Research Article
Print
Research Article
Paravietnura gen. n., a new intriguing genus of Neanurini from the Caucasus (Collembola, Neanuridae, Neanurinae)
expand article infoAdrian Smolis, Nataliya Kuznetsova§
‡ University of Wrocław, Wrocław, Poland
§ Moscow State Pedagogical University, Moscow, Russia
Open Access

Abstract

Paravietnura gen. n. is described and established for two new species of Neanurini from the Caucasus. The new genus is characterized by an unusual combination of features: the fusion of all lateral tubercles on the head into a single mass, the strong reduction of chaetae on the head, the fusion of cephalic tubercles Af and Oc into a transverse band, the absence of labial chaetae f, the presence of microchaetae on furca rudimentary, and the penultimate abdominal tergum with two tubercles separated along the midline. Paravietnura gen. n. strongly resembles Vietnura Deharveng & Bedos, 2000, a monotypic genus up to date known only from Vietnam. The main characteristics of Paravietnura notabilis sp. n. include an ogival labrum, the absence of chaetae A on the head, relatively thick and widely sheathed long macrochaetae, and minute microchaetae without chaetopores on furca rudimentary. Paravietnura insolita sp. n. differs from the previous species in chaetotaxic details, the size of furcal microchaetae, and the shape of the labrum. Short comments on the generic diversity of the tribe in the Western Palaearctic are also provided.

Keywords

Paravietnura notabilis sp. n., Paravietnura insolita sp. n., Russia, springtails, taxonomy

Introduction

The subfamily Neanurinae, with more than 800 recognized species, is certainly one of the richest and most diversified taxa among springtails (Collembola). It is also one of the most recognizable ones within Collembola as its members possess very characteristic cuticular tubercles on the dorsal side of the body and completely lack the furca, the organ typical for most described springtails. Regarding the taxonomy of Neanurinae, all species are classified into six established tribes (Cassagnau 1989). Among them, Neanurini is the second largest, after Paleonurini, with over 170 species belonging to 23 genera (Cassagnau 1989, Hopkin 1997, Deharveng and Bedos 2000, Deharveng et al. 2007, Smolis 2007, 2011, Mayvan et al. 2015). The number of eyes, the shape of mandibles, as well as the number and arrangement of tubercles on the body are typically used to separate the genera within the tribe. In the lateral part of the head, for instance, there are four main types of tubercle arrangements. The commonest situation is with lateral tubercles L and So fused but tubercle Dl separate. In turn, a complete fusion of all lateral tubercles seems to be the rarest option, only observed in two species of Neanurini, Vietnura caerulea Deharveng & Bedos, 2000, and Monobella cassagnaui Deharveng, 1981.

The examination of rich Neanurinae materials from the north-western Caucasus has revealed two unknown species. They belong to the mentioned tribe because of a presence of blue hypodermic pigment, the last abdominal segment bilobed and well developed tubercles on the body. Interestingly, both the undescribed taxa are characterized by the complete fusion of the outlined cuticular structures on the lateral part of head. This and other features of these species support proposal of a new genus within the tribe. Its detailed and illustrated description is provided with comments on the present stage of knowledge on Neanurini diversity in the Western Palaearctic region.

Terminology

Terminology for the description follows that of Deharveng (1983), with rationale for the definition of chaetae categories), Deharveng and Weiner (1984), Smolis and Deharveng (2006) and Smolis (2008).

Abbreviations used

General morphology:

Abd. abdomen,

Ant. antenna,

AOIII sensory organ of antennal segment III,

Cx coxa,

Fe femur,

Scx2 subcoxa 2,

T tibiotarsus,

Th. thorax,

Tr trochanter,

VT ventral tube.

Groups of chaetae on body excluding antennae:

Ag antegenital,

An chaetae of anal lobes,

Fu furcal,

Ve ventroexternal,

Vea ventroexternoanterior,

Vem ventroexternomedial,

Vep ventroexteroposterior,

Vel ventroexternolateral,

Vec ventroexternocentral,

Vei ventroexternointernal,

Vi ventrointernal,

Vl ventrolateral.

Groups of chaetae on antennae:

ap apical,

ca centroapical,

cm centromedial,

cp centroposterior,

d dorsal,

vc ventrocentral,

ve ventroexternal,

vi ventrointernal.

