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Tylenchorhynchus clarus Allen, 1955 (Figure 5, Table 4), Published as part of Azizi, Kourosh, Eskandari, Ali, Karegar, Akbar, Ghaderi, Reza, Elsen, Sven Van Den, Holterman, Martijn & Helder, Johannes, 2022, Analyses of morphological and molecular characteristics of Telotylenchinae from Iran point at the validity of the genera Bitylenchus and Sauertylenchus, pp. 425-446 in Zootaxa 5169 (5) on page 435, DOI: 10.11646/zootaxa.5169.5.2, http://zenodo.org/record/6952625, {"references":["Allen, M. W. (1955) A review of the nematode genus Tylenchorhynchus. University of California Publications in Zoology, 61, 129 - 165."]}

Bitylenchus parvus (Allen, 1955) Jairajpuri, 1982 Syn: Tylenchorhynchus parvus Allen, 1955 Bitylenchus parvus (Allen, 1955) Siddiqi, 1986 (Figure 2, Table 3) Description Female Body straight to curved ventrally after heat fixation. Cuticle annuli 0.9–1 μm wide at mid-body. Lateral field consisting of four longitudinal incisures, often areolated. Lip region continuous to slightly offset, 7–7.7 μm wide at base, 4.0–4.1 μm high, and bearing six or seven transverse annuli. Cephalic framework slightly sclerotized, outer margins extending two or three body annuli posteriorly. Stylet strong, basal knobs laterally directed; the conus almost half of the total stylet length. Dorsal pharyngeal gland orifice located 2.2–2.3 μm behind stylet knobs. Median pharyngeal bulb oval, 10–11 μm wide; basal bulb pyriform. Hemizonid 2–4 annuli anterior to secretoryexcretory pore. Reproductive system didelphic, amphidelphic; spermatheca rounded, with globular sperm. Vagina 12–13 μm long, almost 50–60% of the vulval body width. Intestinal fasciculi present in the intestine and inside the post rectal sac which extends into almost the entire tail cavity. Tail cylindrical, finely striated or sometimes smooth terminus, bearing 60–67 annuli on ventral side. Hyaline region of tail terminus 5–6 μm thick. Phasmids located 25–30 μm posterior to anus, located in the posterior half of the tail. Male Not found. REMARKS Compared to the original description of B. parvus Allen (1955), the Iranian population has a lower c' ratio (11.5–12.1 vs 13–16). No other difference in morphological or morphometric diagnostic characters was found. Although this population shows similarity to Sauertylenchus maximus, it can be distinguished from it by a smaller body and stylet (680–800 vs 940–1062, and 17.7–19.6 vs 20–24 μm), and the number of tail annuli (60–67 vs 25– 45). This species was recovered from the rhizosphere of alfalfa (Medicago sativa) in Zanjan and Qazvin provinces during the present study., Published as part of Azizi, Kourosh, Eskandari, Ali, Karegar, Akbar, Ghaderi, Reza, Elsen, Sven Van Den, Holterman, Martijn & Helder, Johannes, 2022, Analyses of morphological and molecular characteristics of Telotylenchinae from Iran point at the validity of the genera Bitylenchus and Sauertylenchus, pp. 425-446 in Zootaxa 5169 (5) on pages 430-432, DOI: 10.11646/zootaxa.5169.5.2, http://zenodo.org/record/6952625, {"references":["Allen, M. W. (1955) A review of the nematode genus Tylenchorhynchus. University of California Publications in Zoology, 61, 129 - 165.","Jairajpuri, M. S. (1982) Some studies on Tylenchorhynchinae: The subgenus Bitylenchus Filipjev, 1934, with description of Tylenchorhynchus (Bitylenchus) depressus n. sp. and a key to species of Bitylenchus. Mededelingen van de Fakulteit Landbouwwetenschappen Rijksuniversiteit Gent, 47, 765 - 770.","Siddiqi, M. R. (1986) Tylenchida Parasites of Plants and Insects. Commonwealth Institute of Parasitology, Slough, UK, 645 pp."]}

Sauertylenchus maximus (Allen, 1955) Siddiqi, 2000 Syn: Bitylenchus maximus (Allen, 1955) Siddiqi, 1986 Tylenchorhynchus maximus Allen, 1950 Tylenchorhynchus wilskii Kornobis, 1980 (Brzeski, 1998) Bitylenchus wilskii (Kornobis, 1980) Jairajpuri, 1982 Bitylenchus wilskii (Kornobis, 1980) Siddiqi, 1986 (Figure 4, Table 3) Description Female Body curved ventrally to C-shaped after heat fixation. Cuticle with distinct annuli, 1.5–2.3 μm wide at midbody. Lateral field with four longitudinal incisures, outer bands areolated. Lip region offset, hemispherical, 8.0– 9.0 μm wide at base, 4.0–5.0 μm high, and bearing six or seven distinct transverse annuli. Cephalic framework lightly sclerotized, outer margins extending over two or three body annuli. Stylet moderately strong, knobs rounded, slightly directed posteriorly; the conus almost half of the total stylet length. Dorsal pharyngeal gland orifice located 2.0–2.1 μm behind stylet knobs. Median pharyngeal bulb oval, 12–13 μm wide; basal bulb pyriform, abutting anterior intestine. Reproductive system didelphic, amphidelphic, spermatheca elongated, empty. Vagina 10 μm long, 40–60% of the vulval body diameter. Intestinal fasciculi present throughout intestine and inside post rectal sac which enters into about two-thirds of the tail length. Tail cylindrical with a hemispherical, annulated terminus, bearing 33–40 annuli on ventral side. Hyaline portion thickness 6–7 μm. Phasmids located 16–17 μm posterior to anus, in the anterior half of the tail. Male Not found. Remarks Compared to the original description of S. maximus by Allen (1955), the Iranian populations differ by the absence of males. The Iranian populations of S. maximus were recovered from Semnan, Ardabil and West Azerbaijan provinces in the rhizosphere of peach (Prunus persica), grass, hawthorn (Crataegus aronia) and apple (Malus pumila), respectively., Published as part of Azizi, Kourosh, Eskandari, Ali, Karegar, Akbar, Ghaderi, Reza, Elsen, Sven Van Den, Holterman, Martijn & Helder, Johannes, 2022, Analyses of morphological and molecular characteristics of Telotylenchinae from Iran point at the validity of the genera Bitylenchus and Sauertylenchus, pp. 425-446 in Zootaxa 5169 (5) on pages 434-435, DOI: 10.11646/zootaxa.5169.5.2, http://zenodo.org/record/6952625, {"references":["Allen, M. W. (1955) A review of the nematode genus Tylenchorhynchus. University of California Publications in Zoology, 61, 129 - 165.","Siddiqi, M. R. (2000) Tylenchida Parasites of Plants and Insects. 2 nd edition. CABI Publishing, Wallingford, Oxon, UK, 833 p. https: // doi. org / 10.1079 / 9780851992020.0000","Siddiqi, M. R. (1986) Tylenchida Parasites of Plants and Insects. Commonwealth Institute of Parasitology, Slough, UK, 645 pp.","Brzeski, M. W & Dolinski C. M. (1998) Compendium of the genus Tylenchorhynchus Cob, 1913 sensu lato (Nematodea: Belonolaimidae). Russian Journal of Nematology, 6 (2), 189 - 199.","Jairajpuri, M. S. (1982) Some studies on Tylenchorhynchinae: The subgenus Bitylenchus Filipjev, 1934, with description of Tylenchorhynchus (Bitylenchus) depressus n. sp. and a key to species of Bitylenchus. Mededelingen van de Fakulteit Landbouwwetenschappen Rijksuniversiteit Gent, 47, 765 - 770."]}

Bitylenchus dubius (Bütschli, 1873) Filipjev, 1934 Syn: Tylenchus dubius Bütschli, 1873 Anguillulina dubia (Bütschli, 1873) Goody, 1932 Tylenchus (Bitylenchus) dubius (Bütschli, 1873) Filipjev, 1934 Tylenchorhynchus dubius (Bütschli, 1873) Filipjev, 1936 Bitylenchus dubius (Bütschli, 1873) Siddiqi, 1986 (Figure 1, Table 3), Published as part of Azizi, Kourosh, Eskandari, Ali, Karegar, Akbar, Ghaderi, Reza, Elsen, Sven Van Den, Holterman, Martijn & Helder, Johannes, 2022, Analyses of morphological and molecular characteristics of Telotylenchinae from Iran point at the validity of the genera Bitylenchus and Sauertylenchus, pp. 425-446 in Zootaxa 5169 (5) on page 428, DOI: 10.11646/zootaxa.5169.5.2, http://zenodo.org/record/6952625, {"references":["Butschli, O. (1873) Beitrage zur Kenntnis der freilebenden Nematoden. Nova acta Academiae Caesareae Leopoldino-Carolinae Germanicae Naturae Curiosorum, 36, 1 - 124.","Filipjev, I. N. (1934) Harmful and Useful Nematodes in Rural Economy. Moscow, Leningrad, USSR, 440 pp. [In Russian]","Filipjev, I. N. (1936) On the classification of the Tylenchinae. Proceedings of the Helminthological Society of Washington, 3, 80 - 82.","Siddiqi, M. R. (1986) Tylenchida Parasites of Plants and Insects. Commonwealth Institute of Parasitology, Slough, UK, 645 pp."]}

Tylenchorhynchus microconus Siddiqi, Mukherjee and Dasgupta, 1982 (Figurs 6, Table 4), Published as part of Azizi, Kourosh, Eskandari, Ali, Karegar, Akbar, Ghaderi, Reza, Elsen, Sven Van Den, Holterman, Martijn & Helder, Johannes, 2022, Analyses of morphological and molecular characteristics of Telotylenchinae from Iran point at the validity of the genera Bitylenchus and Sauertylenchus, pp. 425-446 in Zootaxa 5169 (5) on page 439, DOI: 10.11646/zootaxa.5169.5.2, http://zenodo.org/record/6952625, {"references":["Siddiqi, M. R., Mukherjee, B. & Dasgupta, M. K. (1982) Tylenchorhynchus microconus n. sp., T. crassicaudatus leviterminalis n. subsp. and T. coffeae Siddiqi and Basir, 1959 (Nematoda: Tylenchida). Systematic Parasitology, 4, 257 - 262. https: // doi. org / 10.1007 / BF 00009626"]}

Bitylenchus serranus Gomez-Barcina, Siddiqi & Castillo, 1992 (Figure 3, Table 3), Published as part of Azizi, Kourosh, Eskandari, Ali, Karegar, Akbar, Ghaderi, Reza, Elsen, Sven Van Den, Holterman, Martijn & Helder, Johannes, 2022, Analyses of morphological and molecular characteristics of Telotylenchinae from Iran point at the validity of the genera Bitylenchus and Sauertylenchus, pp. 425-446 in Zootaxa 5169 (5) on page 433, DOI: 10.11646/zootaxa.5169.5.2, http://zenodo.org/record/6952625, {"references":["Gomez-Barcina, A., Siddiqi, M. R. & Castillo, P. (1992) The genus Bitylenchus Filipjev, 1934 (Nematoda: Tylenchida) with descriptions of two new species from Spain. Journal of the Helminthological Society of Washington, 59, 96 - 110."]}

Trophurus ussuriensis Eroshenko, 1981 (Figure 7, Table 4), Published as part of Azizi, Kourosh, Eskandari, Ali, Karegar, Akbar, Ghaderi, Reza, Elsen, Sven Van Den, Holterman, Martijn & Helder, Johannes, 2022, Analyses of morphological and molecular characteristics of Telotylenchinae from Iran point at the validity of the genera Bitylenchus and Sauertylenchus, pp. 425-446 in Zootaxa 5169 (5) on page 440, DOI: 10.11646/zootaxa.5169.5.2, http://zenodo.org/record/6952625, {"references":["Eroshenko, A. S. (1981) Plant parasitic nematodes of underwood; families Tylenchorhynchidae and Hoplolaimidae (Nematoda). In: Svobodnozhivuschie I fitopatogennye nematody fauny Dal'nego Vostoka Biol. Pedolog. Inst. of Far Eastern Research Center of USSR Academic Science Vladivostok, pp. 22 - 27 + 85 - 92."]}

Diphtherophora tenera Ivanova, 1980 (Fig. 3 D, H, K, N, Table 2) In our study, this population has a smaller body and spear than the type population recovered from Tadjikistan (Ivanova 1980). On the other hand, morphological characters displayed comparable values, in agreement with another population also reported from Iran (Ghaderi et al. 2017). This population was recovered from turf grass in the campus of Shiraz University, at Eram Square, Shiraz, Fars province, Southern Iran., Published as part of Ghaderi, Reza, Hesar, Abas Mokaram, Karegar, Akbar & Pereira, Tiago Jos��, 2020, Molecular phylogeny of Diphtherophora de Man, 1880 (Nematoda: Diphtherophoridae), with description of a new species, pp. 371-385 in Zootaxa 4851 (2) on page 377, DOI: 10.11646/zootaxa.4851.2.10, http://zenodo.org/record/4476997, {"references":["Ivanova, T. S. (1980) [Two new species of nematodes from the family Diphtherophoridae (Micoletzky, 1922) Clark, 1961]. Izvestiya Akademii Nauk Tadzhikskoi SSR, Biologicheskii Nauki, 4, 94 - 97. [In Russian]","Ghaderi, R., Kashi, L., Mirbabaei Karani, H. & Karegar, A. (2017) A new and four known species of Diphtherophora (Nematoda: Diphtherophoridae) from Iran, with a diagnostic compendium of its species. Zootaxa, 4365 (3), 311 - 330. https: // doi. org / 10.11646 / zootaxa. 4365.3.3"]}

Diphtherophora eldarica n. sp. (Figs. 1, 2, Table 1) Description. Female: Body habitus straight to slightly curved ventrally after fixation. Cuticle regularly striated from the anterior end to the level of secretory-excretory pore, but appearing smooth posteriorly; relatively thick, about one-fifth to two-fifths of the body width. Subcuticle layer finely striated along the entire body and about 2 µm thick. Head region offset from the body contour. Outer labial papillae not readily observed. Amphid aperture oval in shape and as wide as about one-third of the head width at its base. Spear typical of the genus, with a sclerotized acute tip. Spear guiding apparatus arched, moderately sclerotized. Pharynx typical of the genus and composed of two parts, the anterior region (corpus) slender and expanding gradually to end as a rounded or sometimes slightly pyriform terminal bulb. Nerve ring located at mid-pharynx region or posterior to it. Secretory-excretory pore opposite the posterior end of the slender part of pharynx or at the beginning of the pharyngeal bulb. Cardia not observed under LM. Reproductive system didelphic, amphidelphic, with branches equally developed, ovaries reflexed with germinal zones directed toward vagina, spermatheca not present at the junction of oviduct and uterus. Sperm cells with rodshaped nuclei throughout the uteri. Vulva a transverse slit as seen in a ventral view. Cuticle invaginated just anterior and posterior to the vulva. Vagina orthogonal to the body axis, with moderately to well-developed musculature. Rectum 14–16 µm or 54–73 % of the anal body width. Cuticle overhanging the anterior lip of the anus. Tail conoid, bent dorsally near tail tip, with a rounded terminus. The tail terminus lacks lateral caudal pores. Male: In general, morphologically similar to female. Body habitus almost straight. Cervical ventromedian papillae absent. No lateral body pores observed in the anterior region of the body. Reproductive system monorchic, extending to near mid-body, and containing numerous rod-shaped sperm. Two ventromedian precloacal supplements were observed, the anteriormost near the proximal end of spicules and the posterior one level with the distal ends of spicules. Spicules with marked capitulum and narrow shaft, expanding again at beginning of the blade part and then gradually tapering to the distal end. Spicular capsule weakly developed. Gubernaculum ventrally arcuate at anterior tip. Tail similar to female, with a rounded terminus. Overhanging cuticle on cloacal lips was not observed, as the cuticle did not separate from the body over a short distance anterior or posterior to cloaca. Diagnosis. The new species is characterized by females with striated cuticle at the anterior end of the body, head offset from the body contour, spear 15–17 µm in length, rod-shaped sperm, overhanging cuticle on the anterior lip of the anus and a conical tail bent dorsally near the terminus. Additionally, males lack ventromedian neck papillae but have two ventromedian precloacal supplements at the level of the spicules. Type host and locality. The specimens were collected from the rhizosphere of pine trees (Pinus eldarica Medw.) in Dalampar valley in Margavar, Urmia city, West Azerbaijan province, Northwestern Iran by A. Mokaram Hesar in September 2015 (GPS coordinates: 37º10’65’’N, 44º52’73’’E). Type material. Holotype, three female paratypes and one male paratype, mounted in pure glycerin on a slide, were deposited in the collection of the Department of Plant Protection, School of Agriculture, Shiraz University, Shiraz, Iran. Etymology. The specific epithet refers to the associated plant species, Pinus eldarica, from the rhizosphere of which the new Diphtherophora species was recovered.

