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Research Article
A new subterranean species of Oncopodura Carl & Lebedinsky, 1905 (Collembola, Entomobryomorpha, Oncopoduridae) from a cave in Northeastern Iran
expand article infoMahmood Mehrafrooz Mayvan‡, Andrea Parimuchová‡, Ľubomír Kováč‡
‡ Pavol Jozef Šafárik University, Košice, Slovakia

Corresponding author: Mahmood Mehrafrooz Mayvan ( mahmood_mehrafrooz@yahoo.com )

Corresponding author: Ľubomír Kováč ( lubomir.kovac@upjs.sk )

Academic editor: Louis Deharveng
© 2024 Mahmood Mehrafrooz Mayvan, Andrea Parimuchová, Ľubomír Kováč.
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: Mehrafrooz Mayvan M, Parimuchová A, Kováč Ľ (2024) A new subterranean species of Oncopodura Carl & Lebedinsky, 1905 (Collembola, Entomobryomorpha, Oncopoduridae) from a cave in Northeastern Iran. Subterranean Biology 49: 31-51. https://doi.org/10.3897/subtbiol.49.118293
ZooBank: urn:lsid:zoobank.org:pub:6530C30E-F67D-4679-9F6C-75C100F7AD10
Open Access

Abstract

A new species of Oncopodura (Collembola, Entomobryomorpha, Oncopoduridae) from the Moghan cave in northeastern Iran, built in a carbonate complex of Kopet Dag mountain range, is described. Oncopodura moghanensis sp. nov. can be distinguished from other congeners by (1) 6 long subequal lobes in PAO, each subdivided into 3–6 fingers, (2) dens with 7 dorsal feather-like macrosetae; at basal part with 1 dorsoexternal and 2 dorsointernal hooks, at the distal part with 1 dorsoexternal and 3 dorsointernal hooks, (3) distal part of manubrium with long feather like macrosetae reaching middle part of dens, and (4) mucro with 4 teeth, apical tooth very sharp, and 2 scales at its basal half. A table with diagnostic characters of species related to the Oncopodura moghanensis sp. nov. and an updated key to the world species of Oncopodura are provided.

Keywords

Cave biology, cave fauna, Kopet Dag, Moghan Cave, springtail, subterranean ecosystem

Introduction

Oncopodura Carl & Lebedinsky, 1905 is an eyeless genus of the family Oncopoduridae (Entomobryomorpha) involving both edaphic and subterranean species. Elongated mucro, the presence of scales on the body, thickened sensilla on Ant IV and the presence of hooks and spines on dens are considered characteristic features of this genus (Yu et al. 2014), which covers 49 species worldwide (Bellinger et al. 2023). Of them, only two species have been recorded from Iran: Oncopodura ambigua Christiansen, 1957 and Oncopodura hamata Carl & Lebedinsky, 1905 (Mehrafrooz Mayvan et al. 2023).

In terms of biodiversity, Iran is located in the Palearctic biogeographic region and about 54 % of its land area is covered with mountains, including mountain ranges of Zagros, Alborz and Kopet Dag (Azizi Jalilian et al. 2020; Mehrafrooz Mayvan et al. 2015). Karst caves, as the most common type of caves in the world, are the most widespread in the mountainous areas of the country (Juberthie et al. 2001; Ghaderi and Karimi 2014; Malek Hosseini et al. 2015; Malek Hosseini et al. 2016; Moldovan et al. 2018). The present paper is based on the recent speleobiological fieldworks carried out in the Moghan Cave situated in the northeast of Iran in the Kopet Dag mountain range that is formed in massive sedimentary limestones (Manafzadeh et al. 2016; Raziei 2022). Few papers have focused on the subterranean arthropods in Iran, but no studies involving cave Collembola have yet been conducted. This study is aimed at the genus Oncopodura Carl & Lebedinsky, 1905, which is highly represented in subterranean habitats with 31 troglobionts altogether (Deharveng and Bedos 2018; Lukić 2019). In this paper, description of a new species of Oncopodura, discovered in the Moghan Cave in the Kopet Dag Mountains, northeastern Iran, is provided together with a table summarizing diagnostic characters of world species of Oncopodura with PAO with more than 4 lobes and an updated key to the world species of the genus.

Material and methods

Cave descriptions

Moghan Cave takes its name from a village Moghan, it is situated 35 km to southwest of Mashhad, Khorasan-e-Razavi province, on the eastern part of Kopet Dag mountains (36°06'59"N, 59°22'06"E longitude) at altitude of 2193 m above sea level (Fig. 1A, B) The cave has two adjacent entrances located on a relatively steep slope of the mountain with the main entrance 11 m long and 4 m high (Fig. 2A–C). The vegetation in the cave surroundings is very sparse and the entire twilight cave zone (50 m in distance) is completely dry. The cave is about 500 m long, with a maximum depth of 44 m (Fig. 3). It is divided into two floors connected by wells, reaching the height up to 25 m. Moreover, Moghan Cave has five halls, with a water pond located at the end of the fifth hall on the second floor, 5.66 × 6.93 m in area and depth of 30–60 cm (Fig. 2D). The specimens of the new species were collected from the water pool in the deepest part of the cave, which had constant air temperature (12 °C) and relative humidity (RH) of over 73 %, measured with a thermo-hygrometer TFA 30.5015 DTH in December 2023.

