(Acari: Cunaxidae) from Iran, with key to species of Lupaeus ... - BioOne

Systematic & Applied Acarology 21(6): 778–790 (2016) http://doi.org/10.11158/saa.21.6.5 Article

ISSN 1362-1971 (print) ISSN 2056-6069 (online)

http://zoobank.org/urn:lsid:zoobank.org:pub:6E4400CC-EC22-4A5E-83F7-FAA3825C9CD6

Two new species and eight new combinations of Pulaeini Berlese (Acari: Cunaxidae) from Iran, with key to species of Lupaeus and Pulaeus in the world SAEID PAKTINAT-SAEIJ1*, TATIANE M.M.G. DE CASTRO2, MOHAMMAD BAGHERI1, MICHAEL SKVARLA3 & GILBERTO J. DE MORAES4 1

Department of Plant Protection, Faculty of Agriculture, University of Maragheh, Maragheh, Iran. E-mails: [email protected]; [email protected] 2 Universidade Estadual de Roraima, Rorainopolis, Roraima, Brazil. E-mail: [email protected] 3 Department of Entomology, 319 AGRI Building, University of Arkansas, Fayetteville, Arkansas, 72701, USA. E-mail: [email protected] 4 CNPq researcher, Depto. de Entomologia e Acarologia, ESALQ-Universidade de São Paulo, 13418-900 Piracicaba, São Paulo, Brazil. E-mail: [email protected] * Corresponding author

Abstract Two new species of Pulaeini, Lupaeus damavandiani Paktinat-Saeij & Castro sp. nov., and Pulaeus aryani Paktinat-Saeij & Castro sp. nov., are described and illustrated from soil, humus and moss in the Mazandaran province, Iran. Also, the status of two genera are considered. Eight new combinations are proposed, namely L. akbari (Bashir & Afzal, 2009) comb. nov., L. anjumi (Bashir & Afzal, 2007) comb. nov., L. ferventis (Taj & Chaudhri, 1990) comb. nov., L. haurio (Taj & Chaudhri, 1991) comb. nov., L. payatopalpus (Corpuz-Raros, 1996) comb. nov., L. stultus (Taj & Chaudhri, 1991) comb. nov., L. walii (Bashir & Afzal, 2009) comb. nov. and Neocunaxoides neopectinatus (Shiba, 1978) comb. nov. P. razanensis Den Heyer, 2013 syn. nov. considered a junior synonym of P. krama (Chaudhri, 1977). Lupaeus clarae (Den Heyer, 1981b) is reported for the first time from Iran. Keys to the Lupaeus and Pulaeus species of the world are provided. Key words: Bdelloidea; Cunaxoidinae; predatory mites; taxonomy; Mazandaran Province

Introduction Bdelloidea comprises a moderately large assemblage of predatory species grouped in two cosmopolitan families: Bdellidae and Cunaxidae. Many species of Cunaxidae are active on plant foliage and prey on phytophagous insects and mites. Cunaxoidinae (e.g. Pulaeus Den Heyer, 1980a, Lupaeus Castro & Den Heyer, 2009, Cunaxoides Baker & Hoffmann, 1948 and Neocunaxoides Smiley, 1975) are cruise predators that search out arthropod prey but also take less active forms such as nemathodes (Walter & Kaplan 1991). These mites have relatively short palpi that are held parallel to their chelicerae as they search for prey (Walter et al. 2009). Up to data, Lupaeus iranensis Den Heyer, 2013, L. lectus Catro & Den Heyer, 2009, L. lenis (Corpuz-Raros, 1996), L. martini (Den Heyer, 1981b), L. sativae Den Heyer, 2013, L. subterraneus (Berlese, 1916), L. valentinae Sergeyenko, 2011, Pulaeus glebulentus Den Heyer, 1980a, P. krama (Chaudhri, 1977) and P. pectinatus (Ewing, 1909) have been previously reported from Iran (Kamali et al. 2001; Akbari et al. 2010; Jalaeian et al. 2011; Beyzavi & Ostovan 2012; Den Heyer et al. 2013; Paktinat-Saeij et al. 2014). In this work, two new species, Lupaeus damavandiani sp. nov. and Pulaeus aryani sp. nov., are described and Lupaeus clarae (Den Heyer, 1981b) is reported as a new record from Iran. 778 © Systematic & Applied Acarology Society