Tubercles:

Af antenno–frontal,

Cl clypeal,

De dorsoexternal,

Di dorsointernal,

Dl dorsolateral,

L lateral,

Oc ocular,

So subocular.

Types of chaetae:

Ml long macrochaeta,

Mc short macrochaeta,

Mcc very short macrochaeta,

me mesochaeta,

mi microchaeta,

ms ,

S or s chaeta s,

bs s–chaeta on Ant. IV,

miA microchaetae on Ant. IV,

iv ordinary chaetae on ventral Ant. IV,

or organite of Ant IV,

brs border s–chaeta on Ant. IV,

i ordinary chaeta on Ant. IV,

mou cylindrical s–chaetae on Ant. IV (“soies mousses”),

x labial papilla x,

L’ ordinary lateral chaeta on Abd. V,

B4, B5 ordinary chaetae on tibiotarsi,

sgv s–chaeta on Ant. III.

Materials and methods

The specimens were cleared in Nesbitt’s fluid, subsequently mounted on slides in Phoera liquid and studied using a Nikon Eclipse E600 phase contrast microscope. Figures were drawn with camera lucida and prepared for publication using Adobe Photoshop CS3.

Institutions of depository of materials

DIBEC Department of Invertebrate Biology, Evolution and Conservation, Institute of Environmental Biology, University of Wrocław, Poland.

MSPU Moscow State Pedagogical University, Institute of Biology and Chemistry, Moscow, Russia.

Taxonomy

Paravietnura gen. n.

Type species

Paravietnura notabilis sp. n. (here designated).

Diagnosis

Blue pigment present on the body. Tubercles on body well developed, free chaetae on abdomen absent. Two pigmented eyes on each side of head. Mouth parts reduced, maxilla styliform, mandible thin and tridentate. Six tubercles on head, with Dl fused to (L + So), Af fused to Oc, and Di fused to De. Chaetotaxy of head strongly reduced, with chaetae C, D, E, Oca, Di2 and De2 absent. Labrum with four or two prelabral chaetae. Labial chaetae f absent. Tubercles (Di+De+Dl) on Abd. V separate along midline. Cryptopygy present as Abd. VI poorly visible from dorsal side. Claw untoothed.

Etymology

The name “Paravietnura” refers to its strong similarity to Vietnura.

Remarks

The following characters: the presence of 2+2 eyes, the fusion of all lateral tubercles into a single mass on the head, the fusion of cephalic tubercles Af and Oc into a transverse band, the absence of cephalic chaetae Di2 and De2 and the presence of strong cryptopygy place Paravietnura gen. n. very close to Vietnura, the genus established by Deharveng and Bedos (2000) for a single Vietnamese species, V. caerulea Deharveng & Bedos, 2000. As both these genera are distributed in separate biogeographical regions, Paravietnura gen. n. in the Western Palaearctic and Vietnura in the Oriental region, it is recognized that this similarity is probably the result of convergence. Furthermore, they differ in a few essential characters, important from the taxonomic point of view: presence/absence of cephalic chaetae Ocp (in Paravietnura gen. n. present, in Vietnura absent), presence/absence of labial chaetae f (absent in Paravietnura gen. n., Figs 18, 19; present in Vietnura, Fig. 21), the number of tubercles on Abd. V (two tubercles (Di+De+Dl) in Paravietnura gen. n.; three tubercles: 2 (De+Dl) and (Di+Di) in Vietnura), and presence/absence of microchaetae on furca rudimentary (present in Paravietnura gen. n.; absent in Vietnura).

Because of the fusion of lateral tubercles on head, Paravietnura gen. n. resembles Monobella cassagnaui Deharveng, 1981, the species belonging to the European genus Monobella Cassagnau, 1979. Nevertheless, this similarity seems to be definitely superficial given the fact that M. cassagnaui differs from Paravietnura gen. n. in a number of characters e.g.: fusion of tubercles (Di+De) on head along midline (separate in Paravietnura gen. n.), presence of chaetae Di2 and De2 on head (absent in Paravietnura gen. n.), fusion of tubercles Di and De on Th. II–III and Abd. I–II (separate in Paravietnura gen. n.), fusion of tubercles Di on Abd. III along midline (separate in Paravietnura gen. n.) and presence of one tubercle (2Di+2De+2Dl) on Abd. V (two tubercles (Di+De+Dl) in Paravietnura gen. n.).

