Nine New Species of the Genus Oligaphorura Bagnall 1949 (Collembola, Onychiuridae) from Russia
- Авторлар: Shveenkova Y.B.1, Antipova M.D.2, Babenko A.B.2
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Мекемелер:
- ”Privolzhskaya lesostep” State Nature Reserve
- The Severtsov Institute of Ecology and Evolution, Russian Academy of Sciences
- Шығарылым: Том 103, № 1 (2024)
- Беттер: 8-32
- Бөлім: ARTICLES
- URL: https://ogarev-online.ru/0044-5134/article/view/258185
- DOI: https://doi.org/10.31857/S0044513424010029
- EDN: https://elibrary.ru/JOKHIF
- ID: 258185
Дәйексөз келтіру
Толық мәтін
Аннотация
Nine new species of the genus Oligaphorura are described based on material from various parts of the Russian Federation. Among them, three species, O. ossetica sp. n., O. anocellata sp. n., and O. octosetosa sp. n., are representatives of the Caucasian fauna. The former one belongs to the marcuzzii-group and differs from other members of the group by an increased number of dorsal and subcoxal pseudocelli. Oligaphorura anocellata sp. n., from the differens-group, shows the same pattern of dorsal pseudocelli as O. hackeri (Christian), but it lacks subcoxal pseudocelli. The latter Caucasian species described here, O. octosetosa sp. n. (the daii-group), is characterized by the presence of eight distal setae on the tibiotarsi, four papillae in the antennal organ, and only one pair of pseudocelli on the second thoracic tergum. Oligaphorura ligni sp. n., from the absoloni-group, is from the southern Primorye, Russian Far East, and it can be distinguished from the probably most similar, Chinese species O. shifangensis Liu et Sun by a longer unguiculus, a smaller postantennal organ, and fewer ventral parapseudocelli. The remaining five species described belong to the groenlandica-group and were found in different parts of the eastern Palaearctic. Oligaphorura yakutica sp. n. is most similar to O. duocellata Babenko et Fjellberg, from the Magadan Region, due to the presence of two pairs of dorsal pseudocelli on the first thoracic segment and a peculiar furcal remnant. Yet it is easily distinguishable from the latter by the complete absence of pseudocelli on the abdominal sterna. Oligaphorura subnuda sp. n., from the Taimyr Peninsula, northern Siberia, differs from the Nearctic O. nuda (Fjellberg) by the presence of pseudocelli on the fourth abdominal sternum. Oligaphorura cavicola sp. n., a species found in a cave in the northern Urals, is characterized by an increased number of dorsal pseudocelli on all abdominal segments (44454) and, unlike such sympatric congeners with the same number of abdominal pseudocelli as O. interrupta (Fjellberg) or O. nuda (Fjellberg), it shows a pseudocellus each side of the thorax I. Oligaphorura neglecta sp. n., widespread in the NE Palaearctic, is very similar to the circumpolar O. groenlandica (Tullberg). These two species can be distinguished by the presence/absence of a cuticular fold in the furcal remnant and different lengths of axial setae on the fifth abdominal terga. Finally, O. primorica sp. n., from the Russian Far East, can be compared to several Korean species described by Weiner (1994). All of them have a unique type of granulation that covers the anterior part of the body and the last abdominal segment, probably representing an isolated regional group of related species. The new species is characterized by the same number of pseudocelli as O. koreana (Weiner) and differs from the latter species (and the other related congeners of the region) in having strongly differentiated dorsal setae. An updated key to all presently known 77 Oligaphorura species is also given.
Негізгі сөздер
Толық мәтін
The most comprehensive identification key of the genus Oligaphorura Bagnall 1949, proposed by Paśnik and Weiner (2017), includes 55 valid species of the genus. During the current decade, the group has been supplemented by new congeners from Alaska (Paśnik, Kaprus’, 2019), Eastern Asia (Liu et al., 2019; Sun et al., 2019) and the East European part of the Palearctic (Shveenkova, Babenko, 2021, 2022). As a result, 68 named species are currently listed at www.collembola.org (Bellinger et al., 1996–2023). Modern ecological studies of soil springtails carried out in the Caucasus (Kuznetsova et al., 2019) and in the Russian Far East (Kuznetsova et al., 2021), as well as the available collections from the northern regions of Russia (Babenko, 2013), have revealed nine additional, still undescribed species of this group. This paper is devoted to their descriptions. Hence, the already long list of the known representatives of the genus Oligaphorura has increased to 77 species, which brings the genus to the third in the subfamily Onychiurinae in terms of species diversity. We also propose an updated identification key for all known Oligaphorura species of the world, based on species-groups identified earlier in the genus (Shveenkova, Babenko, 2021).
All types of the described species are kept in the collection of the Zoology and Ecology Department of the Moscow State Pedagogical University (MSPU).
Abbreviations. Abd. I–VI, abdominal segments; A-, AC-, ABC-, ABD-type, types of labium (Fjellberg, 1999), A–E, labial papillae; A-, T-, B-, C-, M-setae, tibiotarsal setae named according to Deharveng (1983); a-, b-, c-setae, setae on anal valves (Yoshii, 1996); absoloni (A)-, differens (D)-, humicola (H)-, schoetti (S)-type, types of furcal remnant (Shveenkova, Babenko, 2021, 2022); a-, m-, p-setae, setae of anterior, medial, and posterior rows on terga; Ant. I–IV, antennal segments; AIIIO, antennal organ on Ant. III; a'0-, a0-, a0'-, d0-setae, unpaired dorsal setae on head (Jordana et al., 1997; D’Haese, 2003); a- and b-pso, medial pseudocelli on terga (Pomorski, 1996); ms, microsensillum(a); PAO, postantennal organ; pso, pseudocellus(i); d. pso, full number of dorsal pseudocelli; psx, parapseudocellus(i); S, lanceolate sensorial seta(e); Sc, subcoxa(e); Th. I–III, thoracic segments; Ti, tibiotarsus(i); VT, ventral tube.
SPECIES DESCRIPTIONS
Oligaphorura ossetica
Shveenkova, Antipova et Babenko sp. n. (Figs 1, 1–6)
Diagnosis. Species of the marcuzzii-group. Dorsal sensilla on body well-marked. Anal spines absent. Pseudocellar formulae 42/244/5.7.7.10.6 (dorsal), 11/000/1112(1) (ventral), and 233 (subcoxal). Ventral psx hardly visible. AIIIO with 5 papillae. PAO slightly larger than nearest pso, with 3–4 lobes. Labium of ABD-type (papilla E entirely absent). Th. III without lateral ms. Abd. IV with unpaired seta p0. Distal whorls (A+T) of each tibiotarsus with 9 setae. Furcal remnant of S-type, with a small cuticular fold.
Type material. Holotype, female, Russia, Northern Caucasus, North Ossetian State Nature Reserve, upper part of Tsey Gorge, young birch forest with rich herbaceous cover [42.7788°N 43.8646°E], 2244 m alt., soil and litter, 27.07.2021, M. Antipova & A. Babenko leg. Paratypes, female and I instar juvenile, same data as holotype, but mixed forest with pine, birch and willow [42.7830°N, 43.8701°E], 2193 m alt.
Description. Size 0.7–0.8 mm, holotype 0.8 mm. Colour white in alcohol. Body shape typical of genus. Granulation regular, only slightly coarser around dorsal pso (12–13 granules) (Fig. 1, 1).
Fig. 1. Oligaphorura ossetica sp. n: 1 – dorsal chaetotaxy; 2 – ventral chaetotaxy of abdomen; 3 – ventral chaetotaxy of head; 4 – antenna, dorsal view; 4a – ibid., ventral view; 5 – labial palp; 6 – tip of leg III, inner side; 6a – ibid., outer side. Scales, mm: 1–3 – 0.1; 4–6 – 0.01.
Number of pso: 42/244/5.7.7.10.6 (dorsal) and 11/000/1112(1) (ventral) (Figs 1, 1–3) with some variations detected. Ventral psx usually invisible. Upper Sc of legs I–III with 233 pso.
Antennae slightly shorter than head. Ant. IV with many curved sensilla, three of which thickened stronger, subapical organite present, microsensillum located under proximal row of setae almost hidden under papillae of AIIIO (Fig. 1, 4). AIIIO consisting of 5 papillae, 5 guard setae, 2 sensory rods, 2 granulated clubs (external one larger) and lateral ms. Ant. I–II with 8 and 12–13 setae, respectively. Antennal area not marked. PAO located laterally in a cuticular furrow, with 4(3) lobes, its longer axis about as long as 1.2 diameter of nearest pso. Maxilla unmodified. Maxillary outer lobe simple with 1 basal seta and 2 sublobals. Labrum with 4/342 setae. Labium with 6 proximal, 4 basomedian and 5 basolateral setae, terminal sensilla of papillae A, B and D thickened, papilla C and its terminal sensillum short, but also slightly thickened, papilla E entirely absent (Fig. 1, 5). Eleven guard setae present on labial palp, four of them spiniform as usual, seven others also rather short but curved. Their exact assignment uncertain: three guards located on site of reduced papilla E, and other four – next to papillae B and D.
Dorsal setae distinctly differentiated into macro- and microsetae only on last abdominal terga, mutual position of setae symmetrical in general (Fig. 1, 1). S-setae well marked and usually distributed as follows: 11/012/222110 (dorsally), 11/000/000100 (ventrally) and 000 on lower Sc of legs I–III. Head with only one axial unpaired seta a0, d0 being absent as usual for the genus. Setae p1 on head located in front of setae p2. Th. I with 5–6+5–6 dorsal setae. Lateral ms present only on Th. II. Terga of Th. II–Abd. III with 3–4 pairs of axial setae. Abd. IV with unpaired seta p0 (m0?). On Abd. VI a0 and a2 subequal macrosetae, a1 as microsetae. Thoracic sterna with 0–1–1 setae each side of ventral line. Ventral chaetotaxy of head and abdomen as in Figs 1, 2–3.
Upper subcoxae of legs I–III with 4, 4–5, 4–5 setae, respectively. Tibiotarsi I–III with 18–19–17 setae, respectively: distal whorl (T+A) with 9 setae (setae T2 and T3 absent), 7 B-setae (B7 absent on Ti III), unpaired seta M and one seta (or two setae on Ti II) of C-whorl. Unguis with neither inner nor lateral teeth, unguiculus without clear basal lamella, about half as long as inner edge of unguis (Fig. 1, 6). VT with (6)7+(6)7 proximal and 2+2 setae at base. Furcal remnant of S-type, cuticular fold small (Fig. 1, 2). Each lateral anal valve with a0 and 2a1 setae, unpaired valve with a0, 2b1, 2b2 (a1 and b0 absent) and at least 7 setae in c-row. Anal spines absent.