Material and methods Samples of soil and leaf litter were taken from Mazandaran Province. Mites were extracted using a Berlese-Tullgren funnel and specimens were collected in AGA solution (Smiley 1992). Collected specimens were cleared in Nesbitt’s fluid, mounted in Hoyer’s medium (Walter & Krantz 2009), and examined under an interference contrast microscope (Nikon, Eclipse i80) and a phase contrast microscope (Leica, DMLB) at the Department of Plant Protection, University of Maragheh, Iran, and at the Department of Entomology and Acarology, Escola Superior de Agricultura “Luiz de Queiroz” (ESALQ)-Universidade de São Paulo (USP), Brazil. Illustrations were made with the aid of a drawing tube attached to the phase contrast microscope. Measurements were taken with a graded ocular. The length of gnathosoma was measured from the base to the tip of the subcapitulum, the length of the idiosoma from the suture between the gnathosoma and idiosoma to the posterior margin of the idiosoma, the width of the idiosoma at its broadest level and the legs from the ventral insertion of coxae to the base of the pretarsi. The setal nomenclature of Den Heyer & Castro (2008) is followed for the idiosoma except for the propodosomal setae, which follows the notation given by Fisher et al. (2011) and legs follows that of Den Heyer (1981a). The abbreviation of setal names are as follows: Prodorsal setae: anterior trichobothria (at), posterior trichobothria (pt), lateral proterosomal setae (lps), median proterosomal setae (mps). Hysterosomal setae: internal humerals (c1), external humerals (c2), internal dorsals (d1), internal lumbals (e1), internal sacrals (f1), external sacrals (f2), internal clunals (h1), external clunals (h2). Anal region: postanals (ps); genital region: aggenital setae (ag), genital setae (g). Hypognathal setae (hg). Leg setae: attenuate (sharply) solenidion (asl), bluntpointed rod-like solenidion (bsl), trichobothria (T), simple tactile seta (sts), macroseta (ms), duplex setae (dxs), setae between such brackets indicate duplex condition ({}), famulus (fam), dorsoterminal solenidion (dtsl), terminal solenidion (tsl). All measurements are given in micrometers (μm) for the holotype and paratypes (in parentheses).

Results and discussion Tribe Pulaeini Den Heyer, 1980 Type genus: Pulaeus Den Heyer, 1980a

Key to genera of Pulaeini (For adult females; modified from Castro & Den Heyer 2009) 1. Setae f2 present; basifemora I–IV setal formula generally: 4-6-3-1 or 2 sts . . . . . . . . . . . . . . . . . . . . . . . . 2 – Setae f2 absent; basifemora I–IV setal formula generally: 3-5-2-0 sts. . . . . . Neocunaxoides Smiley, 1975 2 (1) Basifemur IV with one seta; famulus on distal half of tarsus I (subapical); tibiae III with transversely striated blunt solenidia; palp tibiotarsus with one or two pointed processes (one ventral, one median); g3 near outer margin or straight row . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lupaeus Castro & Den Heyer, 2009 – Basifemur IV with two setae; famulus on proximal half of tarsus I; tibiae III with non-striated blunt solenidia; palp tibiotarsus with one pointed process; g3 usually in straight row . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pulaeus Den Heyer, 1980a

Genus Lupaeus Castro & Den Heyer, 2009 Type species: Pulaeus martini Den Heyer, 1981b

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Lupaeus damavandiani Paktinat-Saeij & Castro sp. nov. (Figs. 1–15)

FIGURES 1–4. Lupaeus damavandiani Paktinat-Saeij & Castro sp. nov. (female): 1. Dorsal idiosoma, 2. Ventral idiosoma, 3. Gnathosoma: Subcapitulum (ventral), Palp (dorsal), 4. Chelicera.