Paravietnura notabilis sp. n.

Figs 1–7, 8, 19, 20 Table 1, 2, 3

Type material

Holotype: female on slide, Russia, NW Caucasus, Adygeya, Caucasus Nature Reserve, Lagonaki Plateau (‘Kamennoye More’ ridge), 1843 m.alt., litter of rocky pine forest with birch, N44.06159° E40.02103°, 03.07.2014, leg. M. Potapov, N. Kuznetsova, A. Kremenitsa 2014, leg. (MSPU). Paratype: juvenile on slide, ibid., southern slope, 1847 m alt., mixed forest (pine, birch), coniferous litter, N44.06096° E40.02112°, 21.07.2015, leg. M. Potapov, N. Kuznetsova, A. Kremenitsa, L.Vanyavina (DIBEC).

Etymology

The name reflects the notable morphology of this new generotype species.

Diagnosis

Body stumpy and relatively short. Macrochaetae long, thick and widely sheathed. Buccal cone long, labrum ogival. Labrum with two prelabral chaetae. Tubercle (Af +2Oc) on head with chaetae B, Ocm and Ocp, chaetae A absent. Tubercles (Dl+L+So) on head with ten chaetae, chaetae So2 absent. Furca rudimentary with minute and difficult to detect microchaetae, without chaetopores.

Description

General. Body length (without antennae): 0.55 (juvenile) to 0.85 mm (holotype). Colour of the body bluish. 2+2 black eyes of medium size (Fig. 1).

Figures 1–7. 

Paravietnura notabilis sp. n.: 1 habitus and dorsal chaetotaxy (holotype) 2 ventral chaetotaxy of Ant. III 3 dorsal chaetotaxy of Ant. III–IV 4 chaeta Di1 of Abd. IV 5 sensillum of Abd. V 6 chaeta Di1 of Abd. V 7 furca rudimentary.

Chaetal morphology. Dorsal ordinary chaetae of four types: long macrochaetae (Ml), short macrochaetae (Mc), very short macrochaetae (Mcc) and mesochaetae. Long macrochaetae of large length (distinctly longer than length of segment), thick, slightly arc-like or straight, widely sheathed, strongly serrated and apically rounded (Figs 1, 4, 6). Macrochaetae Mc and Mcc morphologically similar to long macrochaetae, but much shorter. Mesochaetae similar to ventral chaetae, thin, smooth and pointed. S–chaetae of tergites thin, smooth, and very short, from three to six times shorter than nearby macrochaetae (Figs 1, 5).

Antennae. Dorsal chaetotaxy of Ant. III–IV as Fig. 3 and Table 2. S–chaetae of Ant. IV long and moderately thickened, S4 and S7 slightly longer than others (Fig. 3). Apical vesicle distinctly bilobed. Ventral chaetotaxy of Ant. III as Fig. 2 and Table 2, sensillum sgv notably long and s-shaped.

Mouthparts. Buccal cone particularly long with labral sclerifications ogival (Fig. 20). Labrum chaetotaxy: 2/2, 4. Labium with three basal, three distal, and three lateral chaetae, papillae x absent (Fig. 19). Maxilla styliform, mandible thin and tridentate.

Dorsal chaetotaxy and tubercles. Chaetotaxy of head as Fig. 1 and Table 1. Chaetotaxy of Th. and Abd. as Table 3. Thorax and abdomen with chaetae De2 and De3 not free (Fig. 1). On Abd. I–III, the line of chaetae De1–chaeta s perpendicular to the dorsomedian line. On Abd. IV chaetae Di1 distinctly longer than on Abd. V (Figs 1, 4, 6). On Abd. V chaetae Di3 present or absent (Fig. 1). Cryptopygy present, strongly developed.

Ventral chaetotaxy. On head, groups Vea, Vem and Vep with 3, 2, 4 chaetae respectively. Group Vi on head with six chaetae. On Abd. IV, furca rudimentary with minute microchaetae (Fig. 7). On Abd. IV, tubercle L with five chaetae. On Abd. V, chaetae L’ absent.

Legs. Chaetotaxy of legs as in Table 3. Claw without internal tooth. On tibiotarsi, chaeta M present and chaetae B4 and B5 relatively short and pointed.

Table 1.