Diphtherophora species phylogeny In total, 26 DNA sequences, specifically covering the D2���D3 domains of the 28S rRNA gene, were used in our molecular phylogenetic analyses: 18 Diphtherophora, two Tylolaimophorus, one representative of Prismatolaimidae, six representatives of Trichodoridae, two representatives of Tripylidae, and three representatives of Trischistomatidae. Our final alignment was 785 bp long and contained 326 conserved sites, 455 variable sites, 415 parsimonyinformative sites, and 40 singletons. Phylogenetic analyses based on both inference methods (i.e., ML and BI) were highly congruent with respect to the monophyly of the genus Diphtherophora, the relationships among Diphtherophora species and outgroups, and the phylogenetic placement of D. eldarica n. sp. (Fig. 4). Phylogenetic trees showed the existence of six strongly supported clades (A���F) within the Diphtherophora group: clade A is represented by D. cf. caudata from Arasbaran (Northwestern Iran) and D. caudata from Maragheh (Northwestern Iran), the latter species recovered in this study; clade B included sequences representing D. cf. obesa and D. tenera from different regions of Northern and Southern Iran, the latter species, however, being paraphyletic; clade C was comprised of D. perplexans from different sites in Northwestern (Takab and Urmia) and Southern Iran (Shiraz), the latter population recovered in this study, and a specimen characterized only as Diphtherophora sp. from CA, USA; clade D was solely represented by the new species D. eldarica n. sp.; clade E harbored uncharacterized Diphtherophora species from different sites in Northern and Northwestern Iran including Arasbaran and Ali Abad; clade F is solely represented by sequences of D. geraerti from Shiraz (Southern Iran). Our phylogenetic analyses strongly supported (ML = 97, BI = 1.0) the new species D. eldarica n. sp. as a sister taxon of clade D, that is, D. perplexans + Diphtherophora sp. (Fig.4). Genetic divergence (p-distance %) within Diphtherophora clades was relatively low, ranging from 0% (clade A, among D. caudata sequences) to 5.2% (variation within clade E, represented by Diphtherophora sp. sequences from Iran; Table 3). On the other hand, genetic divergence between Diphtherophora clades was much higher, ranging from 8.2% (clade A vs. B) to as high as 23.7% (clade C vs. F). For comparison, genetic divergence between any of the Diphtherophora clades and Tylolaimophorus (also considered a representative of Diphtherophoridae) ranged from 25.6% (Tylolaimophorus vs. clade A) to 32.7% (Tylolaimophorus vs. clade F). Similarly, genetic divergence between Diphtherophora clades and trichodorid genera were in the same range (24.1% to 30.1%) as those observed for comparisons with Tylolaimophorus. This pattern is also consistent when comparing the genetic divergence between Diphtherophora clades and other outgroup taxa including genera Prismatolaimus de Man, 1880, Tripyla Bastian, 1865, and Tripylina Brzeski, 1963 (Table 3). 1 Taxon abbreviations: Tylolaimophorus (Tylo.), Trichodorus (Tric.), Paratrichodorus (Para.), Monotrichodorus (Mono.), Prismatolaimus (Prism.), Tripylina (Tripyli.). 2 Taxon represented by only one DNA sequence. With respect to the new species D. eldarica n. sp., genetic divergence ranged from 13.5% (between clade B, D. tenera + D. cf. obesa) to 19.5% (between clade E, Diphtherophora sp.). Although all six clades were strongly supported in our analyses, with ML branch support always greater than 98% and BPP always with maximum support, some relationships among Diphtherophora clades (i.e. some of the deeper nodes) were either weakly supported or not recovered by one of the inference methods (Fig. 4)., Published as part of Ghaderi, Reza, Hesar, Abas Mokaram, Karegar, Akbar & Pereira, Tiago Jos��, 2020, Molecular phylogeny of Diphtherophora de Man, 1880 (Nematoda: Diphtherophoridae), with description of a new species, pp. 371-385 in Zootaxa 4851 (2) on pages 379-382, DOI: 10.11646/zootaxa.4851.2.10, http://zenodo.org/record/4476997, {"references":["de Man, J. G. (1880) Die Einheimischen, frei in der reinen Erde und im sussen Wasser lebende Nematoden [Free-living soil and freshwater nematodes]. Monographisch Bearbeitet. Tijdschrift der Nederlandsche dierk undige Vereniging, 5, 1 - 104. [in German]","Bastian, H. C. (1865) Monograph on the Anguillulidae. Transactions of the Linnean Society of London, 25, 73 - 184. https: // doi. org / 10.1111 / j. 1096 - 3642.1865. tb 00179. x","Brzeski, M. W. (1963) Nematode genera of the family Tripylidae (Nematoda, Enoplida). Acta Zoologica Cracoviensia, 8, 295 - 308."]}

Diphtherophora eldarica n. sp. (Figs. 1, 2, Table 1) Description. Female: Body habitus straight to slightly curved ventrally after fixation. Cuticle regularly striated from the anterior end to the level of secretory-excretory pore, but appearing smooth posteriorly; relatively thick, about one-fifth to two-fifths of the body width. Subcuticle layer finely striated along the entire body and about 2 ��m thick. Head region offset from the body contour. Outer labial papillae not readily observed. Amphid aperture oval in shape and as wide as about one-third of the head width at its base. Spear typical of the genus, with a sclerotized acute tip. Spear guiding apparatus arched, moderately sclerotized. Pharynx typical of the genus and composed of two parts, the anterior region (corpus) slender and expanding gradually to end as a rounded or sometimes slightly pyriform terminal bulb. Nerve ring located at mid-pharynx region or posterior to it. Secretory-excretory pore opposite the posterior end of the slender part of pharynx or at the beginning of the pharyngeal bulb. Cardia not observed under LM. Reproductive system didelphic, amphidelphic, with branches equally developed, ovaries reflexed with germinal zones directed toward vagina, spermatheca not present at the junction of oviduct and uterus. Sperm cells with rodshaped nuclei throughout the uteri. Vulva a transverse slit as seen in a ventral view. Cuticle invaginated just anterior and posterior to the vulva. Vagina orthogonal to the body axis, with moderately to well-developed musculature. Rectum 14���16 ��m or 54���73 % of the anal body width. Cuticle overhanging the anterior lip of the anus. Tail conoid, bent dorsally near tail tip, with a rounded terminus. The tail terminus lacks lateral caudal pores. Male: In general, morphologically similar to female. Body habitus almost straight. Cervical ventromedian papillae absent. No lateral body pores observed in the anterior region of the body. Reproductive system monorchic, extending to near mid-body, and containing numerous rod-shaped sperm. Two ventromedian precloacal supplements were observed, the anteriormost near the proximal end of spicules and the posterior one level with the distal ends of spicules. Spicules with marked capitulum and narrow shaft, expanding again at beginning of the blade part and then gradually tapering to the distal end. Spicular capsule weakly developed. Gubernaculum ventrally arcuate at anterior tip. Tail similar to female, with a rounded terminus. Overhanging cuticle on cloacal lips was not observed, as the cuticle did not separate from the body over a short distance anterior or posterior to cloaca. Diagnosis. The new species is characterized by females with striated cuticle at the anterior end of the body, head offset from the body contour, spear 15���17 ��m in length, rod-shaped sperm, overhanging cuticle on the anterior lip of the anus and a conical tail bent dorsally near the terminus. Additionally, males lack ventromedian neck papillae but have two ventromedian precloacal supplements at the level of the spicules. Type host and locality. The specimens were collected from the rhizosphere of pine trees (Pinus eldarica Medw.) in Dalampar valley in Margavar, Urmia city, West Azerbaijan province, Northwestern Iran by A. Mokaram Hesar in September 2015 (GPS coordinates: 37��10���65������N, 44��52���73������E). Type material. Holotype, three female paratypes and one male paratype, mounted in pure glycerin on a slide, were deposited in the collection of the Department of Plant Protection, School of Agriculture, Shiraz University, Shiraz, Iran. Etymology. The specific epithet refers to the associated plant species, Pinus eldarica, from the rhizosphere of which the new Diphtherophora species was recovered., Published as part of Ghaderi, Reza, Hesar, Abas Mokaram, Karegar, Akbar & Pereira, Tiago Jos��, 2020, Molecular phylogeny of Diphtherophora de Man, 1880 (Nematoda: Diphtherophoridae), with description of a new species, pp. 371-385 in Zootaxa 4851 (2) on pages 373-377, DOI: 10.11646/zootaxa.4851.2.10, http://zenodo.org/record/4476997

Diphtherophora species phylogeny In total, 26 DNA sequences, specifically covering the D2–D3 domains of the 28S rRNA gene, were used in our molecular phylogenetic analyses: 18 Diphtherophora, two Tylolaimophorus, one representative of Prismatolaimidae, six representatives of Trichodoridae, two representatives of Tripylidae, and three representatives of Trischistomatidae. Our final alignment was 785 bp long and contained 326 conserved sites, 455 variable sites, 415 parsimonyinformative sites, and 40 singletons. Phylogenetic analyses based on both inference methods (i.e., ML and BI) were highly congruent with respect to the monophyly of the genus Diphtherophora, the relationships among Diphtherophora species and outgroups, and the phylogenetic placement of D. eldarica n. sp. (Fig. 4). Phylogenetic trees showed the existence of six strongly supported clades (A–F) within the Diphtherophora group: clade A is represented by D. cf. caudata from Arasbaran (Northwestern Iran) and D. caudata from Maragheh (Northwestern Iran), the latter species recovered in this study; clade B included sequences representing D. cf. obesa and D. tenera from different regions of Northern and Southern Iran, the latter species, however, being paraphyletic; clade C was comprised of D. perplexans from different sites in Northwestern (Takab and Urmia) and Southern Iran (Shiraz), the latter population recovered in this study, and a specimen characterized only as Diphtherophora sp. from CA, USA; clade D was solely represented by the new species D. eldarica n. sp.; clade E harbored uncharacterized Diphtherophora species from different sites in Northern and Northwestern Iran including Arasbaran and Ali Abad; clade F is solely represented by sequences of D. geraerti from Shiraz (Southern Iran). Our phylogenetic analyses strongly supported (ML = 97, BI = 1.0) the new species D. eldarica n. sp. as a sister taxon of clade D, that is, D. perplexans + Diphtherophora sp. (Fig.4). Genetic divergence (p-distance %) within Diphtherophora clades was relatively low, ranging from 0% (clade A, among D. caudata sequences) to 5.2% (variation within clade E, represented by Diphtherophora sp. sequences from Iran; Table 3). On the other hand, genetic divergence between Diphtherophora clades was much higher, ranging from 8.2% (clade A vs. B) to as high as 23.7% (clade C vs. F). For comparison, genetic divergence between any of the Diphtherophora clades and Tylolaimophorus (also considered a representative of Diphtherophoridae) ranged from 25.6% (Tylolaimophorus vs. clade A) to 32.7% (Tylolaimophorus vs. clade F). Similarly, genetic divergence between Diphtherophora clades and trichodorid genera were in the same range (24.1% to 30.1%) as those observed for comparisons with Tylolaimophorus. This pattern is also consistent when comparing the genetic divergence between Diphtherophora clades and other outgroup taxa including genera Prismatolaimus de Man, 1880, Tripyla Bastian, 1865, and Tripylina Brzeski, 1963 (Table 3). 1 Taxon abbreviations: Tylolaimophorus (Tylo.), Trichodorus (Tric.), Paratrichodorus (Para.), Monotrichodorus (Mono.), Prismatolaimus (Prism.), Tripylina (Tripyli.). 2 Taxon represented by only one DNA sequence. With respect to the new species D. eldarica n. sp., genetic divergence ranged from 13.5% (between clade B, D. tenera + D. cf. obesa) to 19.5% (between clade E, Diphtherophora sp.). Although all six clades were strongly supported in our analyses, with ML branch support always greater than 98% and BPP always with maximum support, some relationships among Diphtherophora clades (i.e. some of the deeper nodes) were either weakly supported or not recovered by one of the inference methods (Fig. 4).

Diphtherophora geraerti Ghaderi, Kashi, Mirbabaei Karani & Karegar, 2017 (Fig. 3 B, F, J, M, Table 2) The population recovered in this study matches the morphology of the type population (Ghaderi et al. 2017). This population was found from turf grass in the campus of Shiraz University, at Eram Square, Shiraz, Fars province, Southern Iran., Published as part of Ghaderi, Reza, Hesar, Abas Mokaram, Karegar, Akbar & Pereira, Tiago Jos��, 2020, Molecular phylogeny of Diphtherophora de Man, 1880 (Nematoda: Diphtherophoridae), with description of a new species, pp. 371-385 in Zootaxa 4851 (2) on page 377, DOI: 10.11646/zootaxa.4851.2.10, http://zenodo.org/record/4476997, {"references":["Ghaderi, R., Kashi, L., Mirbabaei Karani, H. & Karegar, A. (2017) A new and four known species of Diphtherophora (Nematoda: Diphtherophoridae) from Iran, with a diagnostic compendium of its species. Zootaxa, 4365 (3), 311 - 330. https: // doi. org / 10.11646 / zootaxa. 4365.3.3"]}

The poorly-known nematodes of the genus Diphtherophora are soil inhabitants often found in the vicinity of plant roots. In the present study, we characterize a new species belonging to the genus Diphtherophora, named D. eldarica n. sp., from Iran using both morphological and molecular approaches. The new species is characterized by striated cuticle at the anterior end of the body, head offset from the body contour, spear 15–17 µm in length, rod-shaped sperm, overhanging cuticle on the anterior lip of the anus, and a conical tail bent dorsally near the terminus. Diphterophora eldarica n. sp. also lacks ventromedian neck papillae whereas male specimens bear two precloacal ventromedian supplements at the level of the spicules. Additionally, we provide morphological and molecular data for four known Diphtherophora species including D. geraerti, D. caudata, D. perplexans, and D. tenera collected from soils of different plants and localities in Iran. Using the D2–D3 expansion segments of the 28S ribosomal (rRNA) gene including D. eldarica n. sp. and additional known species from Iran, we explore for the first time species relationships in the genus Diphtherophora within a molecular phylogenetic framework. Our results support: 1) the monophyly of the genus Diphtherophora with respect to the outgroup taxa (Tylolaimophorus and representatives of Trichodoridae), 2) the existence of six strongly supported clades within Diphtherophora, and 3) a sister relationship between D. eldarica n. sp. and a clade formed by D. perplexans and Diphtherophora sp. from the USA. Finally, this study emphasizes the diversity of the genus Diphtherophora in Iran.

Diphtherophora caudata Ivanova, 1958 (Fig. 3 A, E, I, L, Table 1) The population sampled in the present study comes close to other populations representing the species (Ivanova 1958, Ghaderi et al. 2017), except in its slightly smaller body when compared to those two studies (558���633 vs. 650 and 650���720 ��m) and shorter spear than the type population (15���17 vs. 20���21 ��m). The present population was recovered from the rhizosphere of cherry (Cerasus avium (L.) Moench) trees in Maragheh, East Azarbaijan, Northwestern Iran., Published as part of Ghaderi, Reza, Hesar, Abas Mokaram, Karegar, Akbar & Pereira, Tiago Jos��, 2020, Molecular phylogeny of Diphtherophora de Man, 1880 (Nematoda: Diphtherophoridae), with description of a new species, pp. 371-385 in Zootaxa 4851 (2) on page 377, DOI: 10.11646/zootaxa.4851.2.10, http://zenodo.org/record/4476997, {"references":["Ivanova, T. S. (1958) [Soil nematodes of the genus Diphtherophora of the fauna of U. S. S. R.]. Theses of report 8 - 12 December, 1958, All Union belminth Society of Moscow, 1958, 55 - 56. [in Russian]","Ghaderi, R., Kashi, L., Mirbabaei Karani, H. & Karegar, A. (2017) A new and four known species of Diphtherophora (Nematoda: Diphtherophoridae) from Iran, with a diagnostic compendium of its species. Zootaxa, 4365 (3), 311 - 330. https: // doi. org / 10.11646 / zootaxa. 4365.3.3"]}

Diphtherophora perplexans (Van Der Linde, 1938) Goodey, 1951 (Fig. 3 C, G, Table 2) Morphologically, our populations from Iran fit well with the original population described from the USA (Cobb 1913). However, some morphometric characters also had larger ranges. Moreover, fine striations on the cuticle and large plate-like thickenings on the ventral or dorsal sides of the spear guiding apparatus were observed in some individuals, indicating close proximity of P. perplexans with P. pseudoperplexans (van der Linde, 1938) Goodey, 1951. In this study, four populations representing D. perplexans were found: one population was recovered from the rhizosphere of plum (Prunus domestica L.) trees in Mianeh, East Azerbaijan, Northwestern Iran; another population was recovered from the rhizosphere of apricot (Prunus armeniaca L.) trees in Khalkhal, Ardebil province, Northwestern Iran; two additional populations were recovered from turf grass in the campus of Shiraz University, at Eram Square, Shiraz, Fars province, Southern Iran., Published as part of Ghaderi, Reza, Hesar, Abas Mokaram, Karegar, Akbar & Pereira, Tiago Jos��, 2020, Molecular phylogeny of Diphtherophora de Man, 1880 (Nematoda: Diphtherophoridae), with description of a new species, pp. 371-385 in Zootaxa 4851 (2) on page 377, DOI: 10.11646/zootaxa.4851.2.10, http://zenodo.org/record/4476997, {"references":["Van der Linde, W. J. (1938) A contribution to the study of nematodes. Entomology Mem of Department of Agriculture and Forestry, Union of South Africa, 2, 1 - 40.","Goodey, T. (1951) Soil and freshwater nematodes. Methuen, London, 390 pp. https: // doi. org / 10.1097 / 00010694 - 195107000 - 00019","Cobb, N. A. (1913) New nematode genera found inhabiting freshwater and non-brackish soils. Journal of the Washington Academy of Sciences, 3, 432 - 444. https: // doi. org / 10.5962 / bhl. part. 20323"]}

2. Nothotylenchus hexaglyphus Khan & Siddiqi, 1968 (Figure 5) 3 females: L = 751 ± 159 (568–851) µm; stylet = 7.3 ± 0.2 (7–7.5) µm; pharynx = 114 ± 11 (103–125) µm; tail = 55.6 ± 5.4 (50.5–61.5) µm; a = 35.7 ± 4.7 (31.9–41.0); b = 6.5 ± 0.9 (5.5–7.4); c = 13.5 ± 2.2 (11.2–15.5); c′ = 4.1 ± 0.4 (3.7–4.5); V = 82.8 ± 1.4 (81.6–84.3); V′ = 89.6 ± 0.4 (89.2–90.1); PUS/VBW = 1.1 ± 0.3 (0.8–1.3); PUS/V-A = 28.4 ± 7.3 (21.9–36.3) %; V-A/T = 1.3 ± 0.2 (1.1–1.5). 2 males: L = 683, 922 µm; stylet = 7, 7.5 µm; pharynx = 103, 157 µm; tail = 57, 64.5 µm; a = 41.9, 56.2; b = 6.6, 5.9; c = 12.0, 14.3; c′ = 4.5, 4.2; spicules = 21, 20.5 µm. Diagnosis and relationships. Nothotylenchus hexaglyphus is characterised by its medium-sized body, six lateral field incisures, delicate stylet with very small knobs, pyriform or slightly elongated and offset or slightly overlapping basal pharyngeal bulb, posterior position of vulva, very short PUS, long spicules, and rounded tail terminus. Studied populations of N. hexaglyphus are similar to N. affinis, N. geraerti, N. medians, N. taylori and N. tuberosus, and can be distinguished from all of these species by the very small stylet knobs (Brzeski 1991). It differs from N. affinis, N. geraerti and N. medians by its longer spicules (20.5–21 vs. 15–17, 14–17.5 and 15–18 µm, respectively), and from N. taylori and N. tuberosus by smaller PUS/VBW ratio (0.8–1.3 vs. 1.8–3.0 and 2.0–2.4, respectively).

