Endonura Cassagnau in Iran, with a key to species of the genus (Collembola, Neanuridae, Neanurinae)

Adrian Smolis; Morteza Kahrarian; Agata Piwnik; Dariusz Skarżyński

doi:10.3897/zookeys.553.6009

Abstract

Three new species of Endonura are described from Iran. Endonura dichaeta sp. n. can be recognized by an ogival labrum, head without chaetae O and E, chaeta D connected with tubercle Cl, tubercle Dl with five chaetae on head, absence of tubercles Di on thorax I and tubercle (Di+Di) of thorax V with 2+2 chaetae. Endonura ceratolabralis sp. n. is characterized by large body size, reduction of labral chaetotaxy, ogival labrum, head without chaeta O and fusion of tubercles Di and De on first thoracic segment. Endonura persica sp. n. is distinguished from its congeners by a nonogival labrum, absence of chaeta O, tubercles Dl and (L+So) with five and eight chaetae respectively and claw with inner tooth. The key to all species of the genus is given.

Keywords

Springtails, taxonomy, new species

Introduction

Endonura was established by Cassagnau (1979) as one of four subgenera within the genus Neanura MacGilliwray, 1893. Later, Deharveng (1982) raised it to the generic level. At present, Endonura is one of the largest (37 valid species) and most accurately studied genera within the subfamily Neanurinae (Dallai 1983, Deharveng 1979, 1982, Fanciulli and Dallai 2008, Pomorski and Skarżyński 2000, Pozo and Simón 1982, Smolis and Kaprus’ 2003, 2009, Smolis 2006, Smolis et al. 2007, 2011). It is mostly a Palaearctic genus and only one species, E. reticulata (Axelson, 1905), is known from the Nearctic (Alaska, Smolis et al. 2011). According to a recent definition (Smolis 2008), Endonura is characterized by the following characters: 0–2 ocelli, reduced mouth parts with a thin mandible and a styliform maxilla, separate tubercles Di and De on the head, the non-cross-type of chaetotaxy on the head and three or two tubercles on abdomen V. The highest species diversity is observed in Europe (32 from among the 37 known species). However, this may be a false picture because many areas of the Palaearctic have been poorly studied by collembologists. Undoubtedly, one of such regions is Central Asia, but in this case the situation is rapidly and positively changing (Arbea and Kahrarian 2015, Kahrarian 2014, Kahrarian et al. 2013, Mayvan et al. 2015, Shayanmehr et al. 2013, Smolis et al. 2012). In the present paper, three new non-European Endonura from the western part of Iran are described. An updated key to all species of the genus is included.

Terminology

Terminology for the description follows that given in Deharveng (1983), 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: Ag – antegenital, An – chaetae of anal lobes, ap – apical, ca – centroapical, cm – centromedial, cp – centroposterior, d – dorsal, Fu – furcal, vc – ventrocentral, Ve or ve – ventroexternal, Vea – ventroexternoanterior, Vem – ventroexternomedial, Vep – ventroexternoposterior, Vel – ventroexternolateral, Vec – ventroexternocentral, Vei – ventroexternointernal, Vi or vi – ventrointernal, Vl – ventrolateral.

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–microchaeta or microsensillum, S or s – chaeta s, bs – border 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.

Materials and methods

The specimens were cleared in Nesbitt’s fluid, subsequently mounted on slides in Swan’s medium and observed using a phase contrast microscope Nikon E600. Photographs were made using a camera Nikon D5100 mounted on a microscope mentioned above. Photographs were stacked using Helicon Focus 6.2.2. and prepared for publication using Adobe Photoshop CS3. Material is deposited in the Department of Invertebrate Biology, Evolution and Conservation, Institute of Environmental Biology, University of Wrocław, Poland.

Taxonomy

Endonura dichaeta sp. n.

http://zoobank.org/4CBE64B2-069B-4254-AF20-43D26E6CFE10

Figs 1–4, Table 1

Type material

Holotype: adult female on slide, Iran, Osmanevand area, near Golestan village (N33°55', E47°06', 1241 m a.s.l.), litter in oak forest, 13.XII.2013, leg. M. Kahrarian. Paratypes: female, two males and two juveniles on slides, same data as holotype.