Cephalic chaetotaxy of Paravietnura notabilis sp. n., dorsal side.

Tubercle Number of chaetae Types of chaetae Names of chaetae
Cl 4 Ml F
Mc G
(Af+2Oc) 6 Ml B, Ocm
Mc Ocp
(Di+De) 2 Ml Di1, De1
(Dl+L+So) 10 Ml Dl1, Dl5, L1, So1
Mc L4
Mcc Dl4
me So3–6
Table 2.

Antennal chaetotaxy of Paravietnura notabilis sp. n.

Segment, Group Number of chaetae Segment, Group Number of chaetae adult
I 7 IV or, 8 S, i, 12 mou, 6 brs, 2 iv
II 11
IIIve 5 sensilla AO III
5 ap 8 bs, 5 miA
vc 4 ca 2 bs, 3 miA
vi 4 cm 3 bs, 1 miA
d 4 cp 8 miA, 1 brs
Table 3.

Postcephalic chaetotaxy of Paravietnura notabilis sp. n.

Terga Legs
Di De Dl L Scx2 Cx Tr Fe T
Th. I 1 2 1 - 0 3 6 13 19
Th. II 3 2+s 3+s+ms 3 2 7 6 12 19
Th. III 3 3+s 3+s 3 2 8 6 11 18
Sterna
Abd. I 2 3+s 2 3 VT: 4
Abd. II 2 3+s 2 3 Ve: 4; chaeta Ve1 absent
Abd. III 2 3+s 2 3 Vel:4; Fu: 5 me, 4 mi
Abd. IV 2 2+s 3 5 Vel: 4; Vec: 2; Vei: 2; Vl: 4
Abd. V 6–7+s Ag: 3; Vl: 1
Abd. VI 7 Ve: 9–10; An: 2mi

Remarks

See Remarks of Paravietnura insolita sp. n.

Ecological note

The new species seems to be very local and connected with specific climatic or vegetation conditions (rocky pine-birch forest, Fig. 8) since it has been never recorded outside locus typicus despite many investigations conducted in different parts of the Caucasus.

Figure 8. 

Rocky pine-birch forest in Caucasus Nature Reserve, type locality of Paravietnura notabilis sp. n.

Paravietnura insolita sp. n.

Figs 9–18, Table 4, 5, 6

Type material

Holotype: male on slide, Russia, Caucasus, Northern Ossetia, North Ossetia Nature Reserve, surroundings of the village Tsey, Kalpersky ridge, southern slope, 2160 m alt., litter of rocky pine grass forest, 19.8.1977, leg. M. Rudakovsky, N. Kuznetsova (MSPU). Paratype: male on slide, ibid., green moss pine forest, in mosses, 23.9.1980, leg. I. Kuchiev (DIBEC).

Etymology

Its name reflects a later discovery of another species within the genus (Latin word “insolita” means not lonely).

Diagnosis

Body relatively short and squarish. Macrochaetae long, relatively thin and narrowly sheathed. Buccal cone rather short, labrum non ogival. Labrum with four prelabral chaetae. Tubercle (Af +2Oc) on head with chaetae A, B, Ocm and Ocp. Tubercles (Dl+L+So) on head with eleven chaetae, chaetae So2 present. Furca rudimentary with well visible microchaetae, with chaetopores.

Description

General. Body length (without antennae): 1.10 (paratype) to 1.35 mm (holotype). Colour of the body bluish. 2+2 black eyes of medium size (Fig. 9).

Figures 9–21. 

Paravietnura insolita sp. n.: 9 dorsal chaetotaxy of head and Th. I (holotype) 10 dorsal chaetotaxy of Ant. III–IV 11 ventral chaetotaxy of Ant. III 12 dorsal chaetotaxy of Abd. V–VI 13 furca rudimentary 14 chaeta Di1 of Abd. IV 15 chaeta Di1 of Abd. V 16 sensillum of Abd. V 17 labrum 18 labium. Paravietnura notabilis sp. n.: 19 labium 20 labrum. Vietnura caerulea Deharveng and Bedos, 2000: 21 labium.

