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Ditylenchus azarbaijanensis n. sp. is described and illustrated based upon morphological and morphometric characters. The new species is mainly characterized by having six lines in the lateral field, rudimentary postuterine sac (PUS) ca. 0.2 vulval body width long, anteriorly inclined vagina and conical female tail with a pointed terminus, 48-70 μm long (c = 15-21, c􀀀 = 2.4-3.7). It is further characterized by having fine stylet 7.5-10.0 μm long with small knobs, small pyriform pharyngeal bulb offset from the intestine, and males with 26.5- 31.0 μm long spicules.

Labrys khuzestanensis n. sp. (Figs 1–3) Measurements: see Table 1. Female: Body straight to slightly ventrally arcuate after heat relaxation. Cuticle finely annulated, annulus ca 1 μm wide at midbody. Lateral field distinct, having two incisures with smooth margins under SEM. Cephalic region smooth, continuous with the adjacent part of body, 4.5 μm wide and 2.1–2.3 μm high, trapezoid under LM, dorsoventrally flattened, with protuberant large and elongate labial plate under SEM. The plate is laterally extended and dorso-ventrally not constricted. In lateral view under LM, the plate seems as a V-shaped piece (Fig. 2D; hardly visible due to small size). The oral plate small, laterally elongate, containing labial sensillae, with small and elevated oral opening at the center. Amphidial openings as longitudinal slits, beginning near oral plate, laterally extending to the labial plate corners and confined to it. Stylet delicate, slender, conus shorter than the shaft, comprising ca 33% of its total length, the knobs rounded and small, slightly posteriorly sloping. Dorsal pharyngeal gland orifice (DGO) 1–2 μm from the base of the stylet. Procorpus slender, median bulb moderately developed, elongate fusiform with small valvular apparatus, isthmus narrow, slender, pharyngeal bulb small, pyriform and offset. Cardia small, domeshaped. Nerve ring encircling anterior part of isthmus, 57 to 76 μm from anterior end. Excretory pore wide with sclerotized duct, at the distance between the nerve ring and anterior part of pharyngeal bulb. Reproductive system monodelphic-prodelphic, composed of an outstretched ovary occupying 13–15% of total body length, the oocytes mostly arranged in single row. The border of oviduct, spermatheca and the cell arrangement of crustaformeria indistinct. Vagina with thin wall, perpendicular to body axis and straight or slightly anteriorly sloping. Postvulval uterine sac (PUS) ca 0.6 to 1.0 times corresponding body diameter long. Vulva a small transverse slit with very small lateral flaps. Tail elongate filiform, very gradually narrowing toward end, 1.9 to 3.0 times longer than vulva–anus distance, with finely rounded tip. 1 Distance between anterior end of body and center of median pharyngeal bulb as percentage of pharynx length. 2 Postvulval uterine sac/body width Male: Unknown. Type habitat and locality. The rhizospheric soil of a palm tree (Phoenix dactylifera L.) in city of Ahvaz, Khuzestan province, southwestern Iran, collected during 2016. GPS coordinates: 31°28.582′N, 48°37.394′E. Etymology. The specific epithet refers to Khuzestan province, the original geographic point of which the new species was recovered from. Type material. Holotype female and four paratype females were deposited at Nematode Collection of the Faculty of Agriculture, Tarbiat Modares University, Tehran, Iran (the slide code: TM5043). Three paratype females were deposited at Ghent University Museum, Zoology Collections, Ghent, Belgium (UGMD _104411 and UGMD _ 104412). Two paratype females were deposited at the Nematode Collection of the Department of Animal Biology, Plant Biology and Ecology of the University of Jaén, Jaén, Spain) (Collection ID: Iran 011) (one paratype female was used to prepare SEM photos). Diagnosis and relationships. Labrys khuzestanensis n. sp. is mainly characterized by large and prominent labial plate as revealed by SEM and trapezoid cephalic region under LM. It is further characterized by 590–675 μm long females having a finely annulated cuticle under LM, lateral field with two incisures, cephalic region dorsoventrally flattened and smooth, labial plate not constricted at middle, elongate, laterally extending and appearing as V-shaped piece in lateral view, amphidial openings long slits confined to the labial plate, stylet 7.5–9.0 μm long, median bulb elongate fusiform with small valvular apparatus, excretory pore wide, with sclerotized duct and tail elongate filiform with finely rounded tip. Compared to four currently known species of the genus, the new species has a remarkably large and prominent labial plate and trapezoid cephalic region under LM (vs. rounded). The detailed morphological comparisons of the new species with Labrys members are as follows: From L. chinensis Qing & Bert, 2018 by distinctly annulated cuticle under LM (vs. smooth), greater a ratio (53.6– 74.8 vs. 37.7–48.7), lower c ratio (2.6–3.2 vs. 3.5–4.4), greater cʹ ratio (23.5–32.9 vs. 13.3–20.3), more anteriorly located vulva (V = 48.0–54.3 vs. 54–60), greater tail length/vulva to anus distance ratio (1.9–3.0 vs. 1.3–1.7), longer tail (188–237 vs. 132–171 μm) and difference in tail end morphology (finely rounded tip vs. broadly rounded). From L. fujianensis Qiao, Bai, Wang, Hou, Chen, Xiao, Liu, Bert & Qing, 2019a by laterally elongate oral plate (vs. dorsoventrally, after its SEM data) and slightly longer body (590–675 vs. 524–599 μm). From L. filiformis Panahandeh, Abolafia, Pourjam, Jahanshahi Afshar, Giblin-Davis & Pedram, 2018 by its distinct annulated cuticle under LM (vs. smooth), longer body (590–675 vs. 425–463 μm), greater a ratio (53.6–74.8 vs. 38.7–46.2), greater b ratio (5.7–7.4 vs. 4.9–5.7), lower c ratio (2.6–3.2 vs. 3.5–4.0), greater cʹ ratio (23.5– 32.9 vs. 13.2–15.9), more anteriorly located vulva (V = 48.0–54.3 vs. 57.7–61.5), lower Uʹ ratio (74.6–79.8 vs. 79.9–84.1) and longer tail (188–237 vs. 106–127 μm). From L. fuzhouensis Qiao, Bai, He, Chen, Xiao, Cheng, Liu, Braun-Miyara & Qing, 2019b by longer body (590–675 vs. 495–564 μm), greater a ratio (53.6–74.8 vs. 40.6–53.1), longer neck (86–108 vs. 72–80 μm) and lacking male (vs. present). Phylogenetic analyses. To study of the phylogenetic affinities of the new species with other genera/species of the family Tylenchidae, its partial SSU rDNA was used in corresponding phylogenetic analysis. A total number of 75 species/isolates (including the newly obtained SSU sequence of the new species and three aphelenchid+a phelenchoidid outgroups) were selected for these analyses (Bayesian inference and ML trees). The alignment of SSU dataset had 1824 total characters with 981 variable characters. The average nucleotide composition was as follows: 24.5% A, 22.9% C, 28.3% G, and 24.3% T. Figure 4 represents the Bayesian phylogenetic tree inferred using the abovementioned dataset. In this tree, the new species and three previously described species viz. Labrys chinensis (KY776630, KY776631, KY776632, KY776633), L. fuzhouensis (MK039731, MK039732), and L. filiformis (MG686086) formed a clade and Discotylencus lorestanensis Mehrabian, Azizi, Bazgir & Darvishnia, 2017 (KU878152, KU878153) is the closest taxon to these species. The other species of Labrys, L. fujianensis occupied a distant placement to them.

Labrys khuzestanensis n. sp. (Figs 1���3) Measurements: see Table 1. Female: Body straight to slightly ventrally arcuate after heat relaxation. Cuticle finely annulated, annulus ca 1 ��m wide at midbody. Lateral field distinct, having two incisures with smooth margins under SEM. Cephalic region smooth, continuous with the adjacent part of body, 4.5 ��m wide and 2.1���2.3 ��m high, trapezoid under LM, dorsoventrally flattened, with protuberant large and elongate labial plate under SEM. The plate is laterally extended and dorso-ventrally not constricted. In lateral view under LM, the plate seems as a V-shaped piece (Fig. 2D; hardly visible due to small size). The oral plate small, laterally elongate, containing labial sensillae, with small and elevated oral opening at the center. Amphidial openings as longitudinal slits, beginning near oral plate, laterally extending to the labial plate corners and confined to it. Stylet delicate, slender, conus shorter than the shaft, comprising ca 33% of its total length, the knobs rounded and small, slightly posteriorly sloping. Dorsal pharyngeal gland orifice (DGO) 1���2 ��m from the base of the stylet. Procorpus slender, median bulb moderately developed, elongate fusiform with small valvular apparatus, isthmus narrow, slender, pharyngeal bulb small, pyriform and offset. Cardia small, domeshaped. Nerve ring encircling anterior part of isthmus, 57 to 76 ��m from anterior end. Excretory pore wide with sclerotized duct, at the distance between the nerve ring and anterior part of pharyngeal bulb. Reproductive system monodelphic-prodelphic, composed of an outstretched ovary occupying 13���15% of total body length, the oocytes mostly arranged in single row. The border of oviduct, spermatheca and the cell arrangement of crustaformeria indistinct. Vagina with thin wall, perpendicular to body axis and straight or slightly anteriorly sloping. Postvulval uterine sac (PUS) ca 0.6 to 1.0 times corresponding body diameter long. Vulva a small transverse slit with very small lateral flaps. Tail elongate filiform, very gradually narrowing toward end, 1.9 to 3.0 times longer than vulva���anus distance, with finely rounded tip. 1 Distance between anterior end of body and center of median pharyngeal bulb as percentage of pharynx length. 2 Postvulval uterine sac/body width Male: Unknown. Type habitat and locality. The rhizospheric soil of a palm tree (Phoenix dactylifera L.) in city of Ahvaz, Khuzestan province, southwestern Iran, collected during 2016. GPS coordinates: 31��28.582���N, 48��37.394���E. Etymology. The specific epithet refers to Khuzestan province, the original geographic point of which the new species was recovered from. Type material. Holotype female and four paratype females were deposited at Nematode Collection of the Faculty of Agriculture, Tarbiat Modares University, Tehran, Iran (the slide code: TM5043). Three paratype females were deposited at Ghent University Museum, Zoology Collections, Ghent, Belgium (UGMD _104411 and UGMD _ 104412). Two paratype females were deposited at the Nematode Collection of the Department of Animal Biology, Plant Biology and Ecology of the University of Ja��n, Ja��n, Spain) (Collection ID: Iran 011) (one paratype female was used to prepare SEM photos). Diagnosis and relationships. Labrys khuzestanensis n. sp. is mainly characterized by large and prominent labial plate as revealed by SEM and trapezoid cephalic region under LM. It is further characterized by 590���675 ��m long females having a finely annulated cuticle under LM, lateral field with two incisures, cephalic region dorsoventrally flattened and smooth, labial plate not constricted at middle, elongate, laterally extending and appearing as V-shaped piece in lateral view, amphidial openings long slits confined to the labial plate, stylet 7.5���9.0 ��m long, median bulb elongate fusiform with small valvular apparatus, excretory pore wide, with sclerotized duct and tail elongate filiform with finely rounded tip. Compared to four currently known species of the genus, the new species has a remarkably large and prominent labial plate and trapezoid cephalic region under LM (vs. rounded). The detailed morphological comparisons of the new species with Labrys members are as follows: From L. chinensis Qing & Bert, 2018 by distinctly annulated cuticle under LM (vs. smooth), greater a ratio (53.6��� 74.8 vs. 37.7���48.7), lower c ratio (2.6���3.2 vs. 3.5���4.4), greater c�� ratio (23.5���32.9 vs. 13.3���20.3), more anteriorly located vulva (V = 48.0���54.3 vs. 54���60), greater tail length/vulva to anus distance ratio (1.9���3.0 vs. 1.3���1.7), longer tail (188���237 vs. 132���171 ��m) and difference in tail end morphology (finely rounded tip vs. broadly rounded). From L. fujianensis Qiao, Bai, Wang, Hou, Chen, Xiao, Liu, Bert & Qing, 2019a by laterally elongate oral plate (vs. dorsoventrally, after its SEM data) and slightly longer body (590���675 vs. 524���599 ��m). From L. filiformis Panahandeh, Abolafia, Pourjam, Jahanshahi Afshar, Giblin-Davis & Pedram, 2018 by its distinct annulated cuticle under LM (vs. smooth), longer body (590���675 vs. 425���463 ��m), greater a ratio (53.6���74.8 vs. 38.7���46.2), greater b ratio (5.7���7.4 vs. 4.9���5.7), lower c ratio (2.6���3.2 vs. 3.5���4.0), greater c�� ratio (23.5��� 32.9 vs. 13.2���15.9), more anteriorly located vulva (V = 48.0���54.3 vs. 57.7���61.5), lower U�� ratio (74.6���79.8 vs. 79.9���84.1) and longer tail (188���237 vs. 106���127 ��m). From L. fuzhouensis Qiao, Bai, He, Chen, Xiao, Cheng, Liu, Braun-Miyara & Qing, 2019b by longer body (590���675 vs. 495���564 ��m), greater a ratio (53.6���74.8 vs. 40.6���53.1), longer neck (86���108 vs. 72���80 ��m) and lacking male (vs. present). Phylogenetic analyses. To study of the phylogenetic affinities of the new species with other genera/species of the family Tylenchidae, its partial SSU rDNA was used in corresponding phylogenetic analysis. A total number of 75 species/isolates (including the newly obtained SSU sequence of the new species and three aphelenchid+a phelenchoidid outgroups) were selected for these analyses (Bayesian inference and ML trees). The alignment of SSU dataset had 1824 total characters with 981 variable characters. The average nucleotide composition was as follows: 24.5% A, 22.9% C, 28.3% G, and 24.3% T. Figure 4 represents the Bayesian phylogenetic tree inferred using the abovementioned dataset. In this tree, the new species and three previously described species viz. Labrys chinensis (KY776630, KY776631, KY776632, KY776633), L. fuzhouensis (MK039731, MK039732), and L. filiformis (MG686086) formed a clade and Discotylencus lorestanensis Mehrabian, Azizi, Bazgir & Darvishnia, 2017 (KU878152, KU878153) is the closest taxon to these species. The other species of Labrys, L. fujianensis occupied a distant placement to them., Published as part of Panahandeh, Yousef, Pourjam, Ebrahim, Abolafia, Joaqu��n, Roshan-Bakhsh, Ali, Mojerlou, Shideh, Afshar, Farahnaz Jahanshahi & Pedram, Majid, 2019, Labrys khuzestanensis n. sp. (Nematoda, Tylenchidae), a new member of the genus with large labial plate, pp. 267-276 in Zootaxa 4671 (2) on pages 268-272, DOI: 10.11646/zootaxa.4671.2.7, http://zenodo.org/record/3442420, {"references":["Qing, X. & Bert, W. (2018) 3 D printing in zoological systematics: Integrative taxonomy of Labrys chinensis gen. nov., sp. nov. (Nematoda: Tylenchomorpha). Journal of Zoological Systematics and Evolutionary Research, 56, 35 - 47. https: // doi. org / 10.1111 / jzs. 1219","Qiao, K., Bai, M., Wang, Q., Hou, X., Chen, J., Xiao, S., Liu, G., Bert, W. & Qing, X. (2019 a) Unexpected rDNA divergence between two morphologically minimalistic nematodes with description of a new species (Tylenchomorpha: Tylenchidae). Nematology, 21, 57 - 70. https: // doi. org / 10.1163 / 15685411 - 00003195","Panahandeh, Y., Abolafia, J., Pourjam, E., Jahanshahi Afshar, F., Giblin-Davis, R. M. & Pedram, M. (2018) Morphological and Molecular Characterization of Labrys filiformis n. sp. (Rhabditida: Tylenchidae) from Iran. Journal of Nematology, 50, 343 - 354. https: // doi. org / 10.21307 / jofnem- 2018 - 031","Qiao, K., Bai, M., He, Y., Chen, J., Xiao, S., Cheng, X., Liu, G., Braun-Miyara, S. & Qing, X. (2019 b) Description of Labrys fuzhouensis sp. n. and first record of Coslenchus rafiqi (Nematoda: Tylenchidae) from China. Nematology, 21, 693 - 708, https: // doi. org / 10.1163 / 15685411 - 00003246","Mehrabian, F., Azizi, K., Bazgir, E. & Darvishnia, M. (2017) Morphological and molecular characterisation of Discotylenchus lorestanensis sp. n. (Nematoda: Tylenchidae) from Iran. Zootaxa, 4290 (1), 167 - 176. https: // doi. org / 10.11646 / zootaxa. 4290.1.10"]}

Konani, Ehsan, Panahandeh, Yousef, Pourjam, Ebrahim, Álvarez-Ortega, Sergio, Pedram, Majid (2018): Two rare species of tylenchids, Discotylenchus biannulatus n. sp. and Labrys chinensis Qing & Bert, 2018 (Nematoda: Tylenchidae) from western Iran. Zootaxa 4413 (2): 260-270, DOI: https://doi.org/10.11646/zootaxa.4413.2.2

Discotylenchus biannulatus n. sp. (Figs. 1–3) Description. Measurements. See Table 1. Female: Body straight to slightly ventrally curved after heat relaxation. Cuticle finely annulated, annules ca. 1–2 µm wide at mid-body. Lateral fields with four incisures, outer lines partly areolated at anus/tail region in SEM images, 2–3 µm wide at mid-body, approximately 20% of body diameter. Lip region continuous with body contour, 5–6 µm wide and 3–4 µm high, dorso-ventrally flattened, smooth, having two annuli in the proximal end and a prominent perioral disc under SEM. Amphidial openings as longitudinal slits on lateral sides of the head. Stylet delicate, with conus shorter than the shaft comprising ca 32.7% of stylet total length; knobs small, slightly sloping posteriad. Dorsal gland orifice (DGO) 1–2 µm from base of stylet. Procorpus slender, joining to an ellipsoid muscular metacorpus with distinct valvular apparatus, located at 39–43% of total pharynx length; isthmus elongated and narrow; and basal bulb with slight variation in shape, saccate to bottle-shaped, 14–18 µm long and 7–9 µm wide. Cardia small. Nerve ring encircling anterior half of isthmus, 63–68 µm from anterior end. Excretory pore situated at the level of the middle of the isthmus, posterior to hemizonid. Deirids distinct and situated 4–6 annuli posterior to excretory pore. Reproductive system monodelphic-prodelphic, composed of an outstretched ovary occupying 23–26% of the body length, oocytes mainly in a single row, oviduct short, spermatheca rounded and empty, crustaformeria hardly visible, uterus tubular, vagina straight with thin walls; PUS length about 0.6–0.8 times vulval body width, vulva a transverse slit lacking any differentiation. Tail filiform, gradually tapering to a more or less pointed end. Male: not found. Type habitat and locality. Recovered from the rhizosphere of wheat, Giloran region, city of Khorram-Abad, Lorestan province, western Iran. GPS coordinates: N 33 27 ʹ 51.53 and E 48 18 ʹ 17.99. Etymology. The specific epithet refers to two unusual annuli at the anterior end of the cephalic region of the new species. Type material. Holotype and five paratypes deposited at the Ghent University Museum, Zoology Collection, Belgium. Diagnosis and relationships. Discotylenchus biannulatus n. sp. is characterized by its dorso-ventrally flattened smooth lip region having two proximal annuli and a distinct rectangular perioral disc, short longitudinal amphidial slits, lateral field with four incisures, stylet 9–10 µm long, empty spermatheca and short PUS, and filiform female tail with pointed end. With four lines in the lateral field, the new species can be compared with three known species of the genus: D. attenuatus Siddiqi, 1980, D. brevicaudatus and D. discretus. The new species can be distinguished from these species by its smooth cephalic region with two annuli at the anterior end. In addition, Discotylenchus biannulatus n. sp. differs from D. attenuatus by having longer body (675–708 vs 330–400 µm), longer stylet (9–10 vs 6.0–6.5 µm), longer tail (155–161 vs 68–96 µm) and higher cʹ ratio (14.1–16.1 vs 9–12). It can be distinguished from D. brevicaudatus by its dorso-ventrally flattened cephalic region (vs rounded, according to SEM data provided by Yaghoubi et al. 2016), longer body (675–708 vs 320–466 µm), longer stylet (9–10 vs 6–8 µm), more anteriorly located vulva (V = 59.5–60.6 vs 68–73), longer tail (155–161 vs 35–77 µm), smaller c ratio (4.2–4.5 vs 5.9–9.8), higher cʹ ratio (14.1–16.1 vs 4.5–9.6) and filiform tail with pointed end (vs thick with rounded end). Finally, compared to D. discretus, the new species has dorso-ventrally flattened cephalic region (vs rectangular, smooth, according to SEM data provided by Yaghoubi et al. 2016), longer stylet (9–10 vs 7–9 µm), more anteriorly located vulva (V = 59.5–60.6 vs 63–67), longer tail (155–161 vs 80–115 µm), smaller c ratio (4.2–4.5 vs 5.0–6.2), and higher cʹ ratio (14–16 vs 9–11). Remarks. The presently described new species was recovered from an agricultural wheat field usually plowed annually. Detailed SEM studies helped establish its generic identity, but further samplings to get fresh material for molecular phylogenetic studies failed. It has a unique head morphology, however, not reported or illustrated to date for any members of Tylenchidae. Light microscope observations at first indicated the presence of an anterior disk, also confirmed in SEM studies, and all other morphological characters (especially longitudinal amphidial slits) supported its placement under the genus Discotylenchus. Future molecular studies will help determine the phylogenetic affinities of this species.

Discotylenchus biannulatus n. sp. (Figs. 1���3) Description. Measurements. See Table 1. Female: Body straight to slightly ventrally curved after heat relaxation. Cuticle finely annulated, annules ca. 1���2 ��m wide at mid-body. Lateral fields with four incisures, outer lines partly areolated at anus/tail region in SEM images, 2���3 ��m wide at mid-body, approximately 20% of body diameter. Lip region continuous with body contour, 5���6 ��m wide and 3���4 ��m high, dorso-ventrally flattened, smooth, having two annuli in the proximal end and a prominent perioral disc under SEM. Amphidial openings as longitudinal slits on lateral sides of the head. Stylet delicate, with conus shorter than the shaft comprising ca 32.7% of stylet total length; knobs small, slightly sloping posteriad. Dorsal gland orifice (DGO) 1���2 ��m from base of stylet. Procorpus slender, joining to an ellipsoid muscular metacorpus with distinct valvular apparatus, located at 39���43% of total pharynx length; isthmus elongated and narrow; and basal bulb with slight variation in shape, saccate to bottle-shaped, 14���18 ��m long and 7���9 ��m wide. Cardia small. Nerve ring encircling anterior half of isthmus, 63���68 ��m from anterior end. Excretory pore situated at the level of the middle of the isthmus, posterior to hemizonid. Deirids distinct and situated 4���6 annuli posterior to excretory pore. Reproductive system monodelphic-prodelphic, composed of an outstretched ovary occupying 23���26% of the body length, oocytes mainly in a single row, oviduct short, spermatheca rounded and empty, crustaformeria hardly visible, uterus tubular, vagina straight with thin walls; PUS length about 0.6���0.8 times vulval body width, vulva a transverse slit lacking any differentiation. Tail filiform, gradually tapering to a more or less pointed end. Male: not found. Type habitat and locality. Recovered from the rhizosphere of wheat, Giloran region, city of Khorram-Abad, Lorestan province, western Iran. GPS coordinates: N 33 27 �� 51.53 and E 48 18 �� 17.99. Etymology. The specific epithet refers to two unusual annuli at the anterior end of the cephalic region of the new species. Type material. Holotype and five paratypes deposited at the Ghent University Museum, Zoology Collection, Belgium. Diagnosis and relationships. Discotylenchus biannulatus n. sp. is characterized by its dorso-ventrally flattened smooth lip region having two proximal annuli and a distinct rectangular perioral disc, short longitudinal amphidial slits, lateral field with four incisures, stylet 9���10 ��m long, empty spermatheca and short PUS, and filiform female tail with pointed end. With four lines in the lateral field, the new species can be compared with three known species of the genus: D. attenuatus Siddiqi, 1980, D. brevicaudatus and D. discretus. The new species can be distinguished from these species by its smooth cephalic region with two annuli at the anterior end. In addition, Discotylenchus biannulatus n. sp. differs from D. attenuatus by having longer body (675���708 vs 330���400 ��m), longer stylet (9���10 vs 6.0���6.5 ��m), longer tail (155���161 vs 68���96 ��m) and higher c�� ratio (14.1���16.1 vs 9���12). It can be distinguished from D. brevicaudatus by its dorso-ventrally flattened cephalic region (vs rounded, according to SEM data provided by Yaghoubi et al. 2016), longer body (675���708 vs 320���466 ��m), longer stylet (9���10 vs 6���8 ��m), more anteriorly located vulva (V = 59.5���60.6 vs 68���73), longer tail (155���161 vs 35���77 ��m), smaller c ratio (4.2���4.5 vs 5.9���9.8), higher c�� ratio (14.1���16.1 vs 4.5���9.6) and filiform tail with pointed end (vs thick with rounded end). Finally, compared to D. discretus, the new species has dorso-ventrally flattened cephalic region (vs rectangular, smooth, according to SEM data provided by Yaghoubi et al. 2016), longer stylet (9���10 vs 7���9 ��m), more anteriorly located vulva (V = 59.5���60.6 vs 63���67), longer tail (155���161 vs 80���115 ��m), smaller c ratio (4.2���4.5 vs 5.0���6.2), and higher c�� ratio (14���16 vs 9���11). Remarks. The presently described new species was recovered from an agricultural wheat field usually plowed annually. Detailed SEM studies helped establish its generic identity, but further samplings to get fresh material for molecular phylogenetic studies failed. It has a unique head morphology, however, not reported or illustrated to date for any members of Tylenchidae. Light microscope observations at first indicated the presence of an anterior disk, also confirmed in SEM studies, and all other morphological characters (especially longitudinal amphidial slits) supported its placement under the genus Discotylenchus. Future molecular studies will help determine the phylogenetic affinities of this species., Published as part of Konani, Ehsan, Panahandeh, Yousef, Pourjam, Ebrahim, ��lvarez-Ortega, Sergio & Pedram, Majid, 2018, Two rare species of tylenchids, Discotylenchus biannulatus n. sp. and Labrys chinensis Qing & Bert, 2018 (Nematoda: Tylenchidae) from western Iran, pp. 260-270 in Zootaxa 4413 (2) on pages 261-264, DOI: 10.11646/zootaxa.4413.2.2, http://zenodo.org/record/1226931, {"references":["Siddiqi, M. R. (1980) Two new nematode genera, Safianema (Anguinidae) and Discotylenchus (Tylenchidae), with descriptions of three new species. Proceedings of the Helminthological Society of Washington, 47, 85 - 94.","Yaghoubi, A., Pourjam, E., Alvarez-Ortega, S. Liebanas, G., Atighi, M. R. & Pedram, M. (2016) Discopersicus n. gen., a New Member of the Family Tylenchidae Orley, 1880 with Detailed SEM Study on Two Known Species of the Genus Discotylenchus Siddiqi, 1980 (Nematoda; Tylenchidae) from Iran. Journal of Nematology, 48, 214 - 221. https: // doi. org / 10.21307 / jofnem- 2017 - 029"]}

