisting of 37–45 simple vesicles (Fig.
Pso formula dorsally 32/022/33332, ventrally 2/000/0001 (Figs
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.
Chaetotaxy of ventral side of head as in Fig.
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.
The name of the new species refers to the Tuva Republic (Russian Federation), the place where the type specimens were collected.
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).
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 (
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.
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.
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.
PAO of middle length, consisting of 32–36 simple vesicles (Fig.
Pso formula dorsally 32/022/33332, ventrally 2/000/0001 (Figs
Dorsal chaetotaxy rather symmetrical, as in Fig.
Chaetotaxy of ventral side of head as in Fig.
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.
The species is cordially dedicated to Vasilina, a granddaughter of Dr. Sophya Stebaeva.
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).
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) |
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)
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.