Etymology. The species is named after the region of its type locality, the Republic of North Ossetia–Alania.
Affinities. Oligaphorura ossetica sp. n. is a typical member of the marcuzzii-group due to the presence of 4+4 pso at the antennal base and the absence of anal spines. Its labial palps are of the peculiar ABD-type with a reduced papilla E, like those of most other members of the group. Due to the presence of nine distal setae on each tibiotarsus, the new species is similar to O. igori Shveenkova et Babenko 2022, but differs clearly in the number of dorsal and subcoxal pso (42/244/5.7.7.10.6 and 233 in O. ossetica sp. n., vs 43/133/44354 and 222 in O. igori).
Oligaphorura ossetica sp. n. was recently referred to in North Ossetia’s species checklist of Collembola as Oligaphorura sp. 4 (Antipova, Babenko, 2022).
The marcuzzii-group appears to be rather diverse in the Caucasus including the region of the type locality of O. ossetica sp. n. For instance, O. montivaga Shveenkova et Babenko 2022 was recently described from a nearby area of the same republic. Moreover, when describing the latter species, an undescribed form, represented by juvenile specimens and found in the same gorge as O. ossetica sp. n. was mentioned as well (Shveenkova, Babenko, 2022, p. 583). It is characterized by the same type of the labium and 9 distal setae on the tibiotarsi, but has a different pseudocellar formula with a lower pso number. Previously this latter form was referred to as Oligaphorura. sp. n. 2 aff. kremenitsai Shvejonkova et Potapov (Kuznetsova et al., 2019) or Oligaphorura sp. 3 (Antipova, Babenko, 2022).
Oligaphorura anocellata
Shveenkova, Antipova et Babenko sp. n. (Figs 2, 1–6)
Diagnosis. Species of the differens-group. Dorsal sensilla not distinguishable. Anal spines present. Pseudocellar formulae: 32/022/33343 (dorsal), 10(?)/000/0000 (ventral), and 000 (subcoxal). Ventral psx not visible. AIIIO with 5 papillae, PAO slightly larger than nearest pso, with 3–4 lobes. Labium of ABC-type. Th. III without lateral ms. Distal whorls (A+T) of each tibiotarsus with 9 setae. Furcal remnant of D-type.
Type material. Holotype, male, Russia, Northern Caucasus, North Ossetian State Nature Reserve, vicinity of Verkhniy Tsey., 1676 m alt., green moss-rhododendron pine forest [42.7971°N, 43.9242°E], soil and litter, 20.06.2016, N. Kuznetsova & A. Geraskina leg. Paratype, male, same data as holotype.
Description. Holotype length about 0.8 mm. Colour white in alcohol. Body shape typical of the genus. Granulation regular, slightly coarser around dorsal pso (10–11 granules) (Fig. 2, 1).
Number of pso: 32/022/333431 (dorsal) and 10(?)/000/0000 (ventral) (Figs 2, 1–3). Ventral psx invisible. Upper Sc of legs I–III with neither pso nor psx.
Antennae about as long as head. Ant. IV without thickened S-setae, subapical organite present, microsensillum located in proximal row of setae (Fig. 2, 4). AIIIO consisting of 5 papillae, 5 guard setae, 2 sensory rods, 2 granulated clubs (external one larger) and lateral ms. Ant. I–II with 8 and 12–13 setae, respectively. Antennal area slightly marked. PAO located laterally in a cuticular furrow, with 4(3) lobes and slightly larger than nearest pso. Maxilla unmodified. Maxillary outer lobe simple with 1 basal seta and 2 sublobals. Labrum with 4/342 setae. Labial palp with 6 proximal setae and 10 guards, four of which spiniform as usual, remaining six guards longer and curved. Basomedian and basolateral fields of labium with 4 and 5 setae, respectively. Terminal sensilla of papillae A, B and C thickened (Fig. 2, 5).
Fig. 2. Oligaphorura anocellata sp. n.: 1 – dorsal chaetotaxy; 2 – ventral chaetotaxy of abdomen; 3 – ventral chaetotaxy of head; 4 – antenna, dorsal view; 4a – ibid., ventral view; 5 – labial palp with basomedian and basolateral parts of labium (spiniform guards not shown); 6 – tip of leg III, outer side; 6a – ibid., inner side. Scales, mm: 1–3 – 0.1; 4–6 – 0.01.
Dorsal setae poorly differentiated into macro- and microsetae, symmetrical in general (Fig. 2, 1). S-setae not distinct. Head probably with two axial unpaired seta a0 and a'0 (?), d0 absent as usual for the genus. Setae p1 on head located slightly above of setae p2. Th. I with 4(5)+4(5) dorsal setae. Lateral ms present only on Th. II and absent on Th. III. Terga of Th. II–Abd. III each with 3–4 pairs of axial setae. On Abd. VI, setae a0 and a2 approximately same in size, setae a1 half as long as a0. Thoracic sterna with 0–1–1 setae each side of ventral line. Ventral chaetotaxy as in Figs 2, 2–3.
Upper subcoxae of legs I–III with 2, 3, 3 setae, respectively. Tibiotarsal chaetotaxy with 18(17)–18–17 setae: distal whorl (T+A) with 9 setae (setae T2 and T3 absent), 7 B-setae (B7 absent on Ti III), an unpaired seta M and one seta of C-whorl. Unguis with neither inner nor lateral teeth, unguiculus with a broad basal lamella, approximately as long as 0.8–0.9 of inner edge of unguis (Fig. 2, 6). VT with 6+6 proximal setae and 1+1 setae at base. Furcal remnant of D-type (Fig. 2, 2). Each lateral anal valve with a0 and 2a1 setae, unpaired valve with a0, 2b1 (b0, b2 and a1 absent) and 5 setae in c-row. Anal spines present.
Etymology. The name of the new species reflects the absence of subcoxal pso, which is still unique in the genus.
Affinities. Oligaphorura anocellata sp. n., characterized by a slightly reduced number of tibiotarsal setae and the absence of cuticular fold on the sternum of Abd. IV (furcal remnant of the D-type), can be considered as a member of the differens-group. The new species has the same number of dorsal pso as O. hackeri (Christian 1986), i. e. 32/022/33343, but lacks subcoxal pso. The latter character has never been mentioned among congeners.
In the Collembola fauna of Northern Ossetia it was previously known as O. cf. caucasica (Weiner et Kaprus’) (Kuznetsova et al., 2019) or Oligaphorura sp. 1 (Antipova, Babenko, 2022).
Oligaphorura octosetosa
Shveenkova, Antipova et Babenko sp. n. (Figs 3, 1–6)
Diagnosis. Species of the daii-group. Dorsal sensilla on body poorly marked. Anal spines present. Pseudocellar formulae: 32/012/33343 (dorsal), 11/000/0000 (ventral), and 111 (subcoxal). Ventral psx not visible. AIIIO with 4 papillae, PAO 1.5 times larger than nearest pso, with 3–4 lobes. Labium of ABC-type. Th. III without lateral ms. Distal whorls (A+T) of each tibiotarsus with 8 setae. Furcal remnant of D-type.
Type material. Holotype, male, Russia, Western Caucasus, Krasnodar Krai, ~ 45 km E of Sochi, Krasnaya Polyana, Aibga Range, Laura River basin [43.4827°N, 40.5800°E], mixed (fir-beech) forest with maple, ~1490 m alt., litter, 21.06.2018, N. Kuznetsova & A. Saraeva leg. Paratypes, 6 females, 3 males, same data as holotype.
Description. Size 0.50–0.65 mm, holotype 0.62 mm. Colour white in alcohol. Body shape typical of the genus. Granulation regular, slightly coarser around dorsal pso (9–10 granules) (Fig. 3, 1).
Fig. 3. Oligaphorura octosetosa sp. n.: 1 – dorsal chaetotaxy; 2 – ventral chaetotaxy of abdomen; 3 – ventral chaetotaxy of head; 4 – antenna, dorsal view; 4a – ibid., ventral view; 5 – labial palp with basomedian and basolateral parts of labium (spiniform guards not shown); 6 – tip of leg I, outer side; 6a – ibid., inner side. Scales, mm: 1–3 – 0.1; 4–6 – 0.01.
Number of pso: 32/012/33343 (dorsal)and 11/000/0000 (ventral) (Figs 3, 1–3). Each upper Sc of legs I–III with one pso. Ventral psx invisible.
Antennae slightly shorter than head. Ant. IV with two slightly thickened S-setae, subapical organite present, microsensillum located in proximal row of setae (Fig. 3, 4). AIIIO consisting of 4 papillae, 5 guard setae, 2 sensory rods, 2 granulated clubs (external one larger) and lateral ms. Ant. I–II with 8 and 12–13 setae, respectively. Antennal area not marked. PAO located laterally in a cuticular furrow, with 4(3) lobes, about 1.5 times larger than nearest pso. Maxilla unmodified. Maxillary outer lobe simple with 1 basal seta and 2 sublobals. Labrum with 4/342 setae. Labial palp with 6 proximal setae and 10 guards, four of guards spiniform as usual, remaining six long and curved. Basomedian and basolateral fields of labium with 4 and 5 setae, respectively. Terminal sensilla of papillae A, B and C thickened (Fig. 3, 5).
Dorsal setae only slightly differentiated into macro- and microsetae, symmetrical in general (Fig. 3, 1). S-setae poorly marked and distributed as follows: 11/011/00000 (dorsally), invisible on sterna and lower Sc. Head with two axial unpaired setae a0 and a'0; d0 absent as usual for the genus. Setae p1 on head located above of setae p2. Th. I with 4+4 dorsal setae. Lateral ms present only on Th. II. Terga of Th. II–Abd. III each with 3–4 pairs of axial setae. On Abd. VI setae a0 shorter than a2, setae a1 half as long as a2. Thoracic sterna with 0–1–1 setae each side of ventral line. Ventral chaetotaxy of abdomen as in Fig. 3, 2.
