New species of Hexabdella and Neomolgus (Acari ... - Biotaxa

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Jan 29, 2016 - http://zoobank.org/urn:lsid:zoobank.org:pub:9AA85C82-9BBF-4993-80BD-B83EF21DAF53. New species of Hexabdella and Neomolgus (Acari: ...
Zootaxa 4072 (2): 291–300 http://www.mapress.com/j/zt/ Copyright © 2016 Magnolia Press

Article

ISSN 1175-5326 (print edition)

ZOOTAXA

ISSN 1175-5334 (online edition)

http://doi.org/10.11646/zootaxa.4072.2.10 http://zoobank.org/urn:lsid:zoobank.org:pub:9AA85C82-9BBF-4993-80BD-B83EF21DAF53

New species of Hexabdella and Neomolgus (Acari: Prostigmata: Bdellidae) from Iran AMIR HOSSEIN EGHBALIAN1, MOHAMMAD KHANJANI2,5, MOHAMMAD HASSAN SAFARALIZADEH1 & EDWARD A. UECKERMANN3,4 1

Department of Plant Protection, College of Agriculture, Urmia University, Uromia, Iran. E-mails: [email protected], [email protected] 2 Department of Plant Protection, College of Agriculture, Bu–Ali Sina University, Hamedan, Iran. E-mail: [email protected] 3 ARC–Plant Protection Research Institute, Private Bag X134, Queenswood, Pretoria 0121, South Africa. E-mail: [email protected] 4 School of Biological Sciences/Zoology, North–West University, Potchefstroom Campus 2520, South Africa 5 Corresponding author. E-mail: [email protected]

Abstract A new species of Hexabdella Van der Schyff, Theron & Ueckermann (Acariformes: Bdellidae), H. quercusi Eghbalian, Khanjani, Safaralizadeh, Ueckermann sp. nov., and a new species of Neomolgus Oudemans, N. iraniensis Eghbalian, Khanjani, Safaralizadeh, Ueckermann sp. nov., are described and illustrated. A key is provided for adult females of all known species of Hexabdella, as well as for adult females of Neomolgus from Asian and neighbouring countries, including Iran, Japan and the Siberian region of Russia. All specimens were collected from soil and litter under oak trees, Quercus brantii Lindley (Fagaceae), or from soil and litter under wild almond, Amygdalus scoparia L. (Rosaceae), from western Iran. Key words: Bdellinae, predatory mite, Quercus, Neomolgus, Hexabdella, Iran

Introduction Members of the family Bdellidae Dugès are predators of small arthropods such as soft bodied insects, collembolans and mites (Gerson et al. 2003; Atyeo 1960). One of the main subfamilies is Bdellinae, which is characterized by having six ventral subcapitular setae (vh1–vh6) in adults and trichobothria absent on tibia II. Van der Schyff et al. (2004) erected Hexabdella (type species H. denheyeri) to accommodate five newly described species, which can be recognized by lacking a trichobothrium on tarsus IV in contrast to Bdella Latreille. Van der Schyff et al. (2004) transferred Bdella mexicana (Baker & Block) to Hexabdella and Hernandes (2013) transferred Bdella brevitarsis Banks (in Tothill, 1919), a nomen nudum based on unpublished material, to this genus after a revision of the type specimens of Bdellidae in Banks’ collection. Hexabdella currently contains nine species, of which five (H. singula, H. denheyeri, H. maraugia, H. miranda and H. unusoculata) were described from South Africa (Van der Schyff et al. 2004); one species (H. brevitarsis) from Canada (Hernandes 2013); two species (H. cinquaginta and H. mexicana) from South America (Baker & Balock 1944; Hernandes 2013); and H. persiaensis Paktinat Saeej et al., 2014 from Iran. Oudemans (1937) transferred Bdella capillata (Berlese, 1981), as type species for his new genus Neomolgus, which can be distinguished by having more than two setae (6-21) on each chelicera, in contrast with other genera of Odontoscirinae (Atyeo, 1960). N. pygmaeus Shiba (1969) and N. longipalpus Kuznetzov (1984) were described and illustrated from Japan and the Siberian region of Russia, respectively. In this paper, two new species of these genera are described, illustrated from Western Iran. A key to the adult females is provided for all known species of Hexabdella, and for adult females of Neomolgus of the Asian region and the Siberian part of Russia [Iran by Ostovan & Kamali (1995); Japan by Ehara (1961) and Shiba (1969); Siberian region by Thor (1931); Uzbekistan by Kuznetsove and Barilo (1984) and Kuznetsov (1984)]. Accepted by O. Seeman: 9 Dec. 2015; published: 29 Jan. 2016