Figure 1. 

A Razavi Khorasan province in Iran is marked with an orange color; location of the Moghan Cave, where the new Oncopodura species was collected, is marked with an asterisk B cave entrance and surroundings of Moghan Cave in autumn (Photo: M. Mehrafrooz).

Figure 2. 

Moghan Cave, northeast Iran A main entrance of the cave in spring (Photo: M. Mehrafrooz) B second entrance (Photo: M. Mehrafrooz) C two cave entrances, view from inside the cave (Photo: M. Mehrafrooz) D pool at the end of the cave located on the second floor (Photo: Vahid Ashrafi).

Figure 3. 

Ground plan of the Moghan Cave (Sketched by V. Ashrafi).

Methods

For observation in an optical microscope with phase contrast, initial specimens were mounted in Heinz’s medium after clearing in Nesbitt’s fluid. To compare effectivity of different clearing methods, additional specimens were boiled softly in evaporation glass dish with 95% ethyl alcohol on electric cooking plate for approximately 1 min to remove fat from the body. For clearing, the specimens were transported to concave glass dish with 10% water solution of KOH for 1 min and then transported to a dish containing chlorophenol until the specimens were transparent. At the end, specimens were mounted on Swann medium. After drying, the cover glasses were ringed with nail polish or Canada balsam to seal the mounting medium. The specimens were observed in Leica DM 2500 light microscope equipped with phase and DIC contrasts, and a drawing arm.

Abbreviations used

Abd – abdominal segment; Ant – antennal segment; PAO – post antennal organ; PS – pseudopore; S – sensillum; Ti – tibiotarsus; VT – ventral tube.

Repositories of investigated material: CoPJSU – Collembola collection of the Department of Zoology, Institute of Biology and Ecology, Faculty of Science, Pavol Jozef Šafárik University, Košice, Slovakia; ZMFUM – Zoology Museum of Ferdowsi University of Mashhad, Mashhad, Iran.

Results

Species description

Oncopodura moghanensis Mehrafrooz Mayvan & Kováč, sp. nov.

https://zoobank.org/8AC9E1C6-F1F1-4F59-8FB6-CB901436009F
Figs 1, 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, Table 1

Diagnosis

PAO with 6 long, subequal lobes, each lobe secondarily divided into 3–6 fingers. Unguis normal, not elongated, untoothed; unguiculus acuminate. Dorsal side of manubrium with 11+11 setae. Dens at basal part with 1 dorsoexternal and 2 dorsointernal hooks, at the distal part with 1 dorsoexternal and 3 dorsointernal hooks. Mucro with 4 teeth.

Type locality

Iran, Razavi Khorasan province, Mashhad city, Moghan village, Moghan Cave, Coordinates: 36°06'59"N, 59°22'06"E, 2193 m a.s.l.

Type material

Holotype : female on slide [MoCa101], dark zone, the end part of the cave, 2nd cave floor, collected from water pond surface, 28.ix.2022, temperature = 11.9 °C, RH 73.0%, leg. M. Mehrafrooz. Paratypes: 4 specimens on slides [MoCa102, MoCa103, MoCa104, MoCa105], ibidem, hand collecting on a stalagmite, 30.iv.2022, temperature = 12.0 °C, RH 73.4%, leg. M. Mehrafrooz. Holotype and one paratype deposited in UPJŠ; three paratypes deposited in ZMFUM.

Table 1.
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Differential characters of world species of Oncopodura with PAO with more than 4 lobes. Characters: 1, Ant IV medial S number; 2, Ant IV basal S number; 3, Ant IV S form, 1 = long, 2 = pointed, 3 = rounded, 4 = short, 5 = trapezoidal; 4, Ant. III S number; 5, Ant. II S number; 6, PAO lobes number; 7, PAO vesicle fingers number; 8, Basal dens dorsoexternal hooks number; 9, Basal dens dorsointernal hooks number; 10, Basal dens external ciliated macrosetae number; 11, Basal dens internal ciliated macrosetae number; 12, Distal dens dorsoexternal hooks number; 13, Distal dens dorsointernal hooks number; 14, Distal dens external ciliated macrosetae number; 15, Distal dens internal ciliated macrosetae number; 16, Mucro teeth number; 17, Mucronal basal scales: 0 = absent, 1 = present; 18, Ungual basal lamella, 0 = absent, 1 = or < ½ unguis, 2 > ½ unguis; 19, Unguis, 1 = normal, 2 = long. Dif. - number of differential characters in new species versus total number of compared characters. * = differences from new species O. moghanensis sp. nov. “–“= absence of data; “?” = Insufficient or confusing information in original description; S – sensillum.