Nothotylenchus brzeskii n. sp. (Figures 1 & 2) Measurements: Table 1. Female: Body slightly or completely curved ventrally. Cuticular annulation distinct, 1–1.5 µm wide; lateral fields with four incisures, 5.5–7.5 µm wide and 18.8–33.3% of the body width, two outer ones sometimes crenate. Head wide and low, 1.5–2.5 µm high and 6.5–7.5 µm wide, with rounded edges and two or three fine annuli, not offset. Cephalic skeleton weak, the outer margin of basal plate extending two annuli into body. Stylet delicate, conus 33.3–40.0% of total stylet length, knobs small and rounded, 1.5–2 µm wide. DGO 1–1.5 µm posterior to stylet knobs. Median bulb elongated and fusiform, slightly wider than procorpus, 5–6 µm wide, without thickenings of lumen walls. Isthmus cylindrical and more slender than procorpus. Nerve ring surrounding isthmus near middle or in its posterior half, sometimes at the junction of isthmus and basal pharyngeal bulb, 90 (84–93) µm from anterior end. Hemizonid two to three cuticular annuli long, 112 (109–117) µm from anterior end, one to three annuli anterior to S-E pore. S-E pore opposite the anterior or posterior half of the basal bulb. In one specimen two pairs of hemizonid and S-E pore were observed. The first pair located opposite the anterior half and the second pair near the end of basal pharyngeal bulb. Deirids at the level of S-E pore. Basal pharyngeal bulb pyriform and offset from intestine or with slight overlap, up to 7.5 µm, 2.1 times (1.6–2.8) longer than wide, with the two anterior-most intestinal cells appearing hyaline. Reproductive system characteristic of Nothotylenchus: reproductive tract mono-prodelphic, oocytes in single row, cylindrical, inline spermatheca filled with large rounded sperms or sometimes empty, uterus quadricolumellar, vagina 6.5–9.5 µm in length and 25.6–46.6% of VBW. PUS short, 16.5–24 µm long, 0.9 (0.6–1.2) of VBW or 28.1 (20.2–35.3) % V-A and 1.3 (0.9–1.8) ABW. Post-vulval body length 8.3 (7.6–8.9) ABW. Tail thick, conical and narrowed gradually from anus, ventrally arcuate at the end, with rounded to sometimes dull terminus. Male: General morphology, stylet, pharynx and tail shape similar to females. Spicules slightly or completely ventrally arcuate, gubernaculum simple and crescent-shaped, bursa 27–36.5 µm long and enveloping 51.2–74.1% of tail length. Tail with rounded terminus. Habitat & locality. Type population collected in 2014 from rhizosphere of alfalfa in Darab region, Fars province, Iran (GPS coordinates: 28°41.167′N, 54°16.056′E, elevation 1367 m.). Type material. Female holotype, two female paratypes and two male paratypes on three glass slides kept in the nematode collection of the Department of Plant Protection, School of Agriculture, Shiraz University, Iran, and two female paratypes and two male paratypes on two glass slides deposited in the Nematode Collection of the Plantenziektenkundige Dienst, Wageningen, The Netherlands. Diagnosis and relationships. Nothotylenchus brzeskii n. sp. is characterised by a body length of 774–922 µm, lateral fields with four incisures, delicate, short stylet (7–8 µm) with small rounded knobs, pyriform, offset from intestine or slightly overlapping basal pharyngeal bulb, posterior vulva position (V = 83.4–84.4), short PUS (16.5–24 µm), spicules 20.5–23 µm in length, and thick tail with rounded to dull terminus. N. brzeskii n. sp. resembles N. basiri, N. cylindricollis Thorne, 1941, N. cylindricus Khan & Siddiqi, 1968, N. singhi Das & Shivaswamy, 1980, N. thornei Andrássy, 1958 and N. websteri Kumar, 1983. It differs from N. basiri by longer spicules (20.5–23 vs. 13–15 µm), from N. cylindricollis by different tail terminus (rounded to dull vs. pointed) and different basal pharyngeal bulb (pyriform vs. elongate and cylindroid), from N. cylindricus by the longer spicules (20.5–23 vs. 14 µm), and different basal pharyngeal bulb (pyriform vs. elongate and cylindroid), from N. singhi by longer spicules (20.5–23 vs. 13–15 µm) and lower PUS/VBW ratio (0.6–1.2 vs. ˃ 1.5), from N. thornei by different tail terminus (rounded to dull vs. mucronate) and lower c′ value (3.3–4.0 vs. 7.6–9.0) and from N. websteri by longer spicules (20.5–23 vs. 11–13 µm) and different tail terminus (rounded to dull vs. pointed). Etymology. The species is named in honor of Dr. Michal W. Brzeski for his outstanding contributions to the science of nematology., Published as part of Hashemi, Kobra & Karegar, Akbar, 2020, New and known species of Nothotylenchus Thorne, 1941 (Nematoda: Anguinidae) from Iran with an updated list of species, pp. 482-500 in Zootaxa 4729 (4) on pages 483-487, DOI: 10.11646/zootaxa.4729.4.2, http://zenodo.org/record/3752014, {"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.","Khan, A. M. & Siddiqi, M. R. (1968) Three new species of Nothotylenchus (Nematoda: Neotylenchidae) from north India. Nematologica, 14, 369 - 376. https: // doi. org / 10.1163 / 187529268 X 00048","Das, V. M. & Shivaswamy, V. (1980) Paurodontus brassicae n. sp. and Nothotylenchus singhi n. sp. from south India. Proceedings of the Indian Academy of Parasitology, 1, 62 - 65.","Andrassy, I. (1958) Erd-und Susswassernematoden aus Bulgarien Acta Zoologica Academiae Scientiarum Hungaricae, 4, 1 - 88.","Kumar, P. (1983) Nothotylenchus websteri n. sp. (Nematoda: Neotylenchoidea) from Lucknow. Indian Journal of Parasitology, 7, 105 - 107."]}

2. Nothotylenchus hexaglyphus Khan & Siddiqi, 1968 (Figure 5) 3 females: L = 751 ± 159 (568–851) µm; stylet = 7.3 ± 0.2 (7–7.5) µm; pharynx = 114 ± 11 (103–125) µm; tail = 55.6 ± 5.4 (50.5–61.5) µm; a = 35.7 ± 4.7 (31.9–41.0); b = 6.5 ± 0.9 (5.5–7.4); c = 13.5 ± 2.2 (11.2–15.5); c′ = 4.1 ± 0.4 (3.7–4.5); V = 82.8 ± 1.4 (81.6–84.3); V′ = 89.6 ± 0.4 (89.2–90.1); PUS/VBW = 1.1 ± 0.3 (0.8–1.3); PUS/V-A = 28.4 ± 7.3 (21.9–36.3) %; V-A/T = 1.3 ± 0.2 (1.1–1.5). 2 males: L = 683, 922 µm; stylet = 7, 7.5 µm; pharynx = 103, 157 µm; tail = 57, 64.5 µm; a = 41.9, 56.2; b = 6.6, 5.9; c = 12.0, 14.3; c′ = 4.5, 4.2; spicules = 21, 20.5 µm. Diagnosis and relationships. Nothotylenchus hexaglyphus is characterised by its medium-sized body, six lateral field incisures, delicate stylet with very small knobs, pyriform or slightly elongated and offset or slightly overlapping basal pharyngeal bulb, posterior position of vulva, very short PUS, long spicules, and rounded tail terminus. Studied populations of N. hexaglyphus are similar to N. affinis, N. geraerti, N. medians, N. taylori and N. tuberosus, and can be distinguished from all of these species by the very small stylet knobs (Brzeski 1991). It differs from N. affinis, N. geraerti and N. medians by its longer spicules (20.5–21 vs. 15–17, 14–17.5 and 15–18 µm, respectively), and from N. taylori and N. tuberosus by smaller PUS/VBW ratio (0.8–1.3 vs. 1.8–3.0 and 2.0–2.4, respectively)., Published as part of Hashemi, Kobra & Karegar, Akbar, 2020, New and known species of Nothotylenchus Thorne, 1941 (Nematoda: Anguinidae) from Iran with an updated list of species, pp. 482-500 in Zootaxa 4729 (4) on page 491, DOI: 10.11646/zootaxa.4729.4.2, http://zenodo.org/record/3752014, {"references":["Khan, A. M. & Siddiqi, M. R. (1968) Three new species of Nothotylenchus (Nematoda: Neotylenchidae) from north India. Nematologica, 14, 369 - 376. https: // doi. org / 10.1163 / 187529268 X 00048","Brzeski, M. W. (1991) Review of the genus Ditylenchus Filipjev, 1936 (Nematoda: Anguinidae). Revue de Nematologie, 14, 9 - 59."]}

1. Nothotylenchus geraerti Kheiri, 1971 48 females: L = 742 ± 122 (506–1172) µm; stylet = 7.3 ± 0.3 (6.5–8) µm; pharynx = 127 ± 17 (96.5–167) µm; tail = 59.6 ± 8.1 (43.5–77) µm; a = 37.6 ± 4.9 (28.7–50.0); b = 5.8 ± 0.6 (4.6–7.6); c = 12.3 ± 1.4 (9.3–15.6); c′ = 4.9 ± 0.6 (3.7–6.1); V = 81.0 ± 2.3 (74.8–84.6); V′ = 88.3 ± 2.3 (82.0–95.8); PUS/VBW = 1.8 ± 0.4 (1.1–3.0); PUS/V-A = 42.9 ± 11.7 (14.8–77.0) %; V-A/T = 1.3 ± 0.2 (0.9–2.2). 10 males: L = 609 ± 75 (493–738) µm; stylet = 7.2 ± 0.2 (7–7.5) µm; pharynx = 122 ± 17 (98–154) µm; tail = 57.1 ± 7.5 (46.5–68.5) µm; a = 38.5 ± 2.6 (34.7–42.6); b = 4.9 ± 0.2 (4.6–5.3); c = 10.8 ± 1.6 (8.2–13.3); c′ = 5.0 ± 0.8 (4.1–6.3); spicules = 16.9 ± 1.1 (15.5–18.5) µm. Diagnosis and relationships. Nothotylenchus geraerti is characterised by its medium-sized body, six lateral field incisures, delicate stylet with posteriorly-sloping knobs, pyriform or slightly elongate and offset or slightly overlapping basal pharyngeal bulb, posterior vulva position, medium-sized spicules, and variable tail terminus shape (rounded to pointed). The studied populations of N. geraerti are similar to N. affinis, N. hexaglyphus, N. medians, N. taylori and N. tuberosus. The species differs from N. hexaglyphus by its more developed stylet and knobs, greater PUS/VBW ratio (1.1–3.0 vs. 0.5–1.1) and shorter spicules (15.5–18.5 vs. 19–22 μm). The four other species are very similar to N. geraerti, and, as their differentiation is so difficult, we think that these four species may all be synonymous. However, according to other published studies (Kheiri 1971, Husain & Khan 1974, Brzeski 1991, Zeidan & Geraert 1991), N. geraerti differs from N. affinis by its shorter stylet (6.5–8 vs. 8–9 μm), higher head, and greater PUS/VBW average (˃1.5 vs. ˂1.5), from N. medians by higher head, posteriorly slopping knobs (vs. rounded) and less-thick tail (vs. thick), from N. taylori by its relatively shorter spicules (maximum length ˂ 19 vs. ˃ 19 μm), and from N. tuberosus by shorter uterus length/VBW ratio (˂ 3 vs. 3–4) and basal bulb shape (mostly pyriform and offset vs. slightly overlapping). All of the features used to differentiate these four species display inter-population variation, making further studies on their type specimens necessary., Published as part of Hashemi, Kobra & Karegar, Akbar, 2020, New and known species of Nothotylenchus Thorne, 1941 (Nematoda: Anguinidae) from Iran with an updated list of species, pp. 482-500 in Zootaxa 4729 (4) on page 491, DOI: 10.11646/zootaxa.4729.4.2, http://zenodo.org/record/3752014, {"references":["Kheiri, A. (1971) Two new species of Nothotylenchus Thorne, 1941 from Iran and a redescription of N. affinis Thorne, 1941 (Nematoda, Neotylenchidae) with a key to the species of the genus. Nematologica, 16 (1970), 591 - 600. https: // doi. org / 10.1163 / 187529270 X 00810","Husain, S. I. & Khan, A. M. (1974) Three new species of neotylenchid nematodes from north India. Indian Journal of Nematology, 4, 81 - 87.","Brzeski, M. W. (1991) Review of the genus Ditylenchus Filipjev, 1936 (Nematoda: Anguinidae). Revue de Nematologie, 14, 9 - 59.","Zeidan, A. B. & Geraert, E. (1991) The Genus Ditylenchus Filipjev, 1936 in Sudan (Nematoda: Tylenchida). Afro-Asian Journal of Nematology, 1, 5 - 14."]}

Nothotylenchus siddiqii n. sp. (Figures 3 & 4) Measurements: Table 2. Female: Body straight or slightly curved ventrally. Cuticular annulation distinct, about 1 µm wide; lateral fields with six to nine incisures, 5–7 µm wide and occupying 31.2–42.1% of body width. Head low, with rounded margins, not offset, with two or three fine annuli, 1.5–2 µm high and about 5.5 µm wide. Cephalic skeleton moderately developed, outer margin of basal plate extending two to three annuli inside body. Stylet delicate, conus 26.3–39.7% of total stylet length, knobs medium-sized or small and rounded, 1–2 µm wide. DGO 0.5–1 µm posterior to stylet knobs. Median pharyngeal bulb elongated and spindle shape, wider than procorpus, 4.5–6 µm wide, valveless. Isthmus cylindrical and more slender than procorpus. Nerve ring surrounding isthmus almost at its middle, 69 (65.5–71.5) µm from anterior end. Hemizonid about two to three cuticular annuli long, 82 (79.5–88.5) µm from anterior end, two annuli anterior to S-E pore. S-E pore located from posterior half of isthmus to junction of isthmus and basal bulb. Deirids at the level of S-E pore. Basal pharyngeal bulb pyriform to slightly elongate and offset from intestine, length to width ratio 2.3 (1.8–2.8), with the two anterior-most intestinal cells appearing hyaline. Oocytes in single row, spermatheca elongate, filled with round sperms or empty, uterus quadricolumellar, sometimes with a valve between spermatheca and uterus, vagina 4–6 µm in length, 27.0–34.9% of VBW. PUS well developed, 26.5–40 µm, 2.2 (1.9–2.4) VBW, 54.2 (46.3–61.3) % V-A and 3.1 (2.6–3.6) ABW. Post-vulval body length 10.6 (10.5–10.9) ABW. Tail conical with usually rounded or seldom dull terminus. Male: General morphology, stylet, pharynx and tail shape similar to females. Spicules ventrally arcuate, gubernaculum simple and crescent-shaped, bursa 12–23.5 µm long and enveloping 25.7–48.6% of tail length. Tail conical with rounded terminus. Habitat & locality. Type population collected in 2014 from rhizosphere of alfalfa in Darab region, Fars province, Iran (GPS coordinates: 28°41.167′N, 54°16.056′E, elevation 1367 m.). Type material. Female holotype, three female paratypes and six male paratypes on six glass slides kept in the nematode collection of the Department of Plant Protection, School of Agriculture, Shiraz University, Iran, and two female paratypes and three male paratypes on two glass slides deposited in the Nematode Collection of the Plantenziektenkundige Dienst, Wageningen, The Netherlands. Diagnosis and relationships. Nothotylenchus siddiqi n. sp. is characterised by its short body, 573–645 µm in length, six to nine lateral field incisures, short, delicate stylet (6.5–7.5 µm) with rounded knobs, pyriform or slightly elongate and offset basal pharyngeal bulb, posterior vulva position (V = 79.3–81.0), PUS = 26.5–40 µm, short spicules, 14.5–16.5 µm long, and tail with rounded terminus. N. siddiqii n. sp. resembles N. affinis, N. hexaglyphus, N. persicus, N taylori Husain & Khan, 1974; N. geraerti and N. medians. But it can be distinguished from all these species by having 6–9 incisures in lateral fields (vs. 6). Moreover N. siddiqii n. sp. differs from N. affinis by greater PUS/VBW ratio (1.9–2.4 vs. 1.1–1.3), from N. hexaglyphus by shorter spicules (14.5–16.5 vs. 19–22 μm) and greater PUS/VBW ratio (1.9–2.4 vs. 0.5–1.3), from N. taylori by relatively shorter spicules (14.5–16.5 vs. 16.5–20 μm) and from N. persicus by shorter spicules (14.5–16.5 vs. 21–22 μm), position of the S-E pore (posterior half of isthmus to its junction with basal pharyngeal bulb vs. after basal pharyngeal bulb) and the tail terminus shape (rounded vs. pointed). Etymology. The species is named in honor of Dr. Mohammad Rafiq Siddiqi for his outstanding contributions to the science of nematology., Published as part of Hashemi, Kobra & Karegar, Akbar, 2020, New and known species of Nothotylenchus Thorne, 1941 (Nematoda: Anguinidae) from Iran with an updated list of species, pp. 482-500 in Zootaxa 4729 (4) on pages 487-489, DOI: 10.11646/zootaxa.4729.4.2, http://zenodo.org/record/3752014, {"references":["Husain, S. I. & Khan, A. M. (1974) Three new species of neotylenchid nematodes from north India. Indian Journal of Nematology, 4, 81 - 87."]}