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Material and methods Specimens were extracted from soil and litter collected under oak trees, Quercus brantii Lindley (Fagaceae), in the vicinity of Baneh, and wild almond, Amygdalus scoparia L. (Rosaceae), in the vicinity of Kamyaran (Kermanshah Province) Iran, by using Tullgren funnels. These collection sites are located in the Zagrous mountain range, mainly covered by oak trees and wild almond. Specimens were mounted directly on slides in Hoyer’s medium. The slides were oven-dried at 50°C for one week, ringed with nail polish, and examined under an Olympus BX51 phase contrast microscope. Drawings were made by means of a camera lucida. Notations of the idiosomal and leg setae follow those of Kethley (1990) and Den Heyer (1981), respectively. Measurements for the holotypes are given in micrometres (μm), with the range of measurements for the paratypes shown in parentheses. The body length of all specimens was measured from the apex of the hypostome to the posterior margin of the idiosoma, and body width at the level of setae c2. Abbreviations of setae in this study are as follows. Propodosomal setae: internal verticals (vi), external verticals (ve), internal scapular (sci), external scapular (sce). Opisthosomal setae: internal humeral (c1), external humeral (c2), internal dorsal (d1), internal lumbal (e1), internal sacral (f1), external sacral (f2), internal clunal (h1), external clunal (h2). Anal region: postanal (ps1), genital region: aggenital setae (ag), genital setae (g), subcapitular setae (vh1–vh6). Leg setae: solenidia (w, e , f and s), trichobothrium (tr), ventral end seta (VES), dorsal end seta (DES). Subcapitulum: adoral setae (ad) and anteroventral subcapitular setae (avs) adapted from Fisher et al. (2011).

Family Bdellidae Dugès, 1834 Subfamily Bdellinae Grandjean, 1938 Genus Hexabdella Van der Schyff et al. 2004 Type species: Hexabdella denheyeri Van der Schyff et al. 2004