Upper subcoxae of legs I–III with 2, 3, 3 setae, respectively. Tibiotarsal chaetotaxy with 16–16–15 setae: distal whorl (T+A) with 8 setae (setae T2, T3 and A5 absent), 7 B-setae (on Ti III seta B7 absent), and one seta of C-whorl (unpaired seta M absent). Unguis with neither inner nor lateral teeth, unguiculus short, with a narrow basal lamella, about as long as 0.3 of unguis inner edge (Fig. 3, 6). VT with 6+6 proximal setae and 1+1 setae at base. Furcal remnant of D-type (Fig. 3, 2). Each lateral anal valve with a0 and 2a1 setae, unpaired valve with a0, 2b1 (a1, b0 and b2 absent) and 5 setae in c-row. Anal spines present.
Etymology. The name of the new species refers to its most unusual feature, the number of distal setae on the tibiotarsi.
Affinities. Representatives of the daii-group, namely O. daii (Pomorski et al. 1998), O. stojkoae Shvejonkova et Potapov 2012, O. chatyrdagi (Kaprus’ et al. 2002), and O. steposa (Kaprus’ et al. 2002), are all characterized by a pronounced reduction of tibiotarsal chaetotaxy with 5, 6 or 7 distal setae. In this character, O. octosetosa sp. n., which has 8 distal setae on each tibiotarsus, distinctly differs from other species of the group. Moreover, such a number of tibiotarsal setae has not yet been found in any of the congeners. Apart from this, O. octosetosa sp. n., contrary to all other species of the group, has only 4 papillae in AIIIO and one pair of dorsal pso on Th. II (32/012/33343 pso, totally).
Oligaphorura ligni
Shveenkova, Antipova et Babenko sp. n. (Figs 4, 1–6)
Diagnosis. Species of the absoloni-group. Dorsal sensilla poorly marked. Anal spines present. Pseudocellar formulae: 32/133/33343 (dorsal), 11/000/0000 (ventral), 111 (subcoxal). Ventral psx (1+1) present on Abd. II and Abd. IV, as well as on Sc I–III (111). AIIIO with 5 papillae, PAO slightly smaller than (or the same size with) nearest pso, with 3–4 lobes. Labium of AC-type. Th. II–III with lateral ms. Distal whorls (A+T) of each tibiotarsus with 11 setae. Furcal remnant of A-type.
Type material. Holotype, female, Southern Primorye, Chuguev District, National Park “Zov Tigra”, Mount Oblachnaya, Ussuri River valley, mixed forest, 43.6007°N, 134.1930°E, ~550 m alt., rotten wood, 19–20.09.2018, A. Kuprin leg. Paratypes, 5 females, 3 males and 4 juveniles, same data as holotype; 12 females, 3 males and 1 juvenile male, Southern Primorye, Shkotovo District, Mount Falaza, 43.1194°N, 132.7936°E, ~600 m alt., rotten wood with mosses near stream, 08.09.2018, M. Potapov & A. Kuprin leg.
Description. Size 0.75–1.10 mm, holotype 0.75 mm. Colour white in alcohol. Body shape typical of the genus. Granulation regular, slightly coarser around dorsal pso (10–11 granules) (Fig. 4, 1).
Number of pso: 32/133/33343 (dorsal) and 11/000/0000 (ventral) (Figs 4, 1–3). Ventral psx poorly expressed, 0/000/01010 in number (Fig. 4, 2). Each upper Sc of legs I–III with 1 pso and 1 psx.
Fig. 4. Oligaphorura ligni sp. n.: 1 – dorsal chaetotaxy; 2 – ventral chaetotaxy of abdomen; 3 – ventral chaetotaxy of head; 4 – antenna, dorsal view; 4a – ibid., ventral view; 5 – labial palp with basomedian and basolateral parts of labium (spiniform guards not shown); 6 – tip of leg III, anterior side. Scales, mm: 1–4 – 0.1; 5–6 – 0.01.
Antennae about as long as head. Ant. IV with subapical organite, microsensillum located in proximal row of setae (Fig. 4, 4). AIIIO consisting of 5 papillae, 5 guard setae, 2 sensory rods, 2 granulated clubs (external one larger) and lateral ms. Ant. I–II with 9 and 13 setae, respectively. Antennal area not marked. PAO located laterally, in a cuticular furrow, with 4(3) lobes, slightly smaller than (or the same size with) nearest pso. Maxilla unmodified. Maxillary outer lobe simple with 1 basal seta and 2 sublobals. Labral chaetotaxy complete with four prelabral and nine labral setae. Labial palp with 6 proximal setae and 11 guards, four of which spiniform as usual, remaining seven guards long. Basomedian and basolateral fields of labium with 4 and 6 setae, respectively. Terminal sensilla of papillae A and C thickened (Fig. 4, 5).
Dorsal setae poorly differentiated into macro- and microsetae, symmetrical in general (Fig. 4, 1). S-setae expressed only on Th. II–III. Head with two axial unpaired setae: a0 and a0’; d0 absent as usual for the genus. Setae p1 on head located at level with p2. Th. I with 7(9)+7(9) dorsal setae. Lateral ms present on both Th. II–III. Terga of Th. II–Abd. III each with 3–4 pairs of axial setae. Abd. IV usually with unpaired seta m0, more rarely a pair of m1 present. On Abd. VI seta a0 slightly shorter than p0, setae a1 as long as a0 and much shorter than a2. Thoracic sterna with 0–1–1 setae each side of ventral line. Ventral chaetotaxy as in Figs 4, 2–3.
Upper subcoxae of legs I–III with 3–4, 3–5, 3–5 setae, respectively. Tibiotarsal chaetotaxy with 20–20–19 setae, respectively: distal whorl (T+A) with 11 setae, 7 B-setae (B7 absent only on Ti III), an unpaired seta M and one seta of C-whorl. Unguis with neither inner nor lateral teeth, unguiculus with a narrow basal lamella, about as long as inner edge of unguis (1.0–1.1: 1) (Fig. 4, 6). VT with 8–9+8–9 proximal setae and 2+2 setae at base. Furcal remnant of A-type (Fig. 4, 2). Each lateral anal valve with a0 and 2a1 setae, unpaired valve with a0, 2b1, 2b2 (a1 and b0 absent) and 5 setae in c-row. Anal spines present.
Etymology. The name of the species reflects its habitat preference – decaying wood.
Affinities. O. ligni sp. n. belongs to the absoloni-group and is rather similar to O. shifangensis Liu et Sun 2019, described from southwestern China. Both species are characterized by complete distal whorls on tibiotarsi (11 setae), the furcal remnant of A-type, the same numbers of dorsal pso (32/133/33343), and the labium of AC-type. The new species can be distinguished by a longer unguiculus (about as long as the inner edge of the unguis, vs 0.7 of unguis length in O. shifangensis), a smaller PAO (slightly smaller than the nearest pso, vs 1.5–2.0 pso in O. shifangensis), fewer ventral psx (0/000/010100 in the new species, vs 0/000/221101+1m in O. shifangensis). The new species is also characterized by the more pronounced differentiation of the macro- and microsetae on Abd. V: axial setae m1 only slightly shorter than setae a1 and p1. Both a1 and a3 are about twice as long as setae a2 and a4. Contrary, in O. shifangensis setae m1 (microsetae) are clearly shorter than a1 and p1 (mesosetae) and the corresponding mesosetae of a-row (a1 and a3) are about 1.5 times longer than microsetae (a2 and a4).
Oligaphorura yakutica
Shveenkova, Antipova et Babenko sp. n. (Figs 5, 1–7)
Diagnosis. Body shape typical of the genus. Dorsal sensilla indistinct. Anal spines present. Pseudocellar formulae: 32/233/33343 (dorsal), 11/000/0000 (ventral), 111 (subcoxal). Ventral psx invisible, Sc I–III with 111 psx. AIIIO with 5 papillae, PAO slightly larger than nearest pso, with 3–5 lobes. Labium of AC-type. Th. II–III with lateral ms. Distal whorls (A+T) of each tibiotarsus with 11 setae. Furcal remnant of H-type.
Type material. Holotype, male, Russia, Republic of Sakha (Yakutia), vicinity of Ust’-Nera [64.521°N, 143.179°E], lichen larch forest, ~700 m alt., soil and litter, 22.07.1992, N. Kuznetsova & M. Potapov leg. Paratypes, 1 female, 2 males, same data as holotype.
Description. Size 0.98–1.20 mm, holotype 0.98 mm. Colour white in alcohol. Body shape typical of the genus. Granulation regular, not coarser around dorsal pso (12 granules) (Fig. 5, 1).
Number of pso: 32/233/33343 (dorsal) and 11/000/0000 (ventral) (Figs 5, 1–3). Pseudocelli slightly elongated (Fig. 5, 4). Ventral psx invisible. Each upper Sc of legs I–III with one pso.
Fig. 5. Oligaphorura yakutica sp. n.: 1 – dorsal chaetotaxy; 2 – ventral chaetotaxy of abdomen; 3 – ventral chaetotaxy of head; 4 – pso; 5 – antenna, dorsal view; 5a – ibid., ventral view; 6 – labial palp with basomedian and basolateral parts of labium (spiniform guards not shown); 7 – tip of leg I, outer side. Scales, mm: 1–3, 5–0.1; 6–7 – 0.01; 4–0.005.
Antennae somewhat shorter than head. Ant. IV with 10–12 slightly thickened sensilla, subapical organite present, microsensillum located in proximal row of setae (Fig. 5, 5). AIIIO consisting of 5 papillae, 5 guard setae, 2 sensory rods, 2 granulated clubs (external one larger) and lateral ms. Ant. I–II with 8 and 13 setae, respectively. Antennal area not marked. PAO located laterally in a cuticular furrow, with 3–5 lobes, slightly larger than nearest pso. Maxilla unmodified. Maxillary outer lobe simple with 1 basal seta and 2 sublobals. Labrum with 4/342 setae. Labial palp with 6 proximal setae, four spiniform and seven long guards. Basomedian and basolateral fields of labium with 4 and 6 setae, respectively. Terminal sensilla of papillae A and C thickened (Fig. 5, 6).
Dorsal setae poorly differentiated into macro- and microsetae, symmetrical in general (Fig. 5, 1). S-setae indistinct. Head with a single axial unpaired seta a0; d0 absent as usual for the genus. Setae p1 on head in front of setae p2. Th. I with 4(5)+4(5) dorsal setae. Lateral ms present on both Th. II and Th. III. Terga of Th. II–Abd. III each with 3–4 pairs of axial setae. On Abd. VI seta a0 similar in length to setae a2, setae a1 three times shorter than a0. Thoracic sterna without setae. Ventral chaetotaxy of abdomen as in Fig. 5, 2.
