taxonID	type	description	language	source
03AFAC09DD30AA5DFE14FF1AFD38FCC8.taxon	description	(Figures 1 – 3; Tables 1, 2)	en	Vazifeh, Nasir, Niknam, Gholamreza, Jabbari, Habibeh, Fallahi, Amin, Zahedi, Ebrahim, Peña-Santiago, Reyes (2022): Description of Willinema persicum sp. n. (Dorylaimida: Thornenematidae) from Iran, with an updated taxonomy of the genus. Journal of Natural History 56 (13 - 16): 849-863, DOI: 10.1080/00222933.2022.2082895, URL: http://dx.doi.org/10.1080/00222933.2022.2082895
03AFAC09DD30AA5DFE14FF1AFD38FCC8.taxon	diagnosis	Diagnosis The new species is characterised by its 1.42 – 1.89 mm long body, lip region offset by weak constriction and 10 – 13 μm broad, labial and post-labial sclerotisation absent, odontostyle 14 – 18 μm long, neck 270 – 306 μm long, pharyngeal expansion 110 – 137 μm long and occupying 40 – 42 % of total neck length, female genital system mono-opisthodelphic, anterior genital branch reduced to a uterine sac 41 – 133 μm long or 0.8 – 2.9 times the corresponding body diameter, posterior uterus tripartite, vulva a pre-equatorial (V = 39 – 44) transverse slit, tail short and convex-conoid to rounded (19 – 27 μm, c = 61 – 84, c ′ = 0.7 – 1.0), spicules 36 – 41 μm long, and 7 – 9 ventromedian supplements with hiatus.	en	Vazifeh, Nasir, Niknam, Gholamreza, Jabbari, Habibeh, Fallahi, Amin, Zahedi, Ebrahim, Peña-Santiago, Reyes (2022): Description of Willinema persicum sp. n. (Dorylaimida: Thornenematidae) from Iran, with an updated taxonomy of the genus. Journal of Natural History 56 (13 - 16): 849-863, DOI: 10.1080/00222933.2022.2082895, URL: http://dx.doi.org/10.1080/00222933.2022.2082895
03AFAC09DD30AA5DFE14FF1AFD38FCC8.taxon	etymology	Etymology The specific epithet refers to Persia, the former name of Iran. Type locality and habitat North-west Iran, East Azarbaijan province, Sufiyan, Roodghat area (GPS (Global Positioning System) coordinates: 38.256 ° N, 46.159 ° E, altitude 1547 m a. s. l.), in the rhizosphere of common wheat (Triticum aestivum L.). Other locality and habitat North-west Iran, East Azarbaijan province, Arasbaran forests.	en	Vazifeh, Nasir, Niknam, Gholamreza, Jabbari, Habibeh, Fallahi, Amin, Zahedi, Ebrahim, Peña-Santiago, Reyes (2022): Description of Willinema persicum sp. n. (Dorylaimida: Thornenematidae) from Iran, with an updated taxonomy of the genus. Journal of Natural History 56 (13 - 16): 849-863, DOI: 10.1080/00222933.2022.2082895, URL: http://dx.doi.org/10.1080/00222933.2022.2082895
03AFAC09DD30AA5DFE14FF1AFD38FCC8.taxon	materials_examined	Material examined 9 ♀♀ and 6 ♂♂ from two locations and habitats, in acceptable condition. Type material ♀ holotype, ♀♀ and ♂♂ paratypes deposited in the Nematology Collection of the Department of Plant Protection, Faculty of Agriculture, University of Tabriz, Tabriz, Iran, and one paratype ♀ (catalogue number = 682) and one paratype ♂ (catalogue number = 683) deposited in the Nematode Collection of the University of Jaén, Spain.	en	Vazifeh, Nasir, Niknam, Gholamreza, Jabbari, Habibeh, Fallahi, Amin, Zahedi, Ebrahim, Peña-Santiago, Reyes (2022): Description of Willinema persicum sp. n. (Dorylaimida: Thornenematidae) from Iran, with an updated taxonomy of the genus. Journal of Natural History 56 (13 - 16): 849-863, DOI: 10.1080/00222933.2022.2082895, URL: http://dx.doi.org/10.1080/00222933.2022.2082895