Hexabdella quercusi Eghbalian, Khanjani, Safaralizadeh and Ueckermann sp. nov. (Figs. 1–10) Diagnosis. Two pairs of eyes present, fixed cheliceral digit as long as movable digit, chelicerae striated, coxa II with three setae, basifemora II and IV with eight and four setae respectively, genu IV with 5 (+1s) setae, solenidotaxy of tibiae I–IV 3–2–1–0. Female (n= 4). Total body length (including gnathosoma from apex of hypostome to posterior margin of idiosoma) 602 (608–628), body length (excluding gnathosoma) 446 (455–470); width 270 (230–273). Dorsum (Fig. 1). Prodorsum between setae vi and sci with irregularly broken longitudinal striae; prodorsum with two pairs of eyes, diameters of anterior lateral eye 8 (7–8) and posterior lateral eye 5 (4–5). Striae anterior to internal verticals (vi) broken and transverse; all dorsal setae finely serrate; hysterosomal setae not extending to bases of setae next behind (except seta h1). Dorsum of hysterosoma with fine, continuous and transverse striae between setae (c1, d1, e1, f1, h1 and h2), but striae obliquely longitudinal between setae c1–2; seta sce the longest and h2 and f2 the shortest; hysterosomal region with three cupules (ia, im and ip) at level of setae d, e and f1 (Fig. 1). Measurements of dorsal setae as follows: vi 98 (88–93), ve 41 (38–39), sce 69 (65–68), sci 130, c1 47 (50–51), c2 50 (40–45), d1 36 (34–38), e1 35 (33–38), f1 41 (35–36), f2 30 (28–30), h1 47 (43–45), h2 30 (27–30). Distance between dorsal setae: vi–vi 78 (73–75); ve–ve 113 (110–113); vi–ve 33 (30–33); ve–sce 40 (38–40); sci–sci 93 (83– 87); sce–sci 9 (8–10); sci–c1 72 (68–69); sci–c2 102 (103–105); c1–c1 77 (77–78); c1–c2 75 (67–70); c1–d1 60 (58–64); d1–d1 64 (65–68); d1–e1 55 (48–51); e1–e1 80 (70–83); e1–f1 55 (50–54); f1–f1 48 (43–49); f1–f2 33 (33–38); h1–h 24 (27–28); h1–h2 28 (20–27); h2–h2 80 (70–76). Ratio: vi/ve 2.30 (2.40); vi/sce 0.75 (0.68–0.72); sci/sci–sci 1.39 (1.57–1.48); vi/vi–vi 0.1.30 (1.30–1.70), c1/c1–c1 0.61 (0.64–0.65), d1/d1–d1 0.6 (0.50–0.58), e1/ e1–e1 0.44 (0.4–0.54), f1/f1–f1 0.85 (0.73–0.81), h1/h1–h1 1.96 (1.59–1.61), h2/h2–h2 0.34 (0.4–0.43), h1/h2 1.74 (1.43–1.50), c1–c1: d1–d1: e1–e1: f1–f1: 1.60 (1.60–1.80): 1.33 (1.39–1.51): 1.67 (1.63–1.69): 1.00 (1.00).

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FIGURES 1–6: Hexabdella quercusi Eghbalian, Khanjani, Safaralizadeh and Ueckermann sp. nov. (Female): 1. Dorsum of idiosoma; 2. Prodorsal seta vi; 3. Prodorsal seta sce; 4. Gnathosoma; 5. Venter of idiosoma; 6. Ovipositor.

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FIGURES 7–10. Hexabdella quercusi Eghbalian, Khanjani, Safaralizadeh and Ueckermann sp. nov. (Female): 7. Leg I; 8. Leg II; 9. Leg III; 10. Leg IV.

Gnathosoma (Fig. 4). Subcapitulum 155 (148–165) long, width at base 75 (66–78); base of gnathosoma with transverse striae; palp: five–segmented, tibiotarsus with three setae + one solenidion + two long end setae, DES and VES 83 (83) and 108 (106–108) respectively; genu with four setae; telofemur with one seta; basifemur with five setae; trochanter without setae; measurements of palp segments as follows: trochanter 11 (10–12), basifemur 58 (50–53), telofemur 22 (22–23), genu 12 (13–15), tibiotarsus 35 (35–38). Subcapitulum with six pairs of long ventral subcapitular setae (vh1–6), distal pair (vh6) 36 (33–34) about 1.5x length of proximal pair (vh1) 21 (19–21);