Upper subcoxae of legs I–III with 3, 4, 5 setae, respectively. Tibiotarsi with 20–20–19 setae: distal whorl (T+A) with 11 setae, 7 B-setae (B7 absent only on Ti III), an unpaired seta M and one seta of C-whorl. Unguis with neither inner nor lateral teeth; unguiculus with broad basal lamella, slightly shorter than unguis (about 0.9 of its inner edge length) (Fig. 5, 7). VT with 6+6 proximal setae and 1+1 setae at base. Furcal remnant of H-type (Fig. 5, 2). Each lateral anal valve with a0 and 2a1 setae, unpaired valve with a0, 2b1, 2b2 (a1 and b0 absent) and 5 setae in c-row. Small anal spines present.
Etymology. The new species was named after the region of its type locality.
Affinities. Oligaphorura yakutica sp. n. is rather similar to O. duocellata Babenko et Fjellberg 2015, described from the upper reaches of Kolyma River, Magadan Region. Both of these species are characterized by such unusual features as the absence of setae on thoracic sterna, the presence of two pairs of dorsal pso and a few setae on Th. I, as well as a distant position of a- and b-pso on Abd. IV. We tentatively include these two species into the groenlandica-group even though they have a furcal remnant of H-type (in the form of 4 small setae behind a finely granulated area) which is not typical of the group. Oligaphorura yakutica sp. n. can easily be distinguished from O. duocellata by the absence of ventral pso on the abdomen (vs 1111(2) in O. duocellata).
The presence of two pairs of dorsal pso on Th. I is uncommon among Oligaphorura species; previously this character was considered characteristic of only O. quadrituberculata (Börner 1901). This latter species has slightly more numerous dorsal pso, i. e. 32/2(3)3(4)3(4)/33(4)333, and only 3 pso on Abd. IV (vs 4 in the new species), a shorter unguiculus (half of the inner edge of the unguis, vs 3/4 in O. yakutica sp. n.), and the furcal remnant with a cuticular furrow (absent in O. yakutica sp. n.).
One more species, i. e. O. ossetica sp. n., also having 2+2 pso on Th. I, is described in this paper. It belongs to a different species-group and is hardly comparable to O. yakutica sp. n.
Oligaphorura subnuda
Shveenkova, Antipova et Babenko sp. n. (Figs 6, 1–8)
Diagnosis. Species of the groenlandica-group. Body shape as usual for the genus. Anal spines present. Dorsal sensilla not distinguishable. Pseudocellar formulae 32/033/44454 (dorsal), 11/000/0001 (ventral), and 111 (subcoxal). Ventral and subcoxal psx invisible. AIIIO with 5 papillae, granulated clubs distinctly differing in sizes. PAO usually with 3 lobes, slightly smaller than nearest pso. Labial palp of the AC-type. Lateral ms present on both Th. II–III. Thoracic sterna with 0–0–1 setae on each side of the ventral line. Tibiotarsal chaetotaxy complete: distal whorls (A+T) of each Ti with 11 setae. Furcal remnant of S-type, cuticular fold distinct.
Type material. Holotype, female, Russia, Taimyr Peninsula, middle reaches of Pyasina River, former Ust’-Tareya [73.2529°N, 90.5962°E], Dryas punctata association on upper part of river slope, 22.07.2010, O. Makarova & A. Babenko leg. Paratypes, 3 females, 3 males and 2 juveniles, same data as holotype; 2 females and 3 males, same area and date but herbaceous meadow on river slope; 3 females and 2 males, same area, lemming’s colony, 02.08.2010, all O. Makarova & A. Babenko leg.
Description. Size 0.58–0.71 mm, holotype 0.63 mm. Colour white in alcohol. Body cylindrical. Granulation fine and uniform, with 10–12 granules around dorsal pso (Figs 6, 1, 5).
Number of pso stable within available material and expressed as 32/033/44454 (dorsal) and 11/000/0001 (ventral) (Figs 6, 1–2). Ventral and subcoxal psx invisible. Upper Sc of legs I–III with 1 pso each (Fig. 6, 3).
Fig. 6. Oligaphorura subnuda sp. n.: 1 – dorsal chaetotaxy; 2 – ventral chaetotaxy of abdomen; 3 – Th. II–III and Sc II, lateral view; 4 – chaetotaxy of anterior part of head; 5 – antennal pso and PAO; 6 – antennal segments III–IV; 7 – labial palp; 8 – tip of leg I. Scales, mm: 1–2 – 0.1; 3–8 – 0.01.
Antennae slightly shorter than head, Ant. III–IV broad, club-like. Subapical organite present on Ant. IV, microsensillum located in proximal row of setae (Fig. 6, 6). AIIIO consisting of 5 papillae, 5 guard setae, 2 sensory rods, 2 granulated clubs (external one clearly larger) and lateral ms. Antennal area not marked. Ant. I–II with 8 and (12)13 setae, respectively. PAO smaller than nearest pso with 3 subequal lobes as a rule (Fig. 6, 5). Maxillary lamellae unmodified, maxillary outer lobe simple with 1 basal seta and 2 sublobals. Labrum with 4/322 setae. Labial palp with thickened terminal sensilla on papillae A and C, 10 guards (6 long and 4 spiniform) and 6 proximal setae (Fig. 6, 7). Basomedian and basolateral parts of labium with 4 and 5 setae, respectively.
Dorsal chaetotaxy almost symmetrical, especially in medial parts of terga, macrosetae clearly distinguishable only laterally and on abdominal tip (Fig. 6, 1), meso- and microsetae similar in size but differentiated in form: mesosetae usually straight and truncate at tips, microsetae pointed and curved. S-setae not distinguishable as a rule. Head with two axial unpaired setae (a0 and a'0) on frontal part (Fig. 6, 4), d0 being absent as usual for the genus. Setae p1 on head and Th. II–III usually located in front of p2. Th. I with 6–7 dorsal setae on each side. Lateral ms present on both Th. II and Th. III (Figs 6, 3). Terga of Th. II–Abd. III with 3 pairs of axial microsetae and a pair of mesosetae located slightly laterally. Abd. IV–V often with unpaired setae: m0 on Abd. IV and p0 on Abd. V. Setae a0 on Abd. VI slightly shorter than a2, setae a1 almost half as long as a2.
Thoracic sterna with 0–0–1(2) setae each side of ventral line. VT with (6)7+7 proximal and 2+2(1) setae at base. Ventral chaetotaxy as in Fig. 6, 2. Furcal remnant of S-type, usually with 6+6 setae arranged in four irregular rows. Each lateral anal valve with a0 and 2a1 setae, unpaired valve with a0, 2b1, 2b2 (a1 and b0 absent) and 7 setae in c-row.
Upper subcoxae of legs I–III usually with 4, 5, 4 setae, respectively; lower subcoxae with 0, 3, 3 setae; coxae – 3, 8, 8 setae; trochanters – 8, 8, 8 setae and femora – 12, 12, 12 setae. Tibiotarsi with 20–20–19 setae: distal whorls (T+A) complete, totally with 11 setae, 7 B-setae (B7 absent on Ti III), an unpaired seta M within B-whorl and one C-seta on each leg. Unguis with neither inner nor lateral teeth, unguiculus about as long as 3/4 of inner edge of unguis with a narrow but clear basal lamella (Fig. 6, 8). Anal spines rather strong, usually constricted at base, set without papillae.
Etymology. The name of the new species reflects its high similarity with the Nearctic O. nuda (Fjellberg 1987).
Affinities. The species was earlier referred to in the East-Palaearctic fauna as O. sp. aff. nuda (see Babenko, 2013). It is very similar to this Nearctic form indeed, also characterized by the presence of additional pso on each side of abdominal terga and the absence of pso on Th. I. The only characters by which these species can be reliably distinguished are the presence (O. subnuda sp. n.) or absence (O. nuda) of ventral pso on Abd. IV, and the type of the labium (AC- type in the new species, vs A- type in O. nuda (after Fjellberg, 1999)).
Another similar form, O. interrupta (Fjellberg 1987), inhabits more eastern parts of the Palaearctic, ranging from Yakutia to the Magadan Region (Babenko, Fjellberg, 2015). The most common dorsal pseudocellar formula in this species is 32/033/33353, but variations are frequent and specimens with additional pso on some abdominal terga are also observed. The whole formula may be expressed as follows, 32/033/3(4).3(4).3(4).5(4).3(4), which covers that in O. subnuda sp. n. Nevertheless, among the vast available material of O. interrupta, there are no specimens with ventral pso on Abd. IV.
The new species clearly prefers the warmest sites in the area of its type locality, being mainly found on south-facing slopes. Thus, it may well be presumed that the main part of its distributional range encompasses more southern territories than the region where it is still found.
Oligaphorura cavicola
Shveenkova, Antipova et Babenko sp. n. (Figs 7, 1–7)
Diagnosis. Species of the groenlandica-group. Body shape as usual for the genus. Anal spines present. Dorsal sensilla not distinguishable. Pseudocellar formulae 32/133/44454 (dorsal), 11/000/0001 (ventral), and 111 (subcoxal). Ventral and subcoxal psx present. AIIIO with 5 high papillae, granulated clubs distinctly differing in shape and size. PAO with 4 lobes, its longer axis about as long as 1.5–1.7 of nearest pso. Labial palp of the AC-type. Lateral ms present on both Th. II–III. Thoracic sterna with 0–0-(0)1 setae on each side of the ventral line. Tibiotarsal chaetotaxy complete: distal whorls (A+T) of each Ti with 11 setae. Furcal remnant of S-type, cuticular fold distinct.
Type material. Holotype, male, Russia, north of Perm Region, 10 km north of Nyrob, Kolva River basin, Cave Div’ya [60.8097° N, 56.7417° E], grotto Kaban [about 400 m from the entry], 07.01.2007, I. Lavrov leg. Paratypes, 3 females and 4 males, same data as holotype.
Description. Size 1.2–1.6 mm, holotype 1.2 mm. Colour white in alcohol. Body cylindrical (Fig. 7, 1). Granulation fine and uniform, with 12–14 granules around dorsal pso.
