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Aporcelinus paramamillatus sp. n. (Figs 1���2) Material examined. Six females from one location, in good condition. Morphometrics. See Table 1. Description. Female: Moderately slender (a = 21���26) nematodes of medium size, 1.71���2.14 mm long. Habitus visibly curved ventrad upon fixation, C-shaped. Cuticle two-layered, 4 ��m thick at anterior region and mid-body and 4���7 ��m on tail; inner layer thicker and more refractive than the outer layer. Lateral chords 10���14 ��m broad or 12���19 % of mid-body diameter. Body pores often obscure, but two cervical dorsal and two ventral are sometimes distinct at the level of the odontophore. Lip region somewhat angular, offset by weak constriction, 2.6���2.8 times as wide as high and 26���31 % of body diameter at neck base; lips moderately separated, with protruding labial and cephalic papillae. Amphid fovea cup-shaped, its opening 10���12 ��m broad or about one-half of lip region diameter. Cheilostom nearly as long as wide, with no specialization. Odontostyle strong, slightly (1.1���1.2 times) longer than lip region diameter, 5.2���6.5 times as long as wide and 1.1���1.5 % of total body length; aperture 12 ��m or 46���50 % of its total length. Guiding ring simple but distinct and somewhat plicate, located at 12���13 ��m or 0.5 times the lip region diameter from the anterior end. Odontophore rod-like, 1.7 ���2.0 times the odontostyle in length. Pharynx consisting of a slender but muscular anterior section enlarging very gradually into the posterior expansion that is 6.4���8.3 times as long as wide, 2.7���3.4 times the corresponding body diameter long, and occupies 47���56 % of total neck length; gland nuclei located as follows (n = 1): DO = 58, DN = 62, S 1 N 1 = 75, S 1 N 2 = 79, S 2 N = 92. Nerve ring at 163���212 ��m or 31���35 % of total neck length from the anterior end. Pharyngointestinal junction bearing a distinctly developed dorsal lobe; cardia conical, 12��� 18 x 8���15 ��m. Genital system didelphic-amphidelphic, with both branches almost equally and moderately developed, 157���224 ��m long or 9���12 % of total body length: ovaries 57���177 ��m long, surpassing the oviduct-uterus junction; oviduct 65���106 ��m long or 0.9���1.4 times the body diameter, and consisting of a slender portion and a moderately developed pars dilatata; a narrowing separates oviduct and uterus; uterus 71���94 ��m long or 1.0��� 1.6 times the corresponding body diameter long, tripartite, consisting of a short and wider proximal region, a narrower intermediate section with no lumen, and a nearly spherical distal part; uterine egg 90 x 55 ��m, with irregular shell; vagina extending inwards 26���32 ��m or 32���43 % of body diameter, with pars proximalis 16��� 22 x 18���22 ��m and somewhat sigmoid walls surrounded by weak musculature, pars refringens consisting of two drop-shaped pieces 6��� 7 x 4���6 ��m and a combined width of 9���12 ��m, and pars distalis 3���5 ��m long and visibly refractive; vulva a transverse slit. Prerectum 1.8���2.2, rectum 1.0��� 1.2 times the anal body diameter long. Tail conical with finely rounded tip, dorsally and ventrally convex, ending in a short peg-like terminus, occasionally slightly re-curved dorsad; hyaline portion very short; caudal pores two pairs at the anterior part of tail, one subdorsal, the other sublateral. Male: Unknown. Diagnosis. The new species is characterized by its 1.71���2.14 mm long body, lip region offset by weak constriction and 21���22 ��m broad, odontostyle 22���24 ��m at its ventral side or 1.1���1.2 times the lip region diameter, neck 451���503 ��m long, pharyngeal expansion 183���251 ��m long or 46���56 % of total neck length, uterus tripartite and 71���94 ��m long or 1.0��� 1.6 times the corresponding body diameter long, V = 53���54, tail conical (27���34 ��m, c = 50���77, c��� = 0.6���0.9) ending in a peg-like terminal portion and not visibly re-curved dorsad, and male absent. Relationships. This species is very similar to A. mamillatus (Williams, 1959) ��lvarez-Ortega & Pe��a- Santiago, 2011 in its general morphology and many morphometrics, but it can be distinguished by several significant differences: broader lip region (21���22 vs 19���20 ��m), smaller odontostyle (22���24 vs 25���29 ��m, 1.1���1.2 vs 1.3���1.5 times the lip region diameter, n= 9) with shorter aperture (46���50 vs 57���61 % of its total length), more posterior S 2 N (92 vs 83���86) and more posterior vulva (V = 53���54 vs V = 49���52). Type locality and habitat. Northern Vietnam, Cat Ba National Park, Hai Phong Province (GPS coordinates: 20 �� 47 ' 45 '' N and 107 �� 00' 40 ''E, elevation 170 m), in soil of a secondary forest with Cinnamomum sp. as the dominant plant species. One female was collected near the same location in the rhizosphere of Zea mays L. growing in an area of intensive agricultural crops alongside the National Park. Type material. Female holotype and four females paratypes deposited in the nematode collection of the University of Ja��n, Spain. One female paratype in the nematode collection of the Institute of Ecology and Biological Resources (IEBR), Hanoi, Vietnam. Etymology. The specific epithet means ���near mamillatus ��� as the new species resembles A. mamillatus in many aspects., Published as part of Nguyen, Thi Anh Duong, Abolafia, Joaqu��n, Bonkowski, Michael & Santiago, Reyes Pe��a-, 2016, Two new species of the genus Aporcelinus Andr��ssy, 2009 (Nematoda, Dorylaimida, Aporcelaimidae) from Vietnam, pp. 550-560 in Zootaxa 4103 (6) on pages 551-554, DOI: 10.11646/zootaxa.4103.6.5, http://zenodo.org/record/271219, {"references":["Williams, J. R. (1959) Studies on the nematode soil fauna of sugarcane fields in Mauritius. 3. Dorylaimidae (Dorylaimoidea, Enoplida). Mauritius Sugar Industry Research Institute Occasional Paper, 3, 1 - 28."]}