Malenchus geraerti n. sp. Figs 1���3 The new species is named after Prof. Etienne Geraert, the pioneer and outstanding scientist in the taxonomy of Tylenchidae. Measurements, see Table 1. Female. Body slightly ventrally curved after fixation, markedly narrowing from the level of vulva to distal end. Cuticle coarsely annulated, annuli 2.1���2.3 ��m wide at mid-body. Lateral field a plain band (i.e. composed of two incisures), raised in cross section, constricted at the junction with body (Fig. 3B, arrows), originating at about the level of the middle of the stylet shaft in its resting position. Lip region continuous with body contour; not conspicuously narrower than the adjacent body, 5���7 ��m wide, 3���4 ��m high, bearing 3���4 annuli. Amphidial openings S-shaped slits, starting from labial plate, extending posteriorly, markedly narrowing and reaching base of the head (extending to the weakly sclerotized cephalic framework). Stylet shaft moderately developed, conus much finer, knobs directed posteriorly. Dorsal gland orifice (DGO) close to knobs. Excretory pore (ex. pore) at the level of pharyngeal bulb, slightly variable in position, 91���94 ��m from anterior end. Hemizonid 92 ��m (n = 3) and nerve ring 63���75 ��m from anterior end. Pharynx with slender procorpus, muscular, rounded to ellipsoid metacorpus with well-developed valve and corresponding plates at 43���49 ��m from anterior end, slender/narrow isthmus and saccate pharyngeal bulb 13���18 �� 8���11 ��m. Reproductive system mono-prodelphic, ovary outstretched, oocytes mostly in single file, except in germinal zone, oviduct short, spermatheca elongate-rounded, filled with small spheroid sperm cells, offset and sometimes appearing as bilobed (with depression or constriction at middle), crustaformeria and uterus not clearly discernible from each other. Vagina perpendicular to body axis, with moderately sclerotized walls. Small, undifferentiated postvulval uterine sac (PUS) present. Vulva sunken inside body, in the form of a transverse slit, with large epiptygma and no flaps. Prophasmid barely visible, located 14 ��m anterior to vulval opening (n = 1). Vulva���anus distance 64���77 ��m, or 0.8���1.1 times the tail length. Tail conical, tapering gradually to a more or less pointed tip. Male. About half as frequent as females in the studied population. General morphology similar to that of female, except in sexual characters. Testis single, outstretched. Spicules tylenchoid, narrow and slender, slightly arcuate distally, 17���22 ��m long with rounded capitulum, slender shaft and blade narrowing toward distal end. Gubernaculum small, 4���6 ��m. Bursa cloacal, 26���43 ��m long, with smooth margin. Tail similar to that of female. Type Habitat And Locality. Recovered from soil samples collected about rhizosphere of forest trees in forests of Gilan province, northern Iran. GPS coordinates: 37��7��30.719����N, 49��39��25.742����E. Type Material. Five females (holotype and paratypes) and five males deposited in the Nematode Collection of the Faculty of Agriculture, Tarbiat Modares University, Tehran, Iran. Five paratype females and two males deposited at each of the following collections: WANECO collection, Wageningen, The Netherlands, and UGent Nematode Collection of the Nematology Research Unit, Department of Biology, Ghent University, Ghent, Belgium. Diagnosis And Relationships. The new species belongs to the subgenus Malenchus owing to its amphidial opening characteristics (S-shaped in lateral view, wide at anterior end, narrowing distally and reaching to the base of the head), and two lateral lines, forming a plain band under light microscopy. It is further characterized by having short-bodied females, lateral field origin at about the middle of the stylet shaft, muscular, rounded to ellipsoid metacorpus with well-developed valve and valvular plates, small PUS, vulva sunken in body with large epiptygma and no flap, conical tail, tapering gradually to a more or less pointed tip, functional males with 17���22 ��m tylenchoid spicules, adcloacal bursa with smooth margin and tail similar to that of female. Morphologically, the new species is similar to five known Malenchus species: M. fusiformis (Thorne & Malek, 1968) Siddiqi, 1979, M. machadoi (Andr��ssy, 1963) Andr��ssy, 1968, M. pachycephalus Andr��ssy, 1981, M. solovjovae Brzeski, 1989 and M. undulatus but most closely resembles M. pachycephalus forming a morphologically-similar cryptic species pair. Compared to M. fusiformis, a poorly known species of the genus described based only on one female and only known from its type locality (Geraert 2008), the new species has a longer body (439���489 vs 350 ��m), narrower annuli (2.1���2.3 ��m wide at mid-body vs 3 ��m), posteriorly directed stylet knobs (vs rounded) and lacks vulval membranes (vs having obscure lateral membranes). Compared to M. machadoi, the new species has a shorter body (439���489 vs 520���560 ��m), lip region continuous with body contour (vs markedly narrower), annuli 2.1���2.3 ��m wide at mid-body (vs 1.6���1.8 ��m) and median bulb with distinct valves (vs very small or indistinct valves, as described by Andr��ssy 1968). Compared to M. pachycephalus, besides differences in partial SSU and LSU rDNA sequences and divergent placement in the corresponding phylogenetic trees, the new species differs in the nature of the median bulb (welldeveloped with valvular plates vs very weakly developed, without valve), stylet knobs (symmetrical vs slightly asymmetrical), lateral field margin (smooth vs crenate), pharynx length (93���108 vs 86���90 ��m) and lacks vulval flaps (vs small dikes present). Furthermore, the raised lateral fields of the new species are separated from the rest of the body by a sharp constriction (Fig. 3B, arrows); this demarcation is lacking in M. pachycephalus as illustrated by Qing et al. (2017). Compared to M. solovjovae, the new species has a well-developed stylet and corresponding knobs and welldeveloped valve in the median bulb. Compared to M. undulatus, besides differences in SSU and LSU rDNA sequences and distant positions in the corresponding phylogenetic trees, the new species has annuli 2.1���2.3 ��m wide (vs 1.7���1.9), plain lateral field margins (vs deeply crenate), lip region continuous with body contour (vs distinctly narrower), longer stylet (11���13 vs 9.5���10.0 ��m), more posterior vulva (V = 66.0���69.5 vs 63���66) and lacks vulval dikes (vs having dikes 3 annuli long). Compared to M. neosulcus Geraert & Raski, 1986, its sister taxon in our SSU rDNA phylogeny, the new species has 3���4 head annuli (vs 5���6), lateral field with plain margins (vs slightly crenate), well-developed median bulb, valve, and valve plates (vs weakly developed), wider body annuli (2.1���2.3 vs 1.8���2.0 ��m), shorter tail (65���87 vs 86���96 ��m), with smaller c' (5.9���8.1 vs 8���10) and greater c (5.4���6.8 vs 4.7���5.5) values, and differences in SSU sequence (accession numbers KX907138 for the new species and KJ869330 for M. neosulcus). Compared to M. labiatus, its sister taxon in our LSU rDNA phylogeny, the new species has a longer stylet (11��� 13 vs 8���9 ��m), greater V value (66.0���69.5 vs 59���63), shorter tail (65���87 vs 98���112 ��m) and lacks both vulval flaps and a disc-like structure at the lip apex. Molecular Phylogeny. Partial sequencing of the SSU rDNA gene from the new species yielded a single fragment 1066 nt long. Blast searches using the sequence of this fragment revealed the highest coverage with two sequences of M. neosulcus (KJ869330) and M. acarayensis Andr��ssy, 1968 (KJ869331), with identities of 93% and 91%, respectively. A 98% identity was achieved for a short (74% coverage) sequence of M. pachycephalus. Two longer sequences (with 88% coverage) of M. pachycephalus (KX156286 and KX156286) had 92% identity with partial SSU rDNA of the new species. Lower identity and coverage percentages were returned for other sequences. A total of 98 species/isolates of Tylenchoidea ��rley, 1880, along with Aphelenchoididae Skarbilovich, 1947 and two panagrolaimid species as outgroup taxa were selected for our SSU rDNA-based phylogenetic analysis. This dataset comprises 1495 total characters, of which 739 are variable. Figure 4 presents the Bayesian phylogenetic tree inferred using this dataset. The major clade of Malenchus spp. in this tree (based on currently available SSU sequences for the genus, and designated M, in green) has full BPP and moderate (62%) ML BS support. The new species forms a well-supported clade (1.00 BPP / 98% MLBS) with M. neosulcus in this tree. The phylogenetic relationship of these two species (M. geraerti n. sp. and M. neosulcus) with M. acarayensis, M. pachycephalus and an unidentified isolate of Malenchus sp. (KX156289) is not further resolved due to polytomy. Partial sequencing of the LSU rDNA gene (D2/D3 region) from two females of the new species yielded single fragments of 650 (isolate1) and 654 (isolate 2) nt long. A Blast search using these fragments��� sequences indicated no significant identity with any of the currently available sequences deposited in GenBank. A total of 90 species/ isolates of Tylenchoidea (ingroup members) and Aphelenchoididae (as outgroup taxa) were selected for our LSU rDNA gene (D2/D3 region) phylogenetic analysis. This dataset contains 468 total characters, of which 297 are variable. Figure 5 presents the Bayesian tree inferred using this dataset. Species of the genus Malenchus occur in two major clades, A and B, in this tree. Clade A also includes the ambiguous species Filenchus balcarceanus Torres & Geraert, 1996 (sensu Qing et al. 2017), which shares characters of both Filenchus Andr��ssy, 1954 and Malenchus. The monophyly of this clade is highly supported (1.00 BPP / 81% MLBS). Clade B is a strongly supported (1.00 BPP / 99% MLBS) monophyletic group including our new, three known, and three unidentified isolates/species of Malenchus, along with two species of Lelenchus Andr��ssy, 1954. The new species clusters in a subclade containing two species of Lelenchus (KU234169, KP730042, KX156335) and M. labiatus (KP730047)., Published as part of Pedram, Majid, Soleymanzadeh, Mahya, Pourjam, Ebrahim & Mobasseri, Mahyar, 2018, Observations on MaleNChus geraerti n. sp. (Rhabditida: Tylenchidae), a morphological and molecular phylogenetic study, pp. 406-418 in Zootaxa 4369 (3) on pages 408-412, DOI: 10.11646/zootaxa.4369.3.6, http://zenodo.org/record/1135450, {"references":["Thorne, G. & Malek, R. B. (1968) Nematodes of the Northern Great Plains. Part I. Tylenchida (Nemata: Secernentea). South Dakota Agricultural Experiment Station Technical Bulletin 31, 111 pp.","Siddiqi, M. R. (1979) Seven new species in a new nematode subfamily Duosulciinae (Tylenchidae), with proposals for Duosulcius gen. n., Zanenchus gen. n. and Neomalenchus gen. n. Nematologica, 25, 215 - 236. https: // doi. org / 10.1163 / 187529279 X 00244","Andrassy, I. (1963) Freilebende Nematoden aus Angola, I. Einige moosbewohnende Nematoden. Publicacoes Culturais da Companhia de Diamantes de Angola, 66, 55 - 80.","Andrassy, I. (1968) Fauna Paraguayensis. 2. Nematoden aus den Galeriewaldern des Acaray Flusses. Opuscula Zoologica Budapestensis, 8, 167 - 315.","Andrassy, I. (1981) Genera and species of the family Tylenchidae Orley, 1880 (Nematoda). The genus Malenchus Andrassy, 1968. Acta Zoologica Academiae Scientiarum Hungaricae, 27, 1 - 47.","Brzeski, M. W. (1989) Malenchus parvus sp. n., M. solovjovae sp. n. and observations on M. leiodermis (Nematoda, Tylenchidae). Nematologica 34, 47 - 56. https: // doi. org / 10.1163 / 002825988 X 00035","Geraert, E. (2008) The Tylenchidae of the World. Identification of the Family Tylenchidae (Nematoda). Ghent, Belgium, Akademia Press.","Qing, X., Decraemer, W., Claeys, M. & Bert, W. (2017) Molecular phylogeny of Malenchus and Filenchus (Nematoda: Tylenchidae). Zoologica Scripta, 0 0, 1 - 12 https: // doi. org / 10.1111 / zsc. 12236","Geraert, E. & Raski, D. (1986) Unusual Malenchus species (Nematoda: Tylenchidae). Nematologica, 32, 27 - 55. https: // doi. org / 10.1163 / 187529286 X 00020","Orley, L. (1880) Monographie der Anguilluliden. Termeszettudomanyi. Fuzetek, Budapest, 4, 165 pp.","Skarbilovich, T. S. (1947) [Revision of the systematics of the nematode family Anguillulinidae Baylis and Daubney, 1926.] Doklady Akademii Nauk SSR 57, 307 - 308. [In Russian]","Torres, M. S. & Geraert, E. (1996) Tylenchidae from Buenos Aires, Argentina. Nematologica, 42, 42 - 61. https: // doi. org / 10.1163 / 187529296 X 00058","Andrassy, I. (1954) Revision der gattung Tylenchus Bastian, 1865 (Tylenchidae, Nematoda). Acta Zoologica Academic Scientiarum Hungaricae 1, 5 - 42."]}

Enchodorus yeatsi n. sp., a new species of the rare genus Enchodorus is described and illustrated based on its morphological and molecular characters. It was recovered from southern Iran. Females of the new species are characterized by having 1,511.3- to 1,792.5-mm long slender body, angular lip region having prominent papillae, 12- to 13-mm long odontostyle, double guiding ring, simple rod-like odontophore, didelphic–amphidelphic reproductive system, and 102- to 160-mm long elongate-conoid tail with rounded tip. Males of the new species are abundant and functional, characterized by 1,484- to 1,576-mm long body, 40- to 43-mm long spicules, 5 to 6 ventromedian supplements, and ventrally bent elongate conical tail. Compared to the type species, Enchodorus dolichurus, the new species has differences in its tail morphology and V value. These morphological differences and the separation of two species was further supported with basic differences in sequences of 28S rDNA D2/D3 and internal transcribed spacer 1 (ITS1) fragments. Compared to Enchodorus neodolichurus, it has basic differences in tail characters and spicule lengths. Molecular phylogenetic studies using partial sequences of 28S rDNA D2/D3 fragment of the new species and available sequences of Nordiidae members and several other dorylaim species/genera, revealed E. yeatsi n. sp. and E. dolichurus forming a clade with 0.81 Bayesian posterior probability (BPP). This clade forms a sister clade to the clade of Heterodorus sp. and Rhyssocolpus vinciguerrae, again with 0.81 BPP. In ITS1 tree, reconstructed using few available sequences, the new species and E. dolichurus formed a clade with 0.98 BPP.

Data on five known species of the genus Coslenchus are provided. Morphological and morphometric data are given for all studied species. Three of the recovered species were also characterized by molecular phylogenetic data. The species C. leiocephalus was studied for the first time since its original description. Males of the species, C. franklinae and C. oligogyrus were described for the first time and the species C. oligogyrus was reported from Iran for the first time. In molecular phylogenetic studies based on partial sequences of 28S rDNA D2/D3 fragments, all species formed a clade with high Bayesian posterior probability in Bayesian inference, indicating the monophyly of the genus. The clade of Coslenchus spp. formed a highly supported monophyletic group, a sister clade to two species of the genus Aglenchus.

Discopersicus iranicus n. gen., n. comb., previously described from Iran as a new species under the genus Discotylenchus, is illustrated using light microscope and scanning electron microscope (SEM) observations and further studied using molecular characters. SEM studies revealed the newly proposed genus has oblique amphidial apertures on the lateral sides of the lip region. SEM images are also provided for two species of Discotylenchus, namely D. discretus and D. brevicaudatus, as the first SEM study of the genus. These results confirmed longitudinal amphidial aperture type on lateral sides of the lip region in genus Discotylenchus, as noted by Siddiqi while erecting the genus with D. discretus as the type species. Molecular phylogenetic analyses using partial small subunit (SSU) and large subunit (LSU) rDNA sequences revealed the affinity of the genus Discopersicus n. gen. with members of the subfamily Boleodorinae, as supported by morphological characters (mainly, the oblique amphidial opening).

Cryptaphelenchus varicaudatus n. sp. is described and illustrated. It was isolated from bark samples of dead or dying pine (Pinus spp.) trees with bark beetle frass and galleries, in Tehran province. The new species has 275- to 367-mm-long females (amedium-sized species) with distinctly annulated cuticle having three bands in lateral fields, lip region continuous with body contour, delicate knobbed stylet, monodelphic–prodelphic reproductive system with distinct spermatheca, short postvulval uterine sac (PUS), transverse vulval slit with raised posterior lip and body narrowing behind it, sclerotized vagina, simple intestine ending in a blind sac, having no rectum but vestigial anus in some specimens, and distal body end tip (tail tip) with variation in morphology (shape), from sharply or slightly pointed to bluntly rounded. Males of the new species are common, but less frequent than females, characterized by shorter body (235–278 mm long) compared to females, their posterior body end more ventrally bent, arcuate separate spicules with well-developed wide condyles, distinct rostrum having sharp, attenuated tip. The precloacal single supplement (P1) and the distally located pair of caudal papillae close to tail tip were only observed. The new species is morphologically compared with the species of the genus having short PUS and similar body end morphology. In molecular phylogenetic analyses using 1520- and 698-nt-long sequences of small subunit (SSU) and large subunit (LSU) rDNA D2/D3 fragments, the new species formed a clade with two currently available GenBank-derived, unspecified isolates/sequences in SSU and three other isolates/sequences in LSU trees, respectively.

Iranian population of Parasitorhabditis obtusa (Fuchs, 1915) Chitwood & Chitwood, 1950 (Figs. 1; 2) Measurements. See Table 1. ��� continued on the next page Female. Straight to ventrally accurate after heat killing. Cuticle smooth under LM, lacking discernible lateral fields in cross section. Cephalic region continuous with body contour, anteriorly flattened, cephalic framework weak, lips not protruding. Amphidial apertures hardly visible. Stoma well developed, promesostom cylindrical, devoid of teeth, with slight bifurcation at base of the cheilostom with straight parallel walls. Esophastome short. Pharynx rhabditoid, with muscular procorpus, slightly swollen at the junction with isthmus, the latter elongate. Basal bulb pyriform, set off, its valve well-developed. Cardia distinct, intestine simple and anus functional. Excretory pore at the level with the anterior part of pharyngeal bulb or at the middle of isthmus. Nerve ring surrounding middle of the isthmus, anterior to excretory pore, 121-150 ��m from anterior end. Reproductive system monodelphic-prodelphic, occupying 41-64% of the total body length and composed of a reflexed ovary with oocytes arranged usually in two rows. Mature eggs at distal end of genital tract measuring ca. 24 �� 41 ��m. Vagina anteriorly directed, about one third corresponding body diameter long. Vulva a transverse slit without flap or epiptygma. Tail short, conoid, with blunt or narrow tip, with or without a spike or a projection. Male. Common, as frequent as females. General morphology similar to that of female (slightly shorter), except for characters-states associated with sexual differences. Testis single, the genital branch occupies 47-66% of total body length. Spermatocytes arranged in multiple rows. Spicules separated, with straight blade and very short curved head. Gubernaculum simple, rod-shape. Bursa well developed, peloderan, with smooth margin. Genital papillae generally eight or nine pairs in number. Tail conoid with finely rounded tip. Locality and habitat. Recovered from galleries in pine tree barks in different regions of the city of Tehran (Table 2). The described population belongs to the population recovered from the bark of a dead pine trees in Lavizan region. The live females and males were also obtained from the inside of dissected bodies of Orthotomicus erosus individuals collected in association with dead pine trees., Published as part of Valizadeh, Ahmad, Goldasteh, Shila, Rafiei-Karahroodi, Zahra & Pedram, Majid, 2017, First record of the genus Parasitorhabditis Fuchs, 1937 (Rhabditida, Nematoda) from Iran with notes on morphological and molecular characters of the Iranian population of P. obtusa (Fuchs, 1915) Chitwood & Chitwood, 1950, pp. 591-600 in Zootaxa 4353 (3) on pages 592-597, DOI: 10.11646/zootaxa.4353.3.13, http://zenodo.org/record/1066091, {"references":["Fuchs, A. G. (1915) Die Naturgeschichte der Nematoden und einiger anderer Parasiten. 1. Des Ips typographus L. 2. Des Hylobius abietis L. Zool. Jb. (Syst.), 38, 109 - 222.","Chitwood, B. G. & Chitwood, M. B. (1950) An introduction to nematology. Section I. Anatomy. An introduction to nematology. Section I. Anatomy. Baltimore, USA, Monumental Printing Co, 213 pp.","Massey, C. L. (1956) Nematode parasites and associates of the Engelmann spruce beetle (Dendroctonus engelmanni Hopk.)."]}

Valizadeh, Ahmad, Goldasteh, Shila, Rafiei-Karahroodi, Zahra, Pedram, Majid (2017): First record of the genus Parasitorhabditis Fuchs, 1937 (Rhabditida, Nematoda) from Iran with notes on morphological and molecular characters of the Iranian population of P. obtusa (Fuchs, 1915) Chitwood & Chitwood, 1950. Zootaxa 4353 (3): 591-600, DOI: https://doi.org/10.11646/zootaxa.4353.3.13

Adeldoost, Yaser, Heydari, Ramin, Pedram, Majid (2016): YASER ADELDOOST, RAMIN HEYDARI & MAJID PEDRAM (2015) Morphological and molecular characterization of Tylencholaimellus persicus sp. n. (Dorylaimida: Tylencholaimellidae) from Iran. Zootaxa, 4040 (1): 074-082. Zootaxa 4066 (5): 600-600, DOI: 10.11646/zootaxa.4066.5.8

Rotylenchus castilloi n. sp. (Figs 1–4) Measurements. See Table 1. Female: Body habitus open- to closed-C in shape, varying to closed-J or spiral after heat relaxation. Cuticle without longitudinal striations, annuli 2.6–3.3 (2.0± 0.3) wide at mid-body. Lateral field with four smooth equidistant lines, beginning anteriorly at annuli 8–10 as three lines (forming two bands), after 8–11 annuli, central line dividing to form a third band, three bands 11–14 (14.0± 1.5) wide at mid-body, approximately one-fifth as wide as body diameter, external bands regularly areolated in pharynx region, but not on rest of the body. Lip region hemispherical, continuous with body contour, with 7–8 annuli under LM and irregular corncob-like appearance under SEM, 13–16 (14.6 ±1.0) Μm wide, 10–12 (10.8 ± 0.9) Μm high. Labial disc distinct, rounded to somewhat rectangular, separated from rest of lip region, but not elevated. Labial framework strong, outer margin extending 2–3 annuli posteriorly into the body. Stylet well developed, 4–5 times longer than labial region diameter, conus 51.5 –54.0% of total length, basal knobs rounded, slightly flattened anteriorly, 9–11 Μm wide. Dorsal pharyngeal gland orifice at ca. one tenth stylet length behind the knobs. Procorpus cylindrical, 51–68 (58.7 ± 5.4) Μm long, median pharyngeal bulb well developed, oval, 22–30 Μm long and 17–20 Μm diameter, located at 69.1–76.3 (71.6 ± 2.2)% of pharynx length, pharyngeal glands sacciform, overlapping intestine dorsally. Nerve ring enveloping isthmus at mid-point, 152–184 (165.5 ± 8.3) Μm from anterior end. Excretory pore distinct. Hemizonid 1.5 –2.0 annuli long, 1–3 annuli anterior to excretory pore. Reproductive system with two genital branches equally developed, anterior branch 450–660 (576 ± 91.5) Μm long, posterior branch 475–700 (569 ± 84.2) Μm long, ovaries with single row of oocytes, spermatheca rounded, 22–33 Μm in diameter, functional, with spherical sperm. Vulva slightly posterior to mid-body, with protruding double epiptygma. Phasmid pore-like, its location varying from 2–12 Μm anterior, to 2–10 Μm posterior to anus or at the level of anus (1-6 annuli anterior to anus or 1-4 annuli posterior to it). Tail short, variable in shape from rounded to dorsally convex-conoid, rarely bi-lobed, with 8–12 annuli, 0.5–0.7 anal body diameter long, terminus annulated. Male: Common, approximately as abundant as females. Morphology similar to that of female except for sexual dimorphism. Body habitus slightly curved ventrally after heat relaxation. Lip region 11–13 (12.0± 0.7) Μm in diameter, 8–11 (9.4 ± 0.8) Μm high. Lateral field 8–12 (10.2 ±1.0) Μm wide at mid-body, ca one-fifth as wide as body diameter. Stylet and stylet knobs less developed compared to those of female, knobs 6–9 Μm wide. Pharyngeal glands overlapping intestine dorsally. Testis single, anteriorly outstretched, 467–849 (713 ± 127) Μm long. Phasmid distinct, its location varying from 4 Μm anterior to cloaca to 11 Μm posterior to it. Spicules slightly cephalated, ventrally arcuate. Gubernaculum non-protrusible, distal part bent backward. Tail conoid, nearly 1.3–1.8 times cloacal region diameter with hyaline terminus. Bursa well developed, 74–90 Μm long, crenate, enveloping tail terminus. Type locality and habitat. The typel population of Rotylenchus castilloi n. sp. was recovered from a soil sample collected around the rhizosphere of oriental beech tree (Fagus orientalis Lipsky) in Asalem forests, Ardebil province, northwestern Iran (GPS coordinates: N: 37 ° 42 ' 37.373 ", E: 48 ° 53 ' 44.363 ") by Farzad Aliramaji in November, 2013. A second population was recovered from soil samples collected from the rhizosphere of forest trees in Golestan province, northern Iran, during 2014 by Atefeh Talezari. Type specimens. Holotype female, four female, and six male paratypes deposited in the Nematode Collection of the Faculty of Agriculture, Tarbiat Modares University, Tehran, Iran. Two female and two male paratypes deposited at each of the following collections: CABI Europe-UK, Egham, Surrey, UK; USDA Nematode Collection, Beltsville, MD, USA; and WANECO collection, Wageningen (http://www.waneco.eu/). Etymology. The new species is named after Prof. Pablo Castillo for his contribution to the taxonomy of the genus Rotylenchus. Diagnosis and relationships. Rotylenchus castilloi n. sp. is a bisexual species assigned to the species group having a hemispherical lip region, lateral field areolated only in pharynx region, stylet longer than 41 µm and rounded female tail (Castillo & Vovlas 2005). The new species is characterised by having a hemispherical and continuous lip region with irregular corncob-like appearance in SEM, very long stylet (62–68 Μm) vulva located at 49.7–62.2 % with a protruding double epiptygma, rounded to convex-conoid, rarely bi-lobed tail with 8–12 annuli and specific sequences of D 2 -D 3 segments of 28 S and ITS 1 -rRNA genes. Table 1. Morphometrics of Rotylenchus castilloi n. sp. from Iran. All measurements are in μm and in the form: mean ± s.d. (range) a. Abbreviations are defined in Siddiqi (2000). Morphologically, Rotylenchus castilloi n. sp. can be distinguished from all similar species within the genus by its matrix code (as described by Castillo & Vovlas 2005) as follows: A 6, B 1, C 1, D 4, E 4, F 2, G 5, H 2, I 2, J 1, K 2. The new species is morphologically similar to four known species of the genus: R. mesorobustus Zancada, 1985; R. magnus Zancada 1985; R. cazorlaensis Castillo & Gómez Barcina, 1987 and R. jaeni (Castillo, Vovlas, Gómez Barcina & Lamberti, 1993) Vovlas, Subbotin, Troccoli, Liébanas & Castillo, 2008. It differs from R. mesorobustus by its longer body (2258–2919 vs 1300–1480 Μm), shape of the lip region (hemispherical and irregular corncob-like appearance in SEM vs rounded with regular blocks), longer stylet (62–68 vs 47.5–49.5 Μm) and tail (18–30 vs 9.0– 15.5 Μm) and the location of phasmid (varying from anterior to posterior of anus vs 5–8 annuli anterior to anus). The new species differs from R. magnus by its body length (2258–2919 vs 1520–2140 Μm), continuous lip region (vs set off), location of phasmid (varying from anterior to posterior to to anus vs always anterior to anus) and presence of male (vs absence). R. castilloi n. sp. differs from R. cazorlaensis by its body length (2258–2919 vs 1440–2280 Μm), shape of labial region (hemispherical vs truncate), stylet length (62–68 vs 46.5–56.5 Μm), distance of dorsal pharyngeal gland orifice from stylet base (6–8 vs 8.5–11.5 Μm) and location of phasmid (varying from anterior to posterior to anus vs at the level of anus to posterior). When compared with R. jaeni, the new species differs by its body length (2258–2919 vs 1500–2200 Μm), lip region characters (continuous vs set off) and location of phasmid (varying from anterior to posterior to anus vs 0–6 annuli anterior to anus). Molecular phylogenetic relationships. Amplification of the partial D 2 -D 3 expansion segments of 28 S and ITS 1 -rRNA genes for the new species yielded two fragments of 721 and 750 bp long, respectively. The D 2 -D 3 dataset consisted of 33 species/isolates with 577 total characters (after trimming); of these 320 characters were conserved and 253 characters were variable. The average nucleotide composition was as follows: 15.3 % A, 26.2 % C, 36.8 % G and 21.7 % T. The results (number and percentage of similarity and gaps) of pairwise alignment of the new species with the morphologically compared species using D 2 -D 3 region are presented in Table 2. Fig. 5 presents the phylogenetic tree inferred using the D 2 -D 3 dataset. Using Tylenchorhynchus claytoni (EU 368589) as outgroup, two major clades 1 and 2 were inferred. The major clade 1 consisted of several species of Helicotylenchus Steiner, 1945 in a highly supported clade (1.00/ 99) and the major clade 2 (with weak support 0.66 / -) which is divided into two subclades, A and B. The subclade 2 A includes several Rotylenchus spp. and the new species. The species R. striaticeps (Vovlas, Castillo & Lamberti, 1993) Siddiqi, 2000 occupies the most basal position (supported only by Bayesian analysis), but the second most basal species within subgroup 2 A cannot be inferred due to polytomy. In other words, we cannot determine which species is the most closely related to R. striaticeps. This observation can be compared to the phylogenetic relationship of the latter species in our ITS 1 tree (Fig. 6), in which the phylogenetic relation with Helicotylenchus spp. is well supported (1.00/ 96). The subclade 2 B includes five species/isolates of Helicotylenchus, as a sister clade to Rotylenchus dalikhaniensi s. Although the new species and the three morphologically similar species (R. cazorlaensis, R. jaeni and R. magnus) are part of subclade 2 A, the relationships among these four species are not fully resolved due to polytomy. Currently there are no molecular data for R. mesorobustus. Future sequencing of other morphologically similar species (for example, species with long stylet) may provide more insights on phylogenetic relationships of this group of species. The close relationship of the new species with R. eximius Siddiqi, 1964 (although with low branch supports) is surprising as these species differ in some morphological characters such as stylet length and lip region conformation. New species Morphologically compared species/ Similarity Gaps isolates Rotylenchus castilloi n. sp. (KM 251416) Rotylenchus cazorlaensis (EU 373669) 408 / 682 (59.8 %) 193 / 682 (28.3 %) Rotylenchus castilloi n. sp. (KM 251416) Rotylenchus cazorlaensis (EU 373670) 412 / 682 (60.4 %) 191 / 682 (28.0%) Rotylenchus castilloi n. sp. (KM 251416) Rotylenchus cazorlaensis (EU 373671) 415 / 682 (60.9 %) 191 / 682 (28.0%) Rotylenchus castilloi n. sp. (KM 251416) Rotylenchus magnus (EU 373676) 376 / 675 (55.7 %) 236 / 675 (35.0%) Rotylenchus castilloi n. sp. (KM 251416) Rotylenchus magnus (EU 373659) 376 / 675 (55.7 %) 236 / 675 (35.0%) Rotylenchus castilloi n. sp. (KM 251416) Rotylenchus magnus (EU 373659) 392 / 671 (58.4 %) 210 / 671 (31.3 %) Rotylenchus castilloi n. sp. (KM 251416) Rotylenchus magnus (EU 373660) 391 / 670 (58.4 %) 209 / 670 (31.2 %) Rotylenchus castilloi n. sp. (KM 251416) Rotylenchus magnus (EU 373665) 391 / 670 (58.4 %) 209 / 670 (31.2 %) Rotylenchus castilloi n. sp. (KM 251416) Rotylenchus jaeni (EU 373661) 379 / 687 (55.2 %) 250 / 687 (36.4 %) Rotylenchus castilloi n. sp. (KM 251416) Rotylenchus jaeni (EU 373662) 378 / 687 (55.0%) 250 / 687 (36.4 %) The ITS 1 dataset consisted of 43 species/isolates of Rotylenchus, three species of Helicotylenchus and one belonolaim outgroup taxon. The dataset had 836 total characters (after manually trimming) of which 138 characters were conserved and 693 characters were variable. The average nucleotide composition was as follows: 19.6 % A, 26.6 % C, 30.3 % G and 23.8 % T. The results (number and percent of similarity and gaps) of pairwise alignment of the new species with the morphologically compared species using ITS 1 region are presented in Table 3. Fig. 6 presents the phylogenetic tree inferred using the above mentioned dataset. Using Tylenchorhynchus zeae (EF 519711) as outgroup, three major clades were inferred. Clades 1 and 2 include several species of Rotylenchus spp., including the new species in major clade 1, forming a monophyletic group with R. unisexus Sher, 1965 with high (1.00/ 95) support. There are some common morphological characters like shape of lip region, tail in female and presence of double epiptygma which might support affinity of these two species, while both species have basic differences in some key characters like the length of the stylet. Based on current molecular knowledge, the paraphyly of the family Hoplolaimidae and the genus Rotylenchus is well documented (Atighi et al. 2011, 2014; Vo v l a s et al. 2008; Cantalapiedra-Navarrete et al. 2013). Accordingly, we believe more species of the genus need to be sequenced before we can resolve the phylogenetic relationships among the species of the genus Rotylenchus and between the genera Rotylenchus and Helicotylenchus.