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two pairs of short adoral setae near tip of hypostome, avs 10 (11) and ad 9 (8–10). Chelicerae 143 (138–141) long, width 42 (36–38), finely striated and with two dorsal setae (ch1–2), proximal seta 35 (33–35), and distal seta 50 (46–48), respectively, movable chela with one small tooth and fixed chela smooth and attenuate, equal in length to movable chela; distal seta (ch1) short, not extending to base of chela (Fig. 4). Venter (Fig. 5). Striae between coxae I–II and III– IV longitudinal and between coxae II–III finely broken and oblique. One pair of setae between coxae IV. Aggenital region with nine pairs of setae (ag1–9) by genital valves each with nine pairs of setae (g1–9) (Fig. 5); anal region surrounded with oblique continuous striae and three pairs of smooth setae (ps1–3): ps1 33 (33–38), ps2 23 (23–25) and ps3 16 (16–18) long (Fig. 5). Ovipositor (Fig. 6). Ovipositor as indicated in Figure 6; tube like, with 18 smooth setae. Legs (Figs. 7–10). Measurements of legs I-IV: I 255 (238–243), II 231 (215–218), III 272 (250–253), IV 343 (315–338). Setal formulae of leg segments (solenidia in parentheses): coxae I–IV: 5–3–5 (right leg of holotype with 6)–2; trochanters I–IV: 1–1–2–1; basifemora I–IV: 8–8–7–4; telofemora I–IV: 5–5–5–5; genua I–IV: 4ts,1s–4ts,1s– 4ts,1s–5ts,1s; tibiae I–IV: 6ts,3f,1tr–7ts,1f,1bls–7ts,1f–6ts,1tr; tarsi I–IV: 18ts,4w, 1e–16ts,2w,1e–17ts,1tr–15ts,1w (Figs. 7–10). Male and immatures: Unknown. Habitat. Litter under oak trees. Distribution. Only known from Western Iran. Remarks. Hexabdella quercusi Eghbalian, Khanjani, Safaralizadeh and Ueckermann sp. nov. is closely related to H. mexicana (Baker & Balock, 1944), redescribed by Atyeo (1960), by having: tibiae I, IV and tarsus III, each with one trichobothrium; opisthosomal setae not branched distally; two pairs of eyes; palp basifemur with five setae; solenidotaxy of tibiae I–IV 3–2–1–0; it however differs in having: 1) striae between setae vi and sci irregularly broken and longitudinal in the former instead of finely and regularly broken in the latter; 2) fixed digit of equal length as movable digit, instead of two thirds the length of movable digit; 3) genital valve with nine pairs of setae in H. quercusi sp. nov. opposed to eight pairs of setae in H. mexicana; 4) coxa II with three setae in new species while four setae in H. mexicana; 5) subcapitular setae smooth instead of finely serrate; 6) basifemora II and IV with eight and four setae respectively versus seven and five; 7) telofemur IV with five setae instead of four setae; 8) genu IV with 5 (+1σ) setae in H. quercusi whereas 4 (+1σ) in the H. mexicana; 9) tibiae II–IV: 7ts–7ts–6ts vs. 5ts–5ts–5ts; 10) tarsi I–IV 18ts–16ts–17ts–15ts opposed to tarsi I–IV: 14ts–10ts–9ts–8ts. The new species also resembles H. persiaensis Paktinat Saeej, Bagheri, Saboori & Ueckermann, 2014, in having mostly the same leg chaetotaxy, smooth dorsal idiosomal setae and fixed digit of equal length as the movable digit; however it differs by: 1) palpal basifemur with five setae in the former while six setae in the latter; 2) basifemur III with seven setae in the new species whereas six setae in H. persiaensis; 3) genital valve with nine pairs of setae in H. quercusi sp. nov. while eight pairs of setae in H. persiaensis; 4) aggenital region with nine pairs of setae instead of eight; 5) marginal region between dorsal setae d1 and e1 with continuous striae in the former but fine broken striae in the latter; 6) marginal regions between legs II and III with broken striae in new species whereas continuous striae in H. persiaensis; 7) aggenital setae ag 1–5 covered in continuous striae in new species but all aggenital seta, ornamented, on broken striae in H. persiaensis; 8) tarsi I, II and IV with 18, 17 and 15 setae vs 19(20), 16 and 16 setae. Etymology. The specific epithet is derived from Quercus, due to the oak-dominated forest where the new species was collected. Type material. The holotype and three paratype females were collected from soil and litter under oak trees, Quercus brantii Lindl, (Fagaceae), Baneh, Kurdistan Province, Iran, (35° 59' N, 46° 53' E, a.s.l. 1540 m), 17 January 2013, Col. Amir Hossein Eghbalian. Holotype and two paratype females (EHBH15010 and EPBH15011– 12, respectively) are deposited in the Mites Collection of the Acarology Laboratory, University of Bu–Ali Sina, Hamedan, Iran, and one paratype female (slide specimen) will be deposited in the Mites Collection of Arachnida, Plant Protection Research Institute, Pretoria, South Africa.