Nguyen, Thi Anh Duong, Abolafia, Joaquín, Bonkowski, Michael, Santiago, Reyes Peña- (2016): Two new species of the genus Aporcelinus Andrássy, 2009 (Nematoda, Dorylaimida, Aporcelaimidae) from Vietnam. Zootaxa 4103 (6): 550-560, DOI: http://doi.org/10.11646/zootaxa.4103.6.5

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

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

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

Enchodelus saxifragae Popovici, 1995 (Fig. 4) MATERIAL EXAMINED Six females from Suhardu Mic Mountains, three females from Parâng Mountains, two female paratypes and two male paratypes from Retezat Mountains. MEASUREMENTS NEW OBSERVATIONS ON MALE (BASED ON THE TWO PARATYPES AVAILABLE) Adcloacal pair of supplements situated at 13.5 or 13.0 µ m from cloacal aperture posterior ventromedian supplement at 19.5 or 30 µ m from adcloacal pair, ventromedian supplements 9-24 µ m apart. Spicules rather robust, 5.3 or 4.4 times as long as wide and 1.7 anal body diam. long. Lateral guiding piece also robust, distinctly falcate in posterior half, furcate at tip, 14 or 16.5 µ m long and ca 4.0 or 3.3 times as long as wide. DISTRIBUTION Cliff vegetation, Suhardu Mic Mountain, H&abreve;&scedil;ma&scedil; Mountains (Eastern Romanian Carpathians); subalpine meadow, Coasta lui Rus, Parâng Mountains (Southern Romanian Carpathians); cliff vegetation on Albele Peak and the Piatra Iorgovanului area, Retezat Mountains (Southern Romanian Carpathians); sites nos 3, 7-9 in Table 1. REMARKS The females examined perfectly fit the original description (Popovici, 1995) as well as the Iberian specimens recently studied by Guerrero et al. (2008). However, some differences have been noted in male features, such as the peculiar morphology of the lateral guiding pieces, a character which might be a relevant diagnostic feature of this species., Published as part of Ciobanu, Marcel, Popovici, Iuliana, Guerrero, Pablo & Santiago, Reyes Peña-, 2010, Nematodes of the order Dorylaimida from Romania. The genus Enchodelus Thorne, 1939. 3. Species with rounded tail and long odontostyle, pp. 609-618 in Nematology 12 (4) on pages 615-617, DOI: 10.1163/138855410X12628646275925, http://zenodo.org/record/8111712, {"references":["POPOVICI, I. (1995). New species of Tubixaba and Enchodelus (Nematoda: Dorylaimida) from Romania. Nematologica 41, 435 - 448.","GUERRERO, P., LIEBANAS, G. & PENA- SANTIAGO, R. (2008). Nematodes of the order Dorylaimida from Andalucia Oriental, Spain. The genus Enchodelus Thorne, 1939. 2. Description of three known species with rounded tail and long odontostyle. Nematology 10, 451 - 470."]}