Miraeiz, Esmaeil, Heydari, Ramin, Álvarez-Ortega, Sergio, Pedram, Majid, Atighi, Mohammad Reza (2015): Molecular and morphological characterization of Veleshkinema iranicum n. gen., n. sp. (Nematoda: Hexatylina, Sphaerularioidea) from Iran. Zootaxa 4000 (5): 531-546, DOI: http://dx.doi.org/10.11646/zootaxa.4000.5.3

Rotylenchus castilloi n. sp. (Figs 1���4) Measurements. See Table 1. Female: Body habitus open- to closed-C in shape, varying to closed-J or spiral after heat relaxation. Cuticle without longitudinal striations, annuli 2.6���3.3 (2.0�� 0.3) wide at mid-body. Lateral field with four smooth equidistant lines, beginning anteriorly at annuli 8���10 as three lines (forming two bands), after 8���11 annuli, central line dividing to form a third band, three bands 11���14 (14.0�� 1.5) wide at mid-body, approximately one-fifth as wide as body diameter, external bands regularly areolated in pharynx region, but not on rest of the body. Lip region hemispherical, continuous with body contour, with 7���8 annuli under LM and irregular corncob-like appearance under SEM, 13���16 (14.6 ��1.0) ��m wide, 10���12 (10.8 �� 0.9) ��m high. Labial disc distinct, rounded to somewhat rectangular, separated from rest of lip region, but not elevated. Labial framework strong, outer margin extending 2���3 annuli posteriorly into the body. Stylet well developed, 4���5 times longer than labial region diameter, conus 51.5 ���54.0% of total length, basal knobs rounded, slightly flattened anteriorly, 9���11 ��m wide. Dorsal pharyngeal gland orifice at ca. one tenth stylet length behind the knobs. Procorpus cylindrical, 51���68 (58.7 �� 5.4) ��m long, median pharyngeal bulb well developed, oval, 22���30 ��m long and 17���20 ��m diameter, located at 69.1���76.3 (71.6 �� 2.2)% of pharynx length, pharyngeal glands sacciform, overlapping intestine dorsally. Nerve ring enveloping isthmus at mid-point, 152���184 (165.5 �� 8.3) ��m from anterior end. Excretory pore distinct. Hemizonid 1.5 ���2.0 annuli long, 1���3 annuli anterior to excretory pore. Reproductive system with two genital branches equally developed, anterior branch 450���660 (576 �� 91.5) ��m long, posterior branch 475���700 (569 �� 84.2) ��m long, ovaries with single row of oocytes, spermatheca rounded, 22���33 ��m in diameter, functional, with spherical sperm. Vulva slightly posterior to mid-body, with protruding double epiptygma. Phasmid pore-like, its location varying from 2���12 ��m anterior, to 2���10 ��m posterior to anus or at the level of anus (1-6 annuli anterior to anus or 1-4 annuli posterior to it). Tail short, variable in shape from rounded to dorsally convex-conoid, rarely bi-lobed, with 8���12 annuli, 0.5���0.7 anal body diameter long, terminus annulated. Male: Common, approximately as abundant as females. Morphology similar to that of female except for sexual dimorphism. Body habitus slightly curved ventrally after heat relaxation. Lip region 11���13 (12.0�� 0.7) ��m in diameter, 8���11 (9.4 �� 0.8) ��m high. Lateral field 8���12 (10.2 ��1.0) ��m wide at mid-body, ca one-fifth as wide as body diameter. Stylet and stylet knobs less developed compared to those of female, knobs 6���9 ��m wide. Pharyngeal glands overlapping intestine dorsally. Testis single, anteriorly outstretched, 467���849 (713 �� 127) ��m long. Phasmid distinct, its location varying from 4 ��m anterior to cloaca to 11 ��m posterior to it. Spicules slightly cephalated, ventrally arcuate. Gubernaculum non-protrusible, distal part bent backward. Tail conoid, nearly 1.3���1.8 times cloacal region diameter with hyaline terminus. Bursa well developed, 74���90 ��m long, crenate, enveloping tail terminus. Type locality and habitat. The typel population of Rotylenchus castilloi n. sp. was recovered from a soil sample collected around the rhizosphere of oriental beech tree (Fagus orientalis Lipsky) in Asalem forests, Ardebil province, northwestern Iran (GPS coordinates: N: 37 �� 42 ' 37.373 ", E: 48 �� 53 ' 44.363 ") by Farzad Aliramaji in November, 2013. A second population was recovered from soil samples collected from the rhizosphere of forest trees in Golestan province, northern Iran, during 2014 by Atefeh Talezari. Type specimens. Holotype female, four female, and six male paratypes deposited in the Nematode Collection of the Faculty of Agriculture, Tarbiat Modares University, Tehran, Iran. Two female and two male paratypes deposited at each of the following collections: CABI Europe-UK, Egham, Surrey, UK; USDA Nematode Collection, Beltsville, MD, USA; and WANECO collection, Wageningen (http://www.waneco.eu/). Etymology. The new species is named after Prof. Pablo Castillo for his contribution to the taxonomy of the genus Rotylenchus. Diagnosis and relationships. Rotylenchus castilloi n. sp. is a bisexual species assigned to the species group having a hemispherical lip region, lateral field areolated only in pharynx region, stylet longer than 41 ��m and rounded female tail (Castillo & Vovlas 2005). The new species is characterised by having a hemispherical and continuous lip region with irregular corncob-like appearance in SEM, very long stylet (62���68 ��m) vulva located at 49.7���62.2 % with a protruding double epiptygma, rounded to convex-conoid, rarely bi-lobed tail with 8���12 annuli and specific sequences of D 2 -D 3 segments of 28 S and ITS 1 -rRNA genes. Table 1. Morphometrics of Rotylenchus castilloi n. sp. from Iran. All measurements are in ��m and in the form: mean �� s.d. (range) a. Abbreviations are defined in Siddiqi (2000). Morphologically, Rotylenchus castilloi n. sp. can be distinguished from all similar species within the genus by its matrix code (as described by Castillo & Vovlas 2005) as follows: A 6, B 1, C 1, D 4, E 4, F 2, G 5, H 2, I 2, J 1, K 2. The new species is morphologically similar to four known species of the genus: R. mesorobustus Zancada, 1985; R. magnus Zancada 1985; R. cazorlaensis Castillo & G��mez Barcina, 1987 and R. jaeni (Castillo, Vovlas, G��mez Barcina & Lamberti, 1993) Vovlas, Subbotin, Troccoli, Li��banas & Castillo, 2008. It differs from R. mesorobustus by its longer body (2258���2919 vs 1300���1480 ��m), shape of the lip region (hemispherical and irregular corncob-like appearance in SEM vs rounded with regular blocks), longer stylet (62���68 vs 47.5���49.5 ��m) and tail (18���30 vs 9.0��� 15.5 ��m) and the location of phasmid (varying from anterior to posterior of anus vs 5���8 annuli anterior to anus). The new species differs from R. magnus by its body length (2258���2919 vs 1520���2140 ��m), continuous lip region (vs set off), location of phasmid (varying from anterior to posterior to to anus vs always anterior to anus) and presence of male (vs absence). R. castilloi n. sp. differs from R. cazorlaensis by its body length (2258���2919 vs 1440���2280 ��m), shape of labial region (hemispherical vs truncate), stylet length (62���68 vs 46.5���56.5 ��m), distance of dorsal pharyngeal gland orifice from stylet base (6���8 vs 8.5���11.5 ��m) and location of phasmid (varying from anterior to posterior to anus vs at the level of anus to posterior). When compared with R. jaeni, the new species differs by its body length (2258���2919 vs 1500���2200 ��m), lip region characters (continuous vs set off) and location of phasmid (varying from anterior to posterior to anus vs 0���6 annuli anterior to anus). Molecular phylogenetic relationships. Amplification of the partial D 2 -D 3 expansion segments of 28 S and ITS 1 -rRNA genes for the new species yielded two fragments of 721 and 750 bp long, respectively. The D 2 -D 3 dataset consisted of 33 species/isolates with 577 total characters (after trimming); of these 320 characters were conserved and 253 characters were variable. The average nucleotide composition was as follows: 15.3 % A, 26.2 % C, 36.8 % G and 21.7 % T. The results (number and percentage of similarity and gaps) of pairwise alignment of the new species with the morphologically compared species using D 2 -D 3 region are presented in Table 2. Fig. 5 presents the phylogenetic tree inferred using the D 2 -D 3 dataset. Using Tylenchorhynchus claytoni (EU 368589) as outgroup, two major clades 1 and 2 were inferred. The major clade 1 consisted of several species of Helicotylenchus Steiner, 1945 in a highly supported clade (1.00/ 99) and the major clade 2 (with weak support 0.66 / -) which is divided into two subclades, A and B. The subclade 2 A includes several Rotylenchus spp. and the new species. The species R. striaticeps (Vovlas, Castillo & Lamberti, 1993) Siddiqi, 2000 occupies the most basal position (supported only by Bayesian analysis), but the second most basal species within subgroup 2 A cannot be inferred due to polytomy. In other words, we cannot determine which species is the most closely related to R. striaticeps. This observation can be compared to the phylogenetic relationship of the latter species in our ITS 1 tree (Fig. 6), in which the phylogenetic relation with Helicotylenchus spp. is well supported (1.00/ 96). The subclade 2 B includes five species/isolates of Helicotylenchus, as a sister clade to Rotylenchus dalikhaniensi s. Although the new species and the three morphologically similar species (R. cazorlaensis, R. jaeni and R. magnus) are part of subclade 2 A, the relationships among these four species are not fully resolved due to polytomy. Currently there are no molecular data for R. mesorobustus. Future sequencing of other morphologically similar species (for example, species with long stylet) may provide more insights on phylogenetic relationships of this group of species. The close relationship of the new species with R. eximius Siddiqi, 1964 (although with low branch supports) is surprising as these species differ in some morphological characters such as stylet length and lip region conformation. New species Morphologically compared species/ Similarity Gaps isolates Rotylenchus castilloi n. sp. (KM 251416) Rotylenchus cazorlaensis (EU 373669) 408 / 682 (59.8 %) 193 / 682 (28.3 %) Rotylenchus castilloi n. sp. (KM 251416) Rotylenchus cazorlaensis (EU 373670) 412 / 682 (60.4 %) 191 / 682 (28.0%) Rotylenchus castilloi n. sp. (KM 251416) Rotylenchus cazorlaensis (EU 373671) 415 / 682 (60.9 %) 191 / 682 (28.0%) Rotylenchus castilloi n. sp. (KM 251416) Rotylenchus magnus (EU 373676) 376 / 675 (55.7 %) 236 / 675 (35.0%) Rotylenchus castilloi n. sp. (KM 251416) Rotylenchus magnus (EU 373659) 376 / 675 (55.7 %) 236 / 675 (35.0%) Rotylenchus castilloi n. sp. (KM 251416) Rotylenchus magnus (EU 373659) 392 / 671 (58.4 %) 210 / 671 (31.3 %) Rotylenchus castilloi n. sp. (KM 251416) Rotylenchus magnus (EU 373660) 391 / 670 (58.4 %) 209 / 670 (31.2 %) Rotylenchus castilloi n. sp. (KM 251416) Rotylenchus magnus (EU 373665) 391 / 670 (58.4 %) 209 / 670 (31.2 %) Rotylenchus castilloi n. sp. (KM 251416) Rotylenchus jaeni (EU 373661) 379 / 687 (55.2 %) 250 / 687 (36.4 %) Rotylenchus castilloi n. sp. (KM 251416) Rotylenchus jaeni (EU 373662) 378 / 687 (55.0%) 250 / 687 (36.4 %) The ITS 1 dataset consisted of 43 species/isolates of Rotylenchus, three species of Helicotylenchus and one belonolaim outgroup taxon. The dataset had 836 total characters (after manually trimming) of which 138 characters were conserved and 693 characters were variable. The average nucleotide composition was as follows: 19.6 % A, 26.6 % C, 30.3 % G and 23.8 % T. The results (number and percent of similarity and gaps) of pairwise alignment of the new species with the morphologically compared species using ITS 1 region are presented in Table 3. Fig. 6 presents the phylogenetic tree inferred using the above mentioned dataset. Using Tylenchorhynchus zeae (EF 519711) as outgroup, three major clades were inferred. Clades 1 and 2 include several species of Rotylenchus spp., including the new species in major clade 1, forming a monophyletic group with R. unisexus Sher, 1965 with high (1.00/ 95) support. There are some common morphological characters like shape of lip region, tail in female and presence of double epiptygma which might support affinity of these two species, while both species have basic differences in some key characters like the length of the stylet. Based on current molecular knowledge, the paraphyly of the family Hoplolaimidae and the genus Rotylenchus is well documented (Atighi et al. 2011, 2014; Vo v l a s et al. 2008; Cantalapiedra-Navarrete et al. 2013). Accordingly, we believe more species of the genus need to be sequenced before we can resolve the phylogenetic relationships among the species of the genus Rotylenchus and between the genera Rotylenchus and Helicotylenchus., Published as part of Talezari, Atefeh, Pourjam, Ebrahim, Kheiri, Ahmad, Li��banas, Gracia, Aliramaji, Farzad, Pedram, Majid, Rezaee, Saeed & Atighi, Mohammad Reza, 2015, Rotylenchus castilloi n. sp. (Nematoda: Hoplolaimidae), a new species with long stylet from northern Iran, pp. 88-100 in Zootaxa 3931 (1) on pages 90-95, DOI: 10.11646/zootaxa.3931.1.6, http://zenodo.org/record/232446, {"references":["Castillo, P. & Vovlas, N. (2005) Bionomics and identification of the genus Rotylenchus (Nematoda: Hoplolaimidae). Nematology Monographs and Perspectives. Vol. 3. Brill Academic Publishers, Leiden, 377 pp. (Series editors: Hunt, D. J. & Perry, R. N.)","Siddiqi, M. R. (2000) Tylenchida: Parasites of Plants and Insects. 2 nd Edition. CABI Publishing, Wallingford, 833 pp. [UK]","Zancada, M. C. (1985) Rotylenchus magnus sp. n. and R. mesorobustus sp. n. (Nematoda: Tylenchida) from Spain. Nematologica, 31, 134 - 142.","Castillo, P. & Gomez Barcina, A. (1987) Rotylenchus cazorlaensis sp. n. and new record of R. fallorobustus Sher, 1965 (Nematoda: Tylenchida) from south-eastern Spain. Nematologica, 33, 393 - 400. http: // dx. doi. org / 10.1163 / 187529287 X 00056","Vovlas, N., Subbotin, S. A., Troccoli, A., Liebanas, G. & Castillo, P. (2008) Molecular phylogeny of the genus Rotylenchus (Nematoda, Tylenchida) and description of a new species. Zoologica Scripta, 37, 52 - 537. http: // dx. doi. org / 10.1111 / j. 1463 - 6409.2008.00337. x","Siddiqi, M. R. (1964) Rotylenchus eximius n. sp. (Nematoda: Hoplolaiminae) found around almond roots in Tunisia. Nematologica, 10, 101 - 105. http: // dx. doi. org / 10.1163 / 187529264 X 00682","Sher, S. A. (1965) Revision of the Hoplolaimidae (Nematoda) V. Rotylenchus Filipjev, 1936. Nematologica, 11, 173 - 198. http: // dx. doi. org / 10.1163 / 187529265 X 00041","Atighi, M. R., Pourjam, E., Pedram, M., Cantalapiedra-Navarrete, C., Palomares-Rius, J. E. & Castillo, P. (2011) Molecular and morphological characterisations of two new Rotylenchus species from Iran. Nematology, 13, 951 - 964. http: // dx. doi. org / 10.1163 / 138855411 X 571795","Atighi, M. R., Pourjam, M., Ghaemi, R., Pedram, M., Liebanas, G., Cantalapiedra-Navarrete, C., Castillo, P. & Palomares-Rius, J. (2014) Description of Rotylenchus arasbaranensis n. sp. from Iran with discussion on the taxonomic status of Plesiorotylenchus Vovlas, Castillo & Lamberti, 1993 (Nematoda: Hoplolaimidae). Nematology, 16, 1019 - 1045. http: // dx. doi. org / 10.1163 / 15685411 - 00002827","Cantalapiedra-Navarrete, C., Navas-Cortes, J. A., Liebanas, G., Vovlas, N., Subbotin, S. A., Palomares-Rius, J. E. & Castillo, P. (2013) Comparative molecular and morphological characterisations in the genus Rotylenchus: Rotylenchus paravitis n. sp., an example of cryptic speciation. Zoologischer Anzeiger, 252, 246 - 268. http: // dx. doi. org / 10.1016 / j. jcz. 2012.08.002"]}

Tylencholaimellus persicus sp. n. Figs 1 & 2 Measurements. Table 1. Description. Female. Slender nematodes of medium size, slightly curved ventrally upon fixation, tapering gradually towards anterior end. Cuticle dorylaimoid, two layered, outer layer bearing distinct transverse striations. Lateral chord one fourth to one third of mid-body width. Lip region expanded, separated from body contour by a sharp constriction, forming a large disk-like structure, 5.5 ���6.0 ��m wide or ca. 3 times wider than high. Amphidial fovea cup-shaped, its opening a transverse slit ca. one lip region width from anterior end. Odontostyle typical of the genus, ca. 1.8 times longer than odontophore, provided with dorsal stiffening piece covering entire dorsal arm, its aperture one-sixth of odontostyle length. Odontophore with basal knobs. Guiding ring at anterior half of odontostyle while resting. Pharynx composed of the anterior narrow and slender part connecting to the small pharyngeal bulb occupying ca. 20 % of the pharynx, its nuclei not clearly seen. Cardia hemispherical, 5.5���6.5 ��m long and 3.5���4.5 ��m wide. Nerve ring at 60���62 % of the neck length. Intestine coarsely granular. Prerectum 2.2 times, and rectum 0.9���1.2 times anal body diameter long, respectively. Reproductive system mono-opisthodelphic. Anterior genital branch reduced to an AUS, in length, occupying 7���8 % of body length and containing sperm. Posterior branch normal, poorly developed compared to reproductive system of some other dorylaim taxa (especially in the lack of a visible sphincter), 150���241 ��m long or 24.4���27.1 % of body length, composed of a tubular uterus 70���73 ��m long, a 60���65 ��m long oviduct swollen in proximal end and forming a less developed pars dilatata oviductus, containing sperm in almost all examined individuals, and a reflexed ovary, 79���102 ��m long. Sphincter not visible (and maybe undeveloped). Usually a maturing oocyte 40���60 ��m long in size was observed in distal end of the ovary. Vagina perpendicular to body axis, occupying less than half (40 %) of the corresponding body diameter, composed of pars proximalis vaginae ( 10���12 ��m long), short (1.5 ���2.0 ��m long), slightly curved pars distalis vaginae, and pars refringens vaginae apparently absent. Vulva a transverse slit. Tail broadly rounded with two caudal pores on each lateral side. TABLE I. Morphometrics of Tylencholaimellus persicus sp. n. from Iran, in ��m and in the form average �� SD (range). Male. General morphology similar to that of female, except in characters related to the reproductive system. Testes paired, opposed and spermatozoa spindle-shaped, 5.5���6.0��m long and 1 ��m wide. In addition to an adanal pair, there is only one ventromedian supplement outside the spicule range at a distance of 30 ��m anterior to cloacal opening. Spicules moderately sclerotised, dorsal contour irregularly convex, 5.0��� 5.2 times longer than width at wider part in proximal half, ventral contour with shallow hump, but distinct hollow, capitulum small, not offset, 2��� 3 ��m long and distal tip ca. 1.5 ��m wide. Tail dorsally convex with broadly rounded terminus. Diagnosis and relationships. Tylencholaimellus persicus sp. n. is characterized by its 613���885 ��m long body, expanded lip region, separated from body contour by a sharp constriction, forming a large disk-like structure, odontostyle and odontophore respectively 12���14 and 6.5 ���8.0 ��m long, female genital system mono-opisthodelphic with relatively long AUS, 50���80 ��m long, V index of 36.5���41.5, broadly rounded tail, abundant males in population with spicules 22���25 ��m long and one ventromedian copulatory supplement. The new species is compared with five known species of the genus having perioral disc, rounded tail and AUS namely T. coronatus Thorne, 1939, T. incertus Colomba & Vinciguerra, 1979, T. montanus Thorne, 1939, T. ozarkensis Goseco, Ferris, & Ferris, 1975 and T. projectus Siddiqi, 1964. Compared to these taxa, the new species has an expanded lip region separated from body contour by a sharp constriction, forming a large disk-like structure. Detailed comparisons with the above-mentioned species are discussed below. The new species differs from T. coronatus (data from Andr��ssy, 2009) by its shorter odontostyle (12���14 vs 16��� 18 ��m), posteriorly located vulva (V = 36.5���41.5 vs 30���36), smaller c�� ratio (1.1���1.3 vs 1.4���1.6), males with one (vs two) ventromedian supplement and broadly rounded tail end (vs conical, rounded terminus). Compared to T. incertus (probably the closest species, disregarding lip region morphology), the new species differs by its AUS not reaching the pharyngeal bulb and larger c�� [1.1���1.3 vs 0.83 (calculated from drawing)]. In comparison with T. montanus (lectotype and paralectotypes from Goseco et al., 1975), the new species basically differs by its longer odontostyle (12���14 vs 9.6 ��m), longer AUS (50���80 vs 6.4 ��m) and broadly rounded vs bluntly conoid tail end. Compared to T. ozarkensis, another closely related species, the new species has a slightly longer odontostyle (12���14 vs 11 ��m in holotype) and males with one ventromedian supplement (vs lacking). The new species differs from T. projectus (data from Goseco et al., 1975), by its longer basal bulb (30���39 ��m vs 28���32 ��m), more posteriorly located vulva (V = 36.5���41.5 vs 32���35) and broadly rounded tail end (vs conoid, rounded end). Furthermore, the new species is close in morphology to one species of the genus Margollus Pe��a-Santiago, Peralta & Siddiqi, 1993: M. bokanicus Pachideh, Niknam, Jabbari & Pe��a-Santiago, 2015 (although examined individuals of the new species lack the typical labial and postlabial sclerotization characteristic of the genus Margollus). These two taxa can be separated on the basis of the following characters: T. persicu s has an expanded lip region, separated from the rest of the body by a constriction, forming a large disk-like structure vs lip region more or less expanded (only one individual with expanded lip region was found in the type population of M. bokanicus); longer AUS (50���80 vs 26���29 ��m), differences in position of vulva (V = 36.5���41.5 vs 40���47) and the presence of functional males in population vs lacking for M. bokanicus., Published as part of Adeldoost, Yaser, Heydari, Ramin & Pedram, Majid, 2015, Morphological and molecular characterization of Tylencholaimellus persicus sp. n. (Dorylaimida: Tylencholaimellidae) from Iran in Zootaxa 4040 (1), DOI: 10.11646/zootaxa.4040.1.6, http://zenodo.org/record/236629, {"references":["Thorne, G. (1939) A monograph of the nematodes of the superfamily Dorylaimoidea. Capita Zoologica, 8, 1 - 261.","Colomba, G. & Vinciguerra, M. T. (1979) Nematodi d'aqua dolce della Sicilia. Animalia, 6, 89 - 120.","Goseco, C. G., Ferris, V. R. & Ferris, J. M. (1975) Revision in Leptonchoidea (Nematoda: Dorylaimida). Tylencholaimellus, Doryllium, Gerthus n. gen. and Agmodorus in Tylencholaimellidae; and Adenolaimus in Aulolaimoididae. Research Bulletin Agriculture Experimental Station, Purdue University, West Lafayette, No. 928, 1 - 40.","Siddiqi, M. R. (1964) Six new nematode species in the superfamily Dorylaimoidea from India. Labdev Journal of Science and Technology, 2, 136 - 144.","Andrassy, I. (2009) Free-living nematodes of Hungary III (Nematoda errantia). Pedozoologica Hungarica 5. Hungarian Natural History Museum and Systematic Zoology Research Group of the Hungarian Academy of Sciences, Budapest, 608 pp. [series editors: Csuzdi, C. & Mahunka, S.]","Pena-Santiago, R., Peralta, M. & Siddiqi, M. R. (1993) Taxonomy of some new and known species of the genus Tylencholaimellus with a proposal for Margollus gen. n. (Nematode: Dorylaimida). Nematologica, 39, 218 - 233. http: // dx. doi. org / 10.1163 / 187529293 X 00178","Pachideh, A., Niknam, G., Jabbari, H. & Pena-Santiago, R. (2015) Margollus bokanicus n. sp. from Iran, the fourth species of a rare nematode genus (Dorylaimida, Tylencholaimellidae). Journal of Nematology, 47, 67 - 70."]}