Subfamily Odontoscirinae Grandjean, 1938 Genus Neomolgus Oudemans, 1937 Type species: Bdella capillata Berlese 1981

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Neomolgus iraniensis Eghbalian, Khanjani, Safaralizadeh and Ueckermann sp. nov. (Figs. 11–19) Diagnosis. Each chelicera with 21 setae, palp basifemur with seven setae, palp tibiotarsus with 14 setae, genital plate with ten pairs of setae, and aggenital region with three pairs of setae. Female (n= 3). Total body length (including gnathosoma from apex of hypostome to posterior margin of idiosoma) 2025 (1750–1813), body length (excluding gnathosoma) 1500 (1275–1350); width 1000 (862–950). Dorsum (Fig. 11). Dorsum with irregularly broken striae; prodorsum with two pairs of eyes, diameters of anterior eye 50 (40–45) and posterolateral eye 40 (33–38). Striae between setae vi transverse; dorsal setae (c1-h2) smooth, prodorsal setae vi, sci and sce broken on three collected specimens, eyes separated by distance ~ 2 (2–2.4) times diameter of anterior pair eyes, interval between eyes with oblique and longitudinal striae; hysterosomal setae not extending to bases of setae next behind. Between hysterosomal setae (c1, d1, e1) fine, longitudinal and transversal broken striae; between setae f1 longitudinal and setae h1-2 transverse striae; between setae c1–2 with oblique striae; hysterosomal region with three cupules (ia, im and ip) at level of setae d1, e1 and f1 (Fig. 11). Measurements of dorsal setae: vi, sce and sci broken, c1 70 (70–90), c2 65 (65–88), d1 55 (55–88), e1 63 (63–88), f1 58 (75), f2 53 (53–75), h1 63 (63–80), h2 63 (63–88). Distance between dorsal setae: vi–vi 84 (63–69); vi–sce 200 (200–240); sci–sci 185 (185–200); sce–sci 35 (38–48); sce–sce 200 (200); sci–c1 270 (277–288); sci–c2 338 (200–350); c1–c1 375 (180–375); c1–c2 136 (120–140); c1–d1 195 (175); d1–d1 312 (185); d1–e1 200 (185); e1– e1 195 (175); e1–f1 140 (120); f1–f1 64 (80); f1–f2 80 (88); f2–f2 176 (170); f1–h1176 (170); h1–h1 65 (63); h1–h2 100 (123); h2–h2 128 (138). Gnathosoma (Fig. 12). Subcapitulum 425 (462–500) long, width at base 225 (200–250); base of subcapitulum with broken and transverse striae, hypostome smooth; palp five–segmented, palp tibiotarsus with 14 setae [DES and VES 203 (215) and 173 (200) respectively]; genu with four setae; telofemur with one seta; basifemur with seven setae; trochanter lacking setae; measurements of palp segments: trochanter 15 (13), basifemur 290 (355), telofemur 45 (55), genu 55 (53), tibiotarsus 305 (310). Subcapitulum with six pairs of setae (vh1–6), distal pair (vh6) 75(88) more than twice length of