Fig. 7. Oligaphorura cavicola sp. n.: 1 – dorsal chaetotaxy; 2 – ventral chaetotaxy of abdomen; 3 – AIIIO; 4 – sensory rods and granulated clubs of AIIIO; 5 – labial palp; 6 – PAO and antennal pso; 7 – tip of leg III. Scales, mm: 1–2 – 0.1; 3–7 – 0.01.
Number of pso stable within available material and expressed as 32/133/44454 (dorsal) and 11/000/0001 (ventral) (Figs 7, 1–2). Ventral psx usually as 0/000/?11000. Upper Sc of legs I–III with one pso and one psx each, one psx also visible on inner side of each femora.
Antennae about as long as head, Ant. III–IV broad, club-like. Subapical organite present on Ant. IV, microsensillum located in proximal row of setae (Fig. 7, 3). AIIIO consisting of 5 long papillae, 5 guard setae, 2 sensory rods, 2 granulated clubs (external one clearly larger and stronger curved) (Fig. 7, 4) and lateral ms. Antennal area not marked. Ant. I–II with 8 and 12–14 setae, respectively. PAO about as long as 1.5–1.7 of nearest pso with 4 lobes, anterior pair clearly longer (Fig. 7, 6). Maxillary lamellae unmodified, maxillary outer lobe simple with 1 basal seta and 2 sublobals. Labrum with 4/9 setae. Labial palp with thickened terminal sensilla on papillae A and C, 10 guards (6 long and 4 spiniform) and 6 proximal setae (Fig. 7, 5). Basomedian and basolateral parts of labium with 4 and 5 setae, respectively.
Dorsal chaetotaxy more or less symmetrical (Fig. 7, 1), although intraspecific variations frequent; macrosetae well differentiated especially on lateral parts of terga, meso- and macrosetae usually straight and truncate at tips, microsetae pointed and curved. S-setae not distinguishable. Head with one axial unpaired setae (a0) on frontal part, d0 being absent as usual for the genus. Setae p1 on head usually in front of p2. Th. I with 5–6 dorsal setae on each side. Lateral ms present on both Th. II and Th. III. Terga of Th. II–Abd. III usually with 3 pairs of axial microsetae and a pair of mesosetae of similar size located slightly laterally. Chaetotaxy of medial part of Abd. IV unstable; axial mesosetae (a1 and p1) on Abd. V rather long and subequal to microsetae m1, unpaired seta p0 present or absent. Seta a0 on Abd. VI slightly longer than a1, the latter about half as long as a2.
Thoracic sterna with 0-0-(0)1 setae each side of ventral line, sometimes seta present only on one side of Th. III. VT with 6+6 proximal and 2+2(1) setae at base as a rule. Ventral chaetotaxy as in Fig. 7, 2. Furcal remnant of S-type, setae on manubrial field usually arranged in four irregular rows. Each lateral anal valve with a0 and 2a1 setae, unpaired valve with a0, 2a1, 2b1, 2b2 (b0 absent) and usually 7 setae in c-row.
Upper subcoxae of legs I–III with 4(5), 4(5), 4 setae, respectively; lower subcoxae with 0, 3, 3 setae; coxae – 3, 7–9, 9–12 setae; trochanters – 8–9, 8–9, 9–11 setae and femora – 14, 14–15, 11–14 setae. Tibiotarsi with 20–20–19 setae: distal whorls (T+A) complete, totally with 11 setae, 7–7–6 B-setae (B7 absent on Ti III), an unpaired seta M within B-whorl and one C-seta on each leg. Unguis long and narrow with neither inner nor lateral teeth, unguiculus about as long as 0.8 of inner edge of unguis with clear basal lamella (Fig. 7, 7). Anal spines rather strong, slightly constricted at base, set almost without papillae.
Etymology. The name of the new species reflects its cavernicolous style of life.
Affinities. The most characteristic feature of the new species is the increased number of dorsal pso on all abdominal segments, which, however, is not unique for the genus and occurs among a fairly large number of congeners. On the other hand, in most of such cases, additional pso are developed not only on the abdomen, but also on the head and thorax. The basic set of dorsal pso in the anterior part of the body (on the head and thorax) and an increase in their number on abdominal segments is known only for eight described species. Of these, five species, namely O. reversa (Fjellberg 1987), O. nuda (Fjellberg 1987), O. interrupta (Fjellberg 1987), O. arnei Paśnik et Kaprus’ 2019, and O. subnuda sp. n. lack pso on the first thoracic segment, making them quite easy to distinguish from O. cavicola sp. n. It is also fairly easy to distinguish the new species from the Spanish O. alavensis (Simón-Benito et Luciáñez 1994), which has a similar dorsal pso formula, i. e. 32/133/44463, but lacking anal spines.
Only two remaining species, O. uralica (Khanislamova 1986) and O. changbaiensis (Sun et Wu 2012), show a dorsal pso formula completely identical to that of O. cavicola sp. n. Of these, the almost sympatric O. uralica differs significantly in the number of subcoxal (Sc I–III with 233 pso, vs 111 in O. cavicola sp. n.) and ventral pso (2/000/1112, vs 2/000/0001 in O. cavicola sp. n.). The Chinese species, O. changbaiensis, formally seems to be most similar to the new species, although it is characterized by a different type of furcal chaetotaxy (A-type, vs S-type in O. cavicola sp. n.) and therefore was originally described in the genus Micraphorura Bagnall 1949. These two species also differ in a number of other characteristics, for instance, the labial type (ABC, vs AC in O. cavicola sp. n.), the number of ventral pso (2/000/0000, vs 2/000/0001 in O. cavicola sp. n.) and psx on the upper subcoxae (222, vs 111 in O. cavicola sp. n.) and ventrally on the abdomen (2222, vs ?110 in O. cavicola sp. n.).
Formally, O. cavicola sp. n. is also quite similar to O. subnuda sp. n., which is described above. Both species have almost identical dorsal, ventral and subcoxal pso formulas and are found in the central regions of the Palaearctic. Besides the presence of pso on Th. I only in O. cavicola sp. n., these two species differ significantly in size (1.2–1.6 mm in O. cavicola sp. n., vs 0.6–0.7 mm in O. subnuda sp. n.), the differentiation of dorsal setae, the relative size of the PAO, the number of labral setae, and the presence/absence of subcoxal and ventral psx.
Oligaphorura neglecta
Shveenkova, Antipova et Babenko sp. n. (Figs 8, 1–3)
Diagnosis. Species of the groenlandica-group. Body shape as usual for the genus. Anal spines present. Dorsal sensilla present but poorly distinguishable. Pseudocellar formulae 32/133/33343 (dorsal), 11/000/0000 (ventral), 111 (subcoxal). Ventral and subcoxal psx present. AIIIO with 5 papillae, granulated clubs distinctly differing in sizes. PAO usually with 3 lobes, its longer axis clearly longer than nearest pso diameter. Labial palp of the AC-type. Lateral ms present on both Th. II–III. Axial part of Abd. V with setae m1 clearly shorter than a1 and p1. Thoracic sterna with 0–1–1(2) setae on each side of the ventral line. Tibiotarsal chaetotaxy complete: distal whorls (A+T) of each Ti with 11 setae. Furcal remnant of S-type, cuticular fold distinct.
Type material. Holotype, female, Russia, Taimyr Peninsula, middle reaches of Pyasina River, former Ust’-Tareya [73.2529°N, 90.5962°E], Dryas punctata association on pingo, 22.07.2010, O. Makarova & A. Babenko leg. Paratypes, 2 males and 2 juveniles, same data as holotype; 11 females, 6 males and 5 juveniles, same area, nival slope with Cassiope tetragona, 27.07.2010; 5 females, 2 males and 1 juvenile, same area, polygonal mire, 01.08.2010; 4 females, same area, tussock tundra, 28.07.2010; 2 males, same area, zonal tundra association, 27.7.2010; 2 males and 11 juveniles, same area, lemming’s colony, 02.08.2010; 2 females, male and juvenile, same area, Betula nana association, 27.7.2010; all O. Makarova & A. Babenko leg.
Additional material. 2 females, 2 males and juvenile, Novosibirsk Islands, Kotelnyi Island [75.05°N, 140.17°E], slope of valley, 31.07.1994, M. Berezin leg.; 2 females and male, same area and date, nival slope, mosses, A. Babenko leg.; female, Yakutia (Sakha Republic), Suntar-Khayata Mt. Range, upper reaches of Kyubyume River [63.22°N, 139.53°E], 1250 m alt., Salix spp. in floodplain, 30.07.2002, O. Makarova leg.; 3 females and 1 male, same republic, Leno-Vilyui interfluve, environs of Asyma [62.4003°N, 126.7931°E], burnt larch forest, 28.06.2022 and 1 male, same biotope, but 18.08.2022, A. Burnasheva leg.
Description. Size 0.7–1.0 mm (males), 0.9–1.3 (females), holotype 0.89 mm. Colour white in alcohol. Body shape as usual for genus. Granulation fine and uniform, with 10–12 granules around dorsal pso.
Number of pso stable within available material and expressed as 32/133/33343 (dorsal) and 11/000/0000 (ventral). Ventral psx present, their number rather variable: head as a rule with one psx near basolateral part of labium, Abd. I–IV usually with 2222 ventral psx and one psx on each anal lobe (1/000/222201+1m as a whole) (Fig. 8, 1). Upper Sc of legs I–III with one pso each and usually with (1)2–2–2 psx.
Fig. 8. Oligaphorura neglecta sp. n. (1–3) and Oligaphorura groenlandica (4–6): 1, 4 – ventral chaetotaxy of abdomen (1 – male, 4 – female); 2, 5 – sensory rods and granulated clubs of AIIIO; 3, 6 – dorsal chaetotaxy of Abd. V–VI. Scales, mm: 1, 3, 4, 6–0.1; 2, 5–0.01.
Antennae slightly shorter than head, Ant. III–IV broad, club-like. Subapical organite present on Ant. IV, microsensillum located in proximal row of setae. AIIIO consisting of 5 high and narrow papillae, 5 guard setae, 2 sensory rods, 2 granulated clubs (external one clearly larger (Fig. 8, 2)) and lateral ms. Antennal area not marked. Ant. I–II with (8)9 and 14–15 setae, respectively. PAO with 3 lobes as a rule, its longer axis almost 2 times as long as pso diameter. Maxillary lamellae unmodified, maxillary outer lobe simple with 1 basal seta and 2 sublobals. Labrum with 4/522 setae. Labial palp with thickened terminal sensilla on papillae A and C, 10 guards (6 long and 4 spiniform) and 6 proximal setae. Basomedian and basolateral parts of labium with 4 and 5 setae, respectively.