Updated list, identification key and tabular compendium of Nothotylenchus characteristics Lists of Nothotylenchus species were published by Siddiqi (1986, 2000) and Andrássy (2007). In this study, an upto-date species list of the genus Nothotylenchus includes 41 valid species; species that have been synonymized with other species, transferred to other genera, or considered as species inquirendae, species incertae sedis and nomina nuda are compiled in separate lists. Nothotylenchus Thorne, 1941 = Boleodoroides Mathur, Khan & Prasad, 1966 = Boleodorus (Boleodoroides Mathur, Khan & Prasad, 1966) (Khera, 1970) ....Continued next page Tail tip: D = dull, Fili = filiform, FR = finely rounded, m = mucronate, P = pointed, R = rounded;? = no information available; [] = obtained from measured drawing and not given in description;> = slightly more than; Remark: In this table, Bursa/tail% that has been used previously (Brzeski, 1991, 1998; Sturhan & Brzeski, 1991), has been removed, because we observed that this feature is very variable even within a population and it can only be used in case of great differences between species (e.g., species with bursa reaching tail tip vs. adanal forms). Diagnosis (emended after Siddiqi, 2000). Anguinidae. Body vermiform, mature females not or slightly swollen. Body size between 0.4 and 1.2 mm. Cuticle finely striated. Lateral fields usually with four or six (occasionally two, five or more than six) incisures, which may be indistinct. Lip region usually low and flattened, continuous or slightly offset; cephalic framework of six equal sectors, not or slightly sclerotized. Stylet 5–14 µm long with conus less than half its total length, weak to moderately-developed, knobs rounded. Median pharyngeal bulb non-muscular, non-valvate; isthmus not separated from basal bulb by a constriction; pharyngeal glands enclosed in a basal bulb of variable shape and size, offset from intestine or overlapping it for a shorter or longer distance. Vulva at 57–86% of body length. Spermatheca elongate-axial. Post-vulval uterine sac one body width or more long. Tails of both sexes similar, elongate-conoid, rarely filiform, tip pointed to rounded, rarely mucronate or clavate. Deirids present. Phasmids absent. Bursa leptoderan, arising near head of spicules and generally extending to middle of tail, rarely adanal, exceptionally reaching tail end. Type species Nothotylenchus acris Thorne, 1941 Other species Nothotylenchus acutus Khan, 1965 = N. allii Khan & Siddiqi, 1968 = N. srinagarensis Fotedar & Mahajan, 1974 = N. indicus Saxena, Chhabra & Joshi, 1973 = N. paramonovi Gagarin, 1974 N. adasi Sykes, 1980 N. affinis Thorne, 1941 N. andrassy Jalalinasab, Nassaj Hosseini & Heydari, 2018 N. antricolus Andrássy, 1961 N. attenuatus Mulvey, 1969 N. basiri Khan, 1965 N. brzeskii n. sp. N. bhatnagari Tikyani & Khera, 1969 N. bhattii (Das & Bajaj, 2005) comb. n. = Ditylenchus bhattii Das & Bajaj, 2005 N. boroki Gagarin, 1999 N. citri (Varaprasad, Khan & Lal, 1981) Siddiqi, 1986 = Paurodontus citri Varaprasad, Khan & Lal, 1981 N. clavatus Dhanachand & Gambhir, 1991 N. cylindricollis Thorne, 1941 N. cylindricus Khan & Siddiqi, 1968 = N. elongatus Husain & Khan, 1974 N. danubialis Andrássy, 1960 N. drymocolus Rühm, 1956 N. fotedari Mahajan, 1977 N. geraerti Kheiri, 1971 N. goldeni Maqbool, 1982 N. hexaglyphus Khan & Siddiqi, 1968 N. loksai Andrássy, 1959 N. medians Thorne & Malek, 1968 N. oryzae (Mathur, Khan & Prasad, 1966) Siddiqi, 1986 = Boleodoroides oryzae Mathur, Khan & Prasad, 1966 N. persicus Esmaeili, Heydari, Castillo & Palomares-Rius, 2016 N. petilus Massey, 1974 N. phoenixae Esmaeili, Heydari & Ye, 2017 N. siddiqii n. sp. N. similis Thorne & Malek, 1968 N. singhi Das & Shivaswamy, 1980 N. solani (Varaprasad, Khan & Lal, 1981) Siddiqi, 1986 = Paurodontus solani Varaprasad, Khan & Lal, 1981 N. taylori Husain & Khan, 1974 N. tenuis Gagarin, 1999 N. thornei Andrássy, 1958 N. truncatus Eliashvili & Vacheishvili, 1980 N. tuberosus Kheiri, 1971 N. turfus (Yokoo, 1968) Siddiqi, 1986 = Neotylenchus turfus Yokoo, 1968 N. uniformis Truskova & Eroshenko, 1977 N. utschini Gagarin, 1974 N. websteri Kumar, 1983 Species inquirendae N. atypicus (Khera & Chaturvedi, 1977) Siddiqi, 1986 = Boleodorus atypicus Khera & Chaturvedi, 1977 N. buckleyi Das, 1960 N. compactus Massey, 1974 N. exiguus Andrássy, 1958 N. longistylus (Khera & Chaturvedi, 1977) Siddiqi, 1986 = Boleodorus longistylus Khera & Chaturvedi, 1977 N. major Thorne & Malek, 1968 N. montanus Kiknadze & Eliashvilli, 1988 N. parasimilis Massey, 1974 N. petilus Massey, 1974 N. typicus (Husain & Khan, 1968) Siddiqi, 1986 = Boleodorus typicus Husain & Khan, 1968 Ditylenchus robustus Das & Bajaj, 2005 (see remark 1) Ditylenchus triticus Das & Bajaj, 2005 (see remark 2) Species incertae sedis Neotylenchus nitidus Massey, 1969 = N. nitidus (Massey, 1969) Siddiqi, 1986 Nomina nuda N. callidus Izatullaeva, 1967 N. strictus Kapoor, 1982 Species transferred to other genera N. innuptus Andrássy, 1961, to Boleodorus Remarks on some species inquirendae Remark 1: Ditylenchus robustus is described by lateral fields with indistinct incisures, fusiform non-valvate median bulb, elongated, narrow spermatheca, genital tract coiled before crustaformeria, and short PUS. But drawings (Fig. 4A, C & D) show the spermatheca as not elongate and that it seems to be offset and pouch-like, considered by the authors as coiling of genital tract before crustaformeria (Das & Bajaj 2005). Therefore, reexamination and redescription of the species is necessary to determine its taxonomic position. Remark 2: D. triticus is characterised by indistinct lateral fields, fusiform and valveless median bulb, elongateoval spermatheca, filled with large sperms, looped genital tract near spermatheca, and short PUS. The elongated, bipartite and offset spermatheca, filled with small sperms shown in Fig. 3A & C (Das & Bajaj 2005) excludes this species from Ditylenchus or Nothotylenchus, and the disagreement between the description and drawings require it to be considered as species inquirenda. Key for identification of species 1- Lateral fields with two incisures; stylet length = 7.5 µm; V = 76; spicule length = 14 µm..................... N. petilus - Lateral fields with four or more incisures.................................................................. 2 2- Lateral fields with four incisures......................................................................... 3 - Lateral fields with five or more incisures................................................................. 22 3- Average stylet length ˂ 10 µm........................................................................... 4 - Average stylet length> 10 µm.......................................................................... 18 4- Basal pharyngeal bulb always long and cylindrical.......................................................... 5 - Basal pharyngeal bulb usually pyriform or short cylindrical................................................... 9 5- Tail terminus pointed.................................................................................. 6 - Tail terminus rounded; basal pharyngeal bulb cylindrical...................................................... 8 6- V = 90, vagina oblique; V-A about two times VBW; basal pharyngeal bulb cylindrical.................. N. cylindricollis - V = 71–77; vagina perpendicular to body axis; V-A longer.................................................... 7 7- Stylet length = 6.5–7 µm; PUS/VBW = 0.3–0.4.................................................... N. antricolus - Stylet length = 9–10 µm; PUS/VBW> 2........................................................ N. attenuatus 8- Basal pharyngeal bulb without projection; PUS/VBW = 0.8; spicule length = 17–19 µm................... N. bhatnagari - Basal pharyngeal bulb with a small cardiac projection; PUS/VBW = 1.5–2; spicule length = 14 µm.......... N. cylindricus 9- PUS/VBW = 3.4–5.7; tail terminus pointed; bursa covering 1/6 tail................................... N. danubialis - PUS/VBW ≤ 3...................................................................................... 10 10- Spicule length = 20.5–23 µm; V = 83.3–84.4; PUS/VBW = 0.6–1.2; tail thick and usually with rounded tip.. N. brzeskii n. sp. - Spicule length V = 67%; PUS/VBW = 0.6; tail terminus rounded..................................................... N. loksai - V ˃ 70%........................................................................................... 12 12- PUS/VBW = 0.6; tail terminus mucronate.......................................................... N. thornei - PUS/VBW = 1.3–3.0; tail terminus not mucronate.......................................................... 13 13- Body length ≥ 700 µm; tail terminus pointed.............................................................. 14 - Body length ≤ 650 µm; tail terminus variable, pointed, dull or rounded......................................... 15 14- Bursa nearly reaching tail terminus (~100%)......................................................... N. turfus - Bursa extending slightly past middle of tail........................................................... N. acris 15- Spicule length = 11–13 µm; V = 80–82; tail terminus pointed.......................................... N. websteri - Spicule length ≥ 13 µm; V N. acutus - Spicule length = 13–15 µm; tail terminus rounded or dull.................................................... 17 17- Stylet length = 6–8 µm; L = 350–500 µm; basal pharyngeal bulb pyriform.................................. N. basiri - Stylet length = 9 µm, L = 630 µm, basal pharyngeal bulb short cylindrical.................................. N. singhi 18- PUS/VBW N. clavatus - PUS/VBW> 1; tail terminus rounded, dull or pointed....................................................... 19 19- V = 57–66................................................................................. N. truncatus - V = 68–83......................................................................................... 20 20- Spicule length = 13–15 µm; V = 75–79; basal pharyngeal bulb cylindrical................................ N. utschini - Spicule length = 17–24 µm; basal pharyngeal bulb pyriform.................................................. 21 21- V = 68–76.8; spicule length = 21–24 µm............................................................ N. adasi - V = 80.2–83; spicule length = 17–21 µm............................................................ N. bhattii 22- Lateral fields with five incisures; stylet length = 11–12 µm; V = 81–85%; spicule length = 18–21 µm....... N. drymocolus - Lateral fields with six (or more) incisures................................................................. 23 23- Stylet length = 11–12 µm; V = 82–83, PUS/VBW = 0.5–0.7; spicule length = 20–21 µm..................... N. goldeni - Stylet length ≤ 10 µm................................................................................ 24 24- Basal pharyngeal bulb with extension.................................................................... 25 - Basal pharyngeal bulb without extension................................................................. 26 25- Stylet length = 6–8 µm; basal pharyngeal bulb extension short; tail terminus rounded......................... N. solani - Stylet length = 8–10 µm; basal pharyngeal bulb extension long; tail terminus pointed.......................... N. citri 26- Tail terminus clavate; basal pharyngeal bulb cylindrical, set off from the intestine........................... N. oryzae - Tail terminus pointed, dull or rounded................................................................... 27 27- Tail terminus always pointed........................................................................... 28 - Tail terminus rounded, dull or rarely pointed.............................................................. 32 28- Tail terminus filiform; spicule length = 13–14 µm; V = 71–76............................................ N. tenuis - Tail terminus pointed; spicule length ≥ 18 µm; V > 76 µm.................................................... 29 29- Stylet length = 5–7 µm; basal pharyngeal bulb pyriform; PUS/VBW ≤ 1......................................... 30 - Stylet length = 8–9 µm; basal pharyngeal bulb cylindrical; PUS/VBW> 1.2 (two similar species, probably synonymous)..................................................................................................... 31 30- S-E pore located posterior to basal pharyngeal bulb.................................................. N. persicus - S-E pore located at the level of anterior third part of basal pharyngeal bulb.............................. N. phoenixae 31- PUS/VBW = 1.2–1.4; male unknown............................................................ N. uniformis - PUS/VBW = 1.4–2.4; male known............................................................... N. andrassy 32- Stylet length = 10 µm; basal pharyngeal bulb always cylindrical.......................................... N. similis - Stylet length ≤ 9 µm; basal pharyngeal bulb mostly pyriform................................................. 33 33- Maximum spicule length> 19 µm...................................................................... 34 - Maximum spicule length V = 75–80; PUS/VBW = 1.8–3; spicule length = 16.5–20 µm........................................... N. taylori - V = 81.6–86.0; PUS/VBW = 0.5–1.3; spicule length = 19–22 µm.................................... N. hexaglyphus 35- Lateral fields with six to nine incisures; V = 79.3–81.0; PUS/VBW = 1.9–2.4; spicule length = 14.5–16.5 µm.................................................................................................... N. siddiqii n. sp. - Lateral fields with six incisures......................................................................... 36 36- V = 84–86; PUS/VBW N. fotedari - V usually ≤ 84...................................................................................... 37 37- Basal bulb always cylindrical; PUS/VBW = 1.1–1.8; tail terminus finely rounded............................ N. boroki - Basal bulb mostly pyriform or slightly overlapping (four similar species)........................................ 38 38- Uterus length/VBW = 3–4, basal bulb slightly overlapping........................................... N. tuberosus - Uterus length/VBW usually ˂ 3; basal bulb mostly pyriform and offset......................................... 39 39- Head high, stylet knobs sloping posteriorly........................................................ N. geraerti - Head low, stylet knobs rounded......................................................................... 40 40- Stylet length = 8–9 µm, PUS/VBW = 1.1–1.3, tail not thick............................................. N. affinis - Stylet length = 6.5–8 µm, PUS/VBW = 1.3–2.6, tail thick............................................. N. medians, Published as part of Hashemi, Kobra & Karegar, Akbar, 2020, New and known species of Nothotylenchus Thorne, 1941 (Nematoda: Anguinidae) from Iran with an updated list of species, pp. 482-500 in Zootaxa 4729 (4) on pages 491-498, DOI: 10.11646/zootaxa.4729.4.2, http://zenodo.org/record/3752014, {"references":["Siddiqi, M. R. (1986) Tylenchida: Parasites of Plants and Insects. Commonwealth Agricultural Bureaux, Farnham Royal, Slough, 645 pp.","Siddiqi, M. R. (2000) Tylenchida: Parasites of Plants and Insects. CABI Publishing, Wallingford, 833 pp. https: // doi. org / 10.1079 / 9780851992020.0000","Andrassy, I. (2007) Free-living nematodes of Hungary, II (Nematoda errantia). Hungarian Natural History Museum, Budapest, 496 pp.","Thorne, G. (1941) Some nematodes of the family Tylenchidae which do not possess a valvular median esophageal bulb. Great Basin Naturalist, 2, 37 - 85.","Mathur, V. K., Khan, E. & Prasad, S. K. (1966) Boleodoroides oryzae n. g., n. sp. (Nematoda: Boleodorinae) from Bihar, India. Nematologica, 12, 448 - 452. https: // doi. org / 10.1163 / 187529266 X 00950","Khera, S. (1970) Nematodes from the banks of still and running waters. 8. Order Tylenchida. Proceedings of the Zoological Society, Calcutta, 23, 53 - 65.","Brzeski, M. W. (1991) Review of the genus Ditylenchus Filipjev, 1936 (Nematoda: Anguinidae). Revue de Nematologie, 14, 9 - 59.","Brzeski, M. W. (1998) Nematodes of Tylenchina in Poland and Temperate Europe. Muzeum i Instytutu Zoologii, Polska Akademia Nauk (MiIZ PAN), Warszawa, 396 pp.","Sturhan, D. & Brzeski, M. W. (1991) Stem and bulb nematodes, Ditylenchus spp. In: Nickle, W. R. (Ed.), Manual of Agricultural Nematology. Marcel Dekker, Inc. New York, pp. 423 - 464.","Khan, S. H. (1965) Nothotylenchus acutus n. sp. and N. basiri n. sp. (Nematoda: Nothotylenchinae) from North India. Proceedings of the Helminthological Society of Washington, 32, 90 - 93.","Khan, A. M. & Siddiqi, M. R. (1968) Three new species of Nothotylenchus (Nematoda: Neotylenchidae) from north India. Nematologica, 14, 369 - 376. https: // doi. org / 10.1163 / 187529268 X 00048","Fotedar, D. N & Mahajan, R. (1974) Two new nematode species (Nothotylenchidae) from Kashmir. Indian Journal of Nematology, 2 (1972), 169 - 172.","Saxena, P. K., Chhabra, H. K. & Joshi, R. (1973) Nothotylenchus indicus n. sp. (Nematoda: Nothotylenchinae) from rhizosphere of peach trees in the Punjab (India). Zoologischer Anzeiger, 190, 140 - 142.","Gagarin, V. G. (1974) Two new species of the genus Nothotylenchus (Nothotylenchidae: Nematoda) and description of males of Tylocephalus auriculatus and Chronogaster typicus. Trudy Gelan, 24, 30 - 35. [in Russian]","Sykes, G. B. (1980) A new species of Nothotylenchus (Nematoda: Neotylenchoidea) from England. Systematic Parasitology, 1, 237 - 239. https: // doi. org / 10.1007 / BF 00009848","Jalalinasab, P., Nassaj Hosseini, M. & Heydari, R. (2018) Nothotylenchus andrassy n. sp. (Nematoda: Anguinidae) from Northern Iran. Journal of Nematology, 50, 219 - 228. https: // doi. org / 10.21307 / jofnem- 2018 - 025","Andrassy, I. (1961) Zur Taxonomie der Neotylenchiden. Nematologica, 6, 25 - 36. https: // doi. org / 10.1163 / 187529261 X 00243","Mulvey, R. H. (1969) Nematodes of the family Neotylenchidae (Tylenchida: Nematoda) from the Canadian high Arctic. Canadian Journal of Zoology, 47, 1261 - 1268. https: // doi. org / 10.1139 / z 69 - 198","Tikyani, M. G. & Khera, S. (1969) Nothotylenchus bhatnagari n. sp. from the rhizosphere of great millet (Sorghum vulgare Pers). Zoologischer Anzeiger, 182, 87 - 91.","Das, D. & Bajaj, H. K. (2005) New and known species of Ditylenchus Filipjev, 1936 from Haryana, India. Indian Journal of Nematology, 35, 11 - 23.","Gagarin, V. G. (1999) [Nematode fauna in rotten wood of birch, aspen, and pine in Borok (Yaroslavl' district, Central Russia)]. Zoologicheskii Zhurnal, 78, 146 - 157. [in Russian]","Varaprasad, K. S., Khan, E. & Lal, M. (1981) Paurodontus solani n. sp. and P. citri n. sp. (Nematoda: Neotylenchoidea) with a key to the species of Paurodontus Thorne, 1941. Indian Journal of Nematology, 10, 182 - 188.","Dhanachand, C. & Gambhir, R. K. (1991) One new and two known species of Tylenchida from Manipur, India. Current Nematology, 2, 33 - 38.","Husain, S. I. & Khan, A. M. (1974) Three new species of neotylenchid nematodes from north India. Indian Journal of Nematology, 4, 81 - 87.","Andrassy, I. (1960) Beitrage zur kenntnis der freilebenden Nematoden Chinas. Annales Musei Historico-naturalis Hungarici, 52, 201 - 216.","Ruhm, W. (1956) Die Nematoden der Ipiden. Parasitologische Schriftenreihe. Veb Gustav Fischer Verlag, Jena, 437 pp.","Mahajan, R. (1977) Scutellonema petersi n. sp. and Nothotylenchus fotedari n. sp., two new nematodes from India. Rivista di Parassitologia, 38, 334 - 337.","Kheiri, A. (1971) Two new species of Nothotylenchus Thorne, 1941 from Iran and a redescription of N. affinis Thorne, 1941 (Nematoda, Neotylenchidae) with a key to the species of the genus. Nematologica, 16 (1970), 591 - 600. https: // doi. org / 10.1163 / 187529270 X 00810","Maqbool, M. A. (1982) Three new species of the superfamily Neotylenchoidea (Nematoda: Tylenchida) from Pakistan. Journal of Nematology, 14, 317 - 323.","Andrassy, I. (1959) Freilebende nematoden aus Rumanien. Annales Universitatis Scientarum Budapestensis de Rolando Eotvos Nominate, Sectio Biologica, 2, 3 - 27.","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, 1 - 111.","Esmaeili, M., Heydari, R., Castillo, P. & Palomares-Rius, J. E. (2016) Nothotylenchus persicus n. sp. (Nematoda: Anguinidae) from Kermanshah province, Iran. Nematology, 18, 29 - 37. https: // doi. org / 10.1163 / 15685411 - 00002940","Massey, C. L. (1974) Biology and taxonomy of nematode parasites and associates of bark beetles in the United States. Agricultural Handbook No. 446. Department of Agriculture, Washington, 233 pp.","Esmaeili, M., Heydari, R. & Ye, W. (2017) Description of a new anguinid nematode, Nothotylenchus phoenixae n. sp. (Nematoda: Anguinidae) associated with palm date trees and its phylogenetic relations within the family Anguinidae. Journal of Nematology, 49, 268 - 275. https: // doi. org / 10.21307 / jofnem- 2017 - 072","Das, V. M. & Shivaswamy, V. (1980) Paurodontus brassicae n. sp. and Nothotylenchus singhi n. sp. from south India. Proceedings of the Indian Academy of Parasitology, 1, 62 - 65.","Andrassy, I. (1958) Erd-und Susswassernematoden aus Bulgarien Acta Zoologica Academiae Scientiarum Hungaricae, 4, 1 - 88.","Eliashvili, T. S. & Vacheishvili, L. A. (1980) [A new species of the nematode Nothotylenchus truncatus sp. nov. (Nematoda: Tylenchida) from Eastern Georgia]. Soobshcheniia Akademii nauk Gruzinskoi SSR, 98, 177 - 180. [in Russian]","Yokoo, T. (1968) Nematological studies on the yellow patch of green grass of the golf link: II. On the nemic-fauna in the green grass of International Golf Link of Isahaya, Nagasaki Prefecture, with descriptions on new species of Neotylenchus (Nematoda: Neotylenchidae). Agricultural Bulletin, Saga University, 26, 9 - 19.","Truskova, G. M. & Eroshenko, A. S. (1977) [The nematode fauna of herbaceous and ligneous plants in the pine plantations of the Primor'yal]. Trudy Biologicheskogo Instituta Novosibirsk Seriya, 47, 35 - 49. [in Russian]","Kumar, P. (1983) Nothotylenchus websteri n. sp. (Nematoda: Neotylenchoidea) from Lucknow. Indian Journal of Parasitology, 7, 105 - 107.","Khera, S. & Chaturvedi, Y. (1977) Nematodes from tea plantations of Dhera Dun, India. Records of the Zoological Survey of India, 72, 125 - 152.","Das, V. M. (1960) Studies on the nematode parasites of plants in Hyderabad (Andhra Pradesh, India). Zeitschrift fur Parasiten- kunde, 19, 553 - 605. https: // doi. org / 10.1007 / BF 00260158","Husain, S. I. & Khan, A. M. (1968) Paurodontella n. gen. and three new species of nematodes from North India (Nematoda: Neotylenchidae). Nematologica, 13 (1967), 493 - 500. https: // doi. org / 10.1163 / 187529267 X 00274","Massey, C. L. (1969) New species of tylenchs associated with bark beetles in New Mexico and Colorado. Proceedings of the Helminthological Society of Washington, 36, 43 - 52.","Izatullaeva, R. I. (1967) [The helminth fauna of certain flower crops in Kazakhstan]. Materialy Nauchnoy Konferentsii Vsesoyuznogo Obshchestva Gel'mintologov, 5 (1966), 175 - 179. [in Russian]","Kapoor, M. (1982) Taxonomic studies on nematodes of some medicinal and aromatic plants of North India. Thesis, Panjab University, Chandigarh, 418 pp."]}