03AFAC09DD30AA5DFE14FF1AFD38FCC8.taxon	description	Description Adult. Moderately slender nematodes (a = 30 – 39) of medium size, 1.42 – 1.89 mm. Body mostly curved ventrad upon fixation, especially in posterior region, C- or G-shaped. Body cylindrical, visibly tapering towards the anterior end, less so towards the posterior end since the caudal region is short and rounded. Cuticle two-layered, 1.5 – 2.5 μm thick in anterior region, 2 – 4 μm at mid-body and 5 – 7 μm on tail, outer layer distinctly thinner than the inner one and bearing fine transverse striation. Lateral chord very narrow, 3 – 6 μm wide or 6 – 15 % of the corresponding body diameter. Lip region rounded, offset by a weak but perceptible constriction, 2.0 – 2.5 times as broad as high or one-fifth to one-third (21 – 34 %) of body diameter at neck base; lips rather amalgamate, with low papillae; labial and post-labial sclerotisation absent. Amphidial fovea stirrup-shaped, their aperture 5.5 – 7 μm wide, occupying 52 – 64 % lip region diameter. Cheilostom a truncate cone, lacking any differentiation. Odontostyle dorylaimoid, typical of the genus, 1.2 – 1.5 times the lip region diameter and 1.0 – 1.1 % of body length, with aperture 4.5 – 6 μm long or occupying 30 – 38 % of its length. Guiding ring simple, located at 7 – 10 μm from the anterior end or 0.6 – 0.8 times the lip region diameter. Odontophore well demarcated, simple, rod-like, 1.1 – 1.2 times the odontostyle length. Nerve ring encircling the anterior slender part of pharynx, at 113 – 125 μm or 36 – 46 % of neck length from anterior end. Anterior region of pharynx enlarging gradually, basal expansion 4.8 – 6.5 times as long as wide, 2.5 – 3.2 times as long as body diameter at neck base and occupying 40 – 42 % of total neck length; gland nuclei located as follows: DN = 63 – 65, S 1 N 1 = 69 – 73, S 1 N 2 = 76 – 79, S 2 N = 83 – 86. Cardia conical, 15 – 22 × 7 – 11 μm, with a ring-like structure surrounding its junction to pharyngeal base. Female. Reproductive system mono-opistho-ovarian. Anterior branch reduced to a simple uterine sac 41 – 133 μm long or 0.8 – 2.9 times the corresponding body diameter, empty, occupying 2.2 – 9.3 % of body length. Posterior branch well developed, very variable in length, 278 – 407 μm or 15 – 28 % of body length: ovary reflexed, not reaching the oviduct-uterus junction, measuring 50 – 75 μm, with oocytes arranged first in several rows and then in a single row; oviduct joining the ovary subterminally, 120 – 145 μm long, 2.4 – 3.3 body diameter, and consisting of a tubular part and a poorly developed pars dilatata with distinct lumen; oviduct and uterus separated by a marked sphincter; uterus very long and tripartite, 170 – 236 μm or 3.4 – 5.4 body diameters, consisting of a proximal region with visible lumen, an intermediate muscular section with narrow lumen, and nearly spherical pars dilatata close to sphincter; sperm cells present in proximal uterine region. Vagina perpendicular to body axis, extending inwards 18 – 22 μm, up to one-half (36 – 51 %) of body diameter: pars proximalis 11 – 15 × 11 – 14 μm, encircled by circular muscles, pars refringens consisting of two sclerotised triangular to drop-shaped pieces, each measuring 3.3 – 4.5 × 4.5 μm, and very short pars distalis. Vulva a pre-equatorial transverse slit. Prerectum 2.6 – 3.9, rectum 0.9 – 1.5 anal body diameters long. Caudal region short, convex conoid to rounded, with two pairs of caudal pores. Male. Prerectum 4.1 – 5.0, cloaca 1.5 – 1.7 times the body diameter at level of cloacal aperture. Genital system diorchic, with opposed testes. In addition to the ad-cloacal pair, situated at 7 – 8 μm from the cloacal aperture, there is a series of 7 – 9 closely and irregularly spaced (9 – 14 μm apart) ventromedian supplements located outside the range of spicules. Spicules dorylaimoid, robust, sclerotised, 3.6 – 5.1 times as long as wide and 1.2 – 1.7 times the cloacal body diameter; head small, not offset, 3 – 5 μm