Adult female (Figs. 1–8) (n = 2): Length of idiosoma 315 (311); width of idiosoma 226 (240). Length of legs I–IV: 223 (207); 200 (192); 201 (200); 238 (235). Length of tarsi I–IV: 71 (70); 59 (60); 60 (57); 62 (60). Dorsum (Fig. 1). Dorsal shield punctated, sensillae (at and pt) setose (based on paratype, because they are missing in the holotype). Lengths of dorsal setae: at-(90), lps 33 (30), pt-(92), mps 26 (25), c1 25 (24), c2 24 (22), d1 23 (21), e1 24 (23), f1 30 (26), f2 18 (17), h1 24 (27), h2 17 (16). Setae f1 and f2 of each side together on a tiny plate. Distances between dorsal setae: at–at 24 (26);

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lps–lps 64 (60); pt–pt 76 (70); mps–mps 55 (50); mps–c1 27 (25); c1–c1 57 (53); d1–d1 54 (52); e1– e1 40 (38); f1–f1 50 (49).

FIGURES 5–8. Lupaeus damavandiani Paktinat-Saeij & Castro sp. nov. (female): 5. Leg I, 6. Leg II, 7. Leg III, 8. Leg IV.

Venter (Fig. 2). The propodosomal coxal plates completely divided medially; striations posteromedialy to coxae forming a small, ill-defined plate-like area. The integument bears one pair of propodogastral setae and three pairs of hysterogastral setae. One pair of paragenital setae occur laterad to the genital valves. Genital valves with longitudinal rows of papillae; genital setae (g1–g4) subequal in length; setae g3 situated more laterally to row of setae g1, g2 and g4. Gnathosoma (Figs. 3–4). Subcapitulum 120 (117) with four pairs of hg-setae, hg1 14 (13), hg2 15 (15), hg3 31 (27) and hg4 10 (10) and two pairs of adoral setae. Most of the ventral surface of the subcapitulum papillated and with transverse striae line near setae hg4 (Fig. 3). Three-jointed palp (Fig. 3), 68 (65). Trochanter and base of femorogenu papillated; trochanter without setae; femorogenu, 6 sts; tibiotarsus, 5 sts, one bladder like apophysis and two pointed processes. Chelicera (Fig. 4) 123 (121) with cheliceral seta 6 (6). Legs (Figs. 2, 5–8). Leg chaetotaxy as follows: coxae I–IV, 1peg, 3-3-3-3 sts; trochanters I–IV, 1-1-2-1 sts; basifemora I–IV, 4-6-3-1 sts; telofemora I–IV, 5-5-4-3 sts; genua I–IV, {1asl, 1sts}, 3asl, 4sts-2asl, 5sts-1asl, 5sts-1asl, 5sts; tibiae I–IV, 2asl, 5sts-1bsl, 5sts-1bsl, 5sts-1T, 4sts; tarsi I–IV, 3bsl, 1fam, 1dtsl, 2tsl, 19sts-1bsl, 1dtsl, 1tsl, 17sts-1tsl, 15sts-15sts. Adult male (Figs. 9–15). (n = 1). Length of idiosoma 200; width of idiosoma 162. Length of legs I–IV: 173; 170; 164; 188. Length of tarsi I–IV: 60; 49; 48; 51. Dorsum (Fig. 9). Similar to female; lengths of dorsal setae: at 82, lps 27, pt 81, mps 22, c1 21, c2 20, d1 17, e1 17, f1 19, f2 10, h1 20, h2 9. Setae f1 and f2 of each side together on a tiny plate. Distances between dorsal setae: at–at 19; lps–lps 47; pt–pt 52; mps–mps 38; mps–c1 20; c1–c1 47; d1–d1 18; e1–e1 25; f1–f1 27. Venter (Fig. 9). Propodosomal coxal plates fused medially. The integument bears one pair of 2016


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propodogastral setae and one pair of hysterogastral setae. Paragenital setae absent. Genital valves smaller than in females, each bearing four genital setae.