Enchodelus macrodorus (de Man, 1880) Thorne, 1939 (Fig. 3) MATERIAL EXAMINED Five females from Metaliferi Mountains, five females from Bihor Mountains and one female from Trasc&abreve;u Mountains, all in variable but acceptable condition. MEASUREMENTS See Table 3. DISTRIBUTION Hornbeam-beech forest and grassland located at Buce&scedil;- Vulcan, Metaliferi Mountains (Western Romanian Carpathians); mountainous grassland, the Seac&abreve; Valley, Padi&scedil; karst plateau and cliff vegetation at Ghe&tcedil;ar- Sc&abreve;ri&scedil;oara, Bihor Mountains (Western Romanian Carpathians); grassland on a gentle slope at Cheile Turenilor, Trasc&abreve;u Mountains (Western Romanian Carpathians); sites nos 1, 2, 5, 6 and 10 in Table 1. REMARKS The Romanian material fits very well previous descriptions of the type (Guerrero & Peña-Santiago, 2007) and Iberian (Guerrero et al., 2008) populations of this species, in particular concerning its most characteristic features such as its rather anterior vulva and comparatively short (although tripartite) uterus, with poorly developed intermediate region. Nevertheless, some morphometric differences have been noted in agreement with data provided by Popovici (1995) in her earlier report of E. macrodorus in Romania: i ) the three females from hornbeam-beech forest in the Metaliferi Mountains have a longer odontophore (55 vs 43-47 and 42-48 µ m in type and Iberian populations, respectively) and total stylet length (97-100 vs up to 92 µ m); ii ) the four females from Padi&scedil;, in the Bihor Mountains, also have a longer odontophore (50-55 µ m) and total stylet length (92.5-100 µ m) and, moreover, their neck region and pharyngeal expansion are longer too (356-390 vs 303-345 and 288-344 µ m in type and Iberian material, respectively; 144-169 vs 111-130 and 113-138 µ m, respectively); and iii ) three of the four females from the other localities have a somewhat longer prerectum (238, 233, 256 vs 166-212, 112-176 and 205-218 µ m) in type, Iberian and the Romanian specimens studied (Trasc&abreve;u Mountains, Bihor Mountains and the grassland from the Metaliferi Mountains, respectively). The differences observed are herein interpreted as intraspecific geographical variation within a widely distributed taxon in the Holartic region. As mentioned above, Popovici (1995) provided measurements of several Romanian populations, but the true identity of part of this material is herein clarified. The populations collected from Retezat Mountains (Albele and Piatra Iorgovanului), as well as those from Gil&abreve;u, Vl&abreve;deasa and the male from Bihor Mountains belong to E. longispiculus (see Ciobanu et al., 2010b)., Published as part of Ciobanu, Marcel, Popovici, Iuliana, Guerrero, Pablo & Santiago, Reyes Peña-, 2010, Nematodes of the order Dorylaimida from Romania. The genus Enchodelus Thorne, 1939. 3. Species with rounded tail and long odontostyle, pp. 609-618 in Nematology 12 (4) on pages 613-615, DOI: 10.1163/138855410X12628646275925, http://zenodo.org/record/8111712, {"references":["DE MAN, J. G. (1880). Die einheimischen, frei in der reinen Erde und im sussen Wasser lebenden Nematoden. Tijdschrift Nederlandsche Dierkundige Vereeniging 5, 1 - 104.","THORNE, G. (1939). A monograph of the nematodes of the superfamily Dorylaimoidea. Capita Zoologica 8, 1 - 261.","GUERRERO, P., LIEBANAS, G. & PENA- SANTIAGO, R. (2008). Nematodes of the order Dorylaimida from Andalucia Oriental, Spain. The genus Enchodelus Thorne, 1939. 2. Description of three known species with rounded tail and long odontostyle. Nematology 10, 451 - 470.","POPOVICI, I. (1995). New species of Tubixaba and Enchodelus (Nematoda: Dorylaimida) from Romania. Nematologica 41, 435 - 448.","CIOBANU, M., POPOVICI, I., GUERRERO, P. & PENA- SANTIAGO, R. (2010 b). Nematodes of the order Dorylaimida from Romania. The genus Enchodelus Thorne, 1939. 2. Species with rounded tail and medium-sized odontostyle. Nematology 12, 381 - 397."]}