Dorsal chaetotaxy in medial parts of terga almost symmetrical, macrosetae clearly distinguishable only on abdominal tip, setae on other parts of body similar in size but differentiated in form: mesosetae usually more straight and truncate at tips, microsetae pointed and curved. Dorsal S-setae poorly distinguishable from mesosetae, their highest seen number as 2/011/22211. Head with one axial unpaired seta (a0) on frontal part, d0 being absent as usual for the genus. Setae p1 on head in front of p2. Th. I with (7)8–9 dorsal setae on each side. Lateral ms present on both Th. II and Th. III. Terga of Th. II–Abd. III with 3 pairs of axial microsetae and a pair of mesosetae located slightly laterally. Chaetotaxy of medial part of Abd. IV usually more irregular. Axial microsetae m1 on Abd. V clearly shorter than mesosetae a1 and p1 (Fig. 8, 3). Mesoseta a0 on Abd. VI subequal to microsetae a1 and about 1.5 times shorter than macrosetae a2.
Thoracic sterna with 0–1–1(2) setae each side of ventral line. VT with up to 9 proximal setae on each side and usually 2+2 setae at base. Furcal remnant of S-type with clear cuticular fold and 14–16 setae on manubrial field (Fig. 8, 1). Each lateral anal valve with a0 and 2a1 setae, unpaired valve with a0, 2b1, 2b2 and 5 setae in c-row (b0 and a1 absent).
Upper subcoxae of legs I–III with 4, (5)6, (5)6 setae, respectively; lower subcoxae with 0, 3, 4 setae; coxae – 3, 8–11, 12–14 setae; trochanters – 9–10, 9–10, (8)9 setae and femora – 15–18, 16–18, (14)15 setae. Tibiotarsi with 20–20–19 setae: distal whorls (T+A) complete, totally with 11 setae, 7 B-setae (B7 absent on Ti III), an unpaired seta M within B-whorl and one C-seta on each leg. Unguis without inner tooth, lateral ones present only occasionally, unguiculus about as long as 3/4 of inner edge of unguis with a narrow but clear basal lamella. Anal spines rather strong and curved, usually set on low papillae.
Etymology. The name of the new species reflects the fact that, despite being known from the late 80s of the last century, it was described only now.
Affinities. Oligaphorura neglecta sp. n. was first found on the Taimyr Peninsula by A. Fjellberg in the summer of 1989, and till now it was usually considered as a bisexual form of the variable O. groenlandica (Tullberg 1877) (see Fjellberg, 1998; Babenko, Fjellberg, 2015). This point of view seems quite justified, since for some species of the subfamily Onychiurinae the existence of large parthenogenetic and smaller bisexual forms has already been experimentally confirmed (Hale, 1964). The morphological characters commonly used to separate species in the genus Oligaphorura, such as AIIIO and PAO, labrum and labium, number of dorsal, ventral and subcoxal pso, the presence of ms on Th. III, position of “dental” setae on manubrial field, tibiotarsal chaetotaxy, are all identical in both species. The presence of males also does not allow us to reliably distinguish these species. In most of the northern regions of the Holarctic, the true O. groenlandica is only represented by females, but males are still found in some populations. For example, we noted them in the area of Pevek Town, western Chukotka. As a result, apart from a smaller size (0.7–1.3 mm in O. neglecta sp. n., vs 1.3–1.7 mm in O. groenlandica), the only notable characters which truly separate O. neglecta sp. n. from O. groenlandica, are the structure of the furcal remnant (presence vs absence of a cuticular fold, cf. Fig. 8, 1 and Fig. 8, 4), the more pronounced differences in size of AIIIO sensilla (cf. Fig. 8, 2 and Fig. 8, 5) and the differentiation of axial setae on Abd. V: curved microsetae m1 are shorter than straight mesosetae a1 and p1 in the former species (Fig. 8, 3), whereas in O. groenlandica setae m1 are clearly longer than other axial setae (Fig. 8, 6). Apparently, the psx formulas also differ in the species compared, but their rather high variability (see Babenko, Fjellberg, 2015) in both species does not allow them to be reliably separated by this trait. The presence of lateral teeth on the unguis of O. groenlandica, mentioned for this species by Fjellberg (1998), is sometimes also seen in O. neglecta sp. n.
In our opinion, O. neglecta sp. n. is probably not the only cryptic species in the groenlandica complex, which clearly requires additional attention, primarily in relation to the “southern” populations.
Oligaphorura primorica
Shveenkova, Antipova et Babenko sp. n. (Figs 9, 1–7)
Diagnosis. Species of the groenlandica-group. Body shape as usual for the genus; head, thorax and Abd. VI with fields of coarser cuticular granulation; dorsal setae clearly differentiated into long and straight macrochaetae and short and curved microsetae. Anal spines present. Dorsal sensilla invisible. Pseudocellar formulae as 32/033/33343 (dorsal), 11/000/0000 (ventral), and 111 (subcoxal). Ventral and subcoxal psx present. AIIIO with 5 papillae, granulated clubs distinctly differing in sizes. PAO with (3)4 lobes, 1.5–2.0 times longer than nearest pso diameter. Labial palp of the AC-type. Lateral ms present on both Th. II–III. Thoracic sterna with 0–1–1 setae on each side of the ventral line. Tibiotarsal chaetotaxy complete: distal whorls (A+T) of each Ti with 11 setae. Unguis with inner and a pair of lateral teeth. Furcal remnant of S-type, cuticular fold small but distinct.
Type material. Holotype, male, Russia, Southern Primorye, Vladivostok Botanical Garden, coniferous-broad-leaved forest [43.2222°N 131.9936°E], winter traps, 20 November 2016–11 March 2017, A. Komisarenko leg.
Description. Size of a single available specimen (holotype) 1.53 mm. Colour white in alcohol. Body shape as usual for genus. Granulation of dorsal side of body very characteristic with fields of coarser granulation on head, all three thoracic terga and Abd. VI; similar fields developed also on upper subcoxae and Ant. I–II (Fig. 9, 1).
Fig. 9. Oligaphorura primorica sp. n.: 1 – dorsal chaetotaxy; 2 – ventral chaetotaxy of abdomen; 3 – dorsal chaetotaxy of Abd. IV–VI, lateral view; 4 – antennal segments III–IV; 5 – sensory rods and granulated clubs of AIIIO; 6 – PAO and antennal pso; 7 – tip of leg III. Scales, mm: 1–3 – 0.1; 4–7 – 0.01.
Number of pso expressed as 32/033/33343 (dorsal) and 11/000/0000 (ventral) (Figs 9, 1–2), on head two anterior medial pso and one posterior pso set within coarsely granulated part (Figs 9, 1). Ventral psx formula as 0/000/111101m. Upper Sc of legs I–III with one pso and one ventro-lateral psx each.
Antennae about as long as head; Ant. III–IV broad, club-like (Figs 9, 4). Subapical organite present on Ant. IV, microsensillum located in proximal row of setae; dorsal side of segment with two usual types of setae, curved and more straight ones, and also with two slightly thickened sensilla on inner side. AIIIO (Fig. 9, 4–5) consisting of 5 high and thick papillae, 5 guard setae, 2 long and thin sensory rods, 2 granulated clubs (external one curved and clearly larger) and lateral ms. Antennal area clearly marked by tegumental granulation. Ant. I–II with 9 and 16–18 setae, respectively. PAO with (3)4 lobes, its longer axis almost 2 times as long as pso diameter (Fig. 9, 6). Maxillary lamellae unmodified, maxillary outer lobe simple with 1 basal seta and 2 sublobals. Labrum with 4/522 setae, four distal setae clearly longer. Labial palp with thickened terminal sensilla on papillae A and C, 10 guards (6 long and 4 spiniform) and 6 proximal setae. Basomedian and basolateral parts of labium with 4 and 5 setae, respectively.
Dorsal chaetotaxy (Figs 9, 1, 3) in a single available specimen slightly irregular even in medial part of terga, with many additional setae, especially laterally; macrosetae clearly distinguishable, strong and rather long, microsetae short, pointed and curved. Dorsal S-setae undifferentiated. Head with one axial unpaired seta (a0) on frontal part. Setae p1 on head long and clearly set in front of p2. Th. I with 7 dorsal setae on each side. Th. II–Abd. III with a1 as macrosetae. Lateral ms present on both Th. II and Th. III. Mesoseta a0 on Abd. VI subequal to microsetae a1 and about half as long as macrosetae a2.
Thoracic sterna with 0–1–1 setae on each side of ventral line. VT with 9+10 proximal setae, 2+2 setae on anterior side and 2+3 setae at base. Ventral chaetotaxy of abdomen rather abundant (Fig. 9, 2), setae differentiated only on abdominal tip. Furcal remnant of S-type with small, but clear cuticular fold, manubrial field with 19 setae in several irregular rows.
Upper subcoxae of legs I–III with 4, 6, 5 setae, respectively; lower subcoxae with 0, 3, 3–4 setae; coxae – 3, 11, 14 setae; trochanters – 10, 10, 9–10 setae and femora – 19, 18, 16 setae. Tibiotarsi with 20–20–19 setae: distal whorls (T+A) complete, totally with 11 setae, 7 B-setae (B7 absent on Ti III), an unpaired seta M within B-whorl and one C-seta on each leg. Unguis with clear inner tooth and a pair of lateral ones in basal half, unguiculus about as long as 3/4 of inner edge of unguis with narrow basal lamella (Fig. 9, 7). Anal spines conical, set without papillae on coarsely granulated belt of Abd. VI (Fig. 9, 3).
Etymology. The name of the new species is associated with the region of its supposed distribution – i. e. Southern Primorye.