Adeldoost, Yaser, Heydari, Ramin, Pedram, Majid (2015): Morphological and molecular characterization of Tylencholaimellus persicus sp. n. (Dorylaimida: Tylencholaimellidae) from Iran. Zootaxa 4040 (1), DOI: http://dx.doi.org/10.11646/zootaxa.4040.1.6

Talezari, Atefeh, Pourjam, Ebrahim, Kheiri, Ahmad, Liébanas, Gracia, Aliramaji, Farzad, Pedram, Majid, Rezaee, Saeed, Atighi, Mohammad Reza (2015): Rotylenchus castilloi n. sp. (Nematoda: Hoplolaimidae), a new species with long stylet from northern Iran. Zootaxa 3931 (1): 88-100, DOI: http://dx.doi.org/10.11646/zootaxa.3931.1.6

Veleshkinema iranicum n. gen., n. sp. (Figs 1–5) Measurements. See Table 3. Description. Female. Free-living. Medium sized nematodes. Body mostly straight after heat relaxation, sometimes irregularly bent, slightly tapering towards both ends. Cuticle with fine transverse annulations, annuli about 1.2–1.7 Μm wide at mid-body. Lateral field barely visible by light microscopy, not raised, beginning anteriorly at the end of procorpus with three lines (forming two bands) increasing to six areolated equidistant incisures at mid-body, approximately one-fourth as wide as body diameter, gradually decreasing to four at tail region. Deirids at the level of excretory pore. Cephalic region low, anteriorly flat, continuous with body contour, 1- 2 µm high and 6–8 µm wide. Stylet delicate with asymmetrical minute knobs, conus shorter than shaft (28–38 % of total stylet length); DGO opens just posterior to subventral knob. Procorpus non-muscular, metacorpus less developed, isthmus slender, pharyngeal gland lobe dorsally (sometimes dorso-laterally) overlapping intestine about 0.5–2.1 times corresponding body diameter, cardia distinct. Nerve ring circum-pharyngeal, always anterior to excretory pore, located 62–99 µm from anterior end. Excretory pore with sclerotized duct. Hemizonid barely visible, just anterior to excretory pore (5–11 Μm anterior to excretory pore), 2–3 annuli long. Intestine with distinct lumen ending in a distinct rectum. Reproductive system monodelphic-prodelphic, occupying 41–70 % of the body length, ovary outstretched, sometimes reaching dorsal pharyngeal gland, with single row of oocytes. Oviduct cellular, spermatheca rounded, offset and filled with minute spheroid sperm (2–3 µm in diameter). Crustaformeria apparently quadricolumellate, with 8–10 distinct cells in each column, uterus thick-walled, vagina well-sclerotized and sigmoid. Vulva a wide transverse slit with prominent lips and a single epiptygma. Tail conical, with pointed tip, sometimes with a small mucron or bifurcate. 1. Length of conus as percentage of total stylet length. 2. Distance between anterior end of body and center of median pharyngeal bulb as percentage of pharyngeal length. 3. Distance from pharyngo-intestinal junction to end of dorsal gland tip. Male. Free-living. General morphology similar to that of female, slightly shorter and more slender. Genital system monorchic, testis outstretched, occupying 52–61 % of the body length, with developing spermatocytes in single or two rows, never reaching dorsal pharyngeal gland lobe. Spicules tylenchoid, ventrally arcuate with rounded apex. Gubernaculum simple. Bursa annulated, subterminal. Tail similar to that of female. Entomoparasitic generation. Not found. Etymology.The name of the genus refers to Veleshk, the region where the new genus was recovered. The species epithet refers to the country of origin.

Genus Veleshkinema n. gen. Description. Female slender, straight to irregularly curved upon heat relaxation, lacking sexual dimorphism in anterior region. Cuticle with fine transverse annules. Lateral field approximately one-fourth as wide as body diameter at mid-body, barely visible by light microscopy, not raised, with six, areolated equidistant incisures, beginning anteriorly at the end of procorpus with three lines forming two bands, gradually number of incisures increases to six at mid-body, and reduces to four at tail region. Cephalic region low, eight sectored, continuous with body contour, appearing smooth under light microscope, but with 4���5 faint annuli in SEM observations. Labial disc absent. Oral opening circular. Amphidial apertures oblique slits, slightly displaced dorsally, lateral to oral opening. Stylet short, 6.5���7.5 ��m long, delicate with asymmetrical minute rounded knobs with the dorsal knob anterior to the subventral knob. The dorsal gland orifice (DGO) just posterior to subventral knob. Subventral gland orifice (SVO) visible at middle of the less developed metacorpus. Pharynx with non-muscular fusiform median bulb, pharyngeal gland lobe dorsally (rarely dorso-laterally) overlapping intestine for about 0.5���2.1 times corresponding body width, nuclei of pharyngeal glands not clearly visible. Nerve ring enveloping isthmus. Excretory pore with sclerotized duct. Hemizonid barely visible, just anterior to excretory pore. Deirids at the level of excretory pore. Cardia present, small, rounded to triangular. Female reproductive system monodelphicprodelphic, outstretched anteriorly, spermatheca rounded, offset, filled with small, spheroid sperm, crustaformeria quadricolumellate with 8���10 cells in each column, postvulval uterine sac absent, vulva an open transverse slit with prominent lips, anterior lip much more raised than the posterior one, epiptygma single. Tail conoid, terminus narrow, tip sharp, sometimes bifurcated, similar in both sexes. Spicules slightly arcuate; gubernaculum arcuate. Bursa subterminal. Entomoparasitic infective females (if occurring) unknown. Diagnosis and relationships. Based on the above-mentioned morphological and biological characters, Veleshkinema n. gen. belongs to the superfamily Sphaerularioidea and family Sphaerulariidae. Siddiqi (2000) divided the suborder Hexatylina into two superfamilies Sphaerularioidea and Iotonchioidea Goodey, 1953 based on two alternative entomoparasitic generations in host and two or more types of adult in the host���s haemocoel in Iotonchioidea vs one in Sphaerularioidea. Morphologically, the new genus comes close to members of the superfamily Sphaerularioidea. Based on its morphology and our current knowledge of its biology, the new genus belongs to the family Sphaerulariidae and subfamily Sphaerulariinae and can be compared with two genera in the above mentioned subfamily namely: Prothallonema and Sphaerularia and two other neotylenchid genera namely Abursanema and Deladenus Thorne, 1941. Veleshkinema n. gen. differs from Prothallonema by its terminal bulb overlapping intestine dorsally (vs having a terminal stem-like projection), more anterior position of vulva (V = 81���84 vs 88���94 %), postvulval uterine sac absent (vs present) and bursa sub-terminal (vs terminal). It differs from Sphaerularia by having a free-living generation (vs no complete free-living generation), shorter stylet (6.5���7.5 vs 14���18 ��m) with asymmetrical knobs (vs lacking knobs or having thickenings at base of stylet), FIGURE. 1. Veleshkinema iranicum n. gen., n. sp. A: Details of pharynx; B: Anterior end in detail; C: Overlapping and cardia; D: Part of female reproductive system, showing crustaformeria and functional spermatheca; E: Male posterior end; F, G: Female tail; H & I: Male and female entire body. non-raised lateral field with six incisures (vs four conspicuous incisures), postvulval uterine sac absent (vs present), presence of cardia (vs absence), prominent vulval lips (vs flattened), and having of epiptygma (vs lacking). It differs from Abursanema by having stylet knobs (vs absence), pharyngeal gland overlap of intestine (vs a stem-like extension projecting into the lumen of intestine), lateral field with six incisures (vs two), presence of cardia (vs absence) and presence of sub-terminal bursa in males (vs not). The new genus differs from Deladenus by having asymmetrical stylet knobs (vs symmetrical), short pharyngeal gland overlap of intestine (vs very long), more anterior position of vulva (V = 81���84 vs 93���94 %), subterminal bursa (vs terminal) and pharyngo-intestinal junction posterior to nerve ring vs (anterior to nerve ring). Type habitat and locality: Recovered from bark sample of a European nettle tree (Celtis australis L.) in Habib Allah Veleshki shrine in Heydar Abad, west of Gorgan, Golestan province, Iran. GPS coordinates: N 36 �� 85 ', E 54 �� 29 '. No entomoparasitic phase was recovered from specimens of mosquitoes (belonging to the family Ceratopogonidae) that were associated with mosses on bark samples of the tree. Type material. Holotype female, slide NVI001 together with six paratype specimens: three males, four females (Slides NVI001, NVI002) deposited in the Nematode Collection of the Department of Plant Protection, University of Tehran, Karaj, Iran. Two female and one male paratypes deposited in each of the following collections: Nematode Collection of the Faculty of Agriculture, Tarbiat Modares University, Tehran, Iran; CABI Europe-UK, Egham, Surrey, UK; USDA Nematode Collection, Beltsville, MD, USA; WANECO collection, Wageningen (http://www.waneco.eu/)., Published as part of Miraeiz, Esmaeil, Heydari, Ramin, ��lvarez-Ortega, Sergio, Pedram, Majid & Atighi, Mohammad Reza, 2015, Molecular and morphological characterization of Veleshkinema iranicum n. gen., n. sp. (Nematoda: Hexatylina, Sphaerularioidea) from Iran, pp. 531-546 in Zootaxa 4000 (5) on pages 535-538, DOI: 10.11646/zootaxa.4000.5.3, http://zenodo.org/record/232430, {"references":["Siddiqi, M. R. (2000) Tylenchida: Parasites of Plants and Insects. 2 nd Edition. CABI Publishing, 833 pp.","Goodey, T. (1953) On certain eelworms, including Butschli's Tylenchus fungorum, obtained from toad-stools. Journal of Helminthology, 27, 81 - 94. http: // dx. doi. org / 10.1017 / S 0022149 X 00023518","Thorne, G. (1941) Some nematodes of the family Tylenchidae which do not possess a valvular median esophageal bulb. The Great Basin Naturalist, 2, 37 - 85."]}

Genus Veleshkinema n. gen. Description. Female slender, straight to irregularly curved upon heat relaxation, lacking sexual dimorphism in anterior region. Cuticle with fine transverse annules. Lateral field approximately one-fourth as wide as body diameter at mid-body, barely visible by light microscopy, not raised, with six, areolated equidistant incisures, beginning anteriorly at the end of procorpus with three lines forming two bands, gradually number of incisures increases to six at mid-body, and reduces to four at tail region. Cephalic region low, eight sectored, continuous with body contour, appearing smooth under light microscope, but with 4–5 faint annuli in SEM observations. Labial disc absent. Oral opening circular. Amphidial apertures oblique slits, slightly displaced dorsally, lateral to oral opening. Stylet short, 6.5–7.5 Μm long, delicate with asymmetrical minute rounded knobs with the dorsal knob anterior to the subventral knob. The dorsal gland orifice (DGO) just posterior to subventral knob. Subventral gland orifice (SVO) visible at middle of the less developed metacorpus. Pharynx with non-muscular fusiform median bulb, pharyngeal gland lobe dorsally (rarely dorso-laterally) overlapping intestine for about 0.5–2.1 times corresponding body width, nuclei of pharyngeal glands not clearly visible. Nerve ring enveloping isthmus. Excretory pore with sclerotized duct. Hemizonid barely visible, just anterior to excretory pore. Deirids at the level of excretory pore. Cardia present, small, rounded to triangular. Female reproductive system monodelphicprodelphic, outstretched anteriorly, spermatheca rounded, offset, filled with small, spheroid sperm, crustaformeria quadricolumellate with 8–10 cells in each column, postvulval uterine sac absent, vulva an open transverse slit with prominent lips, anterior lip much more raised than the posterior one, epiptygma single. Tail conoid, terminus narrow, tip sharp, sometimes bifurcated, similar in both sexes. Spicules slightly arcuate; gubernaculum arcuate. Bursa subterminal. Entomoparasitic infective females (if occurring) unknown. Diagnosis and relationships. Based on the above-mentioned morphological and biological characters, Veleshkinema n. gen. belongs to the superfamily Sphaerularioidea and family Sphaerulariidae. Siddiqi (2000) divided the suborder Hexatylina into two superfamilies Sphaerularioidea and Iotonchioidea Goodey, 1953 based on two alternative entomoparasitic generations in host and two or more types of adult in the host’s haemocoel in Iotonchioidea vs one in Sphaerularioidea. Morphologically, the new genus comes close to members of the superfamily Sphaerularioidea. Based on its morphology and our current knowledge of its biology, the new genus belongs to the family Sphaerulariidae and subfamily Sphaerulariinae and can be compared with two genera in the above mentioned subfamily namely: Prothallonema and Sphaerularia and two other neotylenchid genera namely Abursanema and Deladenus Thorne, 1941. Veleshkinema n. gen. differs from Prothallonema by its terminal bulb overlapping intestine dorsally (vs having a terminal stem-like projection), more anterior position of vulva (V = 81–84 vs 88–94 %), postvulval uterine sac absent (vs present) and bursa sub-terminal (vs terminal). It differs from Sphaerularia by having a free-living generation (vs no complete free-living generation), shorter stylet (6.5–7.5 vs 14–18 Μm) with asymmetrical knobs (vs lacking knobs or having thickenings at base of stylet), FIGURE. 1. Veleshkinema iranicum n. gen., n. sp. A: Details of pharynx; B: Anterior end in detail; C: Overlapping and cardia; D: Part of female reproductive system, showing crustaformeria and functional spermatheca; E: Male posterior end; F, G: Female tail; H & I: Male and female entire body. non-raised lateral field with six incisures (vs four conspicuous incisures), postvulval uterine sac absent (vs present), presence of cardia (vs absence), prominent vulval lips (vs flattened), and having of epiptygma (vs lacking). It differs from Abursanema by having stylet knobs (vs absence), pharyngeal gland overlap of intestine (vs a stem-like extension projecting into the lumen of intestine), lateral field with six incisures (vs two), presence of cardia (vs absence) and presence of sub-terminal bursa in males (vs not). The new genus differs from Deladenus by having asymmetrical stylet knobs (vs symmetrical), short pharyngeal gland overlap of intestine (vs very long), more anterior position of vulva (V = 81–84 vs 93–94 %), subterminal bursa (vs terminal) and pharyngo-intestinal junction posterior to nerve ring vs (anterior to nerve ring). Type habitat and locality: Recovered from bark sample of a European nettle tree (Celtis australis L.) in Habib Allah Veleshki shrine in Heydar Abad, west of Gorgan, Golestan province, Iran. GPS coordinates: N 36 ° 85 ', E 54 ° 29 '. No entomoparasitic phase was recovered from specimens of mosquitoes (belonging to the family Ceratopogonidae) that were associated with mosses on bark samples of the tree. Type material. Holotype female, slide NVI001 together with six paratype specimens: three males, four females (Slides NVI001, NVI002) deposited in the Nematode Collection of the Department of Plant Protection, University of Tehran, Karaj, Iran. Two female and one male paratypes deposited in each of the following collections: Nematode Collection of the Faculty of Agriculture, Tarbiat Modares University, Tehran, Iran; CABI Europe-UK, Egham, Surrey, UK; USDA Nematode Collection, Beltsville, MD, USA; WANECO collection, Wageningen (http://www.waneco.eu/).

Veleshkinema iranicum n. gen., n. sp. (Figs 1���5) Measurements. See Table 3. Description. Female. Free-living. Medium sized nematodes. Body mostly straight after heat relaxation, sometimes irregularly bent, slightly tapering towards both ends. Cuticle with fine transverse annulations, annuli about 1.2���1.7 ��m wide at mid-body. Lateral field barely visible by light microscopy, not raised, beginning anteriorly at the end of procorpus with three lines (forming two bands) increasing to six areolated equidistant incisures at mid-body, approximately one-fourth as wide as body diameter, gradually decreasing to four at tail region. Deirids at the level of excretory pore. Cephalic region low, anteriorly flat, continuous with body contour, 1- 2 ��m high and 6���8 ��m wide. Stylet delicate with asymmetrical minute knobs, conus shorter than shaft (28���38 % of total stylet length); DGO opens just posterior to subventral knob. Procorpus non-muscular, metacorpus less developed, isthmus slender, pharyngeal gland lobe dorsally (sometimes dorso-laterally) overlapping intestine about 0.5���2.1 times corresponding body diameter, cardia distinct. Nerve ring circum-pharyngeal, always anterior to excretory pore, located 62���99 ��m from anterior end. Excretory pore with sclerotized duct. Hemizonid barely visible, just anterior to excretory pore (5���11 ��m anterior to excretory pore), 2���3 annuli long. Intestine with distinct lumen ending in a distinct rectum. Reproductive system monodelphic-prodelphic, occupying 41���70 % of the body length, ovary outstretched, sometimes reaching dorsal pharyngeal gland, with single row of oocytes. Oviduct cellular, spermatheca rounded, offset and filled with minute spheroid sperm (2���3 ��m in diameter). Crustaformeria apparently quadricolumellate, with 8���10 distinct cells in each column, uterus thick-walled, vagina well-sclerotized and sigmoid. Vulva a wide transverse slit with prominent lips and a single epiptygma. Tail conical, with pointed tip, sometimes with a small mucron or bifurcate. 1. Length of conus as percentage of total stylet length. 2. Distance between anterior end of body and center of median pharyngeal bulb as percentage of pharyngeal length. 3. Distance from pharyngo-intestinal junction to end of dorsal gland tip. Male. Free-living. General morphology similar to that of female, slightly shorter and more slender. Genital system monorchic, testis outstretched, occupying 52���61 % of the body length, with developing spermatocytes in single or two rows, never reaching dorsal pharyngeal gland lobe. Spicules tylenchoid, ventrally arcuate with rounded apex. Gubernaculum simple. Bursa annulated, subterminal. Tail similar to that of female. Entomoparasitic generation. Not found. Etymology.The name of the genus refers to Veleshk, the region where the new genus was recovered. The species epithet refers to the country of origin., Published as part of Miraeiz, Esmaeil, Heydari, Ramin, ��lvarez-Ortega, Sergio, Pedram, Majid & Atighi, Mohammad Reza, 2015, Molecular and morphological characterization of Veleshkinema iranicum n. gen., n. sp. (Nematoda: Hexatylina, Sphaerularioidea) from Iran, pp. 531-546 in Zootaxa 4000 (5) on pages 538-540, DOI: 10.11646/zootaxa.4000.5.3, http://zenodo.org/record/232430

Yaghoubi, Ali, Pourjam, Ebrahim, Pedram, Majid, Siddiqi, Mohammad Rafiq, Atighi, Mohammad Reza (2014): Description of Abursanema iranicum n. gen., n. sp. (Nematoda: Hexatylina, Sphaerularioidea) from Iran and its phylogenetic relationships. Zootaxa 3826 (2): 301-314, DOI: http://dx.doi.org/10.11646/zootaxa.3826.2.1

Trichodorus iranicus sp. n. (Figs 1–3) Measurements. Table 2. Male: Body cylindrical, tapering very slightly anteriorly, J-shaped after heat relaxation. Cuticle moderately swollen, 2–3 µm thick at onchiostyle region and 3 µm thick at mid-body and at anterior lip of anus. Lip region rounded, 8–9 µm wide. Amphidial fovea cup-shaped, its opening a wide slit. Stoma with refractive strengthening rods, approximately 5 µm long. Pharyngostom 42.5 % of pharynx length, onchiostyle well developed, curved, onchium fine, 17–18 µm long, onchiophore well sclerotized, guiding ring at 3.5 times lip region width from anterior end. Pharynx composed of two parts: anterior slender part expanding gradually to form an offset terminal bulb, 20 µm wide× 50 µm long, within which the dorsal gland nucleus and posterior pair of ventrosublateral nuclei are clearly visible, while the anterior pair of ventrosublateral nuclei are much smaller and more difficult to discern. Cardia slightly to moderately developed. Nerve ring 7.5–11.5 µm posterior to the base of pharyngostom. Three distinct cervical papillae present, the anteriormost (CP 1) located at about 50–60 % of pharyngostom length, the second papilla (CP 2) at ca. 5 µm anterior (in paratype male) or posterior (holotype) to the pharyngostom base and the third papilla (CP 3) at or slightly posterior to the middle of the isthmus. Secretory-excretory pore at 13.5 –16.0 µm posterior to CP 3, opposite the isthmus. Reproductive system monorchic, sperm cells with large sausage-shaped nuclei, spicules ventrally curved, slightly sclerotized, their proximal part widened with slightly developed capitulum, middle part indented and provided with a few bristles (two dorsal and two on ventral side in holotype), gubernaculum short. Three precloacal supplements (SP), with the posteriormost (SP 1) located anterior to the proximal end of the spicules. Tail rounded, its length less than one body width at cloacal region, one pair of small postcloacal subventral papillae situated just posterior to the cloacal opening, subventral pair of caudal pores at a short distance from them. Hyaline part of tail 7–9 µm thick. Female: Body straight, cigar-like. General morphology similar to that of male, except for the reproductive system, and with the posterior body region not ventrally curved. Excretory pore opposite isthmus. Anterior and posterior genital tracts of more or less equal length, each with a spermatheca containing sperm cells near the oviduct at the reflexed tip of the ovary. Morphology of sperm cells similar to mature sperm in genital tract of the male. Vagina well developed, 46–57 % of corresponding body width in length, pars proximalis vaginae more or less rhomboid, pars refringens vaginae triangular to rounded-triangular, medium sized (ca. 3.5 µm), 1–2 µm apart from each other; vulva a transverse slit in ventral view. One female with copulatory plug at vulva-vagina level. Post-advulval body pore 33 µm (n= 1) posterior to vulva. Tail short, 3–5 µm in length, and rounded. Caudal pores subventral, just posterior to anus. Juveniles: Some damaged juveniles were found, but were in poor condition and not suitable for morphological study. Code, using the polytomous identification key of Decraemer & Baujard (1998): For male: A = 222 (mean, min, max), B = 22, C = 11, D = 33, E = 12, F = 3, G = 0, H = 20, I = 11, J = 500, K = 33, L = 44, M = 160, N = 11, O = 1, P = 2; for female: A = 200, B = 22, C = 1, D = 1, E = 200, F = 100, G = 2, H = 22, I = 23, J = 11, K = 400, L = 22, M = 1, N = 2, O = 11, P = 11, Q = 1, R = 1, S = 7. Type habitat and localit y. Soil samples collected from the rhizosphere of grasses in forests of Golestan province, GPS coordinates: N 36 ° 34.942, E051° 49.166. Type material. Holotype male and three paratype females deposited in the Nematode Collection of the Faculty of Agriculture, Tarbiat Modares University, Tehran, Iran. One paratype male and two females in the USDA Nematode Collection, Beltsville, MD, USA.