Species/Characters 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 Dif.
Oncopodura ambigua Christiansen, 1957 4 1* 3* 0* 0* 6 1* 0* 1* 0* 1* 1 1* 0* 0* 4 1 2?* 1 13/19
Oncopodura atoyacensis Bonet, 1943 4 0 4* 0* 0* 6 1* 0* 1* 0* 2* 1 1* 0* 0* 4 0* 0 1 12/19
Oncopodura crassicornis Shoebotham, 1911 4 0 3* 0* 0* 6 1* 1 2 0* 1* 1 3 0* 0* 4 1 0 1 8/19
Oncopodura czmur Szeptycki, 1977 4 0 1* 2* 0* 6 1* 0* 3* 2 3 1 2* 1 1 4 1 0 1 7/19
Oncopodura dethieri Janssens & De Bruyn, 2010 4 0 1* 1 0* 6 2-3* 0* 3* – – 1 4* – – 4 0* 0 1 7/15
Oncopodura egerszogensis Loksa, 1961 0* 0 – 1 1 6 3* 1 1* 2 2* 2* 1* 1 1 4 0* 0 2* 8/18
Oncopodura hubbardi Christiansen & Bellinger, 1996 4 1* 1* 2* 2* 5-6* 1* 0* 0* 2 1* 1 1* 1 1 4 1 1* 2* 11/19
Oncopodura itatiaiensis Arlé, 1961 4 0 4* 2* 0* 5* 1* 1 2 1* 1* 1 3 2* 2* 4 1 2?* 1 10/19
Oncopodura kuramaensis Yosii, 1956 4 0 4* 1* 1 6 1* 0* 2 1* 3 1 3 0* 1 4 1 – – 7/18
Oncopodura moghanensis sp. nov. 4 0 2 1 1 6 3-6 1 2 2 3 1 3 1 1 4 1 0 1
Oncopodura pegyi Gruia, 1994 4 0 4* 3* 1 6-8* 1* 0* 2 1* 4* 1 3 1 1 4 1 0 2* 8/19
Oncopodura pelissiei Deharveng, 1988 4 0 3* 3* 2* 6 1* 1 2 2 3 1 3 0* 1 4 1 0 2* 6/19
Oncopodura reyersdorfensis Stach, 1936 3* 0 2 0* 0* 6 1-2* 0* 1* 1* 2* 1 1 0* 0* – 0* 0 2* 12/18
Oncopodura tunica Christiansen & Bellinger, 1980 10* 0 2 0* 2* 5* 1* 0* 1* 1* 2* 1 1* 1 1 4 1 1?* 2* 12/19
Oncopodura yosiiana Szeptycki, 1977 4 0 2 2* 1 6 1* 0* 2 1* 3 1 3 1 1 4 1 0 1 4/19

Description

Body appearance. Length 1.05 mm on average (n = 5, max. length 1.2 mm). Colour white without traces of pigment, body covered by hyaline scales.

Antenna. Antenna nearly as long as head, without apical bulb and scales (Fig. 4A–E). length of Ant I: II: III: IV as 0.02: 0.04: 0.06: 0.08 mm respectively. Ant I with 7 thick dorsal setae and 11 normal setae (smaller than those of Ant II–III). Ant II with 1 broad wrinkled apical sensillum similar to those of Ant III, 1 apical blunt sensillum, 11 dorsal and 6 ventral setae and 4 dorsolateral microsetae. Ant III with sensory organ formed by 2 expanded wrinkled sensilla with 1 blunt, basal sensillum, 8 thick setae at their base, and 10 slender, long setae covered by very soft cilia (Fig. 4B). Ant IV not elongated, with a row of 4 thick and pointed sensilla (Fig. 4C–E), equally distant from each other, and 2 subapical modified setae (Fig. 4D). Ventral side of Ant IV with normal setae, dorsal part with normal and elongated setae.

Figure 4. 

Oncopodura moghanensis sp. nov. A antennal segments I–III B detail of finely ciliated seta C antennal segment IV D detail of two subapical modified setae near the apical sensillum E detail of thick, pointed sensillum. Scale bars: 0.02 mm.

Head. Eyes absent. PAO well developed and located near base of Ant I, very distinct and large, about 24 µm in diameter, with 6 subequal lobes formed around a central circle, each lobe secondarily divided into 3–6 irregular finger-like lobes (Fig. 5A). Labrum with 4 prelabral setae and 4 rows of 2, 3, 5, 4 setae on papillae (Fig. 6A). Left mandible with 4 teeth and right mandible with 3 teeth.

Figure 5. 

Oncopodura moghanensis sp. nov. A postantennal organ B labium and labial triangle C ventral head chaetotaxy. Scale bars: 0.01 mm (A, B); 0.02 mm (C).