4. Ditylenchus equalis Heyns, 1964 8 females: L = 749 (602–861) µm; stylet = 7.4 (7–8) µm; pharynx = 118 (98–145) µm; tail = 73.5 (51–91) µm; a = 44.8 (35.6–57.5); b = 6.4 (5.4–8.2); c = 10.4 (9.1–12.3); c′ = 6.6 (4.2–8.5); V = 80.6 (78.9–82.6); V′ = 89.3 (88.1– 90.5); PUS / VBW = 1.2 (1.0–1.4; in one specimen = 2.1); PUS /V-A = 30.6 (20.9–58.4) %; V-A/T = 1.0 (0.8–1.2). 4 males: L = 537 (494–565) µm; stylet = 7.3 (7–8) µm; pharynx = 116 (104–134) µm; tail = 51.2 (50–53) µm; a = 44.1 (39.1–48.7); b = 4.6 (4.2–4.9); c = 10.5 (9.7–11.1); c′ = 5.1 (4.6–5.5); spicules = 14.9 (14–16) µm. Diagnosis. D. equalis is characterised by four lateral field incisures, unstriated head, delicate, short stylet with rounded knobs, basal pharyngeal bulb pyriform to elongate and usually offset, sometimes with a slight overlap (up to 2 µm), posterior position of vulva, short post-vulval uterine sac, tail tip usually pointed and sometimes rounded, and short spicules. D. equalis is similar to D. deiridus Thorne & Malek, 1968, D. emus Khan, Chawla & Prasad, 1969, D. exilis Brzeski, 1984, D. filimus Anderson, 1983, D. parvus and D. terricolus. It differs from D. deiridus by having PUS (vs. PUS absent), and from D. emus and D. exilis by different shape of tail tip (usually pointed vs. rounded). It can be distinguished from D. filimus by shorter spicules (14–16 vs. 29 μm) and lower V index (78.9–82.6 vs. 81–85), from D. parvus and D. terricolus by greater V index (78.9–82.6 vs. 71–77) and lower PUS/VBW ratio (1.0–1.4 vs. 1.8–4.3 and 2.1–3.2, respectively)., Published as part of Hashemi, Kobra & Karegar, Akbar, 2019, Description of Ditylenchus paraparvus n. sp. from Iran with an updated list of Ditylenchus Filipjev, 1936 (Nematoda: Anguinidae), pp. 85-113 in Zootaxa 4651 (1) on page 92, DOI: 10.11646/zootaxa.4651.1.6, http://zenodo.org/record/3359134, {"references":["Heyns, J. (1964) Aphelenchoides helicus n. sp. and Ditylenchus equalis n. sp., two new soil inhabiting nematodes. South African Journal of Agricultural Science, 7, 147 - 150.","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, 1 - 111.","Khan, E., Chawla, M. L. & Prasad, S. K. (1969) Tylenchus (Aglenchus) indiens n. sp. and Ditylenchus emus n. sp. (Nematoda: Tylenchidae) from India. Labdev Journal of Science and Technology, 7 B, 311 - 314.","Brzeski, M. W. (1984) Three new species of Ditylenchus Filipjev, 1936, and comments on Basiroides longimatricalis Kazachenko, 1975 (Nematoda: Anguinidae). Nematologica, 29, 380 - 389. https: // doi. org / 10.1163 / 187529283 X 00267","Anderson, R. V. (1983) An emended description of Ditylenchus valveus Thorne & Malek, 1968 and description of D. filimus n. sp. (Nematoda: Tylenchidae) from mushroom compost in Canada. Canadian Journal of Zoology, 61, 2319 - 2323. https: // doi. org / 10.1139 / z 83 - 306"]}

7. Ditylenchus myceliophagus Goodey, 1958 348 females: L = 734 (481–1108) µm; stylet = 7.7 (6–9) µm; pharynx = 119 (87–162) µm; anterior end to end of glands = 127 (87–187) µm; tail = 56.0 (33–85) µm; a = 38.1 (22.8–56.0); b = 6.2 (4.3–8.8); c = 13.2 (8.5–18.9); c′ = 4.6 (2.8–7.2); V = 80.8 (73.9–85.3); V′ = 87.6 (82.1–92.5); PUS / VBW = 2.1 (0.8–3.7); PUS /V-A = 45.5 (17.3–69.6) %; V-A/T = 1.5 (0.8–2.4). 201 males: L = 701 (426–939) µm; stylet = 7.7 (6.5–9) µm; pharynx = 120 (86.5–168) µm; anterior end to end of glands = 126 (92–168) µm; tail = 52.8 (39.5–76) µm; a = 40.6 (27.8–56.7); b = 5.9 (3.7–8.4); c = 13.4 (8.6–18.7); c′ = 4.5 (3.2–6.5); spicules = 18.9 (14.5–23.5) µm. Diagnosis. D. myceliophagus is distinctive because of its thin to stout body, six lateral field incisures (sometimes with one to three additional lines), refractive, short crescentic cephalic skeleton, delicate, short stylet with rounded or posteriorly sloping knobs, variable shape of basal pharyngeal bulb (pyriform, cylindrical or elongate), basal pharyngeal bulb offset or with slight to long overlap, up to 66 µm, variable vuvla position and length of postvulval uterine sac, a usually thick tail that suddenly narrows from about half to the last two thirds, with a usually rounded, sometimes dull, tip, and variable spicule length. Unlike the original description of D. myceliophagus (Goodey 1958), the head of different populations in this study was striated. The general shape of tail in the Iranian populations was more diverse than the Brzeski populations (1998) (usually thick and sometimes narrow vs. always thick). The Iranian populations of D. myceliophagus are close to D. acutatus, D. anchilisposomus, D. apus, D. dauniae, D. elegans, D. geraerti, D. medicaginis, D. silvaticus, D. tenuidens, D. triformis and D. valveus. D. myceliophagus can be distinguished from all of these species by its crescentic and refractive cephalic skeleton. In addition, it differs from D. acutatus with a lower PUS / VBW ratio (2.1 (0.8–3.7) vs. 3.5 (2.8–4.1)) and different tail shape (usually thick, with rounded tip vs. thick, with pointed tip), from D. apus by greater PUS / VBW ratio (0.8–3.7 vs. 0.2–0.4) and different shape of basal pharyngeal bulb (pyriform to elongate and offset or with overlap vs. elongate with long overlap), from D. elegans by its shorter body (481–1108 vs. 1030–1370), shorter tail (33–85 vs. 111–149 μm), relatively greater V (73.9–85.3 vs. 71–77), lower PUS / VBW ratio (0.8–3.7 vs. 3.2) and different tail shape (usually thick with rounded tip vs. narrow with pointed tip), from D. medicaginis to some extent by tail shape (usually thick with rounded tip vs. narrow with often pointed to dull tip), from D. silvaticus by different tail shape (usually thick with rounded tip vs. thick with pointed or rounded tip with mucron), from D. tenuidens by striated head (vs. smooth) and different tail tip (usually rounded vs. sharply pointed), from D. triformis by longer spicules (14.5–23.5 vs. 13–15 µm) and six incisures vs. four at anus region, and from D. valveus, to some extent, by tail shape (usually thick with rounded tip vs. narrow with often dull to rounded terminus). In the case of individuals with offset basal pharyngeal bulb or slight overlap over intestine, the basal plate of the cephalic skeleton is the only difference between D. myceliophagus and D. dauniae and D. geraerti, also if there are individuals with long basal bulb overlap over intestine, this feature is again the only difference between D. myceliophagus and D. anchilisposomus., Published as part of Hashemi, Kobra & Karegar, Akbar, 2019, Description of Ditylenchus paraparvus n. sp. from Iran with an updated list of Ditylenchus Filipjev, 1936 (Nematoda: Anguinidae), pp. 85-113 in Zootaxa 4651 (1) on pages 94-95, DOI: 10.11646/zootaxa.4651.1.6, http://zenodo.org/record/3359134, {"references":["Goodey, J. B. (1958) Ditylenchus myceliophagus n. sp. (Nematoda: Tylenchidae). Nematologica, 3, 91 - 96. https: // doi. org / 10.1163 / 187529258 X 00166"]}