Pedram, Majid, Pourjam, Ebrahim (2014): Description of Trichodorus iranicus sp. n. (Diphtherophorina, Trichodoridae) from Iran. Zootaxa 3795 (4): 431-440, DOI: http://dx.doi.org/10.11646/zootaxa.3795.4.3

Abursanema iranicum n. gen., n. sp. (Figs 1 & 2) Measurements. See Table 3. Description. Male. Free-living. Body slightly ventrally arcuate after heat relaxation. Inner cuticle finely annulated, outer cuticle smooth throughout body length, lateral field with two incisures. Deirid at the level of excretory pore. Lip region low, rounded and slightly flattened, continuous with body contour. Stylet short, weak, 6.5���10.5 ��m, without basal knobs, dorsal arm of the shaft slightly longer than the ventral. Orifice of dorsal gland 1���2 ��m behind stylet base. Corpus cylindroid, metacorpus weakly developed, spindle-shaped, without a valvular apparatus, isthmus narrow, 35���47 ��m long. Basal bulb large, pyriform, with stem-like extension projecting into the lumen of intestine. Excretory pore opening at the level of middle of basal bulb, 86���107 ��m from anterior end. Hemizonid located just anterior to excretory pore. Nerve ring enveloping isthmus in its posterior half. Testis single, anteriorly outstretched, gonoduct 159���252 ��m long; sperm spheroid. Spicules paired, ventrally arcuate, with rounded manubrium and pointed distal tip, gubernaculum arcuate. Bursa absent. Phasmid hardly visible, located at 0.75 ���1.0 times body width posterior to cloaca. Tail long, with narrowly rounded or pointed tip. Female. Free-living (infective entomoparasitic female not found). General morphology similar to that of male, except for character states associated with sexual differences. Reproductive system monodelphic-prodelphic, ovary with single row of oocytes, usually reflexed two times at proximal end, spermatheca rounded to ellipsoid, filled with spheroid sperm, crustaformeria short, ca 1.7���2.1 body widths long, vulva a transverse slit, vagina extending into the body a little less than half body width, postvulval uterine sac less than corresponding body width. Vulva-anus distance 46���69 ��m, ca 0.3���0.6 of the tail length. Etymology. The name of the genus is Greek meaning ���nematode without a bursa��� and is neuter in gender. Specific epithet refers to the country of occurrence. Molecular phylogenetic relationships. Partial 18 S rDNA phylogeny. Our 942 bp partial SSU rDNA sequence, GenBank accession number KF 885743, was used to determine the phylogenetic relationships of Abursanema n. gen. with other tylenchid nematodes. A BlastN search of the GenBank sequence database was performed and the closest sequences/species, along with sequences of genera with morphological similarity, were selected for inclusion in the analyses. The dataset was composed of 2493 total characters, of which 2118 characters were variable and 1824 characters were parsimony-informative after manual editing. The average nucleotide composition was as follows: 25.7 % A, 20.6 % C, 26.4 % G and 27.2 % T. Figure 3 presents a Bayesian phylogenetic tree inferred from the multiple alignment of partial SSU sequences of 23 tylenchid taxa, one outgroup taxon and one isolate of the newly recovered/sequenced genus. The BPP and ML BS values are given on the appropriate clades in the form BPP/ML BS. In this tree, the new genus forms a clade with one species of Deladenus Thorne, 1941 (saccession number EU 306345), and two species of Sphaerularia (accession numbers AB 300595 and AB 250213), members of the superfamily Sphaerularioidea (see Siddiqi, 2000). The monophyly of this clade is well supported in both BI and ML methods (1.00 and 0.96 % respectively). The other species of Deladenus used in our analyses (D. proximus Bedding, 1974, accession number JF 304744) does not share a common ancestor with Deladenus sp. (EU 306345) and occupies another position in the tree. This clade forms a sister clade to the remaining members of Hexatylina with low BPP (0.55) and no support under ML. Based on the current paucity of data and the restricted number of Hexatylina species that have been sequenced, the relationships among these taxa cannot be clearly resolved. D 2 /D 3 segment of 28 S rDNA phylogeny. To reconstruct the 28 S rDNA tree, a 706 bp partial sequence of D 2 / D 3 region with accession number KF 885742 was used. Figure 4 presents the phylogenetic tree of 37 tylenchid taxa, one outgroup and one isolate of Abursanema iranicum n. gen., n. sp. The dataset was composed of 941 total characters of which 887 characters were variable and 827 characters were parsimony-informative after manual editing, with an average nucleotide composition of 23.7 % A, 19.6 % C, 29.8 % G and 26.9 % T. As in the 18 S tree, the new genus formed a clade with two species of Sphaerularia (AB 300595 and AB 733665) with high BPP and ML BS support. This makes sense, considering the morphological similarities of these taxa, mostly in the shape of the pharynx. According to Siddiqi (2000), the families Sphaerulariidae and Paurodontidae are synonymous and the grouping of Sphaerularia, as a member of Sphaerulariidae, and the new genus, as a member of Paurodontidae, in our tree supports his opinion. Discussion. Our phylogenetic study, using both partial 18 S rDNA and D 2 /D 3 segment of 28 S rDNA, reveals new insights on the molecular phylogenetic affinities of members of Hexatylina, including Abursanema n. gen. In constructed trees using partial sequences of both SSU (Fig. 3) and LSU (Fig. 4), Abursanema n. gen. (Paurodontidae) clusters with sequenced species of Sphaerularia (Sphaerularidae), supporting the synonymy of Paurodontidae and Sphaerulariidae (Siddiqi, 2000, Chizhov, 2004 and Andr��ssy, 2007). In the partial 18 S tree, Deladenus sp. (EU 306345) forms a clade with Abursanema n. gen. but D. proximus (JF 304744) is placed in a clade with Howardula spp. (AF 519234, JX 291137 and AY 146451) and Bradynema listronoti Zeng, Giblin-Davis, Ye, Belair, Boivin & Thomas, 2007 (DQ 915805). No free-living mycetophagous generation is known for the latter two genera, but one occurs in Deladenus. Based on our result, using of this biological character to classify genera in Hexatylina is questionable. The inferred tree using partial sequences of D 2 /D 3 of 28 S rDNA shows the same result as in the SSU tree. In our 28 S tree, Howardula phyllotretae Oldham, 1933 (DQ 328728) and Parasitylenchus sp. (DQ 328729) form a highly supported clade in both BI and ML methods (1.00/ 99). These genera are currently classified in different families, Allantonematidae Pereira, 1931 and Parasitylenchidae Siddiqi, 1986 respectively, assigned to different superfamilies, Sphaerularioidea and Iotonchioidea Goodey, 1953 (Siddiqi, 1986), respectively. Members of both genera are insect parasites. Thus, parasitism of the insect haemocoel may have evolved independently in both superfamilies (Siddiqi, 2000). We believe that additional molecular data from other genera may give a clearer picture of the relationships among members of Hexatylina. Both partial 18 S and 28 S rDNA trees confirm that Howardula Cobb, 1921 is polyphyletic (see various papers, including Ye et al., 2007). Abursanema n. gen. and Sphaerularia spp. form a highly supported clade in both ML and BI methods, but again these genera have been assigned to different families (Paurodontidae and Sphaerulariidae, respectively), albeit families that may in fact be synonymous. In Paurodontidae, a fungus-feeding generation is well-known but nothing is known of entomoparasitic forms, while in Sphaerulariidae a free-living generation may be present (Siddiqi, 2000). Accordingly, and as noted above for the partial 18 S tree, life cycle should not be considered a basic character for classification of members of the Hextaylina. This proposal conflicts with what was noted by Chizhov et al. (2012) who emphasized the importance of life cycle characteristics in the systematics of hexatylenchid nematodes. The results of our molecular phylogenetic studies with partial sequences of both 18 S rDNA and especially the D 2 /D 3 expansion segment of 28 S rDNA show a close relationship of members of Paurodontidae and Sphaerulariidae, and also highlight the need for data from additional representatives of both families so that their relationships could be more clearly elucidated., Published as part of Yaghoubi, Ali, Pourjam, Ebrahim, Pedram, Majid, Siddiqi, Mohammad Rafiq & Atighi, Mohammad Reza, 2014, Description of Abursanema iranicum n. gen., n. sp. (Nematoda: Hexatylina, Sphaerularioidea) from Iran and its phylogenetic relationships, pp. 301-314 in Zootaxa 3826 (2) on pages 305-309, DOI: 10.11646/zootaxa.3826.2.1, http://zenodo.org/record/254573, {"references":["Thorne, G. (1941) Some nematodes of the family Tylenchidae which do not possess a valvular median esophageal bulb. Great Basin Naturalist, 2, 37 - 85.","Siddiqi, M. R. (2000) Tylenchida: Parasites of Plants and Insects, 2 nd Edition. Wallingford, CABI Publishing, UK, 833 pp.","Bedding, R. A. (1974) Five new species of Deladenus (Neotylenchidae), entomophagous mycetophagous nematodes parasitic in siricid woodwasps. Nematologica, 20, 204 - 225. http: // dx. doi. org / 10.1163 / 187529274 x 00186","Chizhov, V. N. (2004) Entomopathogeneous nematodes from the suborder Hexatylina (Nematoda: Tylenchida). In: Sonin, M. D. (Ed.), Parasitic nematodes of plants and insects, Moscow, Russia, Nauka, pp. 277 - 293.","Andrassy, I. (2007) Free-living nematodes of Hungary, II (Nematoda errantia). Budapest, Hungary, Hungarian Natural History Museum and Systematic Zoology Research Group of the Hungarian Academy of Sciences, 496 pp.","Siddiqi, M. R. (1986) Tylenchida: Parasites of Plants and Insects. Commonwealth Agricultural Bureaux, Farnham Royal, Slough, UK, ix + 645 pp.","Chizhov, V. N., Butorina, N. & Subbotin, S. A. (2012) Entomoparasitic nematodes of the genus Skarbilovinema: S. laumondi and S. lyoni (Nematoda: Tylenchida), parasites of the flies of the family Syrphidae (Diptera), with phylogeny of the suborder Hexatylina. Russian Journal of Nematology, 20, 141 - 155."]}

Trichodorus iranicus sp. n. (Figs 1���3) Measurements. Table 2. Male: Body cylindrical, tapering very slightly anteriorly, J-shaped after heat relaxation. Cuticle moderately swollen, 2���3 ��m thick at onchiostyle region and 3 ��m thick at mid-body and at anterior lip of anus. Lip region rounded, 8���9 ��m wide. Amphidial fovea cup-shaped, its opening a wide slit. Stoma with refractive strengthening rods, approximately 5 ��m long. Pharyngostom 42.5 % of pharynx length, onchiostyle well developed, curved, onchium fine, 17���18 ��m long, onchiophore well sclerotized, guiding ring at 3.5 times lip region width from anterior end. Pharynx composed of two parts: anterior slender part expanding gradually to form an offset terminal bulb, 20 ��m wide�� 50 ��m long, within which the dorsal gland nucleus and posterior pair of ventrosublateral nuclei are clearly visible, while the anterior pair of ventrosublateral nuclei are much smaller and more difficult to discern. Cardia slightly to moderately developed. Nerve ring 7.5���11.5 ��m posterior to the base of pharyngostom. Three distinct cervical papillae present, the anteriormost (CP 1) located at about 50���60 % of pharyngostom length, the second papilla (CP 2) at ca. 5 ��m anterior (in paratype male) or posterior (holotype) to the pharyngostom base and the third papilla (CP 3) at or slightly posterior to the middle of the isthmus. Secretory-excretory pore at 13.5 ���16.0 ��m posterior to CP 3, opposite the isthmus. Reproductive system monorchic, sperm cells with large sausage-shaped nuclei, spicules ventrally curved, slightly sclerotized, their proximal part widened with slightly developed capitulum, middle part indented and provided with a few bristles (two dorsal and two on ventral side in holotype), gubernaculum short. Three precloacal supplements (SP), with the posteriormost (SP 1) located anterior to the proximal end of the spicules. Tail rounded, its length less than one body width at cloacal region, one pair of small postcloacal subventral papillae situated just posterior to the cloacal opening, subventral pair of caudal pores at a short distance from them. Hyaline part of tail 7���9 ��m thick. Female: Body straight, cigar-like. General morphology similar to that of male, except for the reproductive system, and with the posterior body region not ventrally curved. Excretory pore opposite isthmus. Anterior and posterior genital tracts of more or less equal length, each with a spermatheca containing sperm cells near the oviduct at the reflexed tip of the ovary. Morphology of sperm cells similar to mature sperm in genital tract of the male. Vagina well developed, 46���57 % of corresponding body width in length, pars proximalis vaginae more or less rhomboid, pars refringens vaginae triangular to rounded-triangular, medium sized (ca. 3.5 ��m), 1���2 ��m apart from each other; vulva a transverse slit in ventral view. One female with copulatory plug at vulva-vagina level. Post-advulval body pore 33 ��m (n= 1) posterior to vulva. Tail short, 3���5 ��m in length, and rounded. Caudal pores subventral, just posterior to anus. Juveniles: Some damaged juveniles were found, but were in poor condition and not suitable for morphological study. Code, using the polytomous identification key of Decraemer & Baujard (1998): For male: A = 222 (mean, min, max), B = 22, C = 11, D = 33, E = 12, F = 3, G = 0, H = 20, I = 11, J = 500, K = 33, L = 44, M = 160, N = 11, O = 1, P = 2; for female: A = 200, B = 22, C = 1, D = 1, E = 200, F = 100, G = 2, H = 22, I = 23, J = 11, K = 400, L = 22, M = 1, N = 2, O = 11, P = 11, Q = 1, R = 1, S = 7. Type habitat and localit y. Soil samples collected from the rhizosphere of grasses in forests of Golestan province, GPS coordinates: N 36 �� 34.942, E051�� 49.166. Type material. Holotype male and three paratype females deposited in the Nematode Collection of the Faculty of Agriculture, Tarbiat Modares University, Tehran, Iran. One paratype male and two females in the USDA Nematode Collection, Beltsville, MD, USA., Published as part of Pedram, Majid & Pourjam, Ebrahim, 2014, Description of Trichodorus iranicus sp. n. (Diphtherophorina, Trichodoridae) from Iran, pp. 431-440 in Zootaxa 3795 (4) on pages 433-435, DOI: 10.11646/zootaxa.3795.4.3, http://zenodo.org/record/226996, {"references":["Decraemer, W. & Baujard, P. (1998) A polytomous key for identification of species of the family Trichodoridae Thorne, 1935 (Nematoda: Triplonchida). Fundamental and Applied Nematology, 21, 37 - 62."]}

Genus Abursanema n. gen. Description. Body straight or slightly ventrally arcuate after fixation, less than 1mm long. Cuticle thin, outer cuticle smooth, inner cuticle finely annulated. Lateral field narrow, with two incisures. Head low, flattened, framework six-sectored, lateral sectors narrower than submedian ones. Amphidial apertures not clearly observed. Stylet short, without basal knobs, dorsal arm of the shaft slightly longer than the ventral one. Orifice of dorsal pharyngeal gland located close to stylet base, at ca 1���2 ��m distance. Corpus cylindroid, metacorpus spindleshaped, lacking cuticular thickening or valve, isthmus short, terminal bulb large, pyriform, with three pharyngeal glands and a stem-like extension projecting into the lumen of intestine. Excretory pore opening from near base of isthmus to opposite basal bulb. Vulva at 66���75 % of total body length, more than two body widths distance from anus in type species. Ovary single, prodelphic, outstretched anteriorly, usually reflexed twice at proximal end, cells in single row, spermatheca rounded to ellipsoid, filled with spheroid sperm, postvulval uterine sac less than corresponding body width in length. Tail conoid with narrow terminus, similar in both sexes. Spicules simple, slightly arcuate, 18���23 ��m long in type species. Gubernaculum arcuate. Bursa absent. Entomoparasitic infective females not found. Diagnosis and relationships. In the above-mentioned morphological and biological characters, Abursanema n. gen. belongs to the superfamily Sphaerularioidea and family Paurodontidae. The new genus is distinguished from all other valid genera in the family Paurodontidae by the absence of a bursa. It can also be distinguished from Paurodontus by having a stylet without basal knobs (vs stylet with rounded and symmetrical knobs), and by having two incisures in lateral field (vs four or six). The new genus differs from Misticius by the absence of stylet knobs (vs stylet with prominent symmetrical basal knobs), having two incisures in lateral field (vs absence of lateral field), and by having the excretory pore opening near base of isthmus to opposite basal bulb (vs near the stylet base). When compared with Neomisticius, the new genus differs by the absence of stylet knobs (vs stylet with prominent symmetrical basal knobs), excretory pore opening opposite terminal bulb (vs near the base of stylet) and by having a longer tail (vs a short, subcylindrical one with a small projection at its tip). It differs from Paurodontella by the absence of stylet knobs (vs stylet with basal knobs) and the presence of short postvulval uterine sac (vs absence). Abursanema n. gen. is close to Paurodontoides, but differs by the absence of stylet knobs (vs stylet with asymmetrical basal knobs) and by having a head with six sectors (vs eight). From Luella, the new genus differs by having two incisures in the lateral field (vs absence), and structure of the base of the terminal bulb (vs bulb offset and stem-like, basal extension absent). The genus Bealius has a stylet with knobs, a stem-like basal pharyngeal bulb extension projecting into the intestine dorsally, no lateral field or postvulval uterine sac, dissimilar tail shape in female and male, and spicules with a bifurcate tip in comparison with the new genus. Based on molecular data, the new genus has sequences similar to those of Sphaerularia Dufour, 1837. However, it differs from the free-living phase by absence of a bursa in male and by having a stem-like basal extension in pharyngeal bulb projecting into the intestine (vs basal pharyngeal bulb abutting or slightly overlapping the intestine). Based on male characters (especially lacking of bursa), the new genus is similar to Gymnotylenchus Siddiqi, 1961, but is clearly different in having a stylet without basal knobs, extension of the basal pharyngeal bulb into the intestine (vs pharyngeal glands forming an elongate diverticulum extending over intestine dorsally or subdorsally), junction of pharynx and intestine posterior to nerve ring (vs anterior to nerve ring), two incisures in lateral field vs six, and structure of the spicules (tylenchoid vs prominently cephalated) and presence of a gubernaculum (vs absence). Type habitat and locality: Recovered from bark samples of dead Populus alba L. collected in village of Sorkh Abad, Hamedan province, western Iran, during April 2013. Type material: Holotype male, 7 paratype males and 17 females deposited in the Nematode Collection of the Faculty of Agriculture, Tarbiat Modares University, Tehran, Iran. Three male and three female paratypes deposited in each of the following collections: CABI Europe-UK, Egham, Surrey, UK; USDA Nematode Collection, Beltsville, MD, USA; Department of Nematology, Agricultural University, Wageningen, the Netherlands., Published as part of Yaghoubi, Ali, Pourjam, Ebrahim, Pedram, Majid, Siddiqi, Mohammad Rafiq & Atighi, Mohammad Reza, 2014, Description of Abursanema iranicum n. gen., n. sp. (Nematoda: Hexatylina, Sphaerularioidea) from Iran and its phylogenetic relationships, pp. 301-314 in Zootaxa 3826 (2) on pages 304-305, DOI: 10.11646/zootaxa.3826.2.1, http://zenodo.org/record/254573, {"references":["Siddiqi, M. R. (1961) Gymnotylenchus zeae, n. g., n. sp. (Nematoda: Neotylenchidae), a root associate of Zea mays L. (sweet corn) in Aligarh, North India. Nematologica, 6, 59 - 63. http: // dx. doi. org / 10.1163 / 187529261 x 00289"]}

Discotylenchus discretus Siddiqi, 1980 (Figs 4, 5) Measurements. See Table 3. This species was recovered from the rhizosphere of apple (Malus domestica L. Borkh.) trees in the city of Bojnourd, Northern Khorasan province, Iran, during July 2009. The morphological and morphometric characters (see Table 3) are in full agreement with those in original description. The Iranian population of D. discretus is characterized by having a smooth lip region, tapering to an offset, distinct labial disc, lateral field with four incisures, body length 0.54���0.60 mm, stylet 8���9 ��m long, spermatheca round, filled with sperm and tail filiform, 102���112 ��m long., Published as part of Ghaemi, Razieh, Pourjam, Ebrahim, Atighi, Mohammad Reza, Pedram, Majid & Karssen, Gerrit, 2012, First record of the genus Discotylenchus Siddiqi, 1980 (Nematoda: Tylenchidae) from Iran, with description of one new and data on two known species, pp. 72-82 in Zootaxa 3493 on page 81, DOI: 10.5281/zenodo.282451, {"references":["Siddiqi, M. R. (1980) Two new nematode genera, Safianema (Anguinidae) and Discotylenchus (Tylenchidae), with descriptions of three new species. Proceedings of the helminthological Society of Washington, 47, 85 - 94."]}

Discotylenchus iranicus n. sp. (Figs. 1, 2) Measurements. See Table 1. Morphological description. Female. Body almost straight or slightly ventrally arcuate after heat relaxation. Cuticular annuli fine, 0.8���0.9 ��m wide. Lateral field with four incisures, 3.5���4.5 ��m wide at mid-body, approximately one-fifth as wide as body diameter. Lip region continuous and smooth, 6.5���7.5 ��m broad, 3.5 ���5.0 ��m high, tapering to an offset prominent labial disc, measuring 3.5 ��m in diameter. Amphidial aperture minute, appearing as a longitudinal slit in lateral view, originating from the base of labial disc. Stylet long and thin; conus about 32���40 % of total stylet length; knobs rounded, 2.5 ���3.0 ��m wide. Dorsal pharyngeal gland outlet 2���3 ��m from base of stylet knobs. Metacorpus elongate, spindle-shaped with distinct central valvular apparatus and located at 37���41 % of total pharynx length. Isthmus elongated, slender. Basal bulb saccate, 41���43 ��m long and 13���15 ��m wide, not overlapping intestine. Excretory pore located posterior to hemizonid, 89���100 ��m from anterior end. Deirids present, a little (five annuli, n= 1) behind excretory pore level. Nerve ring enveloping isthmus at mid-point, 76���89 ��m from anterior end. Vulva a transverse slit, flaps absent, vagina with thin walls, about one-half times corresponding body diameter, postuterine sac about 0.5���0.9 vulval body width. Reproductive system monodelphic-prodelphic with a single row of oocytes, 31���39 % of body length. Spermatheca almost oval, offset, functional with rounded sperm. Phasmids outside the lateral field, latero-subdorsal in position, 3.5 ��m behind the vulval slit (n= 1). Tail 6.8 ���10.0 times longer than anal body width, not filiform, conoid with a rounded tip. Male. In general morphology similar to female, except for character states associated with sexual differences. Testis single, anteriorly outstretched and 235.6 ��30.0 (185���265) ��m long. Spicules slender, ventrally arcuate. Gubernaculum simple, bursa adanal, 26���30 ��m long. Type habitat and locality. Recovered from the rhizosphere of corn (Zea mays L.) in Kohgiluyeh and Boyer- Ahmad province, south-western Iran during October 2010. Type material. Holotype female, one paratype female and four males deposited in the Nematode Collection of the Faculty of Agriculture, Tarbiat Modares University, Tehran, Iran and one female and one male paratypes at Wageningen Nematode Collection, Plant Protection Service, Wageningen, the Netherlands. Diagnosis and relationships. Discotylenchus iranicus n. sp. is characterized by its continuous and smooth lip region, tapering to an offset prominent labial disc, lateral field with four incisures, stylet length of 14���15 ��m, vulva position at 70.8���76.5 %, tail length of 81���100 ��m, conoid with a rounded tip tail and presence of functional male in population. The new species comes close to Discotylenchus brevicaudatus, D. attenuatus and D. discretus. It differs from D. brevicaudatus by having a longer body (0.68���0.74 vs 0.32���0.39 mm), higher a ratio (35���39 vs 27���33), slightly smaller c ratio (7.0��� 9.2 vs 8.1���9.8), higher c' ratio (6.8���10 vs 4.5 ���6.0), longer stylet (14���15 vs 7���8 ��m), longer tail (81���100 vs 35���47 ��m), position of excretory pore (89���100 vs 55���58 ��m from anterior end) and presence of functional males in population vs absence. From D. attenuatus, it differs by having longer body (0.6���0.7 vs 0.3���0.4 mm), higher a ratio (35���39 vs 25���33), higher c ratio (7.0��� 9.2 vs 4.1 ���5.0), posteriorly located vulva (V = 70.8���76.5 vs 60���64), longer stylet (14���15 vs 6.0��� 6.5 ��m), longer spicules (18���21 vs 11���12 ��m), gubernaculum length (5.0��� 6.5 vs 3.0��� 4.5 ��m) and tail shape (conical with a rounded tip vs attenuated, filiform, with extremely thin end). Finally, the new species differs from D. discretus by having a longer body (0.68���0.74 vs 0.44���0.61 mm), higher c ratio (7.0��� 9.2 vs 5.0��� 6.2), posteriorly located vulva (V= 70.8��� 76.5 vs 63���67), longer stylet (14���15 vs 7.0��� 7.5 ��m), longer spicules (18���21 vs 11���13 ��m), gubernaculum length (5.0��� 6.5 vs 3.5 ���4.0 ��m) and tail shape (conical with a rounded tip vs gradually tapering, filiform). Etymology. The specific epithet refers to Iran, the country where the new species was originally found. TABLE 1. Morphometrics of Discotylenchus iranicus n. sp. All measurements are in ��m, and in the form: mean �� s.d. (range). Holotype Paratype females Paratype males, Published as part of Ghaemi, Razieh, Pourjam, Ebrahim, Atighi, Mohammad Reza, Pedram, Majid & Karssen, Gerrit, 2012, First record of the genus Discotylenchus Siddiqi, 1980 (Nematoda: Tylenchidae) from Iran, with description of one new and data on two known species, pp. 72-82 in Zootaxa 3493 on pages 73-74, DOI: 10.5281/zenodo.282451

Pedram, Majid, Pourjam, Ebrahim, Vinciguerra, Maria T. (2012): Description of a new species of the rare genus Epacrolaimus Andrássy, 2000 (Dorylaimida, Aporcelaimidae) and new data on male of Paraxonchium laetificans (Andrássy, 1956) Altherr & Loof, 1969 (Dorylaimida, Paraxonchiidae) from Iran. Zootaxa 3327: 53-61, DOI: 10.5281/zenodo.210470

Labronemella labiata Andr��ssy, 1985 (Figs 1���4) Measurements. See Table 1. the form average �� standard deviation (range) Description Material examined: Five females and six males, in good state of preservation. Adult: Moderately slender to slender nematodes of medium size, 2.00��� 2.61 mm long. Habitus curved ventrad upon fixation, C-shaped in both sexes. Cuticle two layered: outer layer thin, nearly smooth under LM, but SEM pictures show that very fine transverse striation is present throughout the body; inner layer thicker (3���4 times) than outer one, bearing distinct radial striation; cuticle 4���6 ��m thick at anterior region, 5���7 ��m in mid-body and 6���9 ��m on tail. Lateral chord 14���20 ��m wide at mid-body, occupying one-fifth to one-fourth (21���26 %) of body diameter. Lip region offset by constriction, 3.3���4.3 times as wide as high and one-third to two-fifths (33���43 %) of body diameter at neck base. Its detailed morphology elucidated under SEM: lips completely amalgamated, forming a disc- or sucker-like structure, whose anterior surface (oral field) appears distinctly sunken but with its inner (perioral) area differentiated in six large, separated liplets protruding on oral field; lateral, perioral liplets trapezoidal and visibly larger than subdorsal and subventral ones, which are triangular, so that lip region shows a bi-radial symmetry; inner labial papillae migrated to the margin of labial disc, being located close to outer labial and cephalic papillae. Amphidial fovea funnel-like, its aperture 14���18 ��m or about two-thirds (59���71 %) of lip region width; SEM pictures show that fovea does not bulge on body surface and that aperture is a very narrow slit within the labial constriction. Odontostyle straight and narrow, 9���14 times as long as wide, with slender walls and distinct lumen, 1.1���1.3 times longer than lip region width and 1.13���1.52 % of body length; aperture 9���13 ��m long or up to two-fifths (29���40 %) of its total length. Guiding ring double, at 14���20 ��m or 0.5���0.6 times the lip region width from anterior end. Odontophore rod-like, 1.6���2.1 times the odontostyle. Pharynx consisting of a slender but distinctly muscular anterior part, enlarging gradually; basal expansion 6.5���8.5 times as long as wide or 3.5���4.4 times longer than body diameter at neck base, and occupying up to one-half (45���50 %) of total neck length. Pharyngeal gland nuclei and outlets situated as follows: DN = 52���57, DO = 53���56, S 1 N 1 = 73���75, S 1 N 2 = 77���79, S 2 N = 82���85. Nerve ring at 163���200 ��m or 30���36 % of neck length from anterior end. Cardia well developed, tongue-like, 23 ��m long, 1.5 times as long as wide, surrounded by intestinal tissue that forms a conical extension measuring 40 ��m including the cardia. Female: Genital system didelphic-amphidelphic, with both genital branches equally and very well developed, the anterior 321���479 ��m, the posterior 320���540 ��m long. Ovaries large, occasionally surpassing the sphincter level, the anterior 67���336 ��m, the posterior 69���295 ��m long; oocytes numerous, arranged first in two or more rows and then in one single row. Oviduct 143���279 ��m long or 2.1���3.2 times the body diameter, joining subterminally the ovary and consisting of a tubular part and a well developed, elongate pars dilatata often with visible lumen. Sphincter separating oviduct from uterus, but not very marked. Uterus long but often convoluted and difficult to measure with accuracy, 184���279 ��m long or 2.5���3.6 times the body diameter; it is tripartite since it bears an intermediate, slightly dilated portion with thicker walls and narrower lumen, while the proximal and distal regions are tubular and with thinner walls and distinct lumen. Vagina 33���41 ��m long, extending inwards about one-half (49���54 %) of the corresponding body diameter: pars proximalis 15��� 23 x 20���26 ��m, with slightly sigmoid walls and surrounded by moderately developed musculature; pars refringens with two small drop-shaped, close together, sclerotized pieces measuring 9��� 10 x 3���4 ��m and with a combined width of 6���9 ��m; pars distalis short, 4���7 ��m. Vulva a transverse slit, about 12 ��m long. Prerectum 1.7���2.8 anal body diameters long. Rectum 41���48 ��m long or 1.0��� 1.2 times the anal body diameter. Tail short and rounded to hemispheroid; three pairs of caudal pores at the posterior half of tail, two subventral and the other lateral. Male: Prerectum 2.9���4.7 anal body diameters long. Genital system diorchic, with opposite testes. In addition to the adcloacal pair, situated at 8���11 ��m from cloacal aperture, there is a series of 14���17, contiguous, ventromedian supplements 6���8 ��m apart, the posteriormost of which located at 69���77 ��m from the pre-cloacal pair, out the range of spicules. Spicules dorylaimoid, curved ventrad, 5.5���6.5 times as long as wide and 1.6 ���2.0 times the body diameter at level of cloacal aperture. Lateral guiding pieces 14���18 ��m long, 4���6 times as long as wide. Tail short and rounded; caudal pores four pairs. Diagnosis (emended). This species is characterized by its body 2.00��� 2.66 mm long, lip region sucker-like and 23���26 ��m wide, odontostyle 29���34 ��m long or 1.1���1.3 times the lip region width, neck 521���619 ��m long, pharyngeal expansion 246���300 ��m long or occupying 45���50 % of total neck length, female genital system amphidelphic, uterus long and tripartite, V = 50���56, tail short and rounded (20���30 ��m, c = 84���117, c��� = 0.4���0.8), spicules 65���77 ��m long and 14���17 contiguous ventromedian supplements. Relationships. Labronemella labiata is similar to L. cernowitzensis, from which it differs in having wider lip region (vs 20���22 ��m) and longer odontostyle (vs 25 ��m). It is also morphometrically very close to L. hemicaudata, but it can be easily separated from this by its long, tripartite uterus (vs short and simple). Locality and habitat. Namak Abroud (approx. 36 �� 39 ' N, + 51 �� 18 ' E), northern Iran, where it was collected in soil of a natural forest. Remarks. Above description of Iranian specimens fits very well those of the male and the female studied by Andr��ssy (1985) and Ciobanu et al. (2010), respectively, although some minor morphometric differences have been also noted: slightly wider lip region (vs 23���25 ��m), shorter pharyngeal expansion (vs 300 ��m), shorter tail (vs 27���30 ��m, c��� = 0.6���0.8), and longer spicules (vs 65 ��m). These differences are herein regarded as intraspecific, geographical variations., Published as part of Pe��a-Santiago, Reyes, Abolafia, Joaqu��n & Pedram, Majid, 2012, New data on Labronemella labiata Andr��ssy, 1985 (Dorylaimida: Qudsianematidae) from Iran, with SEM study and a compendium of the genus, pp. 43-54 in Zootaxa 3271 on pages 44-49, DOI: 10.5281/zenodo.280814, {"references":["Andrassy, I. (1985) A dozen new nematode species from Hungary. Opuscula Zoologica Budapestinensis, 19 - 20, 30 - 39.","Ciobanu, M., Popovici, I. & Pena-Santiago, R. (2010) Nematodes of the order Dorylaimida from Romania: two interesting species of subfamily Qudsianematinae Jairajpuri, 1965. Russian Journal of Nematology, 18, 41 - 48."]}