Other material

Two females and male on slide, Iran, Osmanevand area, near Chelkooshk village (N34°03', E47°12', 1516 m a.s.l.), litter in oak forest, 31.I.2014, leg. M. Kahrarian; three juveniles on slide, Iran, Paveh county, near Shabankereh village (N34°52.978', E46°30.760', 1632 m a.s.l.), litter in oak forest, 20.I.2014, leg. M. Kahrarian; two females and juvenile, Iran, Kermanshah county, near Chahar zebra-e-oliya village (N34°13', E46°40', 1592 m a.s.l.), litter in oak forest, 24.I.2014, leg. M. Kahrarian.

Etymology

The species name refers to rare feature within the genus - only two chaetae Di on each side of tubercle (Di+Di) of abdomen V.

Diagnosis

Habitus typical of the genus Endonura. Dorsal tubercles present and well developed, except tubercles Di on th. I. 2+2 unpigmented eyes. Buccal cone long, labrum ogival. Head with chaetae A, B, C, D, F and G. Chaetae O and E absent. Tubercles Cl and Af separate. Tubercle Cl with chaetae D. Tubercles Dl and (L+So) on head with five and eight chaetae respectively. Tubercles De on th. II and III with three and four chaetae respectively. Tubercles L on abd. III and IV with three and six chaetae respectively. Abd. IV and V with eight and three tubercles respectively. Claw without inner tooth. Tibiotarsi with chaetae B4 and B5 short.

Description

Habitus typical of the genus. Body length (without antennae): 0.75–1.55 mm (holotype 1.30 mm). Colour of the body white. 2+2 medium unpigmented eyes (Fig. 1).

Figures 1–4.

Endonura dichaeta sp. n.: 1 head (holotype), dorsal and lateral chaetotaxy 2 ventral sclerification of labrum 3 dorsal chaetotaxy of thorax 4 dorsal chaetotaxy of abdomen III–VI. Arrows indicate the position of eyes.

Types of dorsal ordinary chaetae. Macrochaetae Ml relatively long, strongly thickened, almost cylindrical, arc-like or straight, narrowly sheathed, feebly serrated, apically pointed (Figs 1, 3–4); macrochaetae Mc and Mcc thickened, straight and pointed; mesochaetae and microchaetae short, thin, feebly serrated and pointed.

Head. Labrum ogival, with ventral sclerifications as in Fig. 2. Labrum chaetotaxy 2/2, 4. Labium with four basal, three distal and four lateral chaetae, papillae x absent. Maxilla styliform, mandible thin tridentate. Chaetotaxy of antennae as in Table 1c. Apical vesicle distinct, trilobed. S–chaetae of ant.IV long and moderately thickened. Chaetotaxy of head as in Table 1a, b, and Fig. 1. Chaeta D connected with tubercle Cl. Tubercle Af on head longer than tubercles Oc. Tubercle Dl with five chaetae, chaeta Dl3 absent, chaeta Dl6 as minute microchaeta and hard to detect (Fig. 1). Tubercle (L+So) with eight chaetae, chaetae So2 and L3 absent, chaeta So6 as Mc (Fig. 1). Elementary tubercles BE and CD present. Chaeta A shorter than B.

Table 1a.

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Chaetotaxy of Endonura dichaeta sp. n.: Cephalic chaetotaxy–dorsal side.

Tubercle	Number of chaetae	Types of chaetae	Names of chaetae
Cl	6	Ml Mc	F D, G
Af	6	Ml Mc	A B, C
Oc	3	Ml Mc	Ocm, Ocp Oca
Di	2	Ml Mc	Di1 Di2
De	2	Ml Mc	De1 De2
Dl	5	Ml Mc Mcc mi	Dl1, Dl5 Dl4 Dl2 Dl6
(L+So)	8	Ml Mc me	L1, L4, So1 L2, So6 So3–5

Table 1b.

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Chaetotaxy of Endonura dichaeta sp. n.: Cephalic chaetotaxy–ventral side.

Group	Number of chaetae
Vi	6
Vea	3
Vem	3
Vep	4
labium	11, 0x

Table 1c.

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Chaetotaxy of Endonura dichaeta sp. n.: Chaetotaxy of antennae.