FIGURES 9–15. Lupaeus damavandiani Paktinat-Saeij & Castro sp. nov. (male): 9. Dorsal and ventral view of idiosoma, 10. Gnathosoma: Subcapitulum (ventral view), Palp (dorsal view), 11. Chelicera, 12. Leg I, 13. Leg II, 14. Leg III, 15. Leg IV.

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Gnathosoma (Figs. 10–11). Gnathosoma similar to female. Subcapitulum 95 with four pairs of hg-setae, hg1 12, hg2 13, hg3 28 and hg4 8 and two pairs of adoral setae. Three-jointed palp (Fig. 10), 58. Chelicera (Fig. 11) 105 with cheliceral seta 5. Legs (Figs. 9, 12–15). The leg chaetotaxy differs from that of female as follows: tarsi I–IV, 3bsl, 1fam, 1dtsl, 2tsl, 20sts-1bsl, 1asl, 1dtsl, 1tsl, 16sts-1tsl, 13sts-14sts. Immature stages. Unknown. Remarks The new species resembles Lupaeus iranensis Den Heyer, 2013 and L. lectus Catro & Den Heyer, 2009 by having similar dorsal ornamentation and propodosomal coxal plate completely divided; however, it can be distinguished by having 1) six setae on basifemur II instead of five setae in L. lectus, 2) solenidotaxy of genua I–IV: 4-2-1-1 instead of 3-2-1-1 in L. iranensis and 4-2-1-2 in L. lectus; 3) genital seta g3 near outer margin on genital plate while in L. iranensis is arranged in a straight row. This new species is also very close to L. martini (Den Heyer, 1981b) and L. valentinae Sergeyenko, 2011, but differs from them by having: 1) one solenidion on genua IV (two solenidia in the other species); 2) dorsal shield with subcuticular punctations (smooth in the other species); 3) tarsus I with three solenidia (four solenidia in L. martini). Note Similarly to L. martini, the new species also has a striation pattern after the posteromedian region that resembles a small platelet, but it is not sclerotized and clearly defined as in Neocunaxoides rykei Den Heyer, 1980b. Although in the original description of L. martini is described as a small median platelet, we suggest it is similar to the ill-defined plate-like area present in L. damavandiani. Etymology The species is named in honor of Dr. Mohammad Reza Damavandian (Professor of Entomology, Sari Agricultural Sciences and Natural Resources University, Mazandaran, Iran), professor of the senior author in the graduate period. Type materials Holotype female, one paratype female and one paratype male were collected from samples of soil and rotten leaves under citrus trees, 20 May 2013, Osku-Mahalleh village, Amol city, Mazandaran province, Iran, by Saeid Paktinat-Saeij. The holotype female and one paratype male are deposited in the Acarological Collection, Department of Plant Protection, Faculty of Agriculture, University of Maragheh, Maragheh, Iran, and one paratype female is deposited in the Acarological Collection, Jalal Afshar Zoological Museum, Department of Plant Protection, Faculty of Agriculture, University of Tehran, Karaj, Iran.

Lupaeus clarae (Den Heyer, 1981b) Diagnosis Adult female of this species can be distinguished from other species of Lupaeus by the presence of a dorsal plate almost completely provided with broken striae except for the lateral edges which are smooth; with a small plate-like area between the posteromedian margins; the propodosomal coxal plate is not completely divided; solenidiotaxy of genua I–IV: 1dxs, 3asl-2asl-1asl-2asl. 2016


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Remarks This species is reported for the first time from Iran. Material examined Three females and one male were collected from the soil and rotten leaves under citrus trees, 31 October 2013, Soleyman-Abad village, Tonekabon city, Mazandaran province, Iran, by Saeid Paktinat-Saeij.