Discotylenchus brevicaudatus Brzeski, 1985 (Figs. 3, 5) Measurements. See Table 2. This population was recovered from the rhizosphere of plum (Prunus domestica L.) trees in a garden, close to city of Bojnourd, Northern Khorasan province, north-eastern Iran during May 2009. Morphological and morphometric characters of the Iranian population of D. brevicaudatus agree completely with those in original description (see Table 2), although there is a wider range for some morphometric data compared to the type population i.e. the Iranian population has a longer body (0.42���0.46 vs 0.32���0.39 mm), slightly shorter stylet (6���7 vs 7���8 ��m), longer pharynx (116���132 vs 69���73 ��m), longer tail (57���77 vs 35���47 ��m), smaller c (5.9���7.8 vs 8.1���9.8), higher c' (6.3���9.6 vs 4.5 ���6.0) and slightly smaller V (68.2���71.6 vs 71���73). From a morphological perspective, the basal bulb in Iranian population is long and bottle shaped in dimension of 40���55 �� 7.5���8.5 ��m, whereas in the original population it is pyriform. The male of this species is described for the first time and is similar to females in general morphology except for character states associated with sexual differences. Testis is single, anteriorly outstretched and 91 �� 6 (85���99) ��m long. Spicules slender, ventrally curved. Gubernaculum simple, bursa smooth, adcloacal and 17 �� 2 (15���19) ��m long. Remark. As in the original description (Brzeski, 1985), the females of the Iranian population had sperm in genital tract and the males were also recovered, demonstrating the amphimictic nature of reproduction in this species. TABLE 2. Morphometrics of Discotylenchus brevicaudatus. All measurements are in ��m, and in the form: mean �� s.d. (range). Iranian population Brzeski, 1985, Published as part of Ghaemi, Razieh, Pourjam, Ebrahim, Atighi, Mohammad Reza, Pedram, Majid & Karssen, Gerrit, 2012, First record of the genus Discotylenchus Siddiqi, 1980 (Nematoda: Tylenchidae) from Iran, with description of one new and data on two known species, pp. 72-82 in Zootaxa 3493 on pages 76-77, DOI: 10.5281/zenodo.282451, {"references":["Brzeski, M. W. (1985) Four new species of Tylenchidae (Nematoda). Nematologica, 31, 424 - 432."]}

Epacrolaimus reyesi n. sp. (Figs. 1 & 2) Measurements. See table 1. Paratype females Holotype Paratype males Morphological description. FEMALE. Body very long, rather slender, tapering gradually anteriorly. Cuticle with fine criss-cross lines, comprising a thin outer, very refractive layer and a very thick inner part composed of two or three layers with the same weak refraction, 8–11 µm thick at the level of labial region, 8–13 µm at mid body and 20–22 µm at anus. Cuticle in the neck region crossed by 5–6 ventral and as many dorsal canals. Lip region 2.5 – 3.0 times wider than high, separated from the rest of body by a sharp constriction; lips well separated from each other, ovoid, provided with small inner liplets only slightly protruding. The liplets, when focusing, on their inner surface appear crossed by a kind of longitudinal corrugation (Fig. 1, B & C). Anterior circle of 6 inner labial papillae hardly visible; posterior circle of 10 (outer labial and cephalic) papillae well developed. Amphidial fovea cup shaped, with slit-like aperture bordering lip region base, ca 45 % of lip region diam. wide. Odontostyle small compared to body size, ca 4.5 times longer than wide at base; its aperture (ca 85 % of its length) affects a large part of the dorsal and lateral odontostyle surface, thus assuming an arched aspect and giving a particularly acuminate aspect to the odontostyle; odontophore rod-like and flexible. Guiding ring plicate and variable in position depending on the stylet position. Nerve ring encircling the anterior narrow part of pharynx. Pharynx muscular throughout its length, gradually widening at ca 25 % of its length. The position of the gland nuclei according to Andrássy (1998) are as follows (n= 2): DN: 40.5–42.5 %, AS 1: Not clearly seen, AS 2: 32–36 %; PS 1 and PS 2, at about the same level: 60.0– 63.5 %. Cardia hemispherical to conical, 30–40 µm wide and 27–35 µm high. Prerectum 2.4–2.7 times anal body width long and rectum 0.8–0.9 times anal body width long. Reproductive system amphidelphic with reflexed gonads, the two branches more or less equally developed; each composed of a 350–375 µm long ovary, 487–675 µm long oviduct, separated from uterus by a sphincter. In a single female at the junction between oviduct and ovary, in both branches, a deep constriction separating the ovary from a wide chamber where the most mature oocyte is contained was observed (Fig. 1 H), probably a temporary dilatation due to the passage of the oocyte into the oviduct, since it lacks in the other specimens. Uterus a simple tube, 460–712 µm long, containing 5–6 × 2–3 µm sized sperm and 170–205 × 105 µm sized eggs (1– 3 eggs were observed within the anterior genital tract and 1 in the posterior one); vagina perpendicular to body axis, 50–62 % of corresponding body width long; pars distalis vaginae 15 µm long, more refractive than adjacent cuticle; pars refringens vaginae 20–23 µm long and 10 µm wide with trapezoid sclerotized pieces; pars proximalis vaginae 80–173 µm long and 55–75 µm wide. Vulva a transverse slit surrounded by a slightly wrinkled cuticle. Tail dorsally convex, with rounded terminus, clearly shorter than anal body width long; its hyaline region 12–20 μm thick. MALE. As abundant and common as females and functional. General morphology similar to that of female, except posterior end more ventrally curved. Genital system diorchic, spicules massive, dorylaimoid, 3.8–4.6 times longer than wide. The precloacal pair of supplements located at 25 μm distance from cloacal opening and a series of 9 more or less regularly spaced ventromedian supplements ending at 153–185 μm distance from cloacal opening. Intestine joins prerectum at ca 50 μm distance from the anteriormost supplement. The ventromedian supplements row 202 μm long. Tail similar to that of female. Type habitat and locality. Soil from rhizosphere of forest trees (Fagus orientalis Lipsky), Salaheddin Kola, Mazandaran province, northern Iran. Type material. Holotype female and one paratype male deposited at Nematology Laboratory of Faculty of Agriculture of Tarbiat Modares University, Tehran, Iran. One paratype female and one paratype male deposited at the USDA Nematode Collection, Beltsville, MD, USA and one paratype female at the section of Animal Biology, Department of Biological Geological and Environmental Sciences, of the University of Catania (Italy). Diagnosis and relationships. Epacrolaimus reyesi n. sp. is an amphimictic species characterised by having a long body (7.23–7.87 mm), lip region separated by a sharp constriction by the adjoining body, lips well distinct, liplets provided with longitudinal corrugation on their inner surface, 19–22 µm long odontostyle with a wide, arched, aperture occupying ca 85 % of its length, pharynx expanding at about 25 % of its length, female reproductive apparatus didelphic with reflexed gonad, tail convex with round terminus in both sexes, 9 separated ventromedian supplements in males and 135–145 µm long spicules. The new species is compared with the two other known species of the genus, namely E. declinatoaculeatus, which it resembles more, and E. imperator. Compared to the former, the new species has a different lip region, a wider body (172–178 vs 124–128 µm at midbody region, a = 42.0– 44.5 vs 47–62), shorter (55–67 vs. 63–75 µm) rounded tail vs bluntly conoid, rounded on tip or with a minute peg, shorter spicules (135–145 vs 215 µm) and fewer male copulatory supplements (9 vs 12). Compared to the latter species, E. reyesi n. sp. has a wider body at mid region (172–178 vs 156–160 µm), shorter (55–67 vs 84 µm), rounded (vs convex–conoid, slightly subdigitate on tip) tail, shorter spicules (135–145 vs 202–210 µm), shorter tail in males (58–65 vs 70–86 µm) and fewer male supplements (9 vs 14–16). From both species, E. reyesi n. sp. differs in the longitudinal corrugation of liplets. Etymology. The species is named in honor of Prof. Reyes Peña-Santiago, the well-known authority on the taxonomy of Dorylaimida.

Paraxonchium laetificans (Andrássy, 1956) Altherr & Loof, 1969 (Fig. 3 & 4) Syn. Dorylaimus laetificans Andrássy, 1956 Aporcelaimus laetificans ( Andrássy, 1956) Andrássy, 1958 Drepanodorus laetificans ( Andrássy, 1956) Andrássy, 1958 Paraxonchium striatum Krall, 1958 Paraxonchium magnidens Thorne, 1974 Females Males n 2 4 L (1.48, 1.55) 1.54 ± 0.06 (1.46–1.62) a (23.1, 23.2) 21.5 ± 1.5 (20.5–23.5) b (3.3, 3.4) 3.5 ± 0.2 (3.2–3.6) c (34.5, 42.0) 38.5 ±2.0 (35.5–40.5) c`(0.8, 1.0) 0.8 ± 0.1 (0.7–0.9) V or T (61.5, 62.0) 49.5 ±3.0 (47.0– 52.5) Anterior end to vulva (915, 925) - Diam. at neck base (59, 63) 67 ± 4 (61–70) – at mid-body (64, 65) 72.0± 2.5 (69–75) – at anus (42, 43) 49.0± 4.5 (46–55) Lip region diam. (8.5, 9.0) 9.0±0.0 (9 – 9) Lip region height (4.5, 5.0) 5.0±0.0 (5, 5) Odontostyle length (15, 15) 15.0±0.0 (15 – 15) Odontostyle width at base (3, 3) 3.0± 0.3 (2.5 –3.0) Odontostyle length / Odontostyle width (5.0, 5.0) 5.5 ± 0.5 (5–6) Neck length (432, 460) 455.5 ±7.0 (455–460) Cardia length (16, 21) 14 ± 1 (13–15) Cardia width (13, 15) 13.5 ±1.0 (13–15) Prerectum length (102, 113) 141.0± 7.5 (132–145) Rectum length (40, 43) 53.5 ±3.0 (50–55) Tail length (36, 43) 40.0± 2.5 (37–43) Spicule length - 64.5 ±2.0 (63–67) Ventromedian supplements - 16.5 ± 0.5 (16–17) Lateral accessory pieces - 14.5 ± 0.5 (14–15) Material examined: Two females and four males from forests of Kelardasht, northern Iran; in good condition. Measurements: See Table 2. Morphological description. MALE. Body slightly curved ventrad, L shaped. Body strongly tapering anteriorly, mainly in anterior half of neck. Cuticle 2–3 µm at odontostyle level, varying to 3–5 at mid-body and 5–7 µm on tail, distinctly divided into two layers. Lateral hypodermal chords 15.5–21.5 % of corresponding body width. Lip region rounded to moderately angular, 1.6 –2.0 times as wide as high, separated from the rest of body by a constriction. Amphidial fovea cup-shaped with slit-like opening, occupying 72–75 % of lip region width. Odontostyle 5–6 times longer than wide, with the aperture 54–60 % of its length; odontophore rod-shaped, its base not clearly visible. Nerve ring located at 130–156 µm from anterior end. Pharynx composed of a slender but well muscled anterior part expanding gradually into a basal expansion occupying 42–45 % of total neck length. Cardia conical, its length 0.8 –1.0 times its width. Reproductive system diorchic, dorylaimoid, occupying ca 30.5 % of body length. Spicules dorylaimoid, well sclerotised, moderately slender, 5–6 times longer than wide. Lateral guiding pieces present in two of the four males, more or less cylindrical, with bifurcate tip, ca 15 × 3 μm sized or 5 times longer than wide. Adcloacal pair of supplements located at 9–13 μm from cloacal opening; 16–17 spaced ventromedian supplements running anteriorly, the supplements row ca 120 μm long, the posteriormost located at 70–75 μm from adcloacal pair. Rectum ca 2.5 times longer than prerectum. The latter starts in the range of the supplements row. Tail asymmetrically conical with flat ventral side and a subdigitate mucro, bearing four pores in each side (n= 1). FEMALE. General morphology similar to that of male except for its habitus straight after heat relaxation. Genital system didelphic-amphidelphic, identical to that described in Iranian population by Baniyamuddin et al. (2010). Remarks. P. laetificans was originally described on a single male found in Hungary (Andrássy, 1956). Heyns (1988) redescribed the species reporting data of numerous populations from Europe and Iraq mostly composed by females only, with the exception of a population from Switzerland composed by ten females and a male. In this paper Heyns observed that only one female of the Swiss population was impregnated while none of the other females, in the same and in the other populations observed, most of which were gravid, contained sperm. Moreover he stated that the male spicules lacked the lateral accessory pieces commonly present in dorylaimoid spicules, confirming what originally observed by Andrássy in the type specimen. In a recent study, Baniyamuddin et al. (2010) revised the family Paraxonchiidae and reported a population of this species from Shahrekord, Iran, also in this case composed by females only. The small population object of our study, otherwise, is clearly amphimictic, with males relatively abundant and females containing sperm in their uterus. The morphology and morphometric data of this population agree well with those given in the original description except that for odontostyle length, which, both in males and females, is greater compared to that in the original description (15 µm vs. 11.7 μm); but the data given for the other European populations by Heyns (1988) range from 10.5 to 13 µm and those of the Iranian population reported by Baniyamuddin et al., (2010) from 13 to 14 μm. It is noteworthy that the males in our population may possess (in two males) or lack (in two males) the lateral accessory pieces that usually accompany the spicules; it is possible that these structures are initially present in all males, as usual also in this species, but that they can be lost during the animal's life.

Epacrolaimus reyesi n. sp. (Figs. 1 & 2) Measurements. See table 1. Paratype females Holotype Paratype males Morphological description. FEMALE. Body very long, rather slender, tapering gradually anteriorly. Cuticle with fine criss-cross lines, comprising a thin outer, very refractive layer and a very thick inner part composed of two or three layers with the same weak refraction, 8���11 ��m thick at the level of labial region, 8���13 ��m at mid body and 20���22 ��m at anus. Cuticle in the neck region crossed by 5���6 ventral and as many dorsal canals. Lip region 2.5 ��� 3.0 times wider than high, separated from the rest of body by a sharp constriction; lips well separated from each other, ovoid, provided with small inner liplets only slightly protruding. The liplets, when focusing, on their inner surface appear crossed by a kind of longitudinal corrugation (Fig. 1, B & C). Anterior circle of 6 inner labial papillae hardly visible; posterior circle of 10 (outer labial and cephalic) papillae well developed. Amphidial fovea cup shaped, with slit-like aperture bordering lip region base, ca 45 % of lip region diam. wide. Odontostyle small compared to body size, ca 4.5 times longer than wide at base; its aperture (ca 85 % of its length) affects a large part of the dorsal and lateral odontostyle surface, thus assuming an arched aspect and giving a particularly acuminate aspect to the odontostyle; odontophore rod-like and flexible. Guiding ring plicate and variable in position depending on the stylet position. Nerve ring encircling the anterior narrow part of pharynx. Pharynx muscular throughout its length, gradually widening at ca 25 % of its length. The position of the gland nuclei according to Andr��ssy (1998) are as follows (n= 2): DN: 40.5���42.5 %, AS 1: Not clearly seen, AS 2: 32���36 %; PS 1 and PS 2, at about the same level: 60.0��� 63.5 %. Cardia hemispherical to conical, 30���40 ��m wide and 27���35 ��m high. Prerectum 2.4���2.7 times anal body width long and rectum 0.8���0.9 times anal body width long. Reproductive system amphidelphic with reflexed gonads, the two branches more or less equally developed; each composed of a 350���375 ��m long ovary, 487���675 ��m long oviduct, separated from uterus by a sphincter. In a single female at the junction between oviduct and ovary, in both branches, a deep constriction separating the ovary from a wide chamber where the most mature oocyte is contained was observed (Fig. 1 H), probably a temporary dilatation due to the passage of the oocyte into the oviduct, since it lacks in the other specimens. Uterus a simple tube, 460���712 ��m long, containing 5���6 �� 2���3 ��m sized sperm and 170���205 �� 105 ��m sized eggs (1��� 3 eggs were observed within the anterior genital tract and 1 in the posterior one); vagina perpendicular to body axis, 50���62 % of corresponding body width long; pars distalis vaginae 15 ��m long, more refractive than adjacent cuticle; pars refringens vaginae 20���23 ��m long and 10 ��m wide with trapezoid sclerotized pieces; pars proximalis vaginae 80���173 ��m long and 55���75 ��m wide. Vulva a transverse slit surrounded by a slightly wrinkled cuticle. Tail dorsally convex, with rounded terminus, clearly shorter than anal body width long; its hyaline region 12���20 ��m thick. MALE. As abundant and common as females and functional. General morphology similar to that of female, except posterior end more ventrally curved. Genital system diorchic, spicules massive, dorylaimoid, 3.8���4.6 times longer than wide. The precloacal pair of supplements located at 25 ��m distance from cloacal opening and a series of 9 more or less regularly spaced ventromedian supplements ending at 153���185 ��m distance from cloacal opening. Intestine joins prerectum at ca 50 ��m distance from the anteriormost supplement. The ventromedian supplements row 202 ��m long. Tail similar to that of female. Type habitat and locality. Soil from rhizosphere of forest trees (Fagus orientalis Lipsky), Salaheddin Kola, Mazandaran province, northern Iran. Type material. Holotype female and one paratype male deposited at Nematology Laboratory of Faculty of Agriculture of Tarbiat Modares University, Tehran, Iran. One paratype female and one paratype male deposited at the USDA Nematode Collection, Beltsville, MD, USA and one paratype female at the section of Animal Biology, Department of Biological Geological and Environmental Sciences, of the University of Catania (Italy). Diagnosis and relationships. Epacrolaimus reyesi n. sp. is an amphimictic species characterised by having a long body (7.23���7.87 mm), lip region separated by a sharp constriction by the adjoining body, lips well distinct, liplets provided with longitudinal corrugation on their inner surface, 19���22 ��m long odontostyle with a wide, arched, aperture occupying ca 85 % of its length, pharynx expanding at about 25 % of its length, female reproductive apparatus didelphic with reflexed gonad, tail convex with round terminus in both sexes, 9 separated ventromedian supplements in males and 135���145 ��m long spicules. The new species is compared with the two other known species of the genus, namely E. declinatoaculeatus, which it resembles more, and E. imperator. Compared to the former, the new species has a different lip region, a wider body (172���178 vs 124���128 ��m at midbody region, a = 42.0��� 44.5 vs 47���62), shorter (55���67 vs. 63���75 ��m) rounded tail vs bluntly conoid, rounded on tip or with a minute peg, shorter spicules (135���145 vs 215 ��m) and fewer male copulatory supplements (9 vs 12). Compared to the latter species, E. reyesi n. sp. has a wider body at mid region (172���178 vs 156���160 ��m), shorter (55���67 vs 84 ��m), rounded (vs convex���conoid, slightly subdigitate on tip) tail, shorter spicules (135���145 vs 202���210 ��m), shorter tail in males (58���65 vs 70���86 ��m) and fewer male supplements (9 vs 14���16). From both species, E. reyesi n. sp. differs in the longitudinal corrugation of liplets. Etymology. The species is named in honor of Prof. Reyes Pe��a-Santiago, the well-known authority on the taxonomy of Dorylaimida., Published as part of Pedram, Majid, Pourjam, Ebrahim & Vinciguerra, Maria T., 2012, Description of a new species of the rare genus Epacrolaimus Andr��ssy, 2000 (Dorylaimida, Aporcelaimidae) and new data on male of Paraxonchium laetificans (Andr��ssy, 1956) Altherr & Loof, 1969 (Dorylaimida, Paraxonchiidae) from Iran, pp. 53-61 in Zootaxa 3327 on pages 54-57, DOI: 10.5281/zenodo.210470, {"references":["Andrassy, I. (1998) Once more: the oesophageal gland nuclei in the dorylaimoid nematodes. Opuscula Zoologica Budapestinensis, 31, 165 - 170."]}

Paraxonchium laetificans (Andr��ssy, 1956) Altherr & Loof, 1969 (Fig. 3 & 4) Syn. Dorylaimus laetificans Andr��ssy, 1956 Aporcelaimus laetificans ( Andr��ssy, 1956) Andr��ssy, 1958 Drepanodorus laetificans ( Andr��ssy, 1956) Andr��ssy, 1958 Paraxonchium striatum Krall, 1958 Paraxonchium magnidens Thorne, 1974 Females Males n 2 4 L (1.48, 1.55) 1.54 �� 0.06 (1.46���1.62) a (23.1, 23.2) 21.5 �� 1.5 (20.5���23.5) b (3.3, 3.4) 3.5 �� 0.2 (3.2���3.6) c (34.5, 42.0) 38.5 ��2.0 (35.5���40.5) c`(0.8, 1.0) 0.8 �� 0.1 (0.7���0.9) V or T (61.5, 62.0) 49.5 ��3.0 (47.0��� 52.5) Anterior end to vulva (915, 925) - Diam. at neck base (59, 63) 67 �� 4 (61���70) ��� at mid-body (64, 65) 72.0�� 2.5 (69���75) ��� at anus (42, 43) 49.0�� 4.5 (46���55) Lip region diam. (8.5, 9.0) 9.0��0.0 (9 ��� 9) Lip region height (4.5, 5.0) 5.0��0.0 (5, 5) Odontostyle length (15, 15) 15.0��0.0 (15 ��� 15) Odontostyle width at base (3, 3) 3.0�� 0.3 (2.5 ���3.0) Odontostyle length / Odontostyle width (5.0, 5.0) 5.5 �� 0.5 (5���6) Neck length (432, 460) 455.5 ��7.0 (455���460) Cardia length (16, 21) 14 �� 1 (13���15) Cardia width (13, 15) 13.5 ��1.0 (13���15) Prerectum length (102, 113) 141.0�� 7.5 (132���145) Rectum length (40, 43) 53.5 ��3.0 (50���55) Tail length (36, 43) 40.0�� 2.5 (37���43) Spicule length - 64.5 ��2.0 (63���67) Ventromedian supplements - 16.5 �� 0.5 (16���17) Lateral accessory pieces - 14.5 �� 0.5 (14���15) Material examined: Two females and four males from forests of Kelardasht, northern Iran; in good condition. Measurements: See Table 2. Morphological description. MALE. Body slightly curved ventrad, L shaped. Body strongly tapering anteriorly, mainly in anterior half of neck. Cuticle 2���3 ��m at odontostyle level, varying to 3���5 at mid-body and 5���7 ��m on tail, distinctly divided into two layers. Lateral hypodermal chords 15.5���21.5 % of corresponding body width. Lip region rounded to moderately angular, 1.6 ���2.0 times as wide as high, separated from the rest of body by a constriction. Amphidial fovea cup-shaped with slit-like opening, occupying 72���75 % of lip region width. Odontostyle 5���6 times longer than wide, with the aperture 54���60 % of its length; odontophore rod-shaped, its base not clearly visible. Nerve ring located at 130���156 ��m from anterior end. Pharynx composed of a slender but well muscled anterior part expanding gradually into a basal expansion occupying 42���45 % of total neck length. Cardia conical, its length 0.8 ���1.0 times its width. Reproductive system diorchic, dorylaimoid, occupying ca 30.5 % of body length. Spicules dorylaimoid, well sclerotised, moderately slender, 5���6 times longer than wide. Lateral guiding pieces present in two of the four males, more or less cylindrical, with bifurcate tip, ca 15 �� 3 ��m sized or 5 times longer than wide. Adcloacal pair of supplements located at 9���13 ��m from cloacal opening; 16���17 spaced ventromedian supplements running anteriorly, the supplements row ca 120 ��m long, the posteriormost located at 70���75 ��m from adcloacal pair. Rectum ca 2.5 times longer than prerectum. The latter starts in the range of the supplements row. Tail asymmetrically conical with flat ventral side and a subdigitate mucro, bearing four pores in each side (n= 1). FEMALE. General morphology similar to that of male except for its habitus straight after heat relaxation. Genital system didelphic-amphidelphic, identical to that described in Iranian population by Baniyamuddin et al. (2010). Remarks. P. laetificans was originally described on a single male found in Hungary (Andr��ssy, 1956). Heyns (1988) redescribed the species reporting data of numerous populations from Europe and Iraq mostly composed by females only, with the exception of a population from Switzerland composed by ten females and a male. In this paper Heyns observed that only one female of the Swiss population was impregnated while none of the other females, in the same and in the other populations observed, most of which were gravid, contained sperm. Moreover he stated that the male spicules lacked the lateral accessory pieces commonly present in dorylaimoid spicules, confirming what originally observed by Andr��ssy in the type specimen. In a recent study, Baniyamuddin et al. (2010) revised the family Paraxonchiidae and reported a population of this species from Shahrekord, Iran, also in this case composed by females only. The small population object of our study, otherwise, is clearly amphimictic, with males relatively abundant and females containing sperm in their uterus. The morphology and morphometric data of this population agree well with those given in the original description except that for odontostyle length, which, both in males and females, is greater compared to that in the original description (15 ��m vs. 11.7 ��m); but the data given for the other European populations by Heyns (1988) range from 10.5 to 13 ��m and those of the Iranian population reported by Baniyamuddin et al., (2010) from 13 to 14 ��m. It is noteworthy that the males in our population may possess (in two males) or lack (in two males) the lateral accessory pieces that usually accompany the spicules; it is possible that these structures are initially present in all males, as usual also in this species, but that they can be lost during the animal's life., Published as part of Pedram, Majid, Pourjam, Ebrahim & Vinciguerra, Maria T., 2012, Description of a new species of the rare genus Epacrolaimus Andr��ssy, 2000 (Dorylaimida, Aporcelaimidae) and new data on male of Paraxonchium laetificans (Andr��ssy, 1956) Altherr & Loof, 1969 (Dorylaimida, Paraxonchiidae) from Iran, pp. 53-61 in Zootaxa 3327 on pages 57-59, DOI: 10.5281/zenodo.210470, {"references":["Andrassy, I. (1956) Eine interessante Nematodenfauna der Gerste, Nematologische Notizen. 4. Acta Zoologica Academiae Scientiarum Hungaricae, 2, 307 - 317.","Altherr, E. & Loof, P. A. A. (1969) Paraxonchium Krall, 1958 a valid generic name. Nematologica, 15, 430.","Baniyamuddin, M., Ahmad, W. & Jairajpuri, M. S. (2010) Review of the family Paraxonchiidae Dhanachand & Jairajpuri, 1981 and proposal of Ramphidorylaimus gen. n. Journal of Nematode Morphology and Systematics, 13 (1), 51 - 66.","Heyns, J. (1988) Redescription of Paraxonchium laetificans (Andrassy, 1956) and P. m o n h y s t e r a (Brzeski, 1964) (Nematoda: Dorylaimida). Nematologica, 34, 302 - 313"]}