Figure 6. 

Oncopodura moghanensis sp. nov. A labrum B dorsal head chaetotaxy (dotted circle = inter-antennal macrosetae). Scale bars: 0.01 mm (A); 0.02 mm (B).

Ventral head without scales, chaetotaxy (setal pattern) as in Fig. 5C, all ventral setae smooth. Head dorsally with 4+4 inter-antennal and 5 lateral macrosetae on each side placed around PAO; 1 short and thick seta posterior to PAO. Two rows of 5 and 6 macrosetae, respectively, in front of prelabral setae. Head with scales, dorsal microsetae as in Fig. 6B. Labial triangle with 7 smooth macrosetae (Fig. 5B).

Body chaetotaxy . As in Fig. 7A, B. The chaetotaxy follows the general pattern described for the genus (Szeptycki 1977; Jordana et al. 2012). Mesothorax extended forward slightly over the head, distally with a row of smooth spine-like setae, laterally with 1 ciliated spine-like seta and 1 trichobothrium and 1+1 medial pseudopores. Metathorax with 2+2 trichobothria, 2+2 lateral spine-like setae and 1+1 medial pseudopores. Abdomen I with 1+1 ciliated spine-like setae and 1+1 medial pseudopores, Abd. II with 1+1 lateral smooth spine-like microsetae, medially with 1+1 ciliated spine-like setae, 1+1 trichobothria and 1+1 medial pseudopores, Abd. III laterally with 1+1 smooth spine-like microsetae, medially with 1+1 ciliated spine-like setae and 1+1 pseudopores. Abd IV with 4+4 ciliated macrosetae and 1+1 pseudopores. Abd V with 2+2 anterior sensilla, 5+5 ciliated macrosetae (shorter than Abd IV macrosetae) and 1+1 smooth spine-like microsetae. Abd VI: epiproct with 9+9 ciliated and 1+1 smooth setae, paraproct with 12 ciliated macrosetae and 8 smooth setae.

Figure 7. 

Oncopodura moghanensis sp. nov. A dorsal chaetotaxy of thorax and abdomen B abd V & VI (S = anterior sensilla; circles with a slash represent pseudopores). Scale bars: 0.096 mm (A), 0.01 mm (B)

Legs. Legs without scales. Chaetotaxy as in Fig. 8A–C. Leg I: trochanter with 6 setae, femur with 3 whorls of 4, 4, 4 setae, respectively; whorl 2 with 1 microseta and whorl 4 with 2 microsetae; Ti with 5 whorls of 4, 4, 5, 4, 7 setae, respectively. Leg II: trochanter with 5 setae, femur with 3 whorls of 4, 3, 5 setae, respectively; whorl 3 with 2 microsetae; Ti with 5 whorls of 4, 4, 4, 4 and 6 setae, whorl 2 with 1 stout, clavate (apically expanded) macroseta. Leg III: trochanter with 4 setae, femur with 3 whorls of 3, 3, 5 setae; Ti with 5 whorls of 4, 4, 4, 5, 6 setae respectively, whorl 5 with 1 microseta. Most Ti setae large, acuminate, and extremely finely ciliated; tenent hair slender and acuminate. Unguis (Fig. 9A–C) normal, not elongated, untoothed; inner pretarsal setae long, slightly shorter than the unguiculus, about 1/2 length of internal side of unguis; outer pretarsal setae shorter; unguiculus acuminate.

Figure 8. 

Oncopodura moghanensis sp. nov., chaetotaxy of legs A leg I B leg II C leg III. Scale bars: 0.02 mm.

Figure 9. 

Oncopodura moghanensis sp. nov., detail of foot complex A leg I B leg II C leg III. Scale bar: 0.01 mm.

Ventral tube (collophore) without prominent papillae, with 3+3 setae (2+2 dorsolateral and 1+1 frontal). Tenaculum with 4+4 teeth, setae absent.

Furca. length of manubrium: dens: mucro as 0.046: 0.040: 0.031 mm respectively. Ventral side of manubrium with scales, dorsal side with 3+3 long and 1+1 short smooth setae, 6+6 ciliated axial macrosetae; 1+1 long, feather-like distal macrosetae reaching half of the dens (Fig. 10A). Ventral side of dens with scales; chaetotaxy of dorsal side of dens subdivided into basal and distal part. Basal part with 2 dorsointernal ciliated hooks, 1 dorsointernal smooth conical spine (broad at the base and strongly tapered at the top), 1 dorsoexternal ciliated hook and 5 dorsal ciliated macrosetae (basal tubercle with 2 ciliated macrosetae, of which inner 1 feather-like and thinner than outer one). Distal part of dens with 3 dorsointernal ciliated hooks (1 medial, 2 distal), 1 dorsoexternal ciliated hook, 2 dorsal ciliated macrosetae and 2 dorsal short, smooth medial spine-like setae. Dens apically with 5 smooth ventral setae (Fig. 11A, B). Mucro with 4 teeth: 2 apical, 1 medial and 1 basal; 2 scales at its basal half (Fig. 10B).