Key to the world species of Lupaeus Castro & Den Heyer, 2009 (females) Note. Lupaeus longisetus (Corpuz-Raros, 1996) and Lupaeus poliloensis (Corpuz-Raros, 2007) are known only for the males. Due the presence of one seta on basifemur IV the following species are transferred from Pulaeus to Lupaeus: L. akbari (Bashir & Afzal, 2009) comb. nov., L. anjumi (Bashir & Afzal, 2007) comb. nov., L. ferventis (Taj & Chaudhri, 1990) comb. nov., L. haurio (Taj & Chaudhri, 1991) comb. nov., L. payatopalpus (Corpuz-Raros, 1996) comb. nov., L. stultus (Taj & Chaudhri, 1991) comb. nov. and L. walii (Bashir & Afzal, 2009) comb. nov. 1 – 2 – 3 – 4 – 5 – 6 – 7 – 8 – 9 – 10 – 11 – 12 – 13 – 14 – 15 – 16

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Basifemur I with 3 or 5 setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Basifemur I with 4 setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 (1) Basifemur I with 5 setae; Philippines . . . . . . . . . . . . . . . . . . . . . . . . L. filipinus (Corpuz-Raros, 1996) Basifemur I with 3 setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 (2) Basifemur II with 5 setae; Philippines . . . . . . . . . L. payatopalpus (Corpuz-Raros, 1996) comb. nov. Basifemur II with 2 setae; Pakistan . . . . . . . . . . . . . . . . . . . . . L. walii (Bashir &Afzal, 2009) comb. nov. (1) Basifemur II with 4 sts; telofemur I with 3 sts; genu I with 5 setae; USA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. minutes (Baker & Hoffmann, 1948) Basifemur II with 5 or 6 setae; telofemur I with 5 setae; genu I with 7 to 9 setae . . . . . . . . . . . . . . . . . . . 5 (4) Basifemur II with 5 setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Basifemur II with 6 setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 (5) Genu I with 7 setae; palp tibiotarsus with one pointed process; USA . L. subterraneus (Berlese, 1916) Genu I with 8 or 9 setae; palp tibiotarsus with two pointed processes . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 (6) Genu I with 9 setae; genital setae arranged in straight row, being g2 close to g3, Philippines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. lenis (Corpuz-Raros, 1996) Genu I with 8 or 9 setae; genital seta g3 near outer margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 (7) Genu I with 9 setae; tibia II with 5 setae; Philippines . . . . . . . . . . . L. dentatus (Corpuz-Raros, 1996) Genu I with 8 setae; tibia II with 6 setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 (8) Dorsal shield with subcuticular punctations; Brazil . . . . . . . . . . . L. lectus Castro & Den Heyer, 2009 Dorsal shield with transverse broken striae; Brazil . . . . . . . . . L. lobidorsalis Castro & Den Heyer, 2009 (5) Genital setae arranged in a straight row . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 Genital seta g3 near outer margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 (10) Genu II without solenidion (a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 Genu II with solenidia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 (11) Genu II with one solenidion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 Genu II with two or three solenidia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 (12) Palp tibiotarsus with one pointed process; dorsum with transverse striations near lateral margins, dots randomly distributed medially; Pakistan . . . . . . . . . . . . . . L. haurio (Taj & Chaudhri, 1991) comb. nov. Palp tibiotarsus with two pointed processes; dorsum smooth, internally alveolate; Iran . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. sativae Den Heyer, 2013 (12) Genu II with two solenidia; Iran . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. iranensis Den Heyer, 2013 Genu II with three solenidia; China . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. platygnathus (Bu & Li, 1991) (11) Genu II with 7 sts; Pakistan . . . . . . . . . . . . . . . . . . . . L. stultus (Taj & Chaudhri, 1991) comb. nov. Genu II with 6 sts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 (15) Genu I with one solenidion; Pakistan . . . . . . . . . . . . . L. akbari (Bashir & Afzal, 2009) comb. nov.