long, occupying 7 – 14 % of total spicule length, with its dorsal side slightly longer than the ventral one and slightly curved, ventral hump and hollow poorly demarcated but appreciable; median piece 24 – 26 μm long, occupying 48 – 50 % of spicule length, and reaching the spicule posterior terminus. Lateral guiding piece small, 3 – 6 μm long. Caudal region similar to that of female. Juveniles were not found. Relationships In having comparatively large general size (body 1.42 – 1.89 mm long, neck more than 270 µm long), short tail (c = 61 – 84) and 7 – 9 ventromedian supplements, the new species is easily distinguishable from other members of the genus. It resembles W. eburnense and E. paraparvum, two African (Ivory Coast and Cameroon, respectively) species, in the presence of a reduced (but appreciably long, more than 40 µm) anterior genital branch, differing from these two species in its longer odontostyle (14 – 18 vs up to 14 µm), and more anterior vulva (V = 39 – 44 vs V = 48 – 51 and V = 44 – 48, respectively). There are also differences in c values in both males (c = 61 – 80 vs 38 – 40 and 48 – 57, respectively) and females (c = 62 – 84 vs 38 – 44 and 48 – 57, respectively). Willinema persicum sp. nov. differs from W. eburnense mainly in longer pharyngeal basal expansion (115 - 127 vs 90 – 98 µm and 110 – 137 vs 92 – 93 µm, respectively) in the females and males, greater body width at mid-body (43 – 50 vs 34 – 37 µm, 41 – 45 vs 29 – 34 µm, respectively) in both females and males, and relatively longer spicule (36 – 41 vs 29 – 35 µm) and prerectum (64 – 95 vs 43 – 49 µm, 103 – 115 vs 45 – 61 µm, respectively) in females and males. The new species can be differentiated from W. paraparvum by higher odontostyle length in females and males (15 – 17 vs 12 – 12.5 µm, 14 – 18 vs 12.5 µm, respectively), prerectum length (64 – 95 vs 46 – 51 µm, 103 – 115 vs 74 – 87 µm, respectively), body width at neck base (38 – 45 vs 28 – 31 µm, 37 – 45 vs 33 – 35 µm, respectively) and mid-body (43 – 50 vs 29 – 32 µm, 41 – 45 vs 34.5 – 36.5 µm, respectively).	en	Vazifeh, Nasir, Niknam, Gholamreza, Jabbari, Habibeh, Fallahi, Amin, Zahedi, Ebrahim, Peña-Santiago, Reyes (2022): Description of Willinema persicum sp. n. (Dorylaimida: Thornenematidae) from Iran, with an updated taxonomy of the genus. Journal of Natural History 56 (13 - 16): 849-863, DOI: 10.1080/00222933.2022.2082895, URL: http://dx.doi.org/10.1080/00222933.2022.2082895
03AFAC09DD3BAA58FF58F81FFD69FA96.taxon	diagnosis	Diagnosis (after Carbonell and Coomans, emended) Thornenematidae, Willinematinae. Small- to medium sized nematodes, 0.38 – 1.89 mm long. Lip region continuous or offset by depression or constriction, lacking any sclerotisation. Odontostyle typical dorylaimid. Guiding ring simple. Odontophore rod-like, with no specialisation. Pharynx enlarging gradually, the basal expansion occupying one-third to one-half of the total neck length. Female genital system mono-opistho-ovarian, often didelphic as the anterior branch appears variably developed, sometimes nearly complete but reduced and not functional, vagina with distinct pars refringens, and transverse vulva. Tail similar in the two sexes, short and rounded; convex-conoid to cylindrical. Male with dorylaimid spicules, and 2 – 9 widely spaced ventromedian supplements with hiatus.	en	Vazifeh, Nasir, Niknam, Gholamreza, Jabbari, Habibeh, Fallahi, Amin, Zahedi, Ebrahim, Peña-Santiago, Reyes (2022): Description of Willinema persicum sp. n. (Dorylaimida: Thornenematidae) from Iran, with an updated taxonomy of the genus. Journal of Natural History 56 (13 - 16): 849-863, DOI: 10.1080/00222933.2022.2082895, URL: http://dx.doi.org/10.1080/00222933.2022.2082895