proximal pair (vh1) 33 (35); two pairs of short adoral setae near tip of hypostome, avs 28 (30) and ad 19 (21). Cheliceral length 443 (425–433), width 95 (100–105), with 21 smooth setae (ch1–21), distal setae [ch1] 98 (75) and proximal setae [ch21] 38 (50), and; movable digit longer than fixed digit, movable chelae with two teeth and fixed digit smooth; distal seta (ch1) long extending to base of chela (Fig. 12). Venter (Fig. 13). Aggenital region with three pairs of setae (ag1–3), each genital plate with ten pairs of setae (g1–10) (Fig. 13); anal region with three pairs of smooth setae (ps1–3): ps1 65, ps2 63 and ps3 broken. Ovipositor (Figs. 14-15) Ovipositor with 16 smooth setae and ovipositor gland as depicted in Figs 14-15. Legs (Figs. 16-19). Measurements of leg segments: I 1075 (1050), II 1050 (1115), III 1150 (1125), IV 1350 (1235). Setal formulae of leg segments: coxae I–IV: 5–3–4–2; trochanters I–IV: 1–1–1–1; basifemora I–IV: 11–10– 10–6; telofemora I–IV: 7–7–6–6; genua I–IV: 7ts,11s–6ts,10s–5ts–7ts,3s; tibiae I–16ts,16f,1e,1tr–15ts,5f,1tr– 17ts,1f–15ts, 1f,1tr; tarsi I–IV: 40ts, 1dt,9w, 1e–41ts,1dt, 3w–41ts,1tr–45ts,1tr (Figs. 16-19). Remarks. Neomolgus iraniensis Eghbalian, Khanjani, Safaralizadeh and Ueckermann sp. nov. is closely related to N. pygmaeus Shiba (1969) in having four setae on the papal genu, coxa I with five setae and telofemora II-IV with 7–6–6 setae but differs from the latter in having: 1) Dorsal setae smooth in N. iraniensis opposed to plumose in N. pygmaeus; 2) each chelicera with 21 setae in new species while eight setae in N. pygmaeus; 3) papal basifemur with seven setae in the former while four setae in the latter; 4) palp tibiotarsus with 14 setae in the new species opposed to nine setae in the latter; 5) each genital plate with ten setae vs eight setae; 6) Coxae II-IV with 3– 4–2 vs 4–3–3. Neomolgus iraniensis Eghbalian, Khanjani, Safaralizadeh and Ueckermann sp. nov. is closely related to N. longipalpus Kuznetsov (1984) in having 7–6–6 setae on telofemora II-IV and coxa III-IV with 4–2 setae but differs from the latter in having: 1) each chelicera with 21 setae in the new species instead of 16 (17) setae in N. longipalpus; 2) palp basifemur with seven setae versus 19 setae; 3) palp genu with four setae versus ten setae; 4) palp tibiotarsus with 14 setae versus 27 setae; 5) coxa I-II with 5–3 setae versus 6–4 setae; 6) basifemura I-IV with 11–10–10–6 setae versus 22–17–15–8 setae; 7) telofemur I with seven setae instead of eight setae. Etymology. The specific epithet is derived from Iran, due to the country where the new species was collected.