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Genu I with two or three solenidia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 (16) Genu I with two solenidia; Pakistan . . . . . . . . . . . L. ferventis (Taj &Chaudhri, 1990) comb. nov. Genu I with three solenidia; Pakistan. . . . . . . . . . . . . . . . . . L. anjumi (Bashir &Afzal, 2007) comb. nov. (10) Tibia I with one solenidion, 6 sts; Philippines . . . . . . . . . . . . L. villacarlosae (Corpuz-Raros, 1996) Tibia I with two solenidia, 5 sts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 (18) Tarsus I with three proximal solenidia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 Tarsus I with four proximal solenidia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 (19) Genu IV with one solenidion; Iran . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. damavandiani sp. nov. Genu IV with two solenidia; Ukraine. . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. valentinae Sergeyenko, 2011 (19) Dorsal shield almost completely covered by broken striae (laterally smooth); South Africa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. clarae (Den Heyer, 1981b) Dorsal shield almost completely smooth except some broken striae present around the sensillae bases; South Africa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. martini (Den Heyer, 1981b)

Pulaeus Den Heyer, 1980a Type species Eupalus pectinatus Ewing, 1909.

Pulaeus aryani Paktinat-Saeij & Castro sp. nov. (Figs. 16–25) Adult Female (n = 1): Length of idiosoma 280; width of idiosoma 178. Length of legs I–IV: 230; 223; 215; 258. Length of tarsi I–IV: 72; 61; 62; 63. Dorsum (Figs. 16–17). Dorsal shield smooth, except area between pt and c2 with broken striae, sensillae (at and pt) setose. Lengths of dorsal setae: at 104, lps 38, pt 107, mps 40, c1 29, c2 34, d1 27, e1 28, f1 30, f2 22, h1 36, h2 30. f1 and f2 on tiny platelet. Distances between dorsal setae: at–at 25; lps–lps 78; pt–pt 86; mps–mps 70; mps–c1 30; c1–c1 70; c1–c2 27; d1–d1 68; e1–e1 48; f1–f1 45. Striae between h1 papillated (Fig. 17). Venter (Fig. 18). The propodosomal coxal plates completely divided medially; striations posteromedialy to coxae forming a small, ill-defined plate-like area. The integument bears one pair of propodogastral setae and three pairs of hysterogastral setae. One pair of paragenital setae occur anterolaterad of the genital valves. Genital valves with longitudinal rows of papillae; genital plate with four pairs of genital setae in almost straight longitudinal row. Setae g4 sort of longer than the other setae. g1 14, g2 11, g3 13, g4 16. Gnathosoma (Figs. 19–21). Subcapitulum 115 with four pairs of hg-setae, hg1 20, hg2 20, hg3 33 and hg4 ? and two pairs of adoral setae. The ventral surface of the subcapitulum with transverse lines posteriad setae hg4 (Fig. 19). The surface laterad and anteriad setae hg4 punctated. Threejointed palp (Fig. 20), 80. Surface of palps almost smooth, except proximal half of femorogenu covered by papillae. Trochanter without setae; femorogenu, 6 sts; tibiotarsus, 5 sts, one bladder like apophysis and one pointed process. Chelicera (Fig. 21), 127 with cheliceral seta 8. Legs (Figs. 18, 22–25). Leg chaetotaxy as follows: coxae I–IV, 1peg, 3-3-3-3 sts; trochanters I– IV, 1-1-2-1 sts; basifemora I–IV, 4-6-3-2 sts; telofemora I–IV, 5sts -5sts -3sts, 1ms-2sts, 1ms; genua I–IV, {1asl, 1sts}, 2asl, 4sts-1asl, 5sts-1asl, 5sts-1asl, 5sts; tibiae I–IV, 2asl, 5sts-1bsl, 5sts-1bsl, 5sts-1T, 4sts; tarsi I–IV, 3bsl, 1fam, 1dtsl, 2tsl, 19sts-1bsl, 1dtsl, 1tsl, 19sts-1tsl, 15sts-16sts. Adult male and immature stages and male. Unknown. Remarks This new species differs from other species of Pulaeus by having the dorsal idiosomal shield mostly smooth except an area between the pt and c2 which is broken striae. Despite it resembles P. semistriatus Sergeyenko, 2011 by also having propodosomal coxal plate completely divided and the 2016


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same legs chaetotaxy, differs by having the dorsal idiosomal shield not like P. semistriatus, which is smooth in the anterior and broken striae in the posterior part. Etymology The epithet aryani refers to “Aryan”, the ancient spelling of the word "Iranian", which translates to noble in old Persian.