Figure 10. 

Oncopodura moghanensis sp. nov. A manubrium dorsal side B mucro with basal scale. Scale bars: 0.01 mm.

Figure 11. 

Oncopodura moghanensis sp. nov., dorsal side of dens A dens with dorsolateral hooks and spines B dens with ciliated dorsal macrosetae. Scale bars: 0.01 mm.

Etymology

The name of the new species is derived from the Moghan Cave in Kopet Dag Mountains in northeastern Iran, i.e. the locality where the species was discovered.

Taxonomic remarks

So far, 50 species of Oncopodura have been described. Based on shape of PAO, Oncopodura moghanensis sp. nov. is similar to O. dethieri Janssens & De Bruyn, 2010, O. egerszogensis Loksa, 1961 and O. reyersdorfensis Stach, 1936. The new species is characteristic with 3–6 finger-like lobes in PAO vesicles, while O. dethieri has 2–3 fingers, O. egerszogensis has 3 fingers and O. reyersdorfensis has 1–2 fingers in PAO. Based on dorsal chaetotaxy of dens, O. moghanensis sp. nov. is similar to O. crassicornis Shoebotham, 1911, O. pelissiei Deharveng, 1988 and O. itatiaiensis Arlé, 1961. However, Szeptycki (1977) mentioned more complex chaetotaxy of manubrium for O. crassicornis, when comparing it with O. yosiiana Szeptycki, 1977. The new species has 6 long and subequal lobes in PAO with each lobe secondarily divided into 3–6 fingers, while O. crassicornis and O. pelissiei has 6 simple lobes and O. itatiaiensis 5 simple lobes in PAO. Among other species, O. yosiiana differs from the new species by 6 simple lobes in PAO, manubrium with numerous dorsal setae (16+16 feather-like setae, 6+6 smooth mesosetae, 1+1 smooth microsetae and 1+1 ciliated macrosetae), and dens without basal external hook.

Deharveng (1988) recognized two basic groups of Oncopodura congeners:

  1. O. crassicornis –group: Ant IV with 4 short and thick sensilla (S-setae) arranged in a curved line, thickened sensilla at the base of segment absent; PAO large with 6 elongated lobes; unguis without large basal lamella (external tooth); pretarsal setae long; dental hooks finely scaly-ciliated, lacking strong denticles.
  2. O. tricuspidata –group: Ant IV with 4 thick sensilla (S-setae) arranged in a line, and one thickened sensillum at the base of segment (sometimes absent); PAO small, with 3–4 rounded lobes, or they are absent; unguis with large basal lamella, dental hooks smooth, but with strong denticles on their outer surface.

Recently, numerous species have been described with various combinations of the group-specific characters. In many species of O. crassicornis–group, Ant IV sensilla are in a line and in some species Ant IV basal sensilla is present, such as in O. ambigua Christiansen, 1957 and O. hubbardi Christiansen & Bellinger, 1996. Also, PAO in O. hubbardi, O. itatiaiensis and O. tunica Christiansen & Bellinger, 1980 have 5 lobes and in O. pegyi Gruia, 1994 has 6–8 lobes. Unguis basal lamella is present in species including O. ambigua, O. hubbardi, O. itatiaiensis and O. tunica.

In some species of O. tricuspidata–group, ant IV basal sensilla is absent such as O. cruciata Bonet, 1943, O. equatoriana Thibaud & Najt, 1987, O. hoffi Christiansen & Bellinger, 1980, O. iowae Christiansen, 1961, O. mala Christiansen & Bellinger, 1996, O. meridionalis Cassagnau, 1959, O. puncteola Yosii, 1956 and O. subhoffi Christiansen & Bellinger, 1998. Also, there are some species with 1 single lobe PAO including O. cavernarum Stach, 1934, O. gledensis Baquero et al., 2007, O. mala, O. siquierae Seminario-Cordova et al. 2018 and O. vioreli Gruia, 1989. On the other, ungual basal lamella is absent in some species including O. cruciata, O. equatoriana, O. hoffi, O. iowae, O. mala, O. meridionalis, O. puncteola and O. subhoffi. Thus, practical use of sorting of Oncopodura congeners to tricuspidata- and crassicornis-group has become questionable.

The new species was collected in the deep cave zone but, nevertheless, it does not show morphological adaptations to subterranean environment such as elongation of unguis or increase in the number of sensilla on antennae and body or augmentation of setae over the head, characteristic for some troglomorphic congeners (e.g. Deharveng 1988).

Key to world species of genus Oncopodura

The following key was prepared based on the diagnostic characters of worldwide Oncopodura species (adapted from Absolon and Kseneman 1932; Bonet 1943; Szeptycki 1977; Mari Mutt 1984; Christiansen and Bellinger 1998). This should be considered a tentative key, since the limited data on several important characters in some species included.