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

Thorax, abdomen, legs. Body s–chaeta thin and smooth, shorter than nearby macrochaetae (Figs 3, 4). Chaetotaxy of th. and abd. as in Table 1d and in Figs 3, 4. Tubercles Di on th.I not differentiated. Chaetae De3 on th. III and abd. I–III as Mcc. Chaetae De2 on th. II–III and De3 on th. III connected with tubercle De. Chaetae De3 on abd. I–III connected with tubercle De (Fig. 4). The line of chaetae De1–chaeta s not perpendicular to the dorsomedian line on abd I–III. Furca rudimentary with 2–4 microchaetae. Tubercles Di on abd. V fused, with chaetae Di2 as Mc or Mcc, chaetae Di3 absent (Fig. 4). Chaetae L’ and Vl on abd. V present. IV abd. with 2+2 chaetae Ag. No cryptopygy. Chaetotaxy of legs as in Table 1d.

Table 1d.

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Chaetotaxy of Endonura dichaeta sp. n.: Postcephalic chaetotaxy.

	Di	Terga De	Dl	L	Scx2	Cx	Legs 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: 4–5	Ve1 -	present
abd. III	2	3+s	2	3	Vel:4–5			Fu:5–6me	2–4mi
abd. IV	2	2+s	3	6	Vel: 4	Vec: 2	Vei: 2	Vl: 4
abd. V	(2+2)	5+s			Ag: 2			Vl: 1	L’: 1
abd. VI		7			Ve:13–14			An: 2mi

Remarks

In general appearance (shape of dorsal chaetae, chaetotaxy of central area of head and dorsal side of thorax and abdomen, complete absence of pigmentation and absence of cryptopygy), E. dichaeta sp. n. strongly resembles E. tartaginenis Deharveng, 1980 described from Corsica. Nevertheless, both taxa differ in some essential characters, important from taxonomic point of view: presence/absence of chaetae E on head (dichaeta sp. n. absent, tartaginenis present), number of chaetae Dl on head (dichaeta sp. n. five, tartaginenis six), number of chaetae (L+So) (dichaeta sp. n. eight, tartaginenis nine), presence/absence of elementary tubercle EE on head (dichaeta sp. n. absent, tartaginenis present), presence/absence of tubercles Di on the first thoracic segment (dichaeta sp. n. absent, tartaginenis present), number of chaetae Di on abd. V (dichaeta sp. n. 2+2, tartaginenis 3+3) and presence/absence of tooth on claw (dichaeta sp. n. absent, tartaginenis present). In addition, the new species is characterized by only 2+2 antegenital chaetae (tartaginenis 3+3) and ogival labrum (unknown in tartaginenis), characters rarely observed within the genus.

Endonura ceratolabralis sp. n.

http://zoobank.org/FC09DDF3-EB60-416D-B31C-A290A4E812D5

Figs 5–9, Table 2

Type material

Holotype: adult female on slide, Iran, Osmanevand area, near Markhor village (N33°53', E47°05', 1389 m a.s.l.), litter in oak forest, 13.XII.2013, leg. M. Kahrarian. Paratypes: 3 females on slide, same data as holotype.

Other material

Three females on slide, Iran, Osmanevand area, near Ghader marz village (N34°01.030', E47°12.415', 1682 m a.s.l.), litter in oak forest, 31.I.2014, leg. M. Kahrarian.

Etymology

The species name refers to sharp labral apex which looks like a horn (“cera” in latin).

Diagnosis

Habitus typical of the genus Endonura. Dorsal tubercles present and well developed. 2+2 eyes darkly pigmented. Buccal cone long. Head with chaetae A, B, C, D, E, F and G. Chaeta O absent. Tubercles Cl and Af separate. Tubercles Dl and (L+So) on head with six and nine chaetae respectively. Tuberles Di and De on th. I fused. Tubercles De on th. II and III with three and four chaetae respectively. Tubercles L on abd. III and IV with three and 6–7 chaetae respectively. Abd. IV and V with eight and three tubercles respectively. Claw without inner tooth. Tibiotarsi with chaetae B4 and B5 short.

Description

Habitus typical of the genus. Body length (without antennae): 2.25–2.55 mm (holotype: 2.55 mm). Colour of the body bluish grey. 2+2 medium dark-pigmented eyes (Fig. 5).

Figures 5–9.

Endonura ceratolabralis sp. n.: 5 head (holotype), dorsal and lateral chaetotaxy 6 ventral sclerification of labrum 7 dorsal chaetotaxy of thorax 8 dorsal chaetotaxy of abd. II 9 dorsal chaetotaxy of abdomen IV–VI. Arrows indicate the position of eyes.

Types of dorsal ordinary chaetae. Macrochaetae Ml thickened, relatively long, arc-like or straight, narrowly sheathed, feebly serrated, apically pointed or rarely rounded (Figs 5, 7–9); macrochaetae Mc and Mcc thickened, straight, pointed or apically rounded; mesochaetae and microchaetae short, thin and pointed.