Chaetal morphology. Dorsal ordinary chaetae of four types: long macrochaetae (Ml), short macrochaetae (Mc), very short macrochaetae (Mcc) and mesochaetae. Long macrochaetae of medium length (longer than length of segment), relatively thin, slightly arc-like or straight, narrowly sheathed, strongly serrated and apically rounded (Figs 9, 12, 14, 16). Macrochaetae Mc and Mcc morphologically similar to long macrochaetae, but much shorter. Mesochaetae similar to ventral chaetae, thin, smooth, and pointed. S–chaetae of tergites thin, smooth, and markedly short, three or four times shorter than nearby macrochaetae (Figs 12, 15).

Antennae. Dorsal chaetotaxy of Ant. III–IV as Fig. 10 and Table 5. S–chaetae of Ant. IV long and moderately thickened, S4 and S7 slightly longer than others (Fig. 10). Apical vesicle distinct bilobed. Ventral chaetotaxy of Ant. III as Fig. 11 and Table 5, sensillum sgv notably elongate and s-shaped.

Mouthparts. Buccal cone short with labral sclerifications non-ogival (Fig. 17). Labrum chaetotaxy: 4/2, 4. Labium with three basal, three distal and three lateral chaetae, papillae x absent (Fig. 18). Maxilla styliform, mandible thin and tridentate.

Dorsal chaetotaxy and tubercles. Chaetotaxy of head as Fig. 9 and Table 4. Chaetotaxy of Th. and Abd. as Table 6. Thorax and abdomen with chaetae De2 and De3 not free. On Abd. I–III, the line of chaetae De1–chaeta s perpendicular to the dorsomedian line. On Abd. IV chaetae Di1 longer than on Abd. V (Figs 14, 16). On Abd. V chaetae Di2 present and Di3 absent (Fig. 12). Cryptopygy present, well developed.

Ventral chaetotaxy. On head, groups Vea, Vem and Vep with 3, 2, 4 chaetae respectively. Group Vi on head with six chaetae. On Abd. IV, furca rudimentary with clearly visible microchaetae, each with chaetopore (Fig. 13). On Abd. IV, tubercle L with five chaetae. On Abd. V, chaetae L’ present.

Legs. Chaetotaxy of legs as in Table 6. Claw without internal tooth. On tibiotarsi, chaeta M present and chaetae B4 and B5 relatively short and pointed.

Remarks

Paravietnura insolita sp. n. can be confused with the only other described species in the genus Paravietnura notabilis sp. n., which has thick and widely sheathed long macrochaetae (distinctly thinner and narrowly sheathed in insolita), an ogival labrum (non-ogival in insolita), chaetae A and So2 reduced on the head (present in insolita), chaetae L’ reduced on Abd. V (present in insolita), and the furca rudimentary with minute microchaetae without chaetopores (longer microchaetae with distinct chaetopores in insolita).

Ecological note

Similarly to the previous species, Paravietnura insolita sp. n. seems to be very local and connected with specific climatic and vegetation conditions, probably with mountain xeric pine forest.

Variabilty

We observed an asymmetrical absence of chaeta A in the holotype (Fig. 9).

Table 4.

Cephalic chaetotaxy of Paravietnura insolita sp. n., dorsal side.

Tubercle Number of chaetae Types of chaetae Names of chaetae
Cl 4 MlMc FG
(Af+2Oc) 8 MlMc B, OcmA, Ocp
(Di+De) 2 Ml Di1, De1
(Dl+L+So) 11 MlMcMccme Dl, Dl5, L1, So1L4Dl4, So2So3–6
Table 5.

Antennal chaetotaxy of Paravietnura insolita sp. n.

Segment, Group Number of chaetae Segment, Group Number of chaetae adult
I 7 IVap or, 8 S, i, 12 mou, 6 brs, 2 iv
II 11
IIIve 5 sensilla AO III
5 8 bs, 5 miA
vc 4 ca 2 bs, 3 miA
vi 4 cm 3 bs, 1 miA
d 4 cp 8 miA, 1 brs
Table 6.

Postcephalic chaetotaxy of Paravietnura insolita sp. n.