FIGURES 16–21. Pulaeus aryani Paktinat-Saeij & Castro sp. nov. (female): 16–17. Dorsal idiosoma, 18. Ventral idiosoma, 19. Subcapitulum (ventral), 20. Palp (dorsal), 21. Chelicera.

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FIGURES 22–25. Pulaeus aryani Paktinat-Saeij & Castro sp. nov. (female): 22. Leg I, 23. Leg II, 24. Leg III, 25. Leg IV.

Type materials Holotype female collected from the soil and rotten leaves under citrus trees, 20 May 2013, OskuMahalleh village, Amol city, Mazandaran province, Iran, by Saeid Paktinat-Saeij. The holotype female is deposited in the Acarological Collection, Department of Plant Protection, Faculty of Agriculture, University of Maragheh, Maragheh, Iran.

Key to the world species of Pulaeus Den Heyer, 1980a (females) Note. Available information about P. parapatzuarensis (Shiba, 1978), P. pseudominutus (Shiba, 1978) and P. zaherii (El-Bishlawy & Rakha, 1983) is not sufficient to allow their inclusion in the key, despite the complementary information recently published by Skvarla et al. (2014). Also, P. cinctus (Chaudhri, 1980) can be distinguished by having a divided propodosomal coxal plate, lacks a median platelet between coxae II–III, setae f1 and f2 not located on sclerotized platelets or shields and having four pairs of setae on integument between coxal and genital plates. Pulaeus ranzanensis Den Heyer, 2013 was not included in this key because based on the available literature information no major differences could be observed between this species and Pulaeus krama (Chaudhri, Akbar & Rasool, 1979). Therefore, we suggest P. razanensis Den Heyer, 2013 syn. nov. is junior synonym of P. krama. In addition, Neocunaxoides neopectinatus (Shiba, 1978) comb. nov. described as Cunaxoides neopectinatus Shiba, 1978 was transferred to Pulaeus by Skvarla et al. (2014). In this paper we

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consider it to belong to Neocunaxoides, because of the absence of f2; thus a new combination is now established for this species, Neocunaxoides neopectinatus (Shiba, 1978) comb. nov. 1 – 2 – 3 – 4 – 5 – 6 – 7 – 8 – 9 – 10 – 11 – 12 – 13 – 14 – 15 – 16 – 17 – 18 – 19 – 20 –