Head. Buccal cone very long. Labrum ogival, with ventral sclerifications as in Fig. 6. Labrum chaetotaxy 0/2, 2. Labium with four basal, three distal and four lateral chaetae, papillae x absent. Maxilla styliform, mandible thin with two basal and two apical teeth. Chaetotaxy of antennae as in Table 2c. Apical vesicle distinct, trilobed. S–chaetae of ant.IV of medium length and moderately thickened. Chaetotaxy of head as in Table 2a, b, and Fig. 5. Tubercles Cl and Af separate. Tubercle Af on head longer than tubercles Oc. Chaeta O absent. Chaeta D free. Tubercle Dl with six chaetae, chaeta Dl3 present. Tubercle (L+So) with nine chaetae, chaeta So2 absent and chaeta So3 as Mc (Fig. 5). Elementary tubercles BE and CD present. Chaeta A shorter than B.

Table 2a.

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Chaetotaxy of Endonura ceratolabralis sp. n.: Cephalic chaetotaxy–dorsal side.

Tubercle	Number of chaetae	Types of chaetae	Names of chaetae
Cl	4	Ml Mc	F G
Af	10	Ml Mc	A B, C, D, E
Oc	3	Ml me	Ocm, Ocp Oca
Di	2	Ml Mc	Di1 Di2
De	2	Ml Mc	De1 De2
Dl	6	Ml Mc mi	Dl1, Dl5 Dl2, Dl3, Dl4 Dl6
(L+So)	9	Ml Mc me	L1, L4, So1 L2, L3, So3 So4–6

Table 2b.

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Chaetotaxy of Endonura ceratolabralis sp. n.: Cephalic chaetotaxy–ventral side.

Group	Number of chaetae
Vi	6
Vea	3–4
Vem	3
Vep	4
Labium	11, 0x

Table 2c.

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Chaetotaxy of Endonura ceratolabralis sp. n.: Chaetotaxy of antennae.

Segment, Group	Number of chaetae	Segment, Group	Number of chaetae adult
I	7	IV ap	or, 8 S, i, 12 mou, 6 brs, 2 iv
II	12–14
III ve	5 sensilla AO III
III ve	5		8 bs, 5 miA
vc	4	ca	2 bs, 3 miA
vi	4	cm	3 bs, 1 miA
d	5	cp	8 miA, 1 brs

Thorax, abdomen, legs. Body s–chaetae thin and smooth, shorter than nearby macrochaetae (Figs 7–9). Chaetotaxy of th. and abd. as in Table 2d and in Figs 7–9. Tubercles Di on th.I differentiated and fused with De (Fig. 7). Dorsal side of th. and abd. without free chaetae De. The line of chaetae De1–chaeta s perpendicular to the dorsomedian line on abd I–III. Furca rudimentary with two or without microchaetae. Tubercles Di on abd. V fused, with chaetae Di2 and Di3 as Mc (Fig. 9). Chaetae L’ and Vl on abd. V present. No cryptopygy. Chaetotaxy of legs as in Table 2d.

Table 2d.

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Chaetotaxy of Endonura ceratolabralis sp. n.: Postcephalic chaetotaxwy.

	Di	Terga De	Dl	L	Scx2	Cx	Legs 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: 5–6	Ve1 -	Present
abd. III	2	3+s	2	3	Vel:4–5			Fu:3–6me	0–2mi
abd. IV	2	2+s	3	6–7	Vel: 4	Vec: 2	Vei: 2	Vl: 4
abd. V	(3+3)	5+s			Ag: 3			Vl: 1	L’: 1
abd. VI		7			Ve: 13–14			An: 2mi

Remarks

Because of the very characteristic long and pointed labrum, E. ceratolabralis sp. n. seems to be most similar to E. cretensis (Ellis, 1976) (Crete) and E. gracilirostris Smolis et al. 2007 (Crimea). Nevertheless, the new species can be easily distinguished from these two taxa by the following combination of characters: maximum length of the body without antennae (ceratolabralis sp. n. 2.55 mm; gracilirostris1.45 mm; cretensis 0.8 mm), labral formula (ceratolabralis sp. n. 0/2, 2; gracilirostris 0/2, 4; cretensis 2/2, 4), presence/absence of chaeta O on head (ceratolabralis sp. n. absent, in others present), number of chaetae Dl on head (cretensis three, in others six), number of chaetae (L+So) on head (cretensis seven, in others nine), fusion/separation of tubercles Di and de on th. I (gracilirostris separate, in others fused), number of ordinary chaetae De on th. III (cretensis two, others three), presence/absence of free chaetae on thorax (ceratolabralis sp. n. absent, in others present) and number of chaetae Di on abd. V (cretensis 1-2, others three).