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FIGURES 11–15. Neomolgus iraniensis Eghbalian, Khanjani, Safaralizadeh and Ueckermann sp. nov. (Female): 11. Dorsum of idiosoma; 12. Gnathosoma; 13. Genital plate; 14. Ovipositor; 15. Ovipositor gland.

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FIGURES 16–19. Neomolgus iraniensis Eghbalian, Khanjani, Safaralizadeh and Ueckermann sp. nov., (Female): 16. Leg I; 17. Leg II; 18. Leg III; 19. Leg IV.

Type material. The holotype and two paratype females were collected from soil and litter under wild almond trees, Amygdalus scoparia L. (Rosaceae), "Sirvan river bank", located in Palangan village, Kamyaran, Kermanshah Province, Iran, (35° 03.7' N, 46° 35.97' E, a.s.l. 864 m), 14 IV 2013, by Amir Hossein Eghbalian. The holotype and

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one paratype females (EHBN15014 and EPBH15015, respectively) are deposited in the Mites Collection of the Acarology Laboratory, University of Bu–Ali Sina, Hamedan, Iran, and one paratype female slide will be deposited in the National Collection of Arachnida, Plant Protection Research Institute, Pretoria, South Africa.

Key to adult female Hexabdella of the world (modified from Van der Schyff et al. 2004) and to Neomolgus of Asia (Iran, Japan, Uzbekistan) and the Siberian region of Russia 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. -

Tibia II without trichobothria, each chelicera with two setae . . . . . . . . . . . . . . . . . . Bdellinae…Hexabdella Van der Schyff …2 Tibia II with trichobothria, each chelicera with more than 2 setae . . . . . . . . . . . Odontoscirinae …Neomolgus Oudemans…11 Opisthosomal setae distally branched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Opisthosomal setae smooth or slightly plumose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Dorsal striae coarsely to sparsely broken; seta ps1 smooth; coxa IV without macroseta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Dorsal striae finely broken; seta ps1 branched; coxa IV with a serrated macroseta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. denheyeri Van der Schyff, Theron & Ueckermann Marginal striae of hysterosoma with lobes, visible as dashed lines . . . . . . . . . . . . . . .H. cinquaginta Hernandes, Daud & Feres Marginal striae of hysterosoma smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. singula Van der Schyff, Theron & Ueckermann Eyes present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Eyes absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .H. maraugia Van der Schyff, Theron & Ueckermann Two pairs of eyes present; chelicerae striated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 One pair of eyes present; chelicerae smooth . . . . . . . . . . . . . . . . . . . . . .H. unusoculata Van der Schyff, Theron & Ueckermann Palp basifemur with five or six setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Palp basifemur with four setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. brevitarsis Hernandes Palp basifemur with five setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 Palp basifemur with six setae . . . . . . . . . . . . . . . . . . . . . . . . . . H. persiaensis Paktinat Saeej, Bagheri, Saboori & Ueckermann Solenidotaxy of tibiae I–IV 3–2–1–0 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 Solenidotaxy of tibiae I– IV 1–1–1–0 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. miranda Van der Schyff, Theron & Ueckermann Coxa II with four setae; basifemura II and IV with seven and fivesetae; genu IV with 4 setae (1s); fixed digit two thirds the length of movable digit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .H. mexicana (Baker & Balock) Coxa II with three setae; basifemura II and IV with eight and four setae; genu IV with five setae (1s); fixed digit as long as movable digit. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. quercusi Eghbalian, Khanjani, Safaralizadeh & Ueckermann sp. nov. Palp genu with four setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .12 Palp genu with ten setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. longipalpus Kuznetsov Palp tibiotarsus with 8–14 setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 Palp tibiotarsus with more than 20 setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..N. littoralis (L.) Chelicerae with 7–11 setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Chelicerae with 21 setae . . . . . . . . . . . . . . . . . . . . . . N. iraniensis Eghbalian, Khanjani, Safaralizadeh & Ueckermann sp. nov. Palp basifemur with four setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 Palp basifemur with five to seven setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 Palp tibiotarsus with eight setae (including end setae); chelicerae with seven setae . . . . . . . . . . . N. raptor Kuznetsov & Barilo Palp tibiotarsus with nine setae (including end setae); chelicerae with eight setae . . . . . . . . . . . . . . . . . . . . N. pygmaeus Shiba Prodorsum without distinct dorsal shield . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 Prodorsum with distinct dorsal shield . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. clypeatus (Thor) Chelicerae with nine setae; palp basifemur with 5–6 setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .N. pallipes (L.) Chelicerae with 11 setae; palp basifemur with 7 setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. capillatus (Kramer)

Acknowledgements This paper is a part of Ph. D thesis of the senior author which was financially supported by Vice Research of Urmia University, Iran.

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