Affinities. Despite the limited material (only one specimen of this species was found in three-month trap collections), we venture to describe it, since it is hard to confuse with any of the known species, even taking into account the very possible variability of some characters. First of all, the peculiar coarse granulation of some parts of the body, such as the head, thoracic segments and the tip of the abdomen, should be noted. Enlarged granulations on some segments are certainly not a unique feature of the genus, but a similar type of granulation that covers only the anterior part of the body and the last segment of the abdomen, is only known in four species of the genus, namely O. tottabetsuensis (Yosii 1972), O. koreana (Weiner 1994), O. linderae (Weiner 1994), and O. judithae (Weiner 1994). All of them were described from adjacent East Asian regions: the former species from Hokkaido, and the other three from the Korean Peninsula. Most likely they represent an isolated regional group of related species, and O. primorica sp. n. occupies a somewhat isolated position in it, differing from all species of this group in a noticeable differentiation of the dorsal setae. Among representatives of this group, O. primorica sp. n. shares the same number of dorsal and ventral pso only with O. koreana. It is also possible that an identical dorsal pso formula is also characteristic of O. tottabetsuensis. This latter species also lacks pso on Th. I, whereas the presence of only three pso on Abd. IV is postulated, which, in our opinion, needs confirmation. Apart from weakly differentiated dorsal setae and the absence of teeth from the unguis, these two species, O. koreana and O. tottabetsuensis, differ quite clearly from O. primorica sp. n. in the location of the pso on the head. In O. primorica sp. n. only one pso of the anterior group is located inside the antennal base (within the area of finer granulation), vs two such pso in O. tottabetsuensis, and all three pso in O. koreana. In addition, in the latter species, both posterior pso on the head are located beyond the coarsely granulated field, vs only one pso in O. primorica sp. n. or both pso located “at the margin of the granulate area” (Yosii, 1972, p. 84) in O. tottabetsuensis. The other two species of the group are characterized not only by the presence of pso on the first thoracic segment, but also by a greater number of pso at the end of the abdomen (32/133/33354, totally).
In the Far Eastern regions, only one other species of the genus, O. aborigensis (Fjellberg 1987), is known, which, like O. primorica sp. n., shows a differentiated dorsal chaetom with large macrosetae. Its body granulation, however, is “fine and uniform, not significantly enlarged on Abd.6” (Fjellberg, 1987, p. 285), and it is characterized by such a feature as the absence of sublobal hairs on the maxillary outer lobe which is unique in the genus.
A key to the Oligaphorura species of the world fauna
1. Head with more than 4 postantennal pso – Head with fewer postantennal pso | O. multiperforata (Gruia 1973) [d. pso: 10.9/6.15.15/15.19.18.22.9] 2 | 40. Abd. V with 3+3 dorsal pso – Abd. V with 4+4 dorsal pso | O. jingyueensis (Sun et Wu 2012) [d. pso: 32/133/33353] 41 |
2. Head with 4 postantennal pso – Head with 3 postantennal pso | 3 4 | 41. Abd. IV with ventral pso present – Abd. I–IV without ventral pso | O. pseudoinya (Weiner et Kaprus’ 2014) [d. pso: 32/133/33354] 42 |
3. Th. III with lateral ms; anal spines present; setae on thoracic sterna absent montana-group – Th. III without ms (unknown for O. marcuzzii); anal spines absent; setae on thoracic sterna usually present marcuzzii-group | 13
17 | 42. Abd. I–III with 212 ventral psx; Th. I–III with 011 dorsal S-setae; PAO 1.6–1.7 times as long as the nearest pso – Abd. I–III with 121 ventral psx; Th. I–III with 022 dorsal S-chaetae; PAO about 3 times as long as than the nearest pso | O. inya (Weiner et Kaprus’ 2014) [d. pso: 32/133/33354] O. wanglangensis Sun et Xie 2019 [d. pso: 32/133/33354] |
4. AIIIO with 4 papillae; Abd. IV with 3+3 dorsal pso – AIIIO usually with 5 papillae; if only 4 papillae present, than Abd. IV with 4+4 dorsal pso | O. palissai (Yosii 1971) [d pso: 32/133/33333] 5 | 43. Th. I without pso – Th. I with 1+1 pso | O. sanjiangensis (Sun et Wu 2012) [d. pso: 32/033/33343] 44 |
5. AIIIO with 4 papillae; Abd. I–III with 444 dorsal pso – AIIIO usually with 5 papillae; if only 4 papillae present, than Abd. I–III with 333 dorsal pso | O. gela (Christiansen et Bellinger 1980) [d pso: 31/133/44444] 6 |
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6. Th. I with 2(3)+2(3) pso; furcal remnant with a cuticular fold – Th. I usually with 0–1+0–1 pso; if 2+2 dorsal pso present on Th. I, than furcal remnant without a cuticular fold | O. quadrituberculata (Börner 1901) [d pso: 32/2(3)3(4)3(4)/33(4)333] 7 | 44. Abd. IV with 1+1 ventral pso present – Abd. IV without ventral pso | O. alnus (Fjellberg 1987) [d. pso: 32/133/33343] O. sibirica (Weiner et Kaprus’ 2014) [d. pso: 32/133/33343] |
7. Tibiotarsi with 11 setae in distal whorls (T and A) – Tibiotarsi with 9 or fewer distal setae | 9 8 | 45. Posterior edge of head with pso in triangular arrangement – Posterior pso on head in usual position 46 | O. kurtshevae (Martynova 1981) [d. pso: 33(1+2)/0(1)44/3334(5)4)] |
8. Tibiotarsi with 9 distal setae differens-group – Tibiotarsi with 5–8 distal setae daii-group | 24 35 | 46. Th. III without lateral ms – Lateral ms on Th. III present | 47 49 |
9. Anal spines absent – Anal spines present | 10 11 | 47. AIIIO with 4 papillae – AIIIO with 5 papillae | O. absoloni (Börner 1901) [d. pso: 32/133/33343] 48 |
10. Th. I–III with 133 pso; Sc with 122 pso – Th. I–III with 122 pso; Sc with 111 pso | O. alavensis (Simón-Benito et Luciáñez 1994) [d. pso: 32/133/44463] O. serratotuberculata (Stach 1933) [d. pso: 32/122/33343] | 48. Abd. IV with ventral pso present – Abd. IV without ventral pso | O. pieninensis (Weiner 1988) [d. pso: 32/133/33343] O. gamae (Buşmachiu et Weiner 2013) [d. pso: 32/133/33343] |
11. Furcal area without cuticular fold, D-type alnus-group – Furcal area with or without cuticular fold – A-, S- or H-type | 39 12 | 49. Abd. I–V with 44454 dorsal pso – Abd. I–V with 33343 dorsal pso | 50 51 |
12. Furcal remnant of A-type absoloni-group – Furcal remnant of S- or H-type groenlandica-group | 45 54 | 50. Sc I–III with 111 pso; labium of ABC-type – Sc I–III with 233 pso; labium of AC-type | O. changbaiensis (Sun et Wu 2012) [d. pso: 32/133/44454] O. uralica (Khanislamova 1986) [d. pso: 32/133/44454] |
13. Head with 2+2 pso on posterior edge – Head with 3+3 pso on posterior edge | 14 15 | 51. Th. I without pso – Th. I with 1+1 pso | O. jiguliensis Shveenkova et Babenko 2021 [d. pso: 32/033/33343] 52 |
14. Abd. I–III with 5(6)55 dorsal pso – Abd. I–III with 444 dorsal pso | O. ussurica Shveenkova et Babenko 2019 [d pso: 42/144/5(6)5554] O. kedroviensis Shveenkova et Sun 2019 [d pso: 42/144/44454] | 52. Labium of ABC-type; Th. I–III with 022 S-setae – Labium of AC-type; Th. I–III with 011 S-setae | O. mazeii Shveenkova et Babenko 2021 [d. pso: 32/133/33343] 53 |
15. Abd. I–III with 555 dorsal pso – Abd. I–III with 544 dorsal pso | O. chankaensis Sun et Wu 2012 [d. pso: 43/144/5556(7)4(5–6)] 16 | 53. Unguiculus about as long as inner edge of unguis; PAO slightly smaller than nearest pso; on Abd. V axial setae a1 and p1 long and subequal to setae m1 – Unguiculus shorter, about 0.7 of inner edge of unguis; PAO 1.5–2.0 times larger than nearest pso; on Abd.5 axial microsetae m1 are clearly shorter than mesosetae a1 and p1 | O. ligni sp. n. [d. pso: 32/133/33343] O. shifangensis Liu et Sun 2019 [d. pso: 32/133/33343] |
16. Abd. IV–V with 53 dorsal pso – Abd. IV–V with 64 dorsal pso | O. montana Weiner 1994 [d. pso: 43/144/54453] O. pseudomontana Sun et Wu 2012 [d. pso: 43/144/54464] | 54. Abd. IV with 3+3 dorsal pso; dorsal side of body with fields of coarser granulation on head, thoracic terga and posterior half of Abd. VI; AIIIO with 5 short papillae and 4 guard setae – Abd. IV with 4 or 5 dorsal pso on each side | O. tottabetsuensis (Yosii 1972) [d. pso: 32/033/33333] 55 |
17. Head with 2+2 pso on posterior edge – Head with 3+3 pso on posterior edge | 18 22 | 55. Th. III without lateral ms – Lateral ms on Th. III present | 56 60 |
18. Abd. I–III with 444 dorsal pso O. marcuzzii (Cassagnau 1968) [d. pso: 42/144/44454] – Abd. I–III with different number of dorsal pso 19 |
| 56. Th. I without pso – Th. I with 1+1 pso | O. nataliae (Fjellberg 1987) [d. pso: 32/033/33343] 57 |
19. Tibiotarsi with 11 distal setae O. ambigua Babenko et Fjellberg 2015 [d. pso: 42/133/33354] – Tibiotarsi with 9 or fewer setae in distal whorls 20 |
| 57. Abd. IV with 5+5 dorsal pso – Abd. IV with 4+4 dorsal pso | O. sabulosa Babenko 2007 [d. pso: 32/133/33353] 58 |
20. Tibiotarsi with 9 distal setae – Tibiotarsi with 6 distal setae | O. ossetica sp. n. [d. pso: 42/244/5.7.7.10.6] 21 | 58. Labium of A-type – Labium of ABC-type | O. ursi (Fjellberg 1984) [d. pso: 32/133/33343] 59 |