10. Ditylenchus valveus Thorne & Malek, 1968 9 females: L = 739 (638–907) µm; stylet = 7.2 (7–8) µm; pharynx = 123 (106–153) µm; tail = 66.4 (55–92) µm; a = 38.3 (29.3–51.7); b = 6.1 (5.2–8.0); c = 11.5 (9.8–14.4); c′ = 5.5 (4.5–7.4); V = 80.4 (79.0–82.7); V′ = 88.3 (84.9–89.8); PUS / VBW = 2.0 (1.1–2.6); PUS /V-A = 46.2 (23.5–68.4) %; V-A/T = 1.2 (1.0–1.8). 2 males: L = 625, 816 µm; stylet = 7, 8 µm; pharynx = 122, 128 µm; tail = 59, 74 µm; a = 44.5, 54.1; b = 5.1, 6.4; c = 10.5, 11.1; c′ = 5.4, 6.3; spicules = 18.5, 19 µm. Diagnosis. D. valveus is characterised by six lateral field incisures, delicate, short stylet with round knobs, usually pyriform, but sometimes cylindrical, basal pharyngeal bulb that is usually offset and sometimes with slight overlap (up to 2 µm), posterior position of vulva, usually long post-vulval uterine sac, usually dull and rounded, but sometimes mucronate, tail tip, and spicule length. The Iranian population of D. valveus comes close to D. acutatus, D. apus, D. dauniae, D. elegans, D. geraerti, D. medicaginis, D. myceliophagus, D. silvaticus, D. tenuidens and D. triformis. It can be distinguished from D. acutatus by its lower PUS / VBW ratio (1.1–2.6 vs. 2.8–4.1) and different tail tip (rounded to dull vs. pointed), from D. apus by having greater V and PUS / VBW ratio (79.0–82.7 and 1.1–2.6 vs. 75–76 and 0.2–0.4, respectively) and also by the shape of the basal pharyngeal bulb (pyriform and usually offset vs. elongate with long overlap), from D. dauniae by lower V (79.0–82.7 vs. 83–84) and greater c ′ index (4.5–7.4 vs. 3.2–4.1), from D. elegans by shorter body and tail length (638–907 and 55–92 vs. 1030–1370 and 111–149 µm), greater V (79.0–82.7 vs. 71–77), lower c ′ ratio (4.5–7.4 vs. 7.2–11.3) and different tail tip shape (rounded and dull vs. pointed), from D. geraerti by tail shape (narrow with rounded to dull tip vs. thick with rounded tip) and greater c′ index (4.5–7.4 vs. 3.5–5), from D. myceliophagus by different cephalic skeleton development (moderate vs. crescentic and refractive), from D. silvaticus by different tail shape (narrow with rounded to dull terminus vs. thick with pointed or rounded tip with mucron), from D. tenuidens by striated (vs. smooth) head, position of S-E pore (located between posterior half of isthmus and anterior half of basal bulb vs. half of isthmus), shape of basal bulb (pyriform vs. elongate) and tail tip (rounded to dull vs. pointed), and from D. triformis by longer spicules (18.5–19 vs. 13–15 μm) and having six incisures at anus region (vs. four). The closest species to D. valveus is D. medicaginis (see D. medicaginis diagnosis)., Published as part of Hashemi, Kobra & Karegar, Akbar, 2019, Description of Ditylenchus paraparvus n. sp. from Iran with an updated list of Ditylenchus Filipjev, 1936 (Nematoda: Anguinidae), pp. 85-113 in Zootaxa 4651 (1) on page 96, DOI: 10.11646/zootaxa.4651.1.6, http://zenodo.org/record/3359134, {"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, 1 - 111."]}

9. Ditylenchus triformis Hirschmann & Sasser, 1955 (Figure 4) 9 females: L = 621 (493–729) µm; stylet = 7.0 (7–8) µm; pharynx = 118 (97–131) µm; tail = 56.9 (44–66) µm; a = 39.3 (34.7–46.1); b = 5.3 (4.6–6.0); c = 10.9 (9.1–12.7); c′ = 5.4 (4.7–6.6); V = 78.3 (76.2–80.2); V′ = 86.2 (83.3–88.3); PUS / VBW = 1.7 (1.3–2.7); PUS /V-A = 32.6 (22.2–48.2) %; V-A/T = 1.4 (1.1–1.8). 2 males: L = 462, 581 μm; stylet = 6.5, 7 μm; pharynx = 97, 117 µm; tail = 51, 56 µm; a = 42.4, 44.7; b = 4.8, 5.0; c = 9.0, 10.3; c ′ = 6.9, 5.6; spicules = 14, 16 µm. Diagnosis. D. triformis is characterised by six lateral field incisures that reduced to four at anus level, delicate, short stylet with rounded knobs, elongate basal pharyngeal bulb that is usually offset and sometimes slightly overlapping (up to 6 µm), posterior position of vulva, rather long post-vulval uterine sac, usually rounded, seldom dull or pointed, tail tip, and short spicules. The Iranian population of D. triformis is close to 11 species, including D. acutatus, D. dauniae, D. geraerti, D. medicaginis, D. myceliophagus, D. silvaticus, D. valveus, D. virtudesae, D. apus, D. elegans and D. tenuidens. It differs from all of these species by the reduction of lateral field incisures at the anus region to four. In addition, it can be distinguished from D. virtudesae by longer spicules (14–16 vs. 11 μm), from D. acutatus and D. elegans by lower PUS / VBW ratio (1.3–2.7 vs. 2.8–4.1, 3.2, respectively) and different tail tip shape (usually rounded vs. pointed or dull), and from D. apus by greater PUS / VBW ratio (1.3–2.7 vs. 0.2–0.4) and the nature of the basal pharyngeal bulb (pyriform to elongate, offset or with slight overlap vs. elongated and with long overlap). It differs from D. myceliophagus by different basal plate of cephalic skeleton (weak to moderate vs. refractive and crescentic), from D. silvaticus and D. tenuidens by striated (vs. smooth) head and different tail tip shape (usually rounded vs. pointed), and from D. valveus by shorter spicules (14–16 vs. 16–23 µm). The only difference between D. triformis and the three remaining species is the number of incisures at the anus region., Published as part of Hashemi, Kobra & Karegar, Akbar, 2019, Description of Ditylenchus paraparvus n. sp. from Iran with an updated list of Ditylenchus Filipjev, 1936 (Nematoda: Anguinidae), pp. 85-113 in Zootaxa 4651 (1) on page 96, DOI: 10.11646/zootaxa.4651.1.6, http://zenodo.org/record/3359134, {"references":["Hirschmann, H. & Sasser, J. N. (1955) On the occurrence of an intersexual form in Ditylenchus triformis, n. sp. (Nematoda, Tylenchida). Proceedings of the Helminthological Society of Washington, 22, 115 - 123."]}

1. Ditylenchus anchilisposomus (Tarjan, 1958) Fortuner, 1982 7 females: L = 660 (615–708) µm; stylet = 8.4 (7.5–9) µm; pharynx = 117 (107–139) µm; anterior end to end of glands = 144 (114–170) µm; tail = 55.8 (51–66) µm; a = 42.1 (37.3–46.9); b = 5.7 (4.4–6.5); c = 11.9 (10.6–13.2); c′ = 5.3 (4.8–5.7); V = 80.1 (77.5–82.0); V′ = 87.5 (85.3–88.7); PUS / VBW = 2.0 (1.4–2.6); PUS /V-A = 40.4 (29.2– 46.2) %; V-A/T = 1.4 (1.2–1.7). 3 males: L = 620 (610–632) µm; stylet = 7.8 (7–8) µm; pharynx = 110 (103–114) µm; anterior end to end of glands = 138 (135–139) µm; tail = 55.4 (50–64) µm; a = 44.5 (41.5–47.3); b = 5.7 (5.4–6.0); c = 11.3 (9.9–12.1); c′ = 5.3 (4.7–6.6); spicules = 15.5 (15–16) µm. Diagnosis. D. anchilisposomus is characterised by six lateral field incisures, head with rounded margins and weak cephalic skeleton, delicate medium-sized stylet with small rounded to posteriorly sloping knobs, usually long (seldom pyriform) basal pharyngeal bulb overlapping intestine and measuring 7–41 µm and seldom pyriform, posterior position of vulva, long post-vulval uterine sac, rounded to dull tail tip and short spicules. The Iranian population of D. anchilisposomus is similar to D. dauniae Brzeski & Marinari, 1991, D. geraerti, D. medicaginis Wasilewska, 1965, D. myceliophagus Goodey, 1958, D. silvaticus Brzeski, 1991, D. tenuidens Gritzenko, 1971, D. triformis and D. valveus Thorne & Malek, 1968. It can be distinguished from all of these, except some specimens of D. myceliophagus, by having a long pharyngeal lobe overlapping intestine. The only difference between D. anchilisposomus and D. myceliophagus is the cephalic skeleton (weak to medium and extending two annuli into the body vs. short, crescentic and with refractive margins)., Published as part of Hashemi, Kobra & Karegar, Akbar, 2019, Description of Ditylenchus paraparvus n. sp. from Iran with an updated list of Ditylenchus Filipjev, 1936 (Nematoda: Anguinidae), pp. 85-113 in Zootaxa 4651 (1) on page 90, DOI: 10.11646/zootaxa.4651.1.6, http://zenodo.org/record/3359134, {"references":["Tarjan, A. C. (1958) A new genus, Pseudhalenchus (Tylenchinae: Nematoda), with descriptions of two new species. Proceedings of the Helminthological Society of Washington, 25, 20 - 25.","Fortuner, R. (1982) On the genus Dilylenchus Filipjev, 1936 (Nematoda: Tylenchida). Revue de Nematologie, 5, 17 - 38.","Wasilewska, L. (1965) Ditylenchus medicaginis n. sp., a new parasitic nematode from Poland (Nematoda, Tylenchidae). Bulletin de l'Academie polonaise des Sciences. Classe II. Serie des Sciences Biologiques, 13, 167 - 170.","Goodey, J. B. (1958) Ditylenchus myceliophagus n. sp. (Nematoda: Tylenchidae). Nematologica, 3, 91 - 96. https: // doi. org / 10.1163 / 187529258 X 00166","Brzeski, M. W. (1991) Review of the genus Ditylenchus Filipjev, 1936 (Nematoda: Anguinidae). Revue de Nematologie, 14, 9 - 59.","Gritzenko, V. P. (1971) Ditylenchus tenuidens n. sp. and Aphelenchoides curiolis n. sp. (Nematoda, Tylenchidae and Aphelenchoididae) from Kirgizia. Zoologicheskii Zhurnal, 50, 1402 - 1405.","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, 1 - 111."]}

3. Ditylenchus dipsaci (Kühn, 1857) Filipjev, 1936 191 females: L = 1140 (878–1510) µm; stylet = 10.5 (9–13) µm; pharynx = 177 (143–221) µm; tail = 77.6 (53–105) µm; a = 46.1 (29.5–62.5; in one specimen = 18.5); b = 6.5 (5.0–8.4); c = 14.8 (11.5–25.7); c′ = 5.4 (3.7–7.7); V = 80.4 (75.8–87.3; in one specimen = 69.6); V ′ = 86.3 (74.1–93.9); PUS / VBW = 2.6 (1.5–3.6); PUS /V-A = 43.4 (25.6–77.8) %; V-A/T = 1.8 (0.8–2.4; in one specimen = 3.9). 134 males: L = 1103 (877–1346) µm; stylet = 10.5 (9.5–12) µm; pharynx = 175 (130–210) μm; tail = 76.5 (58–99) μm; a = 48.4 (33.8–66.5); b = 6.3 (5.2–8.1); c = 14.5 (12.0–18.3); c′ = 5.1 (3.5–7.4); spicules = 23.7 (20–28) µm. Diagnosis. D. dipsaci is distinguished by its long, slender body, four lateral field incisures (sometimes with one to three additional lines), well-developed cephalic skeleton, moderately large stylet with distinct rounded knobs, moderately-developed median bulb with distinct valve, variable shape of basal pharyngeal bulb (usually long and cylindrical, sometimes pyriform and shorter, sometimes with a short stem), basal pharyngeal bulb usually with slight intestinal overlap, averaging 5 µm (seldom up to 29 µm) and sometimes offset, posterior position of vulva, long post-vulval uterine sac, thick tail with usually pointed, but sometimes dull tip, and long spicules. The Iranian populations of D. dipsaci are similar to D. angustus (Butler, 1913) Filipjev, 1936, D. gigas Vovlas, Troccoli, Palomares-Rius, De Luca, Liébanas, Landa, Subbotin & Castillo, 2011, D. laurae Skwiercz, Kornobis, Winiszewska, Przybylska, Obrępalska-Stęplowska, Gawlak & Subbotin, 2017, D. solani, D. sturhani Mirbabaei Karani, Eskandari, Ghaderi, Heydari, & Miraeez, 2017 and D. weischeri Chizhov, Borisov & Subbotin, 2010. They differs from D. angustus by relatively longer spicules (20–28 vs. 16–21 μm), less bursa/tail% (24.8–85.2 vs. approximately 100%), different shape of basal pharyngeal bulb (long and cylindrical vs. clavate). It can be distinguished from D. laurae by shorter body (878–1510 vs. 1523–2095 μm), shorter tail (53–105 vs. 104–127 μm), less a index (18.5–62.5 vs. 72.5–103) and PUS / VBW (1.5–3.6 vs. 4.3–5.6), shorter basal pharyngeal bulb (length/width = 1.9–5.8 vs. approximately 10) and the shape of tail tip (usually pointed vs. mucronate), from D. solani by having longer spicules (20–28 vs. 18–20 μm), the shape of basal pharyngeal bulb and tail tip (long and cylindrical and usually pointed vs. pyriform and dull to rounded, respectively), from D. sturhani with longer body 878–1510 vs. 656–865 μm), longer spicules (20–28 vs. 18.5–20.5 μm), greater PUS / VBW (2.6 (1.5–3.6) vs. 1.4 (1.0–1.9)) and different shape of tail tip (usually pointed vs. usually dull), from D. weischeri with greater c′ index (5.4 (3.7–7.7) vs. 3.7 (2.9–4.8)). In the description of D. weischeri, other differences with D. dipsaci such as shorter tail and spicules, greater c index, V-A, V-A/T and PUS length are also noted, that falls within the range of variation of different D. dispaci populations in the present study and are not useful. The closest species to D. dipsaci is D. gigas, which is distinguished only with shorter body length (878–1510 vs. 1270–1932 μm)., Published as part of Hashemi, Kobra & Karegar, Akbar, 2019, Description of Ditylenchus paraparvus n. sp. from Iran with an updated list of Ditylenchus Filipjev, 1936 (Nematoda: Anguinidae), pp. 85-113 in Zootaxa 4651 (1) on page 91, DOI: 10.11646/zootaxa.4651.1.6, http://zenodo.org/record/3359134, {"references":["Kuhn, J. (1857) Uber Das Vorkommen von Anguillulen in erkrankten Bluthenkopfen von Dipsacus fullonum L. Zeitschrift fur wissenschaftliche Zoologie, 9, 129 - 137.","Filipjev, I. N. (1936) On the classification of the Tylenchinae. Proceedings of the Helminthological Society of Washington, 3, 80 - 82.","Butler, E. J. (1913) Disease of rice: An eelworm disease of rice. Bulletin Agricultural Research Institute, Pusa, Bulletin, 34, 1 - 37.","Vovlas, N., Troccoli, A., Palomares-Rius, J. E., De Luca, F., Liebanas, G., Landa, B. B., Subbotin, S. A. & Castillo, P. (2011) Ditylenchus gigas n. sp. parasitizing broad bean: a new stem nematode singled out from the Ditylenchus dipsaci species complex using a polyphasic approach with molecular phylogeny. Plant Pathology, 60, 762 - 775. https: // doi. org / 10.1111 / j. 1365 - 3059.2011.02430. x","Skwiercz, A. T., Kornobis, F. W., Winiszewska, G., Przybylska, A., Obrepalska-Steplowska, A., Gawlak, M. & Subbotin, S. A. (2017) Ditylenchus laurae n. sp. (Tylenchida: Anguinidae) from Poland-a new species of the D. dipsaci complex associated with a water plant, Potamogeton perfoliatus L. Nematology, 19, 197 - 209. https: // doi. org / 10.1163 / 15685411 - 00003040","Chizhov, V. N., Borisov, B. A. & Subbotin, S. A. (2010) A new stem nematode, Ditylenchus weischeri n. sp. (Nematoda: Tylenchida), a parasite of Cirsium arvense (L.) Scop. in the Central Region of the Non-Chernozem Zone Russia. Russian Journal of Nematology, 18, 95 - 102."]}

9. Ditylenchus triformis Hirschmann & Sasser, 1955 (Figure 4) 9 females: L = 621 (493–729) µm; stylet = 7.0 (7–8) µm; pharynx = 118 (97–131) µm; tail = 56.9 (44–66) µm; a = 39.3 (34.7–46.1); b = 5.3 (4.6–6.0); c = 10.9 (9.1–12.7); c′ = 5.4 (4.7–6.6); V = 78.3 (76.2–80.2); V′ = 86.2 (83.3–88.3); PUS / VBW = 1.7 (1.3–2.7); PUS /V-A = 32.6 (22.2–48.2) %; V-A/T = 1.4 (1.1–1.8). 2 males: L = 462, 581 μm; stylet = 6.5, 7 μm; pharynx = 97, 117 µm; tail = 51, 56 µm; a = 42.4, 44.7; b = 4.8, 5.0; c = 9.0, 10.3; c ′ = 6.9, 5.6; spicules = 14, 16 µm. Diagnosis. D. triformis is characterised by six lateral field incisures that reduced to four at anus level, delicate, short stylet with rounded knobs, elongate basal pharyngeal bulb that is usually offset and sometimes slightly overlapping (up to 6 µm), posterior position of vulva, rather long post-vulval uterine sac, usually rounded, seldom dull or pointed, tail tip, and short spicules. The Iranian population of D. triformis is close to 11 species, including D. acutatus, D. dauniae, D. geraerti, D. medicaginis, D. myceliophagus, D. silvaticus, D. valveus, D. virtudesae, D. apus, D. elegans and D. tenuidens. It differs from all of these species by the reduction of lateral field incisures at the anus region to four. In addition, it can be distinguished from D. virtudesae by longer spicules (14–16 vs. 11 μm), from D. acutatus and D. elegans by lower PUS / VBW ratio (1.3–2.7 vs. 2.8–4.1, 3.2, respectively) and different tail tip shape (usually rounded vs. pointed or dull), and from D. apus by greater PUS / VBW ratio (1.3–2.7 vs. 0.2–0.4) and the nature of the basal pharyngeal bulb (pyriform to elongate, offset or with slight overlap vs. elongated and with long overlap). It differs from D. myceliophagus by different basal plate of cephalic skeleton (weak to moderate vs. refractive and crescentic), from D. silvaticus and D. tenuidens by striated (vs. smooth) head and different tail tip shape (usually rounded vs. pointed), and from D. valveus by shorter spicules (14–16 vs. 16–23 µm). The only difference between D. triformis and the three remaining species is the number of incisures at the anus region.