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Telofemora I–IV 5-5-4-3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 Telofemora I–IV not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 (1) Telofemora I–IV 4-5-4-3; median platelet with a seta anteriad to genital opening; USA, South African, Brazil . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. pectinatus (Ewing, 1909) Telofemora I–IV not as above; median platelet anteriad to genital opening when present without seta . . . ................................................................................... 3 (2) Basifemora I–III 4-4-3 and telefemora I–IV 3-3-2-2; USA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. americanus (Baker & Hoffmann, 1948) Basifemora and telofemora not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 (3) Basifemora I–III 4-5-3 and telofemora I–IV 5-4-4-4; USA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. whartoni (Baker &Hoffmann, 1948) Basifemora and telofemora not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 (4) Basifemora I–III 3-6-3 and telefemora I–IV 4-6-4-3; Pakistan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. banksi Bashir & Afzal, 2009 Basifemora and telofemora not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 (5) Basifemora I–III 4-5-5 and telofemora I–IV 4-4-4-3; USA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. patzcuarensis (Baker & Hoffmann, 1948) Basifemora and telofemora not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 (6) Basifemora I–III 3-6-4 and telofemora I–IV 5-6-4-3; Philippines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. samarensis Corpuz-Raros, 2007 Basifemora I–III 4-6-3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 (7) Telofemora I–IV 5-5-3-2; basifemur I with one blunt solenidion; Pakistan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. camar Taj & Chaudhri, 1991 Telofemora not as above and basifemur I without solenidia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 (8) Telofemora I–IV 5-5-3-3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. galumma Taj & Chaudhri, 1991 Telofemora I–IV 5-4-4-3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 (9) Surface of the dorsum shield covered by broken striae; Philippines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. palawanensis Corpuz-Raros, 2007 Surface of the dorsum covered by dots (punctions); Pakistan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. punctatus Bashir, Afzal & Akbar, 2005 (1) Basifemora I–II 4-6. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Basifemora I–II not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 (11) Basifemora I–II 4-7; Philippines . . . . . . . . . . . . . . . . . . . . . . . . . . P. rimandoi Corpuz-Raros, 1996 Basifemora I-II 4-5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 (12) Genital setae (g1–g4) situated in longitudinal row; Philippines . . . P. cebuensis Corpuz-Raros, 2007 Genital seta (g3) situated near outer margin; Ukraine . . . . . . . . . . . . . . . . . . . P. leonidi Sergeyenko, 2011 (11) Genital setae g3 situated more laterally in comparison to the row formed by other g setae; South Africa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. glebulentus Den Heyer, 1980a Genital setae g1–g4 situated in longitudinal row . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 (14) Genua I–IV with 5 sts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 Genua I–IV not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 16 (15) Genu I with four solenidia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 Genu I with five solenidia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 (16) Dorsal idiosomal shield mostly smooth; Pakistan . . . . . . . . . . . . P. osculum Taj & Chaudhri, 1990 Dorsal idiosomal shield not smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 (17) Genu I with 6 sts; Pakistan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. P. desitis Taj & Chaudhri, 1990 Genu I with 4 sts; Pakistan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. verno Taj & Chaudhri, 1990 (16) Genu I with 3 sts; Pakistan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. erinaceus Taj & Chaudhri, 1991 Genu I with 4 sts. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 (19) Dorsal idiosomal shield broken striae in the anterior and smooth in the posterior parts; Uzbekistan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. P. ardeola Barilo 1991 Dorsal idiosomal shield almost broken striae except median area between mps and d1 punctated; Pakistan .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. silicula Taj & Chaudhri, 1991 SYSTEMATIC & APPLIED ACAROLOGY

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21 – 22 – 23 – 24 – 25 – 26 – 27 –

(15) Genu II with one solenidion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 Genu II with two solenidia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 (21) Genu I with two solenidia; China . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. musci Liang, 1985 Genu I with three solenidia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 (22) Dorsal idiosomal shield completely covered by broken striae; Ukraine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. maslovi Sergeyenko, 2011 Dorsal idiosomal shield mostly smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 (23) Dorsal idiosomal shield smooth in anterior and with transverse striae in posterior halves; Ukraine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. semistriatus Sergeyenko, 2011 Dorsal idiosomal shield mostly smooth except an area between the pt and c2 with broken striae; Iran . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. aryani sp. nov. (21) Dorsal idiosomal shield with surface broken striae; Pakistan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. P. krama (Chaudhri, Akbar & Rasool, 1979) Dorsal idiosomal shield with surface punctuated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 (25) Genu I with four solenidia; Brazil . . . . . . . . . . . . . . P. quadrisolenidius Castro & Den Heyer, 2009 Genu I with three solenidia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 (26) Dorsal shield strongly punctated; Brazil. . . . . . . . . . . . . . .. P. myrtaceus Castro & Den Heyer, 2009 Dorsal shield slightly punctated; South African. . . . . . . . . . . . . . . . . . . . P. franciscae Den Heyer, 1981b

Acknowledgements We greatly appreciate the support for this project provided by the Research Divisions of University of Maragheh, Maragheh, Iran. This paper is dedicated to the family of the senior author.

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