1 PAO absent 2
– PAO present 18
2 Ant IV with 4 medial sensilla 3
– Ant IV with more than 4 medial sensilla 16
3 Mucro with two teeth O. bidentata Delamare Deboutteville, 1948 (Ivory coast, soil)*
– Mucro with more than two teeth 4
4 Mucro with 4 teeth 5
– Mucro with more than 4 teeth 11
5 Dens without basal dorsointernal hooks 6
– Dens with at laest 1 basal dorsointernal hooks 7
6 Dens with 1 distal dorsoexternal hook and 1 distal dorsointernal hook. Unguis basal lamella shorter than or equal to half the length of the unguis O. hamata Carl & Lebedinsky, 1905 (Crimea, cave)
– Dens without distal dorsoexternal hook and with 2 distal dorsointernal hooks. Unguis basal lamella longer than half the length of the unguis O. occidentalis Bonet, 1931 (Spain & France, cave)
7 Dens without dorsoexternal hooks O. tiegsi Womersley, 1942 (Australia, decaying log)*
– Dens with dorsoexternal hooks 8
8 Unguis basal lamella shorter than or equal to half the length of the unguis 9
– Unguis basal lamella longer than half the length of the unguis 10
9 Ant II with 7 blunt sensilla. Unguis normal O prietoi Bonet, 1943 (Mexico, cave)
– Ant II with 13 blunt sensilla. Unguis elongated O. jauzioni Deharveng, 1988 (France, cave)
10 Apex of Ant II and III with many blunt sensilla (more than 15). Dens with 2 distal dorsointernal hooks. Basal scale on mucro present O. dura Christiansen & Reddell, 1986 (Mexico, cave)
– Apex of Ant II and III with few blunt sensilla (less than 5). Dens with 1 distal dorsointernal hook. Basal scale on mucro absent O. delhezi Stomp, 1974 (Algeria, cave)
11 Mucro with 5 teeth 12
– Mucro serrated, with numerous teeth (more than 5) 14
12 Ant II with 16–32 blunt sensilla. Ant III with 6–7 blunt sensilla. Basal scale on mucro present O. susanae Christiansen & Reddell, 1986 (Mexico, cave)
– Ant II with 1–2 blunt sensilla. Ant III with 2–8 blunt sensilla. Basal scale on mucro absent 13
13 Unguis basal lamella shorter than or equal to half the length of the unguis. Ant II with 8 blunt sensilla. Dens with 1 basal dorsointernal hook and without distal dorsoexternal hooks O. fadriquei Jordana et al., 2012 (Spain, cave)
– Unguis basal lamella longer than half the length of the unguis. Ant II with 2 blunt sensilla. Dens without basal dorsointernal hooks, with 1 distal dorsoexternal hook O. alpa Christiansen & Bellinger, 1980 (USA, moss and lichens)
14 Dens with 1 basal dorsoexternal hook and 2 distal dorsointernal hooks O. gul Yosii, 1966b (South Korea, cave)
– Dens without basal dorsoexternal hooks, with 3 distal dorsointernal hooks 15
15 Dens with 1 basal and 3 distal external coarsely serrated hooks, almost digitated. Basal scale on mucro absent. Unguis basal lamella present, longer than half the length of the unguis O. kuramotoi Yosii, 1965 (Japan, cave)
– Dens with 1 basal and 3 distal serrated hooks. Basal scale on mucro present. Unguis basal lamella absent O. puncteola Yosii, 1956 (Japan, cave)
16 Apex of Ant III with many blunt setae. Mucro basal scale present. Unguis basal lamella longer than half the length of the unguis 17
– Apex of Ant III with 2 blunt setae. Mucro basal scale absent. Unguis basal lamella shorter than or equal to half the length of the unguis O. lebretoni Deharveng, 1988 (France, cave)
17 Ant IV with 5 medial sensilla and 1 basal blunt sensillum. Dens with 1 basal internal hook. Mucro with strong media lamella O. fenestra Christiansen & Bellinger, 1996 (USA, cave)
– Ant IV with 6 medial sensilla and 2 basal blunt sensilla. Dens without basal dorsointernal hooks. Mucro without median lamella O. frankeae Christiansen & Bellinger, 1998 (USA, cave)
18 PAO with 1 simple lobe 19
– PAO with 3 or more lobes 23
19 Unguis basal lamella absent or if present, its longer than half the length of the unguis 20
– Unguis basal lamella shorter than or equal to half the length of the unguis 21
20 Unguis basal lamella absent. Ant III with 2 blunt sensilla, Ant II with 2 blunt sensilla. Distal part of dens with 2 ciliated macrosetae O. mala Christiansen & Bellinger, 1996 (USA, cave & epigeic)
– Unguis basal lamella longer than half the length of the unguis. Ant III with 1 blunt sensillum, Ant II without blunt sensilla. Distal part of dens without ciliated macrosetae O. siquierae Seminario-Cordova et al., 2018 (Spain, cave)