Endonura persica sp. n.

http://zoobank.org/9CFE5947-62CC-4A3E-ABF7-5B84EA69A21A

Figs 10–13, Table 3

Type material

Holotype: adult female on slide, Iran, Kermanshah area, near Ghaleh shahin village (N34°25.590', E05°12.415', 566 m a.s.l.), litter in willow shrubs, 7.IV.2014, leg. M. Kahrarian. Paratypes: two females, four males and four juveniles on slides, same data as holotype.

Other material

Female on slide, Iran, Kermanshah Province, Halashi County, near Sarfiroozabad village (N34°02', E47°10', 1624 m a.s.l.), litter in oak forest, 15.II.2014, leg. M. Kahrarian; female and male on slide, Iran, Osmanevand area, near Sarjoob village (N33°56', E47°08', 1240 m a.s.l.), litter in oak forest, 13.XII.2013, leg. M. Kahrarian.

Etymology

The species name refers to the historic name of Iran, Persia.

Diagnosis

Habitus typical of the genus Endonura. Dorsal tubercles present and generally well developed, only tubercles Di on th. I weakly differentiated. 2+2 large dark-pigmented eyes. Buccal cone rather short. Head with chaetae A, B, C, D, E , F and G. Chaeta O absent. Tubercles Cl and Af separate. Tubercles Dl and (L+So) on head with five and eight chaetae respectively. Tubercles De on th. II and III with three and four chaetae respectively. Tubercles L on abd. III and IV with four and 6–7 chaetae respectively. Abd. IV and V with eight and three tubercles respectively. Claw with inner tooth. Tibiotarsi with chaetae B4 and B5 long.

Description

Habitus typical of the genus. Body length (without antennae): 0.75–1.90 mm (holotype: 1.10 mm). Colour of the body bluish grey. 2+2 large dark pigmented eyes (Fig. 10).

Figures 10–13.

Endonura persica sp. n.: 10 head and th. I, dorsal and lateral chaetotaxy 11 ventral sclerification of labrum 12 dorsal chaetotaxy of abdomen III–VI (holotype) 13 tibiotarsi and claw of leg III. Arrows indicate the position of eyes.

Types of dorsal ordinary chaetae. Macrochaetae Ml thickened, relatively long, arc–like or straight, narrowly sheathed, feebly serrated, apically rounded or rarely pointed (Figs 10, 12); macrochaetae Mc and Mcc thickened, straight and not pointed; mesochaetae and microchaetae short, thin and pointed.

Head. Buccal cone short. Labrum rounded, with ventral sclerifications as in Fig. 11. Labrum chaetotaxy 4/2, 4. Labium with four basal, three distal and four lateral chaetae, papillae x absent. Maxilla styliform, mandible thin with two basal and two apical teeth. Chaetotaxy of antennae as in Table 3c. Apical vesicle distinct trilobed. S–chaetae of ant.IV of medium length and moderately thickened. Chaetotaxy of head as in Table 3a, b, and Fig. 10. Chaetae D and E free. Tubercles Cl and Af separate. Tubercle Af on head longer than tubercles Oc. Tubercle Dl with five chaetae, chaeta Dl3 absent. Tubercle (L+So) with eight chaetae, chaetae So2 and L3 absent (Fig. 10). Elementary tubercle BE absent. Chaeta A shorter than B.

Table 3a.

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Chaetotaxy of Endonura persica sp. n.: Cephalic chaetotaxy–dorsal side.

Tubercle	Number of chaetae	Types of chaetae	Names of chaetae
Cl	4	Ml Mc	F G
Af	10	Ml Mc Mc or Mcc	B A, C, E D
Oc	3	Ml Mc mi	Ocm Ocp Oca
Di	2	Ml Mcc	Di1 Di2
De	2	Ml Mcc	De1 De2
Dl	5	Ml Mc or Mcc	Dl1, Dl5 Dl2, Dl4, Dl6
(L+So)	8	Ml Mc me or mi	L1, L4, So1 L2 So3–6

Table 3b.