03AFAC09DD3BAA58FF58F81FFD69FA96.taxon	type_taxon	Type species W. parvum (Williams, 1959) Baqri and Jairajpuri 1967	en	Vazifeh, Nasir, Niknam, Gholamreza, Jabbari, Habibeh, Fallahi, Amin, Zahedi, Ebrahim, Peña-Santiago, Reyes (2022): Description of Willinema persicum sp. n. (Dorylaimida: Thornenematidae) from Iran, with an updated taxonomy of the genus. Journal of Natural History 56 (13 - 16): 849-863, DOI: 10.1080/00222933.2022.2082895, URL: http://dx.doi.org/10.1080/00222933.2022.2082895
03AFAC09DD3BAA58FF58F81FFD69FA96.taxon	discussion	Other valid species W. brunetti (Meyl, 1953) Andrássy 1986 = Dorylaimus brunetti Meyl, 1953 = Eudorylaimus brunetti (Meyl, 1953) Andrassy, 1959 W. chilense sp. nov. = Eudorylaimus brunettii apud Andrássy (1967), nec Meyl (1953) W. eburnense Carbonell and Coomans, 1984 W. laopagense Carbonell and Coomans, 1984 W. nanum Carbonell and Coomans, 1984 W. opisthodelphus (Thorne and Swanger, 1936) Andrássy 1986 = Dorylaimus opisthodelphus Thorne and Swanger, 1936 = Eudorylaimus opisthodelphus (Thorne and Swanger, 1936) Andrássy 1959 W. paraparvum Carbonell and Coomans, 1984 W. persicum sp. nov. Species inquirenda W. sulphasae (Tulaganov, 1949) Andrássy 1986 = Dorylaimus sulphasae Tulaganov, 1949 = Eudorylaimus sulphasae (Tulaganov, 1949) Andrássy 1959 Key to species identification	en	Vazifeh, Nasir, Niknam, Gholamreza, Jabbari, Habibeh, Fallahi, Amin, Zahedi, Ebrahim, Peña-Santiago, Reyes (2022): Description of Willinema persicum sp. n. (Dorylaimida: Thornenematidae) from Iran, with an updated taxonomy of the genus. Journal of Natural History 56 (13 - 16): 849-863, DOI: 10.1080/00222933.2022.2082895, URL: http://dx.doi.org/10.1080/00222933.2022.2082895
03AFAC09DD3BAA58FF58F81FFD69FA96.taxon	discussion	Table 2 provides a compendium of the main morphometrics of Willinema species. Comments on some species E. brunettiae: A species repeatedly recorded in Italy (Meyl 1953, 1954), Ivory Coast (Andrássy 1956; Jacobs 1984), Hungary (Andrássy 1958), Venezuela (Loof 1964; Dao 1970), Chile (Andrássy 1967), and Moldova (Bushmakiu et al. 2000; Poiras 2008). Its original description was quite simple, but Andrássy (1956) provided some addditional information and better illustrations that fitted those of the type material (see Table 2). Nevertheless, Andrássy (1967) studied a Chilean population that significantly differed from the previous ones in several remarkable features: larger general size (body 0.62 – 0.70 vs 0.38 – 0.39 mm long in original and other populations), more slender body (a = 33 – 35 vs a = 22 – 30), and much more posterior vulva (V = 52 – 54 vs V = 39 – 45). This material is also easily distinguishable from the other small species of the genus, W. nanum, with shorter body (0.45 – 0.52 mm) too, as this species bears a distinct, 25 – 45 µm long, anterior uterine sac that is nearly absent in Chilean specimens. Taking into consideration that these specimens were well characterised morphologically and are sharply separable from their relatives, they are regarded as belonging to a different species, herein named W. chilense sp. nov. E. opisthodelphus: Only known to occur in its type locality in the USA, Utha, where Thorne and Swanger (1936) found it associated with sod in a mountain summit. The original description is poor in detail, but sufficient to distinguish it from other members of the genus. It is provisionally retained as a valid species. Popovici (1968) studied several Romanian females identified as Eudorylaimus opisthodelphus, but the true identity of this material cannot be confirmed, although it is certainly not conspecific with the American