Dorylaimoides alborzicus sp. n. (Figs 1 and 2; Table 1) Female. Slender nematodes of medium size. Body in the shape of an open C after fixation, tapering gradually towards both ends, more pronounced anteriorly. Cuticle with very fine transverse striations, 1���2 ��m thick between anterior end and odontophore base, 2 ��m at mid body and 2.5 ���3.0 ��m at anus level. Lateral chords 3.5���4.5 ��m wide or 11.0��� 13.5 % of corresponding body width. Lip region 2.1���2.3 times as wide as high, separated from the rest of body by a sharp constriction. Amphids cup-shaped; their slit-like aperture corresponding to 71���75 % of lip region width. Odontostyle typical of genus, asymmetrical, 1.5 ��m thick at base; its aperture 1.5 ���2.0 ��m long. Odontophore arcuate, about 1.5 times as long as odontostyle. Guiding ring simple. Nerve ring located 90���103 ��m from anterior end. Anterior part of pharynx narrow and weakly muscular, posterior expanded part composed of a 50���62 �� 12���14 ��m sized terminal bulb, occupying 25���28 % of total neck length. Gland nuclei located as follows (n= 3): DN: 79.5 ���82.0, AS 1 and AS 2 almost at same level: 23���30 and PS 1 and PS 2 at same level: 46.5���52.5. Cardia conical; body width at this level 3.0��� 3.6 times as wide as lip region width. Female reproductive system monoopisthodelphic; its anterior branch a uterine sac 37���61 ��m long or 1.1���1.7 times body width; vagina cylindrical, extended inward for 15���23 ��m, about 50.0��� 54.5 % of corresponding body width; its wall adjacent to the vulva not clearly distinct from the body cuticle.Vulva a transverse slit. Posterior genital tract 204���310 ��m long, composed of a 60���80 ��m long ovary, 120���176 ��m long oviduct with a well developed pars dilatata, a well developed sphincter and a 70���83 ��m long, thin-walled, simple uterus containing sperm with dimensions of 5.0�� 1.5 ��m; one 92 �� 26 ��m uterine egg observed. Prerectum 60���107 ��m long or 2.8���5.4 times anal body width and rectum 20���25 ��m long or 1.0��� 1.3 times anal body width. Tail straight to ventrally bent, elongate conoid with rounded terminus, characterised by a very thick cuticle and a thin internal core, 2.3 ���3.0 anal body widths in length. Male. Abundant, equal to females in number and functional. Similar to females in general morphology except for reproductive system and by having the posterior end more ventrally bent. Spicules dorylaimoid, 4.7���5.5 times longer than wide with 7���8 ��m long lateral accessory pieces. Male copulatory supplements composed of one precloacal pair at a distance of 8���11 ��m from cloacal opening and a series of 5���6 ventromedian supplementary papillae extending 16���26 ��m from cloacal pair. Tail conoid, ventrally bent with rounded end. Differential diagnosis and relationships. Dorylaimoides alborzicus sp. n. is an amphimictic species with functional males in this population. It is characterised by having the lip region separated from the rest of body by a sharp constriction, body length of 1.1���1.3 mm, 10���11 ��m long odontostyle (from tip of ventral arm to end of dorsal arm), 15���16 ��m long arcuate odontophore, opisthodelphic genital system with anterior uterine sac 37���61 ��m long or 1.1���1.7 times body width, tail 46���62 ��m long (2.3 ���3.0 times anal body width), straight to slightly ventrally bent, elongate conoid with rounded terminus, with a very thick cuticle and a thin internal core tail; males with 31���35 ��m long spicules, 5���6 ventromedian supplements and ventrally bent tail, 39���60 ��m long. The new species resembles 8 other opisthodelphic species of Dorylaimoides with conoid tails: D. arcuatus Siddiqi, 1964, D. bulbosus (Brzeski & Szczygiel, 1961) Szczygiel, 1965, D. modestus Siddiqi, 1965, D. reversus Thorne, 1964, D. saueri Baqri & Jairajpuri, 1969, D. singaporensis Ahmad & Mushtaq, 2004, D. uralicus (Nesterov, 1976) Andr��ssy, 2009, and D. venustus Andr��ssy, 1959. Compared to D. arcuatus, the new species has a somewhat longer body (L = 1.1���1.3 vs. 0.7���1.2 mm), shorter tail (c = 19.0��� 24.5 vs. 10���20), longer odontostyle (10���11 vs. 4.5 ���7.0 ��m long), tail straight or slightly ventrally arcuate vs. markedly ventrally arcuate and longer spicules (31���35 vs. 20���29 ��m). Compared to D. bulbosus, the new species has a lip region separated from the rest of body by a sharp constriction vs. not set off, anterior uterine sac 1.1���1.7 times the corresponding body width vs. about 2, longer odontostyle (10���11 vs. 8.0��� 8.5 ��m), and functional males. Compared to D. modestus, the new species has a longer body (L = 1.1���1.3 vs. 0.72���0.85 mm), lip region separated from the rest of body by a sharp constriction vs. almost continuous, shorter tail (c = 19.0��� 24.5 vs. 12.0��� 13.5) and presence of males. Compared to D. reversus, the new species has a longer body (L = 1.1���1.3 vs. 0.8 mm), shorter tail (c = 19.0��� 24.5 vs. 13), straight to ventrally curved in females vs. dorsally bent. Compared to D. saueri, the new species has longer body (L = 1.1���1.3 vs. 0.8���0.9 mm), shorter tail (c = 19.0��� 24.5 vs. 9���13; c��� = 2.3 ���3.0 vs. 5���6) and longer spicules (31���35 vs. 19���22 ��m). Compared to D. singaporensis, the new species has a longer body (L = 1.1���1.3 vs. 0.78 mm), longer odontostyle (10���11 ��m dorsal arm vs. 5���6 ��m) and longer spicules (31���35 vs. 21���22 ��m). Compared to D. uralicus, the new species has a shorter tail (46���62 vs. 69.5 ��� 92.0 ��m; c = 19.0��� 24.5 vs. 16.1���16.3;) with thin cuticle and thick core, and with males present. Compared to D. venustus, the new species has a lip region separated from the rest of body by a sharp constriction vs. slightly offset, longer odontostyle (10���11 vs. 6���7 ��m), smaller b value (5.0��� 6.5 vs. 6.8���8.8), shorter tail (c = 19.0��� 24.5 vs. 17.8���19.8; c��� = 2.3 ���3.0 vs. 4.2 ���5.0), and more posterior vulva (V = 34.7���36.5 vs. 33.1���33.7). Etymology. The specific epithet refers to the Alborz Mountains, the locality in northern Iran where the species was found. Type habitat and locality. Dorylaimoides alborzicus sp. n. was recovered from soil samples collected about the rhizosphere of unidentified species of grasses growing at 2946 m above sea level altitude, in Azad Kooh, Alborz Mountains, north of Iran, during May 2010. Type specimens. Holotype female, five paratype females and two paratype males deposited in the nematode collection of Tarbiat Modares University, Tehran, Iran. Two paratype females and two paratype males deposited in the nematode collection of the Department of Biology, University of Catania, and two males and two females in separate slides deposited in USDA Nematode Collection, Beltsville, MD, USA. Character Holotype female Females Males n - 9 8 L (��m) 1229 1199 �� 65 1176 �� 70 (1106���1303) (1041���1251) a 37.2 37.0�� 2.3 37.3 ��4.0 (33.7���39.5) (31.5 ���43.0) b 6.1 5.7 �� 0.5 5.7 �� 0.3 (5.0��� 6.5) (5.2 ���6.0) c 23.2 21.5 �� 1.7 22.0�� 2.2 (19.0��� 24.5) (20.3���26.7) c` 2.5 2.7 �� 0.2 2.4 �� 0.2 (2.3 ���3.0) (2.1���2.7) V 35.8 36.0�� 0.5 - (34.7���36.5) - Lip region diam. 8.5 8.5 �� 0.5 8.3 �� 0.5 (8���9) (8���9) Odontostyle dorsal line 10.0 10.5 �� 0.5 10.0�� 0.5 (10���11) (9.5���10.5) Odontostyle ventral line 9.0 9.3 �� 0.7 8.5 �� 0.5 (8���10) (8.0��� 9.5) Odontophore 15.0 15.0�� 0.5 16.0�� 0.5 (15���16) (15.0��� 16.5) Guiding ring from ant. end 6.5 7.3 �� 0.5 7.6 �� 0.5 (6.5 ���8.0) (7.0��� 8.5) Neck length 200.0 211 �� 14 207.5 �� 10.5 (190���232) (192.5 ���222.0) Anterior end to vulva 440.0 429 �� 17 - (400���457) - continued next page Character Holotype female Females Males Body width. at: neck base 30.0 29 �� 1 28.5 �� 1.5 (27���30) (27���31) mid-body 33.0 33 �� 1 31.7 ��2.0 (31���34) (29���34) anus 21.0 21 �� 1 22.0�� 1.5 (20���22) (19���23) Tail length 53.0 56.5 ��5.0 52.5 �� 6.5 (46���62) (39���60) Spicule length - - 33.0�� 1.5 - (31���35) Number of male VM supplements - - - - (5���6) Lateral accessory pieces - - 7.5 �� 0.5 - (7���8) Cardia length 8.0 9.7 �� 1.5 9.7 �� 1.5 (8���12) (8���11) Cardia width 8.0 8.5 �� 0.5 9 �� 1 (8���9) (8���10), Published as part of Pedram, Majid, Pourjam, Ebrahim & Vinciguerra, Maria T., 2011, Description of Dorylaimoides alborzicus sp. n. (Dorylaimida: Nematoda) from Iran, with updated compendium and key to the species of Dorylaimoides, pp. 58-68 in Zootaxa 3022 on pages 59-63, DOI: 10.5281/zenodo.202885, {"references":["Siddiqi, M. R. (1964) Three new species of Dorylaimoides Thorne & Swanger, 1936, with a description of Xiphinema orbum n. sp. (Nematoda: Dorylaimoidea). Nematologica, 9 (1963), 626 - 634.","Brzeski, M. & Szczygiel, A. (1961) Two new species of the subfamily Dorylaiminae (Nematoda, Dorylaimidae). Bulletin de l'Academie Polonaise des Sciences. Serie des Sciences Biologiques, 9, 511 - 514.","Szczygiel, A. (1965) Taxonomic status of Tarjania Brzeski & Szczygiel and redescription of Dorylaimoides bulbosus (Brzeski & Szczygiel, 1961) n. comb. (Nematoda: Leptonchidae). Nematologica, 11, 409 - 412.","Siddiqi, M. R. (1965) Five new species of soil nematodes in the genera Dorylaimoides Thorne & Swanger, 1936, and Discolaimium Thorne, 1939, from India. Nematologica, 11, 100 - 108.","Thorne, G. (1964) Nematodes of Puerto Rico: Belondiroidea new superfamily, Leptonchidae, Thorne, 1935 and Belonenchidae new family (Nemata, Adenophorea, Dorylaimida). Technical Paper Agricultural Experiment Station Puerto Rico, 39, 1 - 51.","Baqri, Q. H. & Jairajpuri, M. S. (1969) Morasia n. gen. and three new species of Dorylaimoides Thorne & Swanger, 1936 (Nematoda: Dorylaimoidea) from India. Nematologica, 15, 408 - 424.","Ahmad, W. & Mushtaq, P. (2004) Five new species of Dorylaimoides Thorne & Swanger (Nematoda: Dorylaimida) from Singapore. International Journal of Nematology, 14, 99 - 110.","Nesterov, P. I. (1976) (New species phytonematodes of the order Dorylaimida occurring in Moldavia and other areas of the USSR). Izvestiya Akademii Nauk Moldavskoi SSR (Biol Khim), 1976, 52 - 60.","Andrassy, I. (2009) Free-living nematodes of Hungary (Nematoda Errantia), III. - In: Pedozoologica Hungarica, 5, 608 pp.","Andrassy, I. (1959) Neue und wenig bekannte Nematoden aus Jugoslawien. Annales Historico Naturales Musei Nationalis Hungarici, 51, 259 - 275."]}

Enchodelus sardashtensis * sp. n. (Figs 2, 3) MATERIAL EXAMINED Ten females from one location. MEASUREMENTS See Table 3. Female Moderately slender nematodes of medium size, 1.29- 1.50 mm long. Body cylindrical, tapering towards anterior end. Habitus curved ventrad upon fixation, adopting an open C-shape. Cuticle two-layered, 2.0-3.0 µ m thick in anterior region, 1.5-3.0 µ m at mid-body and 4-5 µ m on tail, outer layer bearing very fine transverse striations which are more conspicuous at dorsal surface of tail. Lip region moderately angular, offset by a marked depression, 2.3-2.8 times as broad as high, ca one-third of body diam. at neck base, lips mostly amalgamated, labial papillae distinct, visibly protruding above lip region contour. Amphid fovea cup-shaped, opening at level of labial depression, aperture 8-10 µ m or three-fifths to twothirds of lip region diam. Odontostyle typical of genus, comparatively long and slender, 1.5-2.0 µ m diam., ca 11.5-14.0 times as long as wide, 1.5-1.8 times longer than lip region diam. and ca 1.9% of total body length. Odontophore 1.3-1.7 odontostyle lengths long, with well developed basal flanges. Guiding ring ‘double’, situated at 15-17 µ m from anterior end. Pharynx consisting of slender but muscular anterior section enlarging gradually, basal expansion 3.5-3.8 times as long as broad, ca twice as long as body diam. at its level and occupying 35- 38% of total neck length, gland nuclei located as follows: DN = 68-72, AS obscure, PS = 52-59. Nerve ring situated at 125-133 µ m from anterior end. Cardia rounded conoid, (10-17) × (10-13) µ m, almost as long as wide. Distinct dorsal, ovoid to spindle-shaped cellular mass, ca 40 µ m long, present in all specimens between cardia and proximal end of anterior ovary. Green material observed inside intestine. Genital system didelphic-amphidelphic, both branches equally and well developed, anterior 247- 285 µ m and posterior 230-270 µ m long, ovaries usually large, 95-182 µ m long, often reaching and surpassing sphincter level, oviduct 115-150 µ m or 1.9-2.5 body diam. long, joining ovary subterminally, consisting of a tubular part and a well developed pars dilatata. Oviduct and uterus separated by a distinct sphincter. Uterus 95- 145 µ m long, tripartite, i.e., consisting of a wide proximal region with distinct lumen, a more slender intermediate section with very narrow lumen, and a well developed, spheroid distal portion. One or two uterine eggs, (86-97)× (37-51) µ m, observed within one or both genital tracts, with thick shell. Vagina extending inwards two-fifths to one-half of body diam., pars proximalis (19-22) × (18- 20) µ m, with somewhat sigmoid walls and surrounded by weak musculature, pars refringens consisting of (in lateral view) two triangular sclerotisations, measuring (5-6)×(5- 7) µ m; pars distalis 4.0-5.0 µ m long. Vulva a transverse slit. Prerectum 2.5-3.5 anal body diam. long. Rectum 25- 36 µ m or almost as long as anal body diam. Tail rounded conoid, ventrally almost straight, dorsally convex, a few large and distinct saccate bodies present in ventral region. Two pairs of caudal pores, one subdorsal, one practically lateral (in one female three pores observed). * The species name refers to Sardasht, the geographical region from where it was collected. Male Unknown. TYPE HABITAT AND LOCALITY Soil in the rhizosphere of grasses, Shalmash village, Sardasht, western Azarbayjan, north-western Iran. TYPE MATERIAL Female holotype and five female paratypes deposited in the Nematode collection of the Faculty of Agriculture, University of Tarbiat Modarre, Tehran, Iran. Five female paratypes deposited with the Nematode Collection of Departamento de Biología Animal, Biología Vegetal y Ecología, University of Jaén, Spain and two paratype females deposited with USDA Nematode Collection, Beltsville, MD, USA. DIAGNOSIS AND RELATIONSHIPS The new species is characterised by its body length of 1.3-1.5 mm, lip region offset by marked depression and 13-15 µ m broad, odontostyle 22-27 µ m long or 1.5- 1.8 times the lip region diam., neck 290-310 µ m long, pharyngeal expansion 100-110 µ m long or 35-38% of total neck length, a dorsal cellular mass present between cardia and proximal end of anterior ovary, uterus tripartite and 95-145 µ m long, V = 44-48, female tail rounded conoid (19-23 µ m, c = 61-76, c&lcedil; = 0.7-0.8) and bearing saccate bodies, and male unknown. Having a comparatively small size (L * 1.5 mm), narrow lip region (up to 15 µ m) and short odontostyle (< 30 µ m), E. sardashtensis sp. n. is very similar to E. altherri Vinciguerra & De Francisci, 1973 and E. arcticus Nesterov, 1976, two close, poorly described, species (see also remarks), only known to occur in their corresponding type locations (cf., Guerrero et al., 2008). It differs from E. altherri in its shorter odontophore (35-37 vs 42-45 µ m) and total stylet length (58-65 vs 67-72 µ m), longer neck (290-310 vs 236-276 µ m), more anterior vulva (V = 44- 48 vs 50-52), shorter female tail (c = 61-76 vs 55-58), and male absent vs present, and from E. arcticus in its longer odontostyle (ca 1.5-1.8 vs 1.2 times the lip region diam.), relatively shorter pharyngeal expansion (35-38 vs ca 43% of total neck length), more anterior vulva (V = 44-48 vs 49-51), shorter female tail (19-23 vs 23-31 µ m, c = 61- 76 vs c = 48-53) with vs without saccate bodies, and male absent vs present. MOLECULAR STUDY Ribosomal DNA near-full-length small subunit 18S gene of E. sardashtensis sp. n. was sequenced and deposited in GenBank under accession number HQ404367. Sequencing on internal transcribed spacer and partial 5.8S gene was not successful. Blast search revealed this sequence is unique without identical match with any other species in GenBank. The 18S of this species had 99% identity with E. macrodorus voucher M53 from Iran ( FJ042953) with 3 bp differences over 1190 bp aligned sequences, Enchodelus sp. (AY284793) with 5 bp differences over 1311 bp aligned sequences, Enchodelus sp. (AY284792) with 9 bp differences over 1311 bp aligned sequences, Enchodelus sp. (EP207247) with 12 bp differences over 1311 bp aligned sequences, and 96% identity with E. veletensis voucher M34 from Iran ( EU477379) with 38 bp differences and 3 gaps over 1193 bp aligned sequences. Phylogenetic analysis revealed that E. sardashtensis sp. n. is closest to E. macrodorus M53 (Fig. 4) and is in the round-tail clade (Fig. 5). This 18S fragment is very conserved; E. sardashtensis sp. n. has identical sequence with E. babakicus, E. longispiculus, E. macrodorus and E. hopedorus. REMARKS Some (although minimal) doubts persist as to the true identity of this Iranian population due to the fact that the original descriptions of its most similar species, E. altherri and E. arcticus, lack many details necessary to carry out a detailed comparison. Nevertheless, the differences observed, especially those concerning the length of odontostyle/odontophore/total stylet, pharynx length and vulva position, are significant enough to support the proposal of a new taxon.

Enchodelus groenlandicus (Ditlevsen, 1927) Thorne, 1939 = Dorylaimus (Doryllium) groenlandicus Ditlevsen, 1927 = Dorylaimellus groenlandicus (Ditlevsen, 1927) Thorne & Swanger, 1936 (Fig. 1) MATERIAL EXAMINED Four females collected from one location. MEASUREMENTS See Table 1. DISTRIBUTION Collected in March, 2010 from Ahar Village, Tehran Province, Iran, in the rhizosphere of grasses. REMARKS The examined specimens fit very well with the recent redescription of this species by Guerrero et al. (2008a), although some observed morphometric differences are considered as intraspecific variation. The Iranian population has a slightly longer pharyngeal expansion (140-162 vs 111-142 µ m) and shorter female tail (19-24 vs 22-37 µ m; c = 73-104 vs 53-83; c&lcedil; = 0.4-0.6 vs 0.6-0.8). Moreover, taking into account the tripartite nature of the uterus in this species, as illustrated by Guerrero et al. (2008a), the Iranian population has a less developed pars distalis than Iberian populations, although the pars dilatata oviductus is more developed than in the Iberian specimens and female tail lacks saccate bodies (vs some very small bodies observed). One of the four females observed had two vulvae. Its morphometrics are: L = 1.89 mm, a = 23.9, b = 5.1, c = 82, c&lcedil; = 0.5, V 1 = 34.0, V 2 = 41.5, lip region diam. = 22 µ m, odontostyle length = 45 µ m, odontophore length = 53 µ m, neck length = 372 µ m, pharyngeal expansion length = 155 µ m, anal body diam. = 45 µ m and tail length = 23 µ m. This is the first report of the species in Iran., Published as part of Pedram, Majid, Pourjam, Ebrahim, Robbins, Robert T., Ye, Weimin & Santiago, Reyes Peña-, 2011, Description of one new, and new data on two known, species of Enchodelus Thorne, 1939 (Dorylaimida: Nordiidae) from Iran, pp. 729-740 in Nematology 13 (6) on page 730, DOI: 10.1163/138855410X545786, http://zenodo.org/record/8114679

Enchodelus sardashtensis * sp. n. (Figs 2, 3) MATERIAL EXAMINED Ten females from one location. MEASUREMENTS See Table 3. Female Moderately slender nematodes of medium size, 1.29- 1.50 mm long. Body cylindrical, tapering towards anterior end. Habitus curved ventrad upon fixation, adopting an open C-shape. Cuticle two-layered, 2.0-3.0 µ m thick in anterior region, 1.5-3.0 µ m at mid-body and 4-5 µ m on tail, outer layer bearing very fine transverse striations which are more conspicuous at dorsal surface of tail. Lip region moderately angular, offset by a marked depression, 2.3-2.8 times as broad as high, ca one-third of body diam. at neck base, lips mostly amalgamated, labial papillae distinct, visibly protruding above lip region contour. Amphid fovea cup-shaped, opening at level of labial depression, aperture 8-10 µ m or three-fifths to twothirds of lip region diam. Odontostyle typical of genus, comparatively long and slender, 1.5-2.0 µ m diam., ca 11.5-14.0 times as long as wide, 1.5-1.8 times longer than lip region diam. and ca 1.9% of total body length. Odontophore 1.3-1.7 odontostyle lengths long, with well developed basal flanges. Guiding ring ‘double’, situated at 15-17 µ m from anterior end. Pharynx consisting of slender but muscular anterior section enlarging gradually, basal expansion 3.5-3.8 times as long as broad, ca twice as long as body diam. at its level and occupying 35- 38% of total neck length, gland nuclei located as follows: DN = 68-72, AS obscure, PS = 52-59. Nerve ring situated at 125-133 µ m from anterior end. Cardia rounded conoid, (10-17) × (10-13) µ m, almost as long as wide. Distinct dorsal, ovoid to spindle-shaped cellular mass, ca 40 µ m long, present in all specimens between cardia and proximal end of anterior ovary. Green material observed inside intestine. Genital system didelphic-amphidelphic, both branches equally and well developed, anterior 247- 285 µ m and posterior 230-270 µ m long, ovaries usually large, 95-182 µ m long, often reaching and surpassing sphincter level, oviduct 115-150 µ m or 1.9-2.5 body diam. long, joining ovary subterminally, consisting of a tubular part and a well developed pars dilatata. Oviduct and uterus separated by a distinct sphincter. Uterus 95- 145 µ m long, tripartite, i.e., consisting of a wide proximal region with distinct lumen, a more slender intermediate section with very narrow lumen, and a well developed, spheroid distal portion. One or two uterine eggs, (86-97)× (37-51) µ m, observed within one or both genital tracts, with thick shell. Vagina extending inwards two-fifths to one-half of body diam., pars proximalis (19-22) × (18- 20) µ m, with somewhat sigmoid walls and surrounded by weak musculature, pars refringens consisting of (in lateral view) two triangular sclerotisations, measuring (5-6)×(5- 7) µ m; pars distalis 4.0-5.0 µ m long. Vulva a transverse slit. Prerectum 2.5-3.5 anal body diam. long. Rectum 25- 36 µ m or almost as long as anal body diam. Tail rounded conoid, ventrally almost straight, dorsally convex, a few large and distinct saccate bodies present in ventral region. Two pairs of caudal pores, one subdorsal, one practically lateral (in one female three pores observed). * The species name refers to Sardasht, the geographical region from where it was collected. Male Unknown. TYPE HABITAT AND LOCALITY Soil in the rhizosphere of grasses, Shalmash village, Sardasht, western Azarbayjan, north-western Iran. TYPE MATERIAL Female holotype and five female paratypes deposited in the Nematode collection of the Faculty of Agriculture, University of Tarbiat Modarre, Tehran, Iran. Five female paratypes deposited with the Nematode Collection of Departamento de Biología Animal, Biología Vegetal y Ecología, University of Jaén, Spain and two paratype females deposited with USDA Nematode Collection, Beltsville, MD, USA. DIAGNOSIS AND RELATIONSHIPS The new species is characterised by its body length of 1.3-1.5 mm, lip region offset by marked depression and 13-15 µ m broad, odontostyle 22-27 µ m long or 1.5- 1.8 times the lip region diam., neck 290-310 µ m long, pharyngeal expansion 100-110 µ m long or 35-38% of total neck length, a dorsal cellular mass present between cardia and proximal end of anterior ovary, uterus tripartite and 95-145 µ m long, V = 44-48, female tail rounded conoid (19-23 µ m, c = 61-76, c&lcedil; = 0.7-0.8) and bearing saccate bodies, and male unknown. Having a comparatively small size (L * 1.5 mm), narrow lip region (up to 15 µ m) and short odontostyle (< 30 µ m), E. sardashtensis sp. n. is very similar to E. altherri Vinciguerra & De Francisci, 1973 and E. arcticus Nesterov, 1976, two close, poorly described, species (see also remarks), only known to occur in their corresponding type locations (cf., Guerrero et al., 2008). It differs from E. altherri in its shorter odontophore (35-37 vs 42-45 µ m) and total stylet length (58-65 vs 67-72 µ m), longer neck (290-310 vs 236-276 µ m), more anterior vulva (V = 44- 48 vs 50-52), shorter female tail (c = 61-76 vs 55-58), and male absent vs present, and from E. arcticus in its longer odontostyle (ca 1.5-1.8 vs 1.2 times the lip region diam.), relatively shorter pharyngeal expansion (35-38 vs ca 43% of total neck length), more anterior vulva (V = 44-48 vs 49-51), shorter female tail (19-23 vs 23-31 µ m, c = 61- 76 vs c = 48-53) with vs without saccate bodies, and male absent vs present. MOLECULAR STUDY Ribosomal DNA near-full-length small subunit 18S gene of E. sardashtensis sp. n. was sequenced and deposited in GenBank under accession number HQ404367. Sequencing on internal transcribed spacer and partial 5.8S gene was not successful. Blast search revealed this sequence is unique without identical match with any other species in GenBank. The 18S of this species had 99% identity with E. macrodorus voucher M53 from Iran ( FJ042953) with 3 bp differences over 1190 bp aligned sequences, Enchodelus sp. (AY284793) with 5 bp differences over 1311 bp aligned sequences, Enchodelus sp. (AY284792) with 9 bp differences over 1311 bp aligned sequences, Enchodelus sp. (EP207247) with 12 bp differences over 1311 bp aligned sequences, and 96% identity with E. veletensis voucher M34 from Iran ( EU477379) with 38 bp differences and 3 gaps over 1193 bp aligned sequences. Phylogenetic analysis revealed that E. sardashtensis sp. n. is closest to E. macrodorus M53 (Fig. 4) and is in the round-tail clade (Fig. 5). This 18S fragment is very conserved; E. sardashtensis sp. n. has identical sequence with E. babakicus, E. longispiculus, E. macrodorus and E. hopedorus. REMARKS Some (although minimal) doubts persist as to the true identity of this Iranian population due to the fact that the original descriptions of its most similar species, E. altherri and E. arcticus, lack many details necessary to carry out a detailed comparison. Nevertheless, the differences observed, especially those concerning the length of odontostyle/odontophore/total stylet, pharynx length and vulva position, are significant enough to support the proposal of a new taxon., Published as part of Pedram, Majid, Pourjam, Ebrahim, Robbins, Robert T., Ye, Weimin & Santiago, Reyes Peña-, 2011, Description of one new, and new data on two known, species of Enchodelus Thorne, 1939 (Dorylaimida: Nordiidae) from Iran, pp. 729-740 in Nematology 13 (6) on pages 734-737, DOI: 10.1163/138855410X545786, http://zenodo.org/record/8114679