21 Mucro with 4 teeth, basal scale present 22
– Mucro with 5 teeth, basal scale absent O. gledensis Baquero et al., 2007 (Spain, cave)
22 Unguis basal lamella longer than half the length of the unguis. Ant IV without basal blunt sensilla, Ant II with 2 blunt sensilla O. vioreli Gruia, 1989 (Romania, cave)
– Unguis basal lamella shorter than or equal to half the length of the unguis. Ant IV with 1 basal blunt sensillum, Ant II with 3 blunt sensilla O. cavernarum Stach, 1934 (Italy and Slovenia, cave)
23 PAO with 3 lobes 24
– PAO with 4 or more lobes 26
24 Mucro with 4 teeth. Ant IV with 4 medial sensilla and 1 basal blunt sensillum, Ant III with 2–3 blunt sensilla 25
– Mucro with 5 teeth. Ant IV with 5 medial sensilla and 4 blunt sensilla. Ant III with 3 blunt sensilla O. equatoriana Thibaud & Najt, 1987 (Ecuador, cave)
25 Unguis basal lamella longer than half the length of the unguis. Dens with 1 basal and 2 distal dorsointernal hooks. Ant III with 2 blunt sensilla O. tricuspidata Cassagnau, 1964 (France, cave)
– Unguis basal lamella absent. Dens with 2 basal and 3 distal dorsointernal hooks. Ant III with 3 blunt sensilla O. hoffi Christiansen & Bellinger, 1980 (USA, cave)
26 PAO with 4 lobes 27
– PAO with 5 or more lobes 36
27 Mucro with 4 teeth 28
– Mucro with more than 4 teeth 35
28 Dens with 1 basal dorsoexternal hook. Ant IV with 4 sensilla along curved line O. hyleana Arlé, 1961 (Brazil, litter in forest)
– Dens without basal dorsoexternal hooks. Ant IV with 4–7 sensilla in a line 29
29 Ant IV with 7 sensilla 30
– Ant IV with 4 sensilla 31
30 Ant IV with 1 basal sensillum. Ant III with 4 and Ant II with 2 blunt sensilla, respectively. Dens with 1 distal dorsointernal hook. Basal scale on mucro present. Unguis basal lamella present, shorter than or equal to half the length of the unguis O. nana Thibaud & Najt, 1987 (Ecuador, cave)
– Ant IV without basal sensilla. Ant III with 1 blunt sensillum, Ant II without blunt sensilla. Dens with 3 distal dorsointernal hooks. Basal scale on mucro absent. Unguis basal lamella absent O. meridionalis Cassagnau, 1959 (Spain, beech humus)
31 Unguis basal lamella longer than half the length of the unguis 32
– Unguis basal lamella absent or if present, its shorter than or equal to half the length of the unguis 33
32 Ant IV with 1 basal sensillum. Ant III with 1 and Ant II with 7 blunt sensilla, respectively. Mucro longer than dens. Dens with 2 distal dorsointernal hooks O. jugoslavica Absolon & Kseneman, 1932 (Bosnia & Herzegovina, Croatia and Chile, cave)
– Ant IV without basal sensilla. Ant III without blunt sensilla, Ant II with 2 blunt sensilla. Mucro equal or shorter than dens. Dens with 1 distal dorsointernal hook O. arecibena Mari Mutt, 1984 (Puerto Rico, cave)
33 Dens with 1 distal dorsointernal hook. Ant IV without basal sensilla O. iowae Christiansen, 1961 (USA, cave)
– Dens with 2 distal dorsointernal hooks. Ant IV with 1 basal sensillum 34
34 Ant III with 1 and Ant II with 5 blunt sensilla, respectively. Dens with 2 basal and 1 distal dorsointernal hooks, all are normally serrated. Unguis basal lamella absent O. cruciata Bonet, 1943 (USA, cave)
– Ant II and III without blunt sensilla. Dens with 2 basal and 1 distal dorsointernal hooks, all strongly dentated. Unguis basal lamella shorter than or equal to half the length of the unguis O. indica Yosii, 1966a (India, cave)
35 Mucro with 5 teeth. Ant IV without basal sensilla; Ant III without and Ant. II with 2 blunt sensilla, respectively. Dens with 1 basal and 2 distal dorsointernal hooks. Unguis basal lamella absent O. japonica Yosii, 1956 (Japan, cave)
– Mucro with numerous teeth, serrated. Ant IV with 1 basal sensillum; Ant III with 2 and Ant II with 4–5 blunt sensilla, respectively. Dens with 2 basal and 3 distal dorsointernal hooks. Unguis basal lamella shorter than or equal to half the length of the unguis O. subhoffi Christiansen & Bellinger, 1998 (USA, cave)
36 PAO with 5 lobes 37
– PAO with ≥ 6 lobes 39
37 Unguis basal lamella longer than half the length of the unguis. Dens with 1 distal dorsointernal hook and without basal dorsoexternal hooks. Ant II with 2 blunt sensilla 38