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Chaetotaxy of Endonura persica sp. n.: Cephalic chaetotaxy–ventral side.

Group	Number of chaetae
Vi	6
Vea	3–4
Vem	3
Vep	4
labium	11, 0×

Table 3c.

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Chaetotaxy of Endonura persica sp. n.: Chaetotaxy of antennae.

Segment, Group	Number of chaetae	Segment, Group	Number of chaetaev adult
I	7	IV ap	or, 8 S, i, 12 mou, 6 brs, 2 iv
II	12–14
III ve	5 sensilla AO III
III ve	5		8 bs, 5 miA
vc	4	ca	2 bs, 3 miA
vi	4	cm	3 bs, 1 miA
d	5	cp	8 miA, 1 brs

Thorax, abdomen, legs. Body s–chaetae fine and smooth, distinctly shorter than nearby macrochaetae (Fig. 12). Chaetotaxy of th. and abd. as in Table 3d and in Figs 10, 12. Tubercles Di on th.I differentiated or not. Chaetae De2 on th. II–III and De3 on th. III free. Chaetae De3 on abd. I–III free (Fig. 12). The line of chaetae De1–chaeta s parallel to the dorsomedian line on abd. I–III. Furca rudimentary without microchaetae. Tubercles Di on abd. V fused, with chaetae Di2 as Mcc and Di3 as mi (Fig. 12). Chaetae Vl on abd. V present. Cryptopygy slightly developed. Chaetotaxy of legs as in Table 3d. Tibiotarsi with rather long chaetae B4 and B5. Claw with inner tooth (Fig. 13).

Table 3d.

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Chaetotaxy of Endonura persica sp. n.: Postcephalic chaetotaxy.

	Di	Terga De	Dl	L	Scx2	Cx	Legs 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: 5–6	Ve1 -	present
abd. III	2	3+s	2	4	Vel: 5			Fu:5–10me	0 mi
abd. IV	2	2+s	3	6–7	Vel: 4	Vec: 2	Vei: 2	Vl: 4
abd. V	(3+3)	8+s			Ag: 3			Vl: 1	L’: 1
abd. VI		7			Ve: 13–14			An: 2mi

Remarks

In general appearance and presence of inner tooth on claw, characters rarely observed within the genus, E. persica sp. n. strongly resembles to E. dentifera Smolis et al. 2007 (described from Crimea). However, the new species can be reliably separated from Crimean species with the following characters: number of chaetae Dl on head (persica sp. n. five, dentifera six), number of chaetae (L+So) on head (persica sp. n. eight, dentifera ten), presence/absence of tubercles Di on the first thoracic segment (persica sp. n. present, dentifera absent) and number of chaetae L of abd. IV (persica sp. n. 6–7 chaetae, dentifera 8–9).

Key to the genus Endonura

In 1982, Deharveng, in his PhD thesis, elevated Endonura to the generic level and prepared a key to the genus that comprised 23 species. Nowadays, including the taxa described herein, the genus contains 40 members and is the second largest of the tribe Neanurini, after Deutonura Cassagnau, 1979. Moreover, after the publication of Deharveng’s paper (date), a few species were redescribed and one taxon was synonymised (Smolis and Kaprus’ 2003, Smolis 2008, Smolis et al. 2007, 2011). Considering these facts, the preparation of an updated key to all species of the genus seemed to be highly recommended.