Enchodelus longispiculus Guerrero, Liébanas & Peña-Santiago, 2008 MATERIAL EXAMINED Thirty-nine females and 27 males from three localities. MEASUREMENTS See Table 2. MOLECULAR STUDY Ribosomal DNA near-full-length small subunit 18S gene, internal transcribed spacer and partial 5.8S gene of E. longispiculus were sequenced and deposited in GenBank under the accession number HM851184. Blast search of the 2562 nucleotides yielded the highest match with other Enchodelus species. On the 18S, E. longispiculus had an identical sequence with E. babakicus voucher M32 from Iran (FJ042952). The 18S of this species had 99% identity with E. macrodorus voucher M53 from Iran (FJ042953) with 2 bp differences, Enchodelus sp. (AY284793) with 5 bp differences, Enchodelus sp. (AY284792) with 9 bp differences, Enchodelus sp. (EP207247) with 15 bp differences, and 97% identity with E. veletensis voucher M34 from Iran (EU477379) with 35 bp differences and six gaps. However, the ITS1 region of E. longispiculus had considerable variation compared with three Iranian species including E. babakicus voucher M32 (FJ042950, identities = 601 / 725 (82.9%), gaps = 27 / 725 (3.7%)), E. macrodorus voucher M53 (FJ042951, identities = 568 / 729 (77.9%), gaps = 85 / 729 (11.7%)), and E. veletensis M34 (EU477380, identities = 353 / 672 (52.5%), gaps = 70 / 672 (10.4%)). These significant differences indicate that they are all distinct species. No ITS sequence data is available to compare for other Enchodelus species from GenBank. Figure 4 is a phylogenetic tree from a multiple alignment with 1693 total characters. This tree includes some dorylaimid species with a high match from the blast search result in GenBank. This tree is consistent with our previous results (Pedram et al., 2009a, b). Enchodelus longispiculus is identical to E. babakicus and they are in the same clade with E. sardashtensis sp. n., E. macrodorus and three other unidentified Enchodelus spp. with 100% support. This clade in the Nordiidae is sister to Eudorylaimus Andrássy, 1959 in Qudsianematidae, Epidorylaimus Andrássy, 1986 in Qudsianematidae and Prodorylaimus Andrássy, 1959 in Dorylaimidae. Enchodelus veletensis voucher M34 from Iran is not monophyletic with above-mentioned Enchodelus species. Figure 5 is a phylogenetic tree from a multiple alignment with only 638 total characters, so that all species in Enchodelus from GenBank can be included to examine their relationships. This tree is also consistent with our previous results (Pedram et al., 2009a, b) with two significant clades, viz. conical tail vs rounded tail (the clade to which E. longispiculus belongs). This tree shows that the 3 &lcedil; portion of 18S is highly conserved. Many Enchodelus species share identical DNA sequences although they can be easily differentiated morphologically. * Total range of several populations (taken from Guerrero et al., 2008a). DISTRIBUTION Found in three localities, in the rhizosphere of grasses, in Kaleibar, Eastern Azarbayjan, Chorreh village, Roudbar, Gilan province, and Astara, Gilan province, Iran. Collected in July, August and November, 2009. REMARKS The available data on E. longispiculus, i.e., its original description from the Iberian Peninsula (Guerrero et al., 2008b) and its subsequent report from Romania (Ciobanu et al., 2010), suggest that it might be a widely distributed species. It shows important intraspecific variations, especially in some morphometric features, e.g., body size (L = 1.07-1.87 mm), odontostyle length (24-34 µ m), vulva position (V = 41-51), etc. The three Iranian populations of the species herein studied fit well with those previously known, but it is morphometrically closer to Romanian material in its body size. Moreover, the range of some measurements is wider than that observed so far (although almost always overlaps it) in such parameters as lip region 13-15 µ m (vs 14-18 µ m in previous studies), body diam. at mid-body 40-55 µ m (vs 47-71 µ m) and tail length 15-37 µ m (vs 17-29 µ m). This is the second report of the species in Asia, although the first one in Uzbekistan by Tulaganov and Usmanova (1978) is rather problematic since no description or illustration was provided (cf. Andrássy, in lit.)., Published as part of Pedram, Majid, Pourjam, Ebrahim, Robbins, Robert T., Ye, Weimin & Santiago, Reyes Peña-, 2011, Description of one new, and new data on two known, species of Enchodelus Thorne, 1939 (Dorylaimida: Nordiidae) from Iran, pp. 729-740 in Nematology 13 (6) on pages 730-734, DOI: 10.1163/138855410X545786, http://zenodo.org/record/8114679

Enchodelus babakicus * n. sp. (Figs 1-4) MEASUREMENTS See Table 1. DESCRIPTION Female Moderately slender nematodes of medium size, 1.21- 1.56 mm long. Habitus slightly curved ventrad when heat relaxed, adopting an open C-shape. Body cylindrical, tapering gradually towards both ends but more so anteriorly. Cuticle two-layered, external layer ca 1.0-2.0 µ m thick along entire body, internal layer 2.0-3.0 µ m thick in anterior region, 1.0-1.5 µ m at mid-body and 5.0-7.0 µ m on tail. Cuticle with fine but distinct transverse striations, especially visible at neck and tail regions. Lateral cords granular, forming 15-20% of body diam. at mid-body. Lip region rounded, clearly separated from adjacent body by a distinct constriction, 2.4 times as broad as high or ca one-third of body diam. at neck base. Lips amalgamated, labial and cephalic papillae scarcely projecting above lip contour. Amphidial fovea stirrup shaped, aperture slitlike, located at level of labial constriction, occupying 57- 67% of lip diam. Cheilostom cylindrical, with distinct indentation resembling an additional guiding ring, as described by Coomans (1985) in species of Californidorus and Enchodelus. Odontostyle 2.24-2.34 lip diam. long, well sclerotised with furcate base, 2.5-3.0 µ m in diam. or 13.6-15.3 times as long as broad, guiding ring double, located at 26-30 µ m or 1.3-1.6 lip diam. from anterior end. More or less spherical cavity with weakly sclerotised walls observed just posterior to guiding ring. Odontophore flanged, 1.0-1.2 times longer than odontostyle. Nerve ring located 130-143 µ m from anterior end. Pharynx composed of slender but muscular anterior portion and a muscular expanded part occupying 35.5-38.5% of total neck length, pharyngeal gland nuclei located as follows: DN = 69.0 ± 1.5 (67-72); S 1 N 1 and S 1 N 2 not observed; S 2 N 1 = 54.8 ± 1.0 (53-56); S 2 N 2 = 56.0 ± 1.9 (53-58). Very delicate membrane surrounding pharyngeal expansion. In some specimens, this structure apparently extending along ventral and/or dorsal side of pharyngeal expansion. Cardia hemispherical, about as broad as long, 12-13 × 11-13 µ m in size. Reproductive system amphididelphic, both branches equally developed. Ovaries reflexed, short, not reaching sphincter level, measuring 39- 63 µ m long, oocytes first in two or more rows, then in one row, distal part of ovary consisting of a well developed blind sac, similar to that described by Coomans (1964) in Discolaimus and Longidorus species. Oviduct joining ovary subterminally, close to proximal end of blind sac. Oviduct consisting of tubular part and well developed pars dilatata. Distinct sphincter visible between oviduct and uterus. Uterus 130-175 µ m or 1.9-2.5 body diam. long, tripartite, consisting of a distal spheroid pars dilatata, a slender intermediate portion and a wider proximal part, these three regions generally showing only weak development and obscured in some individuals by intestinal contents. Vagina extending inwards 51-60% of corresponding body diam. Pars proximalis vaginae 21.5 × 23.5 µ m, pars refringens composed of two trapezoidal, well sclerotised, pieces with a combined width of 16.5 × 4.0 µ m, pars distalis vaginae 6.3 µ m long (holotype). Sperm not observed in female genital tract. Prerectum 4.5-8.5 times and rectum 0.9-1.6 anal body diam. long. Tail short, rounded conoid, small saccate bodies observed along ventral surface, cuticle at terminus forming 42-57% of total tail length, two pairs of caudal pores located at middle of tail, one subdorsal, the other lateral. Male General morphology similar to that of female except for posterior region curved more ventrad. Reproductive system composed of two testes, anterior 179-262 and posterior 167-257 µ m long. Sperms ellipsoid, measuring 8.0-9.5 × 3.5 µ m. Spicules dorylaimoid, 1.3-1.8 anal diam. long and 3.9-4.8 times longer than wide, lateral accessory pieces paired, more or less cylindrical with bifurcate end, measuring 10.5-12.0×2.5 µ m. Series of 10-14 spaced ventromedian supplements starting 24- 32 µ m from adcloacal pair with posteriormost supplement located within range of retracted spicules, adcloacal pair located at 7.0-11.5 µ m from cloacal opening. Tail short conoid, dorsally convex, slightly ventrally concave with bluntly rounded terminus. Cuticle along ventral surface lacking saccate bodies. * The specific epithet is derived from Babak Fort, a historic location in the East Azarbaijan province of Iran and where the new species was found. M. Pedram et al. Juveniles Body length gradually increasing with stage. Comparison of length of functional odontostyle and replacement odontostyle identified four juvenile stages. Lip region in J1 flat, in J2 offset by a less marked constriction than in J3, J4 and adults. Lip region becoming progressively broader from J1 to J4. Genital primordium at vulval region observed in some J4. Tail in J1 conical, relatively much longer than in subsequent stages (c&lcedil; = 3), tail in J2- J4 similar in shape to that of female although becoming relatively shorter with stage (c&lcedil; = 1.0, 0.6-0.9 and 0.5-0.7 in J2, J3 and J4, respectively). TYPE MATERIAL Holotype female, ten paratype females, ten paratype males and paratype juveniles deposited in the Nematode Collection of the Faculty of Agriculture, University of Tabriz, Tabriz, Iran; two paratype females and two paratype males in separate slides deposited at CABI UK- Centre, Surrey, UK, and USDA Nematode Collection, Beltsville, MD, USA. TYPE HABITAT AND LOCALITY Rhizosphere of grasses from natural grasslands, Kaleibar City, mountains of Babak Fort, north-west Iran. DIAGNOSIS AND RELATIONSHIPS The new species is characterised by its 1.21-1.56 mm body length, lip region offset by marked constriction and 17.0-19.0 µ m broad, odontostyle 40-45 µ m long, odontophore flanged and 37-51 µ m long, neck region 283- 413 µ m long, pharyngeal expansion 100-136 µ m long or 35.5-38.5% of total neck length, amphididelphic female genital system, tripartite uterus, vulva pre-equatorial (V = 44-49), tail short, rounded (16-22 µ m long, c = 61- 93, c&lcedil; = 0.5- 0.7 in female; 19-30 µ m long, c = 49-72, c&lcedil; = 0.5- 0.9 in male), male as frequent as female with 49-61 µ m long spicules and 10-14 spaced ventromedian supplements, the posteriormost of which being within the range of the retracted spicules. Because of the morphology of the tail and the odontostyle length, E. babakicus n. sp. can be classified un- der the E. macrodorus -group as defined by Guerrero et al. (2008a). This group is characterised by the presence of a rather long odontostyle (> 35 µ m), odontophore with well developed flanges, tripartite uterus and rounded tail, and comprises the species described above together with E. distinctus Ahmad & Jairajpuri, 1980, E. groenlandicus (Ditlevsen, 1927) Thorne, 1939, E. macrodorus (de Man, 1880) Thorne, 1939, E. microdoroides Baqri & Jairajpuri, 1974 and E. saxifragae Popovici, 1995. In having a lip region offset by a strong constriction, the new species can be distinguished from E. groenlandicus, E. macrodorus and E. microdoroides. Furthermore, it can be separated from E. groenlandicus by its shorter female body (1.21-1.56 vs 1.57-2.50 mm), shorter odontostyle, odontophore and total stylet (40-45, 37-51 and 79-95 µ m, respectively, vs 44-53, 45-55 and 94-108 µ m), shorter and less developed uterus and frequent presence of males vs absence; from E. macrodorus, probably the closest relative, apart from the deep constriction at the lip region, the new species differs by its slightly more posteriorly located vulva (V = 44-49 vs 37-47), slightly longer uterus (130-175 vs 56-143 µ m or 1.9-2.5 vs 0.9-2.0 body diam.), female tail more conoid (vs almost hemispherical) and greater number of ventromedian supplements (10-14 vs 6-12); and from E. microdoroides by its slightly longer female body (1.21-1.56 vs 0.9-1.3 mm), broader lip region (17.0-19.0 vs 13.0-14.0 µ m), guiding ring located more anterior (26-30 vs 37-39 µ m from anterior end) and males with longer spicules (49-61 vs 45-50 µ m). The new species resembles the remaining two species, namely E. distinctus and E. saxifragae, in having the lip region offset by a deep constriction. However, it can be distinguished from E. distinctus by its shorter female body (1.21-1.56 vs 1.85 mm), longer odontostyle (40-45 vs 36 µ m), pre-equatorial vulva (V = 44-49 vs 53) and males frequent vs absent. Finally, the new species differs from E. saxifragae by the much shorter female body length (1.21-1.56 vs 1.80-2.38 mm), female genital system with less developed uterus, differently shaped pars refringens vaginae (trapezoidal vs arcuate drop-shaped), shorter tail (16-22 vs 22-40 µ m) and males with slightly shorter spicules (49-61 vs 56-70 µ m) and fewer ventromedian supplements (10-14 vs 13-16). In the key provided by Guerrero et al. (2008), E. babakicus n. sp. can either be keyed out together with E. microdoroides in step 1, due to the small size of some individuals, or with E. saxifragae in step 3. However, it can be easily distinguished from both by the characteristics listed above.

Enchodelus macrodorus (de Man, 1880) Thorne, 1939 = Dorylaimus macrodorus de Man, 1880 = Dorylaimus (Doryllium) macrodorus (de Man, 1880) Ditlevsen, 1928 = Dorylaimellus macrodorus (de Man, 1880) Thorne & Swanger, 1936 (Fig. 5) MEASUREMENTS See Table 1. REMARKS The morphometrics of the Iranian population are within the accepted values for the species as stated by Guerrero et al. (2008a), with only a small broadening of some measurement ranges such as odontophore length (46-57 vs 42-55 µ m), body diam. (68-92 vs 56-86 µ m; a = 17-24 vs 19-32) and tail length (17-23 vs 22-37 µ m; c&lcedil; = 0.4-0.6 vs 0.5-0.7). The following features have been considered of special relevance when comparing this species with E. babakicus n. sp.: i) labial constriction absent vs present; ii) vulva strikingly more anterior (V = 41.5-44.5 vs 44- 49); iii) slightly shorter uterus (119-149 vs 130-175 µ m or 1.6-2.2 vs 1.9-2.5 times corresponding body diam.); iv) males not found vs common (all Enchodelus males found in the sample had a lip morphology typical of E. babakicus n. sp. which had a sex ratio of 1.23 females per male). This evidence supports the suggestion by Guerrero et al. (2008a) that E. macrodorus males are very rare. In that study, 30 records of E. macrodorus were analysed, 12 of these being considered as reasonably confirmed although only three reported the presence of males. Detailed descriptions of E. macrodorus males are not available in the literature and there are no data on important measurements, such as spicule length, which could support a more accurate diagnosis. Individuals identified as E. macrodorus were collected from the same sample as E. babakicus n. sp. and represent the first report of this species from Iran., Published as part of Pedram, Majid, Niknam, Gholamreza, Guerrero, Pablo, Ye, Weimin & Robbins, Robert T., 2009, Morphological and molecular characterisation of Enchodelus babakicus n. sp. and E. macrodorus Thorne, 1939 (Nematoda: Nordiidae) from Iran, pp. 895-907 in Nematology 11 (6) on page 902, DOI: 10.1163/156854109X430563, http://zenodo.org/record/8114788

Enchodelus babakicus * n. sp. (Figs 1-4) MEASUREMENTS See Table 1. DESCRIPTION Female Moderately slender nematodes of medium size, 1.21- 1.56 mm long. Habitus slightly curved ventrad when heat relaxed, adopting an open C-shape. Body cylindrical, tapering gradually towards both ends but more so anteriorly. Cuticle two-layered, external layer ca 1.0-2.0 µ m thick along entire body, internal layer 2.0-3.0 µ m thick in anterior region, 1.0-1.5 µ m at mid-body and 5.0-7.0 µ m on tail. Cuticle with fine but distinct transverse striations, especially visible at neck and tail regions. Lateral cords granular, forming 15-20% of body diam. at mid-body. Lip region rounded, clearly separated from adjacent body by a distinct constriction, 2.4 times as broad as high or ca one-third of body diam. at neck base. Lips amalgamated, labial and cephalic papillae scarcely projecting above lip contour. Amphidial fovea stirrup shaped, aperture slitlike, located at level of labial constriction, occupying 57- 67% of lip diam. Cheilostom cylindrical, with distinct indentation resembling an additional guiding ring, as described by Coomans (1985) in species of Californidorus and Enchodelus. Odontostyle 2.24-2.34 lip diam. long, well sclerotised with furcate base, 2.5-3.0 µ m in diam. or 13.6-15.3 times as long as broad, guiding ring double, located at 26-30 µ m or 1.3-1.6 lip diam. from anterior end. More or less spherical cavity with weakly sclerotised walls observed just posterior to guiding ring. Odontophore flanged, 1.0-1.2 times longer than odontostyle. Nerve ring located 130-143 µ m from anterior end. Pharynx composed of slender but muscular anterior portion and a muscular expanded part occupying 35.5-38.5% of total neck length, pharyngeal gland nuclei located as follows: DN = 69.0 ± 1.5 (67-72); S 1 N 1 and S 1 N 2 not observed; S 2 N 1 = 54.8 ± 1.0 (53-56); S 2 N 2 = 56.0 ± 1.9 (53-58). Very delicate membrane surrounding pharyngeal expansion. In some specimens, this structure apparently extending along ventral and/or dorsal side of pharyngeal expansion. Cardia hemispherical, about as broad as long, 12-13 × 11-13 µ m in size. Reproductive system amphididelphic, both branches equally developed. Ovaries reflexed, short, not reaching sphincter level, measuring 39- 63 µ m long, oocytes first in two or more rows, then in one row, distal part of ovary consisting of a well developed blind sac, similar to that described by Coomans (1964) in Discolaimus and Longidorus species. Oviduct joining ovary subterminally, close to proximal end of blind sac. Oviduct consisting of tubular part and well developed pars dilatata. Distinct sphincter visible between oviduct and uterus. Uterus 130-175 µ m or 1.9-2.5 body diam. long, tripartite, consisting of a distal spheroid pars dilatata, a slender intermediate portion and a wider proximal part, these three regions generally showing only weak development and obscured in some individuals by intestinal contents. Vagina extending inwards 51-60% of corresponding body diam. Pars proximalis vaginae 21.5 × 23.5 µ m, pars refringens composed of two trapezoidal, well sclerotised, pieces with a combined width of 16.5 × 4.0 µ m, pars distalis vaginae 6.3 µ m long (holotype). Sperm not observed in female genital tract. Prerectum 4.5-8.5 times and rectum 0.9-1.6 anal body diam. long. Tail short, rounded conoid, small saccate bodies observed along ventral surface, cuticle at terminus forming 42-57% of total tail length, two pairs of caudal pores located at middle of tail, one subdorsal, the other lateral. Male General morphology similar to that of female except for posterior region curved more ventrad. Reproductive system composed of two testes, anterior 179-262 and posterior 167-257 µ m long. Sperms ellipsoid, measuring 8.0-9.5 × 3.5 µ m. Spicules dorylaimoid, 1.3-1.8 anal diam. long and 3.9-4.8 times longer than wide, lateral accessory pieces paired, more or less cylindrical with bifurcate end, measuring 10.5-12.0×2.5 µ m. Series of 10-14 spaced ventromedian supplements starting 24- 32 µ m from adcloacal pair with posteriormost supplement located within range of retracted spicules, adcloacal pair located at 7.0-11.5 µ m from cloacal opening. Tail short conoid, dorsally convex, slightly ventrally concave with bluntly rounded terminus. Cuticle along ventral surface lacking saccate bodies. * The specific epithet is derived from Babak Fort, a historic location in the East Azarbaijan province of Iran and where the new species was found. M. Pedram et al. Juveniles Body length gradually increasing with stage. Comparison of length of functional odontostyle and replacement odontostyle identified four juvenile stages. Lip region in J1 flat, in J2 offset by a less marked constriction than in J3, J4 and adults. Lip region becoming progressively broader from J1 to J4. Genital primordium at vulval region observed in some J4. Tail in J1 conical, relatively much longer than in subsequent stages (c&lcedil; = 3), tail in J2- J4 similar in shape to that of female although becoming relatively shorter with stage (c&lcedil; = 1.0, 0.6-0.9 and 0.5-0.7 in J2, J3 and J4, respectively). TYPE MATERIAL Holotype female, ten paratype females, ten paratype males and paratype juveniles deposited in the Nematode Collection of the Faculty of Agriculture, University of Tabriz, Tabriz, Iran; two paratype females and two paratype males in separate slides deposited at CABI UK- Centre, Surrey, UK, and USDA Nematode Collection, Beltsville, MD, USA. TYPE HABITAT AND LOCALITY Rhizosphere of grasses from natural grasslands, Kaleibar City, mountains of Babak Fort, north-west Iran. DIAGNOSIS AND RELATIONSHIPS The new species is characterised by its 1.21-1.56 mm body length, lip region offset by marked constriction and 17.0-19.0 µ m broad, odontostyle 40-45 µ m long, odontophore flanged and 37-51 µ m long, neck region 283- 413 µ m long, pharyngeal expansion 100-136 µ m long or 35.5-38.5% of total neck length, amphididelphic female genital system, tripartite uterus, vulva pre-equatorial (V = 44-49), tail short, rounded (16-22 µ m long, c = 61- 93, c&lcedil; = 0.5- 0.7 in female; 19-30 µ m long, c = 49-72, c&lcedil; = 0.5- 0.9 in male), male as frequent as female with 49-61 µ m long spicules and 10-14 spaced ventromedian supplements, the posteriormost of which being within the range of the retracted spicules. Because of the morphology of the tail and the odontostyle length, E. babakicus n. sp. can be classified un- der the E. macrodorus -group as defined by Guerrero et al. (2008a). This group is characterised by the presence of a rather long odontostyle (> 35 µ m), odontophore with well developed flanges, tripartite uterus and rounded tail, and comprises the species described above together with E. distinctus Ahmad & Jairajpuri, 1980, E. groenlandicus (Ditlevsen, 1927) Thorne, 1939, E. macrodorus (de Man, 1880) Thorne, 1939, E. microdoroides Baqri & Jairajpuri, 1974 and E. saxifragae Popovici, 1995. In having a lip region offset by a strong constriction, the new species can be distinguished from E. groenlandicus, E. macrodorus and E. microdoroides. Furthermore, it can be separated from E. groenlandicus by its shorter female body (1.21-1.56 vs 1.57-2.50 mm), shorter odontostyle, odontophore and total stylet (40-45, 37-51 and 79-95 µ m, respectively, vs 44-53, 45-55 and 94-108 µ m), shorter and less developed uterus and frequent presence of males vs absence; from E. macrodorus, probably the closest relative, apart from the deep constriction at the lip region, the new species differs by its slightly more posteriorly located vulva (V = 44-49 vs 37-47), slightly longer uterus (130-175 vs 56-143 µ m or 1.9-2.5 vs 0.9-2.0 body diam.), female tail more conoid (vs almost hemispherical) and greater number of ventromedian supplements (10-14 vs 6-12); and from E. microdoroides by its slightly longer female body (1.21-1.56 vs 0.9-1.3 mm), broader lip region (17.0-19.0 vs 13.0-14.0 µ m), guiding ring located more anterior (26-30 vs 37-39 µ m from anterior end) and males with longer spicules (49-61 vs 45-50 µ m). The new species resembles the remaining two species, namely E. distinctus and E. saxifragae, in having the lip region offset by a deep constriction. However, it can be distinguished from E. distinctus by its shorter female body (1.21-1.56 vs 1.85 mm), longer odontostyle (40-45 vs 36 µ m), pre-equatorial vulva (V = 44-49 vs 53) and males frequent vs absent. Finally, the new species differs from E. saxifragae by the much shorter female body length (1.21-1.56 vs 1.80-2.38 mm), female genital system with less developed uterus, differently shaped pars refringens vaginae (trapezoidal vs arcuate drop-shaped), shorter tail (16-22 vs 22-40 µ m) and males with slightly shorter spicules (49-61 vs 56-70 µ m) and fewer ventromedian supplements (10-14 vs 13-16). In the key provided by Guerrero et al. (2008), E. babakicus n. sp. can either be keyed out together with E. microdoroides in step 1, due to the small size of some individuals, or with E. saxifragae in step 3. However, it can be easily distinguished from both by the characteristics listed above., Published as part of Pedram, Majid, Niknam, Gholamreza, Guerrero, Pablo, Ye, Weimin & Robbins, Robert T., 2009, Morphological and molecular characterisation of Enchodelus babakicus n. sp. and E. macrodorus Thorne, 1939 (Nematoda: Nordiidae) from Iran, pp. 895-907 in Nematology 11 (6) on pages 896-902, DOI: 10.1163/156854109X430563, http://zenodo.org/record/8114788