Terga Legs
Di De Dl L Scx2 Cx Tr Fe T
Th. I 1 2 1 - 0 3 6 13 19
Th. II 3 2+s 3+s+ms 3 2 7 6 12 19
Th. III 3 3+s 3+s 3 2 8 6 11 18
Sterna
Abd. I 2 3+s 2 3 VT: 4
Abd. II 2 3+s 2 3 Ve: 5; chaeta Ve1 present
Abd. III 2 3+s 2 3 Vel: 4; Fu: 5–6 me, 8 mi
Abd. IV 2 2+s 3 5 Vel: 4; Vec: 2; Vei: 2; Vl: 4
Abd. V 6+s Ag: 2; Vl: 1, L‘: 1
Abd. VI 7 Ve: 11–12; An: 2mi

Discussion

As noted in the Introduction, the tribe Neanurini, containing more than 170 species in 23 genera, is the second largest within the subfamily Neanurinae. Paradoxically, in spite of such a large number of known species belonging to the tribe, knowledge about its distribution and diversity seems to be still incomplete and far from satisfactory. For example, the largest generic and species diversity of Neanurini occurs in the Western Palaearctic, where currently 18 genera and nearly 150 species are known. It should be mentioned that the tribe in this region absolutely dominates and constitutes more than 80% of all genera and species of Neanurinae. The other four Neanurinae tribes are represented only by single native species (Paranurini, Sensillanurini), single introduced species (Lobellini), and a few genera with relatively small numbers of species (Paleonurini). However, presently the picture of the generic distribution of Neanurini is highly uneven as its seven genera are known exclusively from Europe (Albanura Deharveng, 1982; Cansilianura Dallai & Fanciulli, 1983; Catalanura Deharveng, 1983; Lathriopyga Caroli, 1910; Monobella Cassagnau, 1979; Neanurella Cassagnau, 1968; Pumilinura Cassagnau, 1979), and the next seven taxa (Balkanura Cassagnau, 1979, Cryptonura Cassagnau, 1979; Deutonura Cassagnau, 1979; Endonura Cassagnau, 1979; Neanura MacGillivray, 1893; Protanura Börner, 1906; Thaumanura Börner, 1932) are present both in Europe and in areas around it, e.g., Asia Minor, the Middle East, the Caucasus, or North Africa. That means that as many as 14 genera of the tribe, of 18 known in the western part of the Palaearctic, are present in Europe. Fortunately, to better understand the distributional pattern of Neanurini, a substantial number of studies dedicated to poorly investigated areas outside the continent have been undertaken during the last three decades. They have resulted the descriptions of four unknown genera, i.e. Caucasanura Kuznetsova & Potapov, 1988; Edoughnura Deharveng, Hamra-Kroua & Bedos, 2007; Ghirkanura Kuznetsova & Potapov, 1988; Persanura Mayvan, Shayanmehr, Smolis & Skarżyński, 2015; and many new species and records of known taxa (e.g., Kuznetsova and Potapov 1988, Deharveng et al. 2007, 2015, Smolis and Kaprus’ 2009, Mayvan et al. 2015, Smolis et al. 2011, 2012, 2016a, b, c, 2017). These papers, including the work presented here, show that the diversity of Neanurini in some regions of the Western Palaearctic is still underestimated, and that this diversity can be crucial to assessing the history of this megadiverse tribe of Nenurinae.

Acknowledgments

The work was financially supported by the Russian Foundation for Basic Research (grant 04-16-01228) and the Institute of Environmental Biology, Faculty of Biological Science, University of Wrocław, Poland (project no. 1076/Ś/IBŚ/2017). We are grateful to Prof. Pietro Paolo Fanciulli and Prof. Gang Jiang who reviewed the manuscript. We are also much indebted to the Editor Prof. Louis Deharveng for helpful remarks.