– Unguis basal lamella shorter than or equal to half the length of the unguis. Dens with 3 distal internal hooks and 1 basal dorsoexternal hook. Ant II without blunt sensilla O. itatiaiensis Arlé, 1961 (Brazil, litter)
38 Unguis with well-developed tunica. Ant IV with 10 median sensilla and without basal sensilla. Ant III without blunt sensilla. Dens with 1 basal dorsointernal hook O. tunica Christiansen & Bellinger, 1980 (USA, cave)
– Unguis without tunica. Ant IV with 4 median sensilla and 1 basal sensillum. Ant. III with 2 blunt sensilla. Dens without basal dorsointernal hooks O. hubbardi Christiansen & Bellinger, 1996 (USA, cave)
39 PAO with 6 simple lobes 40
– PAO with 6 lobes subdivided into 2 or more fingers 47
40 Mucro with 4 teeth 41
– Mucro with 5 teeth O. czmur Szeptycki, 1977 (North Korea, forest litter)
41 Unguis basal lamella present 42
– Unguis basal lamella absent 44
42 Unguis basal lamella longer than half the length of the unguis. PAO with 6 lobes 43
– Unguis basal lamella shorter than or equal to half the length of the unguis. PAO with 6–8 lobes O. pegyi Gruia, 1994 (Romania, cave)
43 Ant IV without basal sensilla. Both Ant III and Ant II with 1 blunt sensillum. Dens with 2 basal and 3 distal dorsointernal hooks O. kuramaensis Yosii, 1956 (Japan, edaphic)
– Ant IV with 1 basal sensillum. Ant III and II without blunt sensilla. Dens with 1 basal and 1 distal dorsointernal hook O. ambigua Christiansen, 1957 (Lebanon, soil)*
44 Dens with 2 basal and 3 distal dorsointernal hooks. Basal scale on mucro present 45
– Dens with 1 basal and 1 distal dorsointernal hook. Basal scale on mucro absent O. atoyacensis Bonet, 1943 (Mexico, epigeic)
45 PAO lobes relatively long O. pelissiei Deharveng, 1988 (France, cave)
– PAO lobes short 46
46 Basal tubercle of dens with 2 ciliated macrosetae. Ant III with 2 and Ant II with 1 blunt sensillum, respectively. Posterior setae at male genital plate feathery O. yosiiana Szeptycki, 1977 (Japan, North Korea and USA, epigeic)
– Basal tubercle of dens with 1 ciliated simple macroseta and 1 ciliated leaf-shaped macroseta. Ant III with 1 blunt sensillum, Ant II without blunt sensilla. Posterior setae at male genital plate smooth, pointed O. crassicornis Shoebotham, 1911 (England, France, Poland and Hungry, eipigeic and cave)
47 Basal scale on mucro present. PAO lobes subdivided into 3–6 fingers. Ant IV with 4 medial sensilla. Dens with 1 basal and 1 distal dorsoexternal hook. Dens with 3 basal and 3 distal dorsointernal hooks O. moghanensis sp. nov. (Iran, Cave)
– Basal scale on mucro absent 48
48 Some PAO lobes simple and some lobes subdivided into 2 fingers. Ant IV with 4 medial sensilla. Dens without basal dorsoexternal hooks, with 1 distal dorsoexternal hook. Dens with 2 basal and 2 distal dorsointernal hooks O. reyersdorfensis Stach, 1936 (Poland, cave)
– PAO lobes subdivided into 2–3 fingers. Ant IV with 4 medial sensilla. Dens without basal dorsoexternal hooks, with 1 distal dorsoexternal hook. Dens with 3 basal and 4 distal dorsointernal hooks O. dethieri Janssens & De Bruyn, 2010 (Belgium, cave)
– PAO lobes subdivided into 3 fingers. Ant IV without medial sensilla. Dens with 1 basal and 2 distal dorsoexternal hooks. Dens with 1 basal and 1 distal dorsointernal hook O. egerszogensis Loksa, 1961 (Hungary, cave)*

Acknowledgements

The study was supported from the Slovak Research and Development Agency, project APVV-21-0379, and from the Slovak Scientific Grant Agency, project VEGA 1/0438/22. The first author was supported from the postdoctoral program of the Faculty of Science, Pavol Jozef Šafárik University in Košice, Slovakia.

We would like to thank professional speleologist Vahid Ashrafi for preparing the map and photo of the cave. Also, we are grateful to Hamid Nezamdoost, Hamid Sharifi, Kourosh Hosseini and Kavoshgaran Caving Club in Mashhad for assistance during the expeditions to Moghan Cave.

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1 considered as species inquirenda, due to insufficient or doubtful data in an original description.
2 considered as species inquirenda, due to insufficient or doubtful data in an original description.
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