ones as their general size is much larger (body 1.9 – 2.06 vs 1.3 mm, respectively) and the vulva much more posterior (V = 49 – 50.5 vs V = 42). E. sulphasae: The true identity of this species remain obscure. Its original (and only available) description by Tulaganov (1949) is very poor in detail and raises reasonable doubts about the nature of some relevant traits. Thus, the female genital system was originally described as being diovarian, with a very anterior vulva position (V = 35), an exceptional combination in dorylaimid taxa where a very anterior vulva is always associated with the reduction of the anterior genital branch. Actually, Andrássy (1987, p. 306) was aware of this apparent inconsistency and stated that the species should be regarded as mono-opistho-ovarian, justifying its transference to Willinema. Some other features of W. sulphasae, in particular its small size and its subcylindrical tail (c’ = 1.5) with a terminal hyaline layer, resemble the pattern observed in some members of the genus Tylencholaimellus Cobb in Cobb, 1915, but the pharyngeal expansion occupies about one-half of the total neck length (vs very short, bulb-like in Tylencholaimellus representatives). Therefore, the species is herein considered a species inquirenda. Notes on its phylogeny Willinema is a dorylaimid genus whose evolutionary affinities have not been elucidated yet. Leaving aside some general traits (odontostyle relatively strong, typical dorylaimoid pharynx), two morphological features were especially relevant to address its classification, namely the short and rounded tail of both sexes, and its monoopistho-ovarian female genital system. In a revision of the family Thornenematidae, Andrássy (1987) created the subfamily Willinematinae to accommodate Willinema and Sclerolabia (see above), both rounded-tailed and mono-opistho-ovarian genera that differ in the absence and presence, respectively, of labial and postlabial sclerotisations. In their evaluation of the taxonomical status of the family Thornenematidae, Coomans and Carbonell (1988) concluded that only three thornenematid genera should be grouped under a subfamilar (rather than familiar) taxon, Thornenematinae; meanwhile, they considered the relationships of other genera, Willinema among them, uncertain, although they should be evaluated within a ‘ wider concept of the family Dorylaimidae’ (p. 384). The new Iranian species herein described perfectly fits the known morphological pattern of Willinema representatives and confirms the two most relevant synapomorphies of the genus, namely the rounded tail in both sexes and the monoopistho-ovarian condition of the female genital system. The rounded tail is a remarkable trait shared with members of the subfamily Labronematinae Peña-Santiago and Álvarez-Ortega, 2014 in Dorylaimidae. Actually, Willinema species much resemble the members of the genus Crassolabium, if we accept the diovarian condition of the latter, a relevant difference indeed since mono-ovarian species do not occur within the components of the family Dorylaimidae as hitherto defined. It is unfortunate that the authors failed in their attempt to sequence the new species herein described, as molecular analyses will certainly provide new insights into the phylogeny of Willinema.	en	Vazifeh, Nasir, Niknam, Gholamreza, Jabbari, Habibeh, Fallahi, Amin, Zahedi, Ebrahim, Peña-Santiago, Reyes (2022): Description of Willinema persicum sp. n. (Dorylaimida: Thornenematidae) from Iran, with an updated taxonomy of the genus. Journal of Natural History 56 (13 - 16): 849-863, DOI: 10.1080/00222933.2022.2082895, URL: http://dx.doi.org/10.1080/00222933.2022.2082895