References

  • Cassagnau P (1989) Les Collemboles Neanurinae; éléments pour une synthèse phylogénétique et biogéographique. In: Dallai R (Ed.) 3rd International Seminar on Apterygota, Siena, 171–182.
  • Deharveng L (1983) Morphologie évolutive des Collemboles Neanurinae en particulier de la ligne Neanurinae. Travaux de Laboratoire d’Écobiologie des Arthropodes Édaphiques, Toulouse 4: 1–63.
  • Deharveng L, Weiner WM (1984) Collemboles de Corée du Nord III–Morulinae et Neanurinae. Travaux de Laboratoire d’Écobiologie des Arthropodes Édaphiques, Toulouse 4: 1–61.
  • Deharveng L, Bedos A (2000) Vietnura caerulea new genus, new species, from Vietnam: first record of the Palearctic tribe Neanurini in tropical Asia (Collembola: Neanuridae). The Raffles Bulletin of Zoology 48(2): 209–214.
  • Deharveng L, Hamra-Kroua S, Bedos A (2007) Edoughnura rara n. gen., n. sp., an enigmatic genus of NeanurinaeCollembola from the Edough Massif (Algeria). Zootaxa 1652: 57–61.
  • Deharveng L, Moloud SA, Bedos A (2015) A new species of Deutonura (Collembola: Neanuridae: Neanurinae) from Algeria, with revised diagnosis of the genus and key to western Palaearctic species. Zootaxa 4000(4): 464–472. https://doi.org/10.11646/zootaxa.4000.4.5
  • Hopkin SP (1997) Biology of the springtails (Insecta: Collembola). Oxford University Press, Oxford, 330 pp.
  • Kuznetsova NA, Potapov MB (1988) New data on the taxonomy of springtails of the family Neanuridae and Hypogastruridae (Collembola). Zoologichesky Zhurnal 67(12): 1833–1844.
  • Mayvan MM, Shayanmehr M, Smolis A, Skarżyński D (2015) Persanura hyrcanica, a new genus and species of Neanurinae (Collembola: Neanuridae) from Iran, with a key to genera of the tribe Neanurini. Zootaxa 3918: 552–558. https://doi.org/10.11646/zootaxa.3918.4.4
  • Smolis A (2007) Kalanura–a new genus of Neanurini (Collembola, Neanuridae, Neanurinae) from Siberia, with decription of four new species. Zootaxa 1511: 1–16.
  • Smolis A (2008) Redescription of four Polish Endonura Cassagnau, 1979 (Collembola, Neanuridae, Neanurinae), with a nomenclature of the ventral chaetae of antennae. Zootaxa 1858: 9–36.
  • Smolis A (2011) Xylanura oregonensis, a new genus and species of saproxylic springtail (Collembola: Neanuridae: Neanurinae) from North America, with a key to genera of the tribe Neanurini. The Pan-Pacific Entomologist 87(1): 15–26. https://doi.org/10.3956/2010-06.1
  • Smolis A, Deharveng L (2006) Vitronura mascula, a new species of Neanurinae (Collembola: Neanuridae) from northern Vietnam, with a key to the species of the genus. Revue suisse de Zoologie 113: 263–268. https://doi.org/10.5962/bhl.part.80349
  • Smolis A, Kaprus’ IJ (2009) Redescription of Endonura cretensis (Ellis, 1976) (Collembola: Neanuridae: Neanurinae) with a new record from Israel. Annales Zoologici 59: 1–6. https://doi.org/10.3161/000345409X432538
  • Smolis A, Deharveng L, Kaprus’ IJ (2011) Studies on the non-European Endonura Cassagnau, 1979 (Collembola, Neanuridae, Neanurinae). Zootaxa 3004: 45–56.
  • Smolis A, Falahati A, Skarżyński D (2012) The genus Cryptonura Cassagnau, 1979 (Collembola, Neanuridae, Neanurinae) in Iran. Zootaxa 3530: 51–58.
  • Smolis A, Kahrarian M, Piwnik A, Skarżyński D (2016a) Endonura Cassagnau in Iran, with a key of the genus (Collembola, Neanuridae, Neanurinae). ZooKeys 553: 53–71. https://doi.org/10.3897/zookeys.553.6009
  • Smolis A, Kuznetsova N (2016b) Remarkable diversity of the genus Endonura Cassagnau, 1979 (Collembola: Neanuridae: Neanurinae) in the Caucasus. Zootaxa 4200(1): 47–82. https://doi.org/10.11646/zootaxa.4200.1.2
  • Smolis A, Skarżyński D, Kahrarian M, Kaprus IJ (2016c) Redescription of Protanura papillata Cassagnau & Delamare Deboutteville, 1955 (Collembola, Neanuridae, Neanurinae), with new records from Middle East, and with supplemented diagnosis and key to the genus. Zootaxa 4092(2): 293–300. http://doi.org/10.11646/zootaxa.4092.2.11
  • Smolis A, Shayanmehr M, Kuznetsova N, Lafooraki EY (2017) Three new remarkable species of the genus Endonura Cassagnau, 1979 from the Middle East and Central Asia (Collembola, Neanuridae, Neanurinae, Neanurini). ZooKeys 673: 135–151. https://doi.org/10.3897/zookeys.673.12084