21. Th. I–III with 133 pso – Th. I–III with 144 pso | O. humicola Shvejonkova et Potapov 2012 [d. pso: 42/133/33354] O. kremenitsai Shvejonkova et Potapov 2012 [d. pso: 42/144/33354] | 59. Sc I–III with 111 pso – Sc I–III with 333 pso | O. psammophila Shveenkova et Babenko 2021 [d. pso: 32/133/33343] O. tuvinica Potapov et Stebaeva 1997 [d. pso: 32/133/33343] |
22. Tibiotarsi with 9 distal setae – Tibiotarsi with 11 distal setae | O. igori Shveenkova et Babenko 2022 [d. pso: 43/133/44354] 23 | 60. Th. I without pso – Th. I with dorsal pso | 61 68 |
23. Th. I–III with 133 pso – Th. I–III with 144 pso | O. tatianae Shveenkova et Babenko 2022 [d. pso: 43/133/44(5)364] O. montivaga Shveenkova et Babenko 2022 [d. pso: 43/144/44354] | 61. Medial pair of pso on Abd. II–III set behind p-setae; thoracic sterna with ventral setae – Medial pso on Abd. II–III set in usual position in front of p-setae; setae on thoracic sterna present or absent | O. reversa (Fjellberg 1987) [d. pso: 32/033/44454] 62 |
24. Anal spines in a form of spiniform setae – Anal spines of normal shape | O. pseudoraxensis (Nosek et Christian 1983) [d. pso: 32/133/33343] 25 | 62. Ventral abdominal pso present at least on Abd. IV – Abd. I–IV without ventral pso | 63 64 |
25. Abd. VI with a band of coarser granulation – Tergum of Abd. VI with homogenous granulation | O. differens (Bagnall 1949) [d. pso: 32/133/33343] 26 | 63. Abd. I–V with 01110(1) ventral pso; Sc I–III with 222 pso – Abd. I–V with 00010 ventral pso; Sc I–III with 111 pso | O. arnei Paśnik et Kaprus’ 2019 [d. pso: 32/033/34(5)443] O. subnuda sp. n. [d. pso: 32/033/44454] |
26. Abd. I–III with 222 dorsal pso – Abd. I–III with other set of dorsal pso | O. melittae (Christian 1993) [d. pso: 32/022/22243] 27 | 64. Thoracic sterna with 0–0–0 or 0–0–1 setae – Thoracic sterna with 0–1–1 or 0–1–2 setae | 65 66 |
27. Th. III with lateral ms present – Th. III without lateral ms | 28 32 | 65. Labium of A-type; without setae on thoracic sterna – Labium of AC-type; ventral setae sometimes present only on Th. III | O. nuda (Fjellberg 1987) [d. pso: 32/033/44454] O. interrupta (Fjellberg 1987) [d. pso: 32/033/3(4).3(4).3(4).5(4).3(4)] |
28. Ventral abdominal pso present at least on Abd. IV – Abd. I–IV without ventral pso | 29 30 | 66. Dorsal granulation almost uniform or enlarged only on head and Abd. VI, thoracic sterna with 0–1–1 ventral setae – Dorsal side of body with distinct fields of coarser granulation on head, thoracic terga and Abd. VI; thoracic sterna with 0–1–1 or 0–1–2 ventral setae | O. pingicola (Fjellberg 1987) [d. pso: 32/033/33343] 67 |
29. Abd. I–IV with 1111 ventral pso; Sc I–III with 122 pso – Abd. I–IV with 0001 ventral pso; Sc I–III with 111 pso | O. olenae (Weiner et Kaprus’ 2014) [d. pso: 32/133/33343] O. irinae (Thibaud et Taraschuk 1997) [d. pso: 32/133/33343] | 67. All dorsal setae more or less similar in length; thoracic sterna with 0–1–2 ventral setae; three pso of anterior group is located inside the antennal base (within the area of finer granulation) – Dorsal setae clearly differentiated into long macrochaetae and short microsetae; thoracic sterna with 0–1–1 ventral setae; only one pso of anterior group on head is located inside the antennal base | O. koreana (Weiner 1994) [d. pso: 32/033/33343] O. primorica sp. n. [d.pso: d. pso: 32/033/33343] |
30. AIIIO with 4 papillae – AIIIO with 5 papillae | O. sophyae (Weiner et Kaprus’ 2014) [d. pso: 32/133/33343] 31 | 68. Th. I with 2(1)+2(1) pso and only 4(5)+4(5) setae; furcal remnant without cuticular furrow (Н-type) – Th. I with 1+1 pso and more setae; furcal remnant of S-type with clear cuticular furrow (except O. groenlandica having H-type) | 69 70 |
31. Th. I without pso – Th. I with 1+1 pso | O. raxensis (Gisin 1961) [d. pso: 32/033/33343] O. imosolica Shveenkova et Babenko 2021 [d. pso: 32/133/33343] | 69. Abd. I–IV with 1111(2) ventral pso; Sc I–III with 23(2)3 pso – Abd. I–IV without ventral pso; Sc I–III with 111 pso | O. duocellata Babenko et Fjellberg 2015 [d. pso: 32/(1)233/33343] O. yakutica sp. n. [d. pso: 32/233/33343] |
32. Th. I–III with 133 pso – Th. I–III with 022 pso | O. caucasica (Weiner et Kaprus’ 2014) [d. pso: 32/133/33343] 33 | 70. Ventral pso present at least on Abd. IV – Abd. I–IV without ventral pso | 71 72 |
33. Abd. IV with 1+1 ventral pso present – Ventral pso on abdomen absent | O. eremia (Kaprus’ et al. 2002) [d. pso: 32/022/22343] 34 | 71. Abd. I–IV with 1111 ventral pso – Abd. I–IV with 0001 ventral pso | O. judithnajtae Weiner et Paśnik 2017 [d. pso: 32/133/33343] O. cavicola sp. n. [d. pso: 32/133/44454] |
34. Sc I–III with 111 pso – Sc I–III without pso; the same number of dorsal pso | O. hackeri (Christian 1986) [d. pso: 32/022/33343] O. anocellata sp. n. [d. pso: 32/022/33343] | 72. Abd. IV with 4+4 dorsal pso – Abd. IV with 5+5 dorsal pso | 73 75 |
35. Anal spines absent; tibiotarsi with 7 distal setae – Anal spines present | O. stojkoae (Shvejonkova et Potapov 2012) [d. pso: 32/133/33343] 36 | 73. Labium of A-type; furcal remnant with cuticular fold (S-type) – Labium of AC-type; sternum of Abd. IV with or without cuticular fold | O. schoetti (Lie-Pettersen 1896) [d. pso: 32/133/33343] 74 |
36. Lateral ms on Th. III present; tibiotarsi with 5 distal setae – Lateral ms on Th. III absent; tibiotarsi with more distal setae | 37 37 | 74. Furcal remnant with cuticular fold (S-type); axial setae m1 on Abd. V shorter than setae a1 and p1 – Furcal remnant without cuticular fold (H-type); axial setae m1 on Abd. V clearly longer than setae a1 and p1 | O. neglecta sp. n. [d. pso: 32/133/33343] O. groenlandica (Tullberg 1877) [d. pso: 32/133/33343] |
37. Th. II–III with 33 dorsal pso – Th. II–III with 22 dorsal pso | O. chatyrdagi (Kaprus’ et al. 2002) [d. pso: 32/033/33343] O. steposa (Kaprus’ et al. 2002) [d. pso: 32/022/33343] | 75. Abd. V with 3+3 dorsal pso; maxillary outer lobe without sublobal setae – Abd. V with 4 or 5 dorsal pso on each side | O. aborigensis (Fjellberg 1987) [d. pso: 32/133/33353] 76
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38. Th. II–III with 22 dorsal pso; tibiotarsi with 6 distal setae – Th. II–III with 12 dorsal pso; tibiotarsi with 8 distal setae | O. daii (Pomorski et al. 1998) [d. pso: 32/022/33343] O. octosetosa sp. n. [d. pso: 32/012/33343] | 76. One of three postantennal pso located outside of antennal area; setae p1 and p2 on head set in a line – Two postantennal pso located outside of antennal area; posterior edge of head with setae p1 set in front of p2 | O. judithae (Weiner 1994) [d. pso: 3(2+1)2/133/33354] O. linderae (Weiner 1994) [d. pso: 3(1+2)2/133/33354(5)] |
39. Abd. IV with 5+5 dorsal pso – Abd. IV with 4+4 dorsal pso | 40 43 |
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ACKNOWLEDGMENTS
We express our sincere thanks to Olga Makarova (Institute of Ecology & Evolution RAS), Natalia Kuznetsova, Mikhail Potapov (MSPU), Anna Geraskina (Center for Forest Ecology and Productivity RAS), Anna Saraeva (Forest Research Institute, Karelian Research Center, Petrozavodsk), Alexander Kuprin (Federal Scientific Center of the East Asia Terrestrial Biodiversity, FEB RAS), Alexander Komisarenko (Vladivostok), Albina Burnasheva (IBPC SB RAS, Yakutsk), Mikhail Berezin (Moscow Zoo), and Igor’ Lavrov (a speleologist from Perm) for their extensive collections, as well as to Anna Bokova (MSPU) for the initial sorting of samples and other cooperation. We are also much indebted to Sergei Golovatch (Institute of Ecology & Evolution RAS) for his editing an early English draft.
AUTHOR CONTRIBUTIONS
The study is the result of the joint work of all of its co-authors. At the same time, the figures for O. anocellata sp. n., O. octosetosa sp. n., O. ligni sp. n. and O. yakutica sp. n. were prepared by Yu. Shveenkova, for O. subnuda sp. n., O. cavicola sp. n., O. neglecta sp. n. and primorica sp. n. by A. Babenko, and for O. ossetica sp. n. by M. Antipova.
FUNDING
Two last authors were partially financed through the Russian Science Foundation (RSF) grant 22-24-00162.
COMPLIANCE WITH ETHICAL STANDARDS
The article does not contain any studies involving animals in experiments performed by the authors.
CONFLICT OF INTEREST
The authors of this work declare that they have no conflicts of interest.
1 one of the two known specimens of O. anocellata sp. n. (para-type) has a pso on one side of Th. I.
Авторлар туралы
Yu. Shveenkova
”Privolzhskaya lesostep” State Nature Reserve
Хат алмасуға жауапты Автор.
Email: jushv@mail.ru
Ресей, Penza, 440031
M. Antipova
The Severtsov Institute of Ecology and Evolution, Russian Academy of Sciences
Email: antimar.2410@gmail.com
Ресей, Moscow, 119071
A. Babenko
The Severtsov Institute of Ecology and Evolution, Russian Academy of Sciences
Email: lsdc@mail.ru
Ресей, Moscow, 119071
Әдебиет тізімі
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