8. Ditylenchus parvus Zell, 1988 19 females: L = 690 (541–813) µm; stylet = 7.2 (6.5–8) µm; pharynx = 121 (110–141) µm; tail = 70.6 (53–102); a = 41.5 (35.3–49.6); b = 5.7 (4.8–6.9); c = 9.9 (6.9–11.0); c′ = 6.3 (4.8–7.9); V = 73.5 (70.8–75.2); V′ = 81.9 (80.4– 83.0); PUS / VBW = 2.7 (1.0–3.6); PUS /V-A = 35.6 (17.8–45.8) %; V-A/T = 1.6 (1.1–1.9). 7 males: L = 595 (442–717) µm; stylet = 7.0 (6.5–7.5) µm; pharynx = 111 (95–128) µm; tail = 63.9 (44–87.5) µm; a = 44.7 (34.7–57.8); b = 5.3 (4.4–6.0); c = 9.5 (8.2–10.3); c′ = 6.2 (4.6–8.2); spicules = 14.6 (12–17). Diagnosis. D. parvus is characterised by four lateral field incisures, delicate, short stylet with rounded knobs, pyriform basal pharyngeal bulb that is offset or slightly overlapping (up to 4 µm), position of vulva, variable length of post-vulval uterine sac, usually pointed and sometimes mucronate tail tip, and short spicules. The Iranian populations of D. parvus in this study in comparison to those of Brzeski (1998) have a slightly overlapping basal pharyngeal bulb (vs. offset) and stylet knobs sometimes sloping backwards (vs. rounded). D. parvus is similar to D. emus, D. equalis, D. exilis and D. terricolus. It can be distinguished from D. emus by the lower V value (70.8–75.2 vs. 79–81) and pointed tail tip (vs. rounded), from D. equalis by lower V (70.8–75.2 vs. 77–84) and shorter range of spicule length (12–17 vs. 14–25 μm), and from D. exilis by lower V (70.8–75.2 vs. 77–81) and greater PUS / VBW ratio (1.0–3.6 vs. 0.6–1.0). D. parvus bears the most resemblance to D. terricolus. The differences between these two species are the structure of median and basal pharyngeal bulbs and the shape of the tail tip (Brzeski 1991). The median bulb is more elongated in the second species, although intermediate forms were seen in individuals of some populations in this study. Also, the basal pharyngeal bulb slightly overlaps the intestine in the second species, which was observed in this study’s populations and another Iranian population (Karegar et al. 1995). On the other hand, according to Brzeski (1991), the tail tip in the first species is pointed, and in the second species is variable, with a thick terminus, while in the populations of this study, a wide variety of tail shapes, from rounded to pointed or mucronate, was observed. Since the populations of this study were generally more similar to D. parvus, they are retained under this name until further investigations can be made. It should be noted that two individuals from one population, in the present study have characteristics outside the range of other populations, which include a longer tail (91–102 vs. 53–80 μm), as well as lower PUS/VBW and PUS/V-A ratios (1.0–1.3 and 17.8–23.4 % v s. 1.7–3.6 and 28.7–45.8 %, respectively). In terms of characteristics, these two individuals were more similar to D. equalis, but due to the value of V (70.8–71.6) being similar to D. parvus and the greater stability of this character, they were placed in D. parvus., Published as part of Hashemi, Kobra & Karegar, Akbar, 2019, Description of Ditylenchus paraparvus n. sp. from Iran with an updated list of Ditylenchus Filipjev, 1936 (Nematoda: Anguinidae), pp. 85-113 in Zootaxa 4651 (1) on page 95, DOI: 10.11646/zootaxa.4651.1.6, http://zenodo.org/record/3359134, {"references":["Zell, H. (1988) Nematodes eines Buchenwaldbodens 11. Die Anguiniden (Nematoda, Anguinoidea). Carolinea, 46, 99 - 114.","Brzeski, M. W. (1998) Nematodes of Tylenchina in Poland and temperate Europe. Muzeum i Instytutu Zoologii, Polska Akademia Nauk (MiIZ PAN), Warszawa, 396 pp.","Brzeski, M. W. (1991) Review of the genus Ditylenchus Filipjev, 1936 (Nematoda: Anguinidae). Revue de Nematologie, 14, 9 - 59."]}

Cacopaurus pestis Thorne, 1943 Gharakhani et al. 2007: 4♀: L = 186 (183���190) ��m; a = 11.3 (10.2���12.7); b = 1.6 (1.5���1.7); St = 81.3 (78���89) ��m; V = 90 (88���91) 7♂: L = 246 (227���274) ��m; a = 30.5 (25.7���36.3); b = 3 (2.7���3.5); Spicules = 15.7 (13���17) ��m Associated plants & localities: Poplar from West Azarbaijan (Sturhan 1977), white poplar and walnut from Mazandaran and Zanjan; pine from Mazandaran (Barooti 1992), pear, walnut, hazelnut and poplar from East Azarbaijan and Ardabil (Barooti 1998), junipers from Qazvin (Razzaz Hashemi 2002, Razzaz Hashemi & Kheiri 2003), walnut from central and northern provinces of Iran (Barooti 2006), walnut from Kerman (Gharakhani et al. 2007 [F]), walnut from Kurdistan (Bahmani et al. 2012, 2013), walnut from West Azarbaijan (Ahmadi et al. 2014), Published as part of Ghaderi, Reza, Karegar, Akbar, Miraeiz, Esmaeil & Hesar, Abbas Mokaram, 2019, An updated and annotated checklist of the Tylenchulidae (Nematoda: Criconematoidea) of Iran, pp. 205-229 in Zootaxa 4545 (2) on page 206, DOI: 10.11646/zootaxa.4545.2.3, http://zenodo.org/record/2618787, {"references":["Thorne, G. (1943) Cacopaurus pestis n. g., n. spec. (Nematoda: Criconematidae), a destructive parasite of the walnut, Juglans regia. Linn. Proceedings of the Helminthological Society of Washington, 10, 78 - 83.","Gharakhani, A., Pourjam, E. & Karegar, A. (2007) Some plant parasitic nematodes (Criconematoidea and Longidoridae) in Kerman province orchards. Iranian Journal of Plant Pathology, 43, 372 - 397.","Sturhan, D. (1977) First record of the Persian sessile nematode, Cacopaurus pestis Thorne, in Iran. Nematologia Mediterranea, 5, 125 - 126.","Barooti, S. (1992) Distribution and host range of Cacopaurus pestis from Iran. Nematologica, 38, 399.","Barooti, S. (1998) The plant nematode fauna of cultivated soil of East-Azarbaijan, Ardabil and Moghan. Applied Entomology and Phytopathology, 66, 32 - 35. [79 - 98, in Persian with English summary]","Razzaz Hashemi, S. R. (2002) Identification of important parasitic nematodes of some of the forest trees and shrubs in Qazvin province. Proceedings of 15 th Iranian Plant Protection Congress. Uol. II. Plant diseases and weeds, Kermanshah, Iran, 2002, 263 - 264.","Razzaz Hashemi, S. R. & Kheiri, A. (2003) Identification of nematodes of parasite in Aras forest province of Qazvin. The Journal of Research and Development of Protection of forests and rangelands, 1, 37 - 57. [in Persian]","Barooti, S. (2006) Identification of plant parasitic and predator nematodes fauna of walnut trees in north and central of Iran. Proceedings of 17 th Iranian Plant Protection Congress. Uol. II. Plant diseases and weeds, Karadj, Iran, 2006, 320.","Bahmani, J., Khozeini, F., Barooti, S. & Rezaee, S. (2012) Identification of nematode fauna associated with walnut in Sanandaj. Proceedings of 20 th Iranian Plant Protection Congress. Uol. II. Plant diseases and weeds, Shiraz, Iran, 2012, 747.","Bahmani, J., Khozeini, F., Barooti, S., Rezaee, S. & Ghaderi, R. (2013) Plant-parasitic nematodes associated with walnut in the Sanandej region of west Iran. Journal of Plant Protection Research, 53, 404 - 408. https: // doi. org / 10.2478 / jppr- 2013 - 0060","Ahmadi, B., Niknam, G., Pachide, A. & Kheiri, A. (2014) Report of Cacopaurus pestis and Trophotylenchulus asoensis from West Azarbaijan and Kurdistan provinces, Iran. 21 th Iranian Plant Protection Congress. Uol. II. Plant diseases and weeds, Urmia, Iran, 2014, 318."]}

Paratylenchus variabilis Raski, 1975 Ghaderi et al. 2014: 10♀: L = 323 (306���337) ��m; a = 22.2 (18.7���25.3); b = 3.9 (3.7���4.1); c = 17.9 (15.6���20.5); c' = 2.3 (1.8���2.6); St = 14.1 (13.0���15.2) ��m; V = 84 (83���86) Associated plant and locality: Turfgrass from Kohgiloyeh & Boyer-Ahmad (Ghaderi et al. 2014 [P]), Published as part of Ghaderi, Reza, Karegar, Akbar, Miraeiz, Esmaeil & Hesar, Abbas Mokaram, 2019, An updated and annotated checklist of the Tylenchulidae (Nematoda: Criconematoidea) of Iran, pp. 205-229 in Zootaxa 4545 (2) on page 214, DOI: 10.11646/zootaxa.4545.2.3, http://zenodo.org/record/2618787, {"references":["Ghaderi, R., Kashi Nahanji, L. & Karegar, A. (2014) Contribution to the study of the genus Paratylenchus Micoletzky, 1922 sensu lato (Nematoda: Tylenchulidae). Zootaxa, 3841 (2), 151 - 187. https: // doi. org / 10.11646 / zootaxa. 3841.2.1"]}

Paratylenchus dianthus Jenkins & Taylor, 1956 (Fig. 2)

Paratylenchus obtusicaudatus Raski, 1975 Ghaderi et al. 2014: 10♀: L = 275 (254���295) ��m; a = 21.9 (19.5���23.2); b = 4.0 (3.6���4.3); c = 18.2 (15.5���21.2); c' = 2.2 (1.9���2.5); St = 20.8 (19.3���22.0) ��m; V = 82 (80���83) 2♂: L = 230, 246 ��m; a = 25.0, 25.1; c = 18.8, 20.0; c' = 1.6, 1.9; Spicules = 13.8, 15.0 ��m Associated plant and locality: Sugarcane from Khuzestan (Nasiri et al. 2012, Ghaderi et al. 2014 [P]), Published as part of Ghaderi, Reza, Karegar, Akbar, Miraeiz, Esmaeil & Hesar, Abbas Mokaram, 2019, An updated and annotated checklist of the Tylenchulidae (Nematoda: Criconematoidea) of Iran, pp. 205-229 in Zootaxa 4545 (2) on page 211, DOI: 10.11646/zootaxa.4545.2.3, http://zenodo.org/record/2618787, {"references":["Ghaderi, R., Kashi Nahanji, L. & Karegar, A. (2014) Contribution to the study of the genus Paratylenchus Micoletzky, 1922 sensu lato (Nematoda: Tylenchulidae). Zootaxa, 3841 (2), 151 - 187. https: // doi. org / 10.11646 / zootaxa. 3841.2.1","Nasiri, M., Karegar, A., Taherkhani, K. & Moazen Rezamahaleh, H. (2012) Identification and distribution of the most important plant parasitic nematodes in the sugarcane fields of Khuzestan province, Iran. Abstracts of 20 th Iranian Plant Protection Congress. Uol. II. Plant diseases and weeds, Shiraz, Iran, 2012, 714."]}

Paratylenchus conicephalus van den Berg, Eskandari, Tiedet & Karegar, 2011 van den Berg et al. 2011: 17♀: L = 455 (365���501) ��m; a = 20 (16.���22.8); b = 4.1 (3.5���4.3); c = 12.4 (10.2���18); c' = 4.1 (2.6���5.5); St = 32.5 (29.5���35) ��m; V = 81 (79���83) Ghaderi & Karegar 2013: 11♀: L = 386 (361���409) ��m; a = 23.7 (22.3���24.7); b = 3.9 (3.5���4.0); c = 15.4 (12.2���17.0); c' = 3.2 (2.8���3.7); St = 29.2 (27.8���31.2) ��m; V = 80 (80���82) Associated plants and localities: Prangos ferulacea from Kurdistan (van den Berg et al. 2011 [F]), grapevine from Kurdistan (Ghaderi & Karegar 2013 [P], Ghaderi et al. 2012, 2014 [P]), Published as part of Ghaderi, Reza, Karegar, Akbar, Miraeiz, Esmaeil & Hesar, Abbas Mokaram, 2019, An updated and annotated checklist of the Tylenchulidae (Nematoda: Criconematoidea) of Iran, pp. 205-229 in Zootaxa 4545 (2) on page 208, DOI: 10.11646/zootaxa.4545.2.3, http://zenodo.org/record/2618787, {"references":["van den Berg, E., Eskandari, A., Tiedt, L. R. & Karegar, A. (2011) Paratylenchus conicephalus sp. n. from Iran with notes on P. similis Khan, Prasad & Mathur, 1967 (Nematoda: Paratylenchidae). Nematology, 13, 529 - 537. https: // doi. org / 10.1163 / 138855410 X 528280","Ghaderi, R. & Karegar, A. (2013) Some species of Paratylenchus (Nematoda: Tylenchulidae) from Iran. Iranian Journal of Plant Pathology, 49, 49 - 52. [in Persian with English summary]","Ghaderi, R., Kashi Nahanji, L. & Karegar, A. (2012) Some species of the superfamily Criconematoidea (Nematoda: Tylenchomorpha) from Iran. Proceedings of 20 th Iranian Plant Protection Congress. Uol. II. Plant diseases and weeds, Shiraz, Iran, 2012, 731."]}

Paratylenchus colinus (Huang & Raski, 1986) Brzeski, 1998 Syn: Gracilacus colina Huang & Raski, 1986 Syn: Gracilpaurus colinus (Huang & Raski, 1986) Ganguly & Khan, 1990, Published as part of Ghaderi, Reza, Karegar, Akbar, Miraeiz, Esmaeil & Hesar, Abbas Mokaram, 2019, An updated and annotated checklist of the Tylenchulidae (Nematoda: Criconematoidea) of Iran, pp. 205-229 in Zootaxa 4545 (2) on page 208, DOI: 10.11646/zootaxa.4545.2.3, http://zenodo.org/record/2618787, {"references":["Huang, C. S. & Raski, D. J. (1986) Four new species of Gracilacus Raski, 1962 (Criconematoidea: Nemata). Revue de Nematologie, 9, 347 - 356.","Brzeski, M. W. (1998) Nematodes of Tylenchina in Poland and temperate Europe. Muzeum I Instytut Zoologii PAN, Warszawa, 397 pp."]}

Paratylenchus Micoletzky, 1922 Syn: Gracilacus Raski, 1962, Published as part of Ghaderi, Reza, Karegar, Akbar, Miraeiz, Esmaeil & Hesar, Abbas Mokaram, 2019, An updated and annotated checklist of the Tylenchulidae (Nematoda: Criconematoidea) of Iran, pp. 205-229 in Zootaxa 4545 (2) on page 207, DOI: 10.11646/zootaxa.4545.2.3, http://zenodo.org/record/2618787, {"references":["Micoletzky, H. (1922) Die Freilebenden Erd-Nematoden. Archiv fur Naturgeschichte, Berlin, A, 87 (1921), 1 - 650.","Raski, D. J. (1962) Paratylenchidae n. fam. with descriptions of five new species of Gracilacus n. g. and an emendation of Cacopaurus Thorne, 1943, Paratylenchus Micoletzky, 1922 and Criconematidae Thorne, 1943. Proceedings of the Helminthological Society of Washington, 29, 189 - 207."]}