1	Head with fusion of tubercles Af and Cl	2
–	Head with separation of tubercles Af and Cl	7
2.	Chaeta O on head present	3
	Chaeta O on head absent	4
3	Tubercles Di on th. I present and fused with De, tubercle (Di+Dl+L) on abd. V with nine chaetae	E. poinsotae Deharveng, 1980 (France, Corsica)
–	Tubercles Di on th. I absent, tubercle (Di+Dl+L) on abd. V with seven chaetae	E. ichnusae Dallai, 1983 (Italy, Sardinia)
4	Tubercles De on abd. I–III with four chaetae	5
–	Tubercles De on abd. I–III with three chaetae	E. granulata (Cassagnau & Delamare Deboutteville, 1955) (Lebanon)
5	Tubercles Di and De on th. I fused, cryptopygy strongly developed	E. gladiirostra Smolis & Kaprus’, 2003 (Israel)
–	Tubercles Di and De on th. I separate, cryptopygy absent or weakly developed	6
6	Chaeta E on head present, Tubercle Dl on head with four chaetae	E. tyrrhenica Dallai, 1983 (Italy, Sardinia)
–	Chaeta E on head absent, Tubercle Dl on head with six chaetae	E. pejai Deharveng, 1980 (France, Corsica)
7	Tubercle Af on head equal or shorter than tubercles Oc	8
–	Tubercle Af on head longer than tubercles Oc	9
8	Labrum with ventral sclerifications ogival and without prelabral chaetae	E. gracilirostris Smolis et al., 2007 (Crimea, Moldova)
–	Labrum with ventral sclerifications nonogival and with prelabral chaetae	E. taurica (Stach, 1951) (Crimea)
9	Chaeta O on head present	10
–	Chaeta O on head absent	26
10	Eyes completely absent	11
–	Eyes present	12
11	Tubercles Di on th. I present	E. arbasensis Deharveng, 1979 (France, Spain)
–	Tubercles Di on th. I absent	E. caeca (Gisin, 1963) (Bosnia and Herzegovina)
12	Anterior eye present and located outside tubercle Oc	E. asiatica Smolis et al., 2011 (Kyrgyzstan)
–	Anterior eye present or absent, if present located within tubercle Oc	13
13	Anterior eye present	14
–	anterior eye absent	E. immaculata Deharveng, 1980 (France, Corsica)
14	Claw with inner tooth, tibiotarsi with long chaetae B4 and B5	15
–	Claw without tooth, tibiotarsi with short chaetae B4 and B5	16
15	Tubercle Dl on head with three chaetae, tubercles Di on th. II–III with two chaetae	E. tetrophtalma (Stach, 1929) (Hungary)
	Tubercle Dl on head with five chaetae, tubercles Di on th. II–III with three chaetae	E. lusatica (Dunger, 1966) (Germany, Poland, Ukraine)
16	Chaeta E on head absent	17
–	Chaeta E on head present	18
17	Tubercle Cl on head with chaetae D, elementary tubercle DF present	E. colorata (Gama, 1964) (Portugal)
–	Tubercle Cl on head without chaetae D, elementary tubercle DF absent	E. cantabrica (Deharveng, 1979) (Spain)
18	Tubercle Dl on head with six chaetae	19
–	Tubercle Dl on head with less number of chaetae	25
19	Tubercles Di on head present	20
–	Tubercles Di on head absent	E. dalensi Deharveng, 1979 (Andorra, France, Spain, Italy)
20	Body white	21
–	Body blue or bluish–grey	22
21.	Tubercle (L+So) on head with nine chaetae, macrochaetae thin and pointed	E. deharvengi Cassagnau & Péja, 1979 (Greece)
–	Tubercle (L+So) on head with eight chaetae, macrochaetae thickened and blunt	E. levantica Smolis et al., 2011 (Israel)
22	Tubercle De on th. III with two ordinary chaetae	E. gladiolifer (Cassagnau, 1954) (Algeria, Spain)
–	Tubercle De on th. III with three ordinary chaetae	23
23	Tubercle Cl on head with chaetae D, furca rudimentary with microchaetae	E. alavensis Pozo & Simon, 1982 (Spain)
–	Tubercle Cl on head without chaetae D, furca rudimentary without microchaetae	24
24	Tubercle (L+So) on head with nine chaetae, free chaeta L on abd. IV present	E. quadriseta Cassagnau & Péja, 1979 (Greece, Turkey, Crimea)
–	Tubercle (L+So) on head with ten chaetae, free chaeta L on abd. IV absent	E. reticulata (Axelson, 1905) (Finland; Russia; Sweden; United States, Alaska)
25	Tubercle Dl on head with four chaetae, tubercles Di and De on th. I separate	E. occidentalis (Deharveng, 1979) (Spain)
–	Tubercle Dl on head with three chaetae, tubercles Di and De on th. I fused	E. cretensis (Ellis, 1976) (Greece, Israel)
26	Cryptopygy strong and complete, tubercles of abd. VI invisible in dorsal view	E. ludovicae (Denis, 1948) (France, Corsica)
–	Cryptopygy absent or weak, tubercles of abd. VI well or partially visible in dorsal view	27
27	Body bluish–grey	28
–	Body white	30
28	Claw with inner tooth, labrum chaetotaxy 4/2, 4	29
–	Claw without inner tooth, labrum chaetotaxy 0/2, 2	E. ceratolabralis sp. n. (Iran)
29	Tubercle Dl on head with five chaetae, tubercles Di on th. I present	E. persica sp. n. (Iran)
–	Tubercle Dl on head with six chaetae, tubercles Di on th. I absent	E. dentifera Smolis et al., 2007 (Crimea)
30	Chaeta C on head absent	31
–	Chaeta C on head present	33
31	Macrochaetae Di1 on abd. V distinctly thickened and club–like	E. baculifer Deharveng, 1979 (Portugal)
–	Macrochaetae Di1 on abd. V slightly thickened and cylindrical	32
32	Eyes present, tubercles Di on th. I present	E. transcaucasica (Stach, 1951) (Georgia)
–	Eyes absent, tubercles Di on th. I absent	E. carpatica Smolis, 2006 (Poland)
33	Tubercle Cl on head with chaetae D, elementary tubercle DF present	34
–	Tubercle Cl on head without chaetae D, elementary tubercle DF absent	35
34	Chaeta E on head present, tubercle Dl on head with six chaetae	E. tartaginenis Deharveng, 1980 (France, Corsica)
–	Chaeta E on head absent, tubercle Dl on head with five chaetae	E. dichaeta sp. n. (Iran)
35	Chaeta E on head present	E. urotuberculata Pomorski & Skarżyński, 2000 (Bulgaria)
–	Chaeta E on head absent	36
36	Chaeta L4 on head free, eyes absent or present unpigmented	37
–	Chaeta L4 within tubercle (L+So), eyes present and pigmented	39
37	Abd. V with two tubercles	E. incolorata (Stach, 1951) (Poland, Ukraine, Romania)
–	Abd. V with three tubercles	38
38	Abd. IV with eight tubercles, macrochaetae Ml relatively short	E. tatricola (Stach, 1951) (Poland, Slovakia)
–	Abd. IV with five tubercles, macrochaetae Ml long	E. dudichi (Loksa, 1967) (Hungary, Poland, Slovakia)
39	Tubercle Dl on head with six chaetae, tubercle L on abd. III with three chaetae	E. centaurea Cassagnau & Péja, 1979 (Greece)
–	Tubercle Dl on head with five chaetae, tubercle L on abd. III with four chaetae	E. saleri Fanciulli & Dallai, 2008 (Italy)

Discussion

Considering the data presented here and those obtained from the literature (Mayvan et al. 2015, Shayanmehr et al. 2013, Smolis et al. 2012), Neanurinae fauna of Iran comprises ten species and seven genera: Bilobella aurantiaca (Caroli, 1912), Cryptonura persica Smolis et al., 2012, C. maxima Smolis et al., 2012, Deutonura decolorata (Gama & Gisin, 1964) (Gisin 1964), Endonura ceratolabralis sp. n., E. dichaeta sp. n., E. persica, sp. n., Neanura muscorum (Templeton, 1835), Persanura hyrcanica Mayvan et al. 2015, Thaumanura echinata (Kos, 1940). It should be noted, however, that until now only the western part of Iran has been roughly studied. Although future research may change the present picture of the subfamily diversity in the studied country and region, some preliminary conclusions can be drawn. The first is related to the higher systematic pattern and composition of Neanurinae of Iran. This fauna consists almost exclusively of members of the tribe Neanurini, the most diverse and dominant among Neanurinae in the western Palaearctic. To date, none of the Lobellini and Paranurini genera have been found in Iran, although they are numerous and widely distributed in south, south-east and east Asia. The second conclusion seems to be more expected, Endonura species from Iran resemble those known from south-east Europe. It suggests their close affinity and the historical connection between these faunas. The third conclusion sheds light on the distribution and the history of this genus. Most Endonura species were recorded from Mediterranean and temperate zones of Europe, where they live predominantly in forests. It is worth saying that the greatest diversity of the genus through the continent is more or less correlated to the areas of land that have never been subjected to glaciations. Till now, the occurrence of only a few species is documented outside Europe, especially in the Middle East (Smolis and Kaprus 2003, 2009; Smolis et al. 2011). The recent and present discoveries of Endonura species in Kyrgyzstan (Smolis et al. 2011) and Iran significantly expand the list of species and also our knowledge on the genus. Undoubtedly, diverse forest habitats of the coastal and montane regions of Iran and adjacent countries hide a rich fauna of Neanurinae. We therefore hope that a more comprehensive study in the future will allow us to present a better picture of the distribution of Endonura in Iran and the near East.

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