Tylenchulus musicola Tanha Maafi, Amani, Stanley, Inserra, Van den Berg & Subbotin, 2012 Tanha Maafi et al. 2012: 12 Mature ♀: L = 287 (240–370) µm; a = 4.8 (3.7–8.2); b = 3.1 (2.5–4.0); St = 10 (8.0–12)µm 31 J 2: L = 306 (278–334) µm; a = 30.1 (27.7–33.4); b = 3.5 (3.2–4.0); St = 11.1 (10–12)µm 5♂: L = 310 (286–326) µm; a = 27.2 (23.8–32.5); b = 3.5 (3.3–3.7); St = 8.5 (8–9) µm; Spicules = 15.7 (14– 18) µm Associated plant and locality: Banana from Sistan & Baluchistan (Tanha Maafi et al. 2010 [M], 2012 [F, M])

Paratylenchus holdemani Raski, 1975 Mirghasemi et al. 2016: 12♀: L = 285 (243-305) ��m; a = 21.5 (18.4-25.4); b = 3.9 (3.6-4.3); c = 13.1 (10.4-15.9); c' = 2.6 (2.5-3.1); St = 22.7 (22-24) ��m; V = 82 (81-84) 2♂: L = 311-317 ��m; a = 26.5-28.3; b = 3.7-4.2; c = 11.7-14.4; c' = 2.4-3.3; Spicules = 16-17 ��m Associated plant and localities: Tea from Gilan (Mirghasemi et al. 2014a, 2016 [F]), Published as part of Ghaderi, Reza, Karegar, Akbar, Miraeiz, Esmaeil & Hesar, Abbas Mokaram, 2019, An updated and annotated checklist of the Tylenchulidae (Nematoda: Criconematoidea) of Iran, pp. 205-229 in Zootaxa 4545 (2) on page 209, DOI: 10.11646/zootaxa.4545.2.3, http://zenodo.org/record/2618787, {"references":["Mirghasemi, S. N., Jamali, S., Seraji, A. & Pedramfar, H. (2016) Some plant parasitic nematodes of the infraorder Tylenchomorpha from tea plantations of Iran. Journal of Plant Protection, 30, 22 - 30.","Mirghasemi, S. N., Seraji, A. & Jamali, S. (2014 a) Identification of plant parasitic nematodes of tea populations (Camellia sinensis) in Iran. Proceedings of 21 th Iranian Plant Protection Congress. Uol. II. Plant diseases and weeds, Shiraz, Iran, 2014, 311."]}

Paratylenchus peraticus (Raski, 1962) Siddiqi & Goodey, 1964 (Fig. 4) Syn: Gracilacus peratica Raski, 1962

Paratylenchus idalimus (Raski, 1962) Siddiqi & Goodey, 1964 Syn: Gracilacus idalima Raski, 1962 Associated plants and localities: Grapevine from West Azarbaijan (Barooti 1987), bean from Markazi (Mohammad Deimi et al. 2008b)

Paratylenchus idalimus (Raski, 1962) Siddiqi & Goodey, 1964 Syn: Gracilacus idalima Raski, 1962 Associated plants and localities: Grapevine from West Azarbaijan (Barooti 1987), bean from Markazi (Mohammad Deimi et al. 2008b), Published as part of Ghaderi, Reza, Karegar, Akbar, Miraeiz, Esmaeil & Hesar, Abbas Mokaram, 2019, An updated and annotated checklist of the Tylenchulidae (Nematoda: Criconematoidea) of Iran, pp. 205-229 in Zootaxa 4545 (2) on page 210, DOI: 10.11646/zootaxa.4545.2.3, http://zenodo.org/record/2618787, {"references":["Raski, D. J. (1962) Paratylenchidae n. fam. with descriptions of five new species of Gracilacus n. g. and an emendation of Cacopaurus Thorne, 1943, Paratylenchus Micoletzky, 1922 and Criconematidae Thorne, 1943. Proceedings of the Helminthological Society of Washington, 29, 189 - 207.","Siddiqi, M. R. & Goodey, J. B. (1964) The status of the genera and subfamilies of the Criconematidae (Nematoda); with a comment on the position of Fergusobia. Nematologica, 9 (1963), 363 - 377.","Barooti, S. (1987) A list of plant parasitic nematodes of Iran (collected between 1956 - 1986). Ministry of Agriculture, Tehran, 39 pp.","Mohammad Deimi, A., Ghalandar, M. & Barooti, S. (2008 b) Study of plant parasitic nematodes on bean in Khomein. Proceedings of 18 th Iranian Plant Protection Congress. Uol. II. Plant diseases and weeds, Hamadan, Iran, 2008, 559."]}

Paratylenchus salubris Raski, 1975 (Fig. 6), Published as part of Ghaderi, Reza, Karegar, Akbar, Miraeiz, Esmaeil & Hesar, Abbas Mokaram, 2019, An updated and annotated checklist of the Tylenchulidae (Nematoda: Criconematoidea) of Iran, pp. 205-229 in Zootaxa 4545 (2) on page 213, DOI: 10.11646/zootaxa.4545.2.3, http://zenodo.org/record/2618787

Paratylenchus neoprojectus Wu & Hawn, 1975 Ghaderi & Karegar 2013: 15♀: L = 387 (344���416) ��m; a = 22.3 (18.8���24.7); b = 4.0 (3.7���4.3); c = 15.9 (13.9���18.1); c' = 2.9 (2.7���3.2); St = 27.4 (24.9���29.3) ��m; V = 83 (81���86) Associated plants and localities: Apricot from Kurdistan and shade trees from East Azarbaijan (Ghaderi & Karegar 2013 [F], Ghaderi et al. 2012, 2014 [P]), alfalfa from Yazd (Ghaderi et al. 2014 [P]), Published as part of Ghaderi, Reza, Karegar, Akbar, Miraeiz, Esmaeil & Hesar, Abbas Mokaram, 2019, An updated and annotated checklist of the Tylenchulidae (Nematoda: Criconematoidea) of Iran, pp. 205-229 in Zootaxa 4545 (2) on page 211, DOI: 10.11646/zootaxa.4545.2.3, http://zenodo.org/record/2618787, {"references":["Wu, L. Y. & Hawn, E. J. (1975) Paratylenchus neoprojectus n. sp. (Paratylenchinae: Nematoda) from alfalfa fields in Alberta, Canada. Canadian Journal of Zoology, 53, 1841 - 1843. https: // doi. org / 10.1139 / z 75 - 217","Ghaderi, R. & Karegar, A. (2013) Some species of Paratylenchus (Nematoda: Tylenchulidae) from Iran. Iranian Journal of Plant Pathology, 49, 49 - 52. [in Persian with English summary]","Ghaderi, R., Kashi Nahanji, L. & Karegar, A. (2012) Some species of the superfamily Criconematoidea (Nematoda: Tylenchomorpha) from Iran. Proceedings of 20 th Iranian Plant Protection Congress. Uol. II. Plant diseases and weeds, Shiraz, Iran, 2012, 731.","Ghaderi, R., Kashi Nahanji, L. & Karegar, A. (2014) Contribution to the study of the genus Paratylenchus Micoletzky, 1922 sensu lato (Nematoda: Tylenchulidae). Zootaxa, 3841 (2), 151 - 187. https: // doi. org / 10.11646 / zootaxa. 3841.2.1"]}

Paratylenchus similis Khan, Prasad & Mathur, 1967 (Fig. 7) Syn: Paratylenchus italiensis Raski, 1975 Syn: Paratylenchus tateae Wu & Townshend, 1973, Published as part of Ghaderi, Reza, Karegar, Akbar, Miraeiz, Esmaeil & Hesar, Abbas Mokaram, 2019, An updated and annotated checklist of the Tylenchulidae (Nematoda: Criconematoidea) of Iran, pp. 205-229 in Zootaxa 4545 (2) on page 213, DOI: 10.11646/zootaxa.4545.2.3, http://zenodo.org/record/2618787, {"references":["Khan, E., Prasad, S. K. & Mathur, V. K. (1967) Two new species of the genus Paratylenchus Micoletzky, 1922 (Nematoda: Criconematidae) from India. Nematologica, 13, 79 - 84. https: // doi. org / 10.1163 / 187529267 X 00959","Wu, Y. L. & Townshend, J. L. (1973) Paratylenchus tateae n. sp. (Paratylenchinae, Nematoda). Canadian Journal of Zoology, 51, 109 - 111. https: // doi. org / 10.1139 / z 73 - 017"]}

Paratylenchus arculatus Luc & de Guiran, 1962 Syn: Paratylenchus nainianus Edward & Misra, 1963, Published as part of Ghaderi, Reza, Karegar, Akbar, Miraeiz, Esmaeil & Hesar, Abbas Mokaram, 2019, An updated and annotated checklist of the Tylenchulidae (Nematoda: Criconematoidea) of Iran, pp. 205-229 in Zootaxa 4545 (2) on page 207, DOI: 10.11646/zootaxa.4545.2.3, http://zenodo.org/record/2618787, {"references":["Luc, M. & De Guiran, G. (1962) Deux nouveaux Paratylenchus (Nematoda-Criconematidae) de Cote d'ivoire. Nematologica, 7, 133 - 138. https: // doi. org / 10.1163 / 187529262 X 00864","Edward, J. C. & Misra, S. L. (1963) Paratylenchus nainianus n. sp. (Nematoda: Criconematidae) from Uttar Pradesh, India. Nematologica, 9, 215 - 217. https: // doi. org / 10.1163 / 187529263 X 00386"]}

Paratylenchus nanus Cobb, 1923 Nouri et al. 2006: 10♀: L = 401 (363���430) ��m; a = 23.4 (18.6���26.5); b = 4.2 (4.0���4.3); c = 14.2 (12.9���16); c' = 3.0 (2.7���3.6); St = 31.8 (29���34) ��m; V = 81 (78���83) Associated plants and localities: Apricot from Zanjan (Nouri et al. 2004, 2006 [F]), walnut from central and northern regions of Iran (Barooti 2006), peach, nectarine, cherry and hazelnut from Ardabil (Rafiee et al. 2010), citrus from Mazandaran (Divsalar et al. 2011), fruit trees from East Azarbaijan (Solouki et al. 2012), peanut from Gilan (Mirghasemi et al. 2014b), grapevine from Kermanshah (Esmaeili & Heydari 2017), Published as part of Ghaderi, Reza, Karegar, Akbar, Miraeiz, Esmaeil & Hesar, Abbas Mokaram, 2019, An updated and annotated checklist of the Tylenchulidae (Nematoda: Criconematoidea) of Iran, pp. 205-229 in Zootaxa 4545 (2) on page 211, DOI: 10.11646/zootaxa.4545.2.3, http://zenodo.org/record/2618787, {"references":["Cobb, N. A. (1923) Notes on Paratylenchus, a genus of nemas. Journal of the Washington Academy and Sciences, 13, 254 - 257.","Nouri, S., Kheiri, A. & Karegar, A. (2006) Some plant parasitic nematodes from orchards in Zanjan region. Iranian Journal of Plant Pathology, 42, 1 - 4. [1 - 17, in Persian with English summary]","Nouri, S., Kheiri, A. & Karegar, A. (2004) Identification of plant parasitic nematodes fauna (Tylenchida) of fruit trees in Zanjan region. Abstracts of 16 th Iranian Plant Protection Congress. Uol. II. Plant diseases and weeds, Tabriz, Iran, 2004, 434.","Barooti, S. (2006) Identification of plant parasitic and predator nematodes fauna of walnut trees in north and central of Iran. Proceedings of 17 th Iranian Plant Protection Congress. Uol. II. Plant diseases and weeds, Karadj, Iran, 2006, 320.","Rafiee, S., Pourjam, E. & Niknam, G. (2010) Some nematode species associated with fruit trees in Moghan region. Journal of Plant Protection, 24, 69 - 79. [in Persian]","Divsalar, N., Jamali, S., Pedramfar, H. & Taheri, H. (2011) Identification of plant-parasitic nematodes associated with the rhizosphere of citrus in east Gilan and west Mazandaran. Journal of Plant Protection, 25, 168 - 177. [in Persian]","Solouki, V., Niknam, G. & Kheiri, A. (2012) Identification of plant parasitic nematodes in orchard rhizosphere of Marand and suburbs in East Azarbaijan province, Iran. Proceedings of 20 th Iranian Plant Protection Congress. Uol. II. Plant diseases and weeds, Shiraz, Iran, 2012, 692.","Mirghasemi, S. N., Jamali, S. & Sharifi, K. (2014 b) Identification of plant parasitic nematodes of peanut (Arachis hypogaea) fields in Guilan province. Proceedings of 21 th Iranian Plant Protection Congress. Uol. II. Plant diseases and weeds, Urmia, Iran, 2014, 315.","Esmaeili, M. & Heydari, R. (2017) Some species of Paratylenchus Micoletzky, 1922 (Nematoda: Tylenchulidae) from vineyards in Kermanshah province, western Iran. Journal of Crop Protection, 6, 259 - 271."]}

Paratylenchus labiosus Anderson & Kimpinski, 1977 Bahmani et al. 2014: 8♀: L = 333 (309���363) ��m; a = 22.8 (19.9���25.0); b = 3.5 (3.1���3.9); c = 12.2 (9.9���14.9); c' = 3.0 (2.7���2.4); St = 16.7 (15.5���17.5) ��m; V = 81 (79���83) 1♂: L = 362 ��m; a = 29.2; c = 11.2; c' = 3.5; Spicules = 20.6 ��m Associated plant and locality. Walnut from Kurdistan (Bahmani et al. 2013, 2014 [F]), Published as part of Ghaderi, Reza, Karegar, Akbar, Miraeiz, Esmaeil & Hesar, Abbas Mokaram, 2019, An updated and annotated checklist of the Tylenchulidae (Nematoda: Criconematoidea) of Iran, pp. 205-229 in Zootaxa 4545 (2) on page 210, DOI: 10.11646/zootaxa.4545.2.3, http://zenodo.org/record/2618787, {"references":["Anderson, R. V. & Kimpinski, J. (1977) Paratylenchus labiosus n. sp. (Nematoda: Paratylenchidae) from Canada. Canadian Journal of Zoology, 55, 1992 - 1996. https: // doi. org / 10.1139 / z 77 - 258","Bahmani, J., Barooti, S. & Ghaderi, R. (2014) On occurrence of Paratylenchus labiosus Anderson & Kimpinski, 1977 (Nematoda: Tylenchulidae) in Iran, with discussion on the validity of the species. Journal of Crop Protection, 3, 273 - 281.","Bahmani, J., Khozeini, F., Barooti, S., Rezaee, S. & Ghaderi, R. (2013) Plant-parasitic nematodes associated with walnut in the Sanandej region of west Iran. Journal of Plant Protection Research, 53, 404 - 408. https: // doi. org / 10.2478 / jppr- 2013 - 0060"]}

Paratylenchus perminimus Siddiqi, 1996 (Fig. 5), Published as part of Ghaderi, Reza, Karegar, Akbar, Miraeiz, Esmaeil & Hesar, Abbas Mokaram, 2019, An updated and annotated checklist of the Tylenchulidae (Nematoda: Criconematoidea) of Iran, pp. 205-229 in Zootaxa 4545 (2) on page 212, DOI: 10.11646/zootaxa.4545.2.3, http://zenodo.org/record/2618787, {"references":["Siddiqi, M. R. (1996) Paratylenchus recisus sp. n. and P. perminimus sp. n. (Criconematina: Paratylenchidae). Afro-Asian Journal of Nematology, 6, 55 - 58."]}

Trophotylenchulus asoensis (Minagawa, 1983) Siddiqi, 1999 Gharakhani et al. 2007: 20♀: L = 435 (316���552) ��m; a = 11.1 (8.9���14.7); b = 3.8 (2.8���4.9); c = 12.9 (10.2���15.6); c' = 3.2 (2.6���4.2); St = 13.2 (11���15) ��m; V = 77 (73���82) 5♂: L = 490 (453���518) ��m; a = 34 (32.4���36); b = 4.3 (3.9���4.9); c = 10 (9.2���10.6); St = 9.2 (8���10.5) ��m; Spicules = 20 (18���22) ��m Associated plants and localities: Plane tree from Kerman (Gharakhani et al. 2007 [F], grasses from Kurdistan (Ahmadi et al. 2014), Published as part of Ghaderi, Reza, Karegar, Akbar, Miraeiz, Esmaeil & Hesar, Abbas Mokaram, 2019, An updated and annotated checklist of the Tylenchulidae (Nematoda: Criconematoidea) of Iran, pp. 205-229 in Zootaxa 4545 (2) on page 222, DOI: 10.11646/zootaxa.4545.2.3, http://zenodo.org/record/2618787, {"references":["Minagawa, N. (1983) Descriptions of two new species of nematode genus Trophonema Raski, 1957 (Tylenchida: Tylenchulidae). Applied Entomology and Zoology, 18, 90 - 97. https: // doi. org / 10.1303 / aez. 18.90","Siddiqi, M. R. (1999) Trophotylenchulus colbrani sp. n. and T. cunctus sp. n. (Criconematina: Tylenchulidae) from Mbalmayo forest in Cameroon. International Journal of Nematology, 8 (1998), 179 - 184.","Gharakhani, A., Pourjam, E. & Karegar, A. (2007) Some plant parasitic nematodes (Criconematoidea and Longidoridae) in Kerman province orchards. Iranian Journal of Plant Pathology, 43, 372 - 397.","Ahmadi, B., Niknam, G., Pachide, A. & Kheiri, A. (2014) Report of Cacopaurus pestis and Trophotylenchulus asoensis from West Azarbaijan and Kurdistan provinces, Iran. 21 th Iranian Plant Protection Congress. Uol. II. Plant diseases and weeds, Urmia, Iran, 2014, 318."]}