Enchodelus carpaticus ** sp. n. (Figs 1, 2) MATERIAL EXAMINED Fourteen females from Mehedin&tcedil;i Mountains, mostly in good condition, some becoming visibly flattened. MEASUREMENTS See Table 2. DESCRIPTION Female Moderately slender nematodes of medium size, 1.59- 1.87 mm long. Habitus after fixation somewhat curved ventrad, adopting an open C shape. Body cylindrical, tapering towards both ends, but more so anteriorly. Cuticle 3.0-5.0 µ m thick in anterior region, 3.5-5.5 µ m at mid-body and 7.0-9.0 µ m on tail, its outer layer with very fine transverse striations and much thinner than inner one, especially at tail level. Lateral chord 6-8 µ m wide or occupying ca one-tenth (8-12%) of mid-body diam., lacking any particular differentiation. Lateral pores obscure, two or three dorsal and ventral moderately coarse pores present at odontostyle/odontophore level, other more or less distinct ventral pores present throughout body. Lip region more or less rounded, offset by a distinct depression, 2.4-3.1 times as broad as high, ca one-fourth (flattened specimens) to one-third (26-34%) of body diam. at neck base. Lips amalgamated, labial and cephalic papillae visible but barely protruding above cephalic contour. Amphid fovea cup-shaped, opening at level of labial depression, occupying 9.5-12.0 µ m or one-half to three-fifths of lip region diam. Cheilostom almost cylindrical, with thick walls. Odontostyle comparatively slender and long, 14-21 times as long as wide and 2.1- 2.5 times longer than lip region diam. or 2.3-2.8% of total body length, aperture small, 3.0-4.5 µ m long or 7-10% of total length. Odontophore 1.0-1.2 times as long as odontostyle, with very well developed flanges. Guiding ring double, located at 25-31 µ m or 1.4-1.7 lip region diam. from anterior end. Pharynx consisting of a slender but muscular anterior portion enlarging gradually, basal expansion 4.6-6.4 times as long as wide, 2.0 (flattened specimens) to 2.9 times body diam. at neck base, occupying ca two-fifths (39-45%) of total neck length. Pharyngeal gland nuclei located as follows: DN = 63-65; S 1 N obscure; S 2 N = 82-86. Cardia rounded conoid, nearly as long as wide, (8-13) × (9-14) µ m. Acell mass or chord, ca one body diam. long, present in dorsal position between cardia level and proximal end of anterior ovary. Genital system didelphic-amphidelphic, both branches equally and well developed, anterior 355- 474 µ m, posterior 337-456 µ m long. Ovaries relatively short, anterior 61-77 µ m, posterior 90-155 µ m long, in general not reaching sphincter level, oocytes first in two or more rows, then in one row. Oviduct 107-148 µ m long or 1.3-2.2 corresponding body diam., consisting of a slender portion with prismatic cells and a moderately developed pars dilatata with visible lumen. Sphincter prominent, located between oviduct and uterus. Uterus typically tripartite, i.e., consisting of a wider proximal region with distinct lumen followed by a narrower intermediate portion with narrow lumen surrounded by cluster of hyaline cells, ending with well developed spheroid pars dilatata distalis, 135-195 µ m or 2.0-2.9 corresponding body diam. long. Uterine eggs, observed only in one female, (90-92) × (40-44) µ m. Only one female observed to contain sperm inside uterus. Vagina extending inwards 37-47 µ m or one-half to three-fifths of body diam., pars proximalis longer than broad, (14-25) × (11-17) µ m, with straight walls, enveloped by distinct circular musculature, pars refringens with (in lateral view) two well developed, trapezoidal sclerotised pieces measuring (7.5-9.5) × (7.5- 9.5) µ m and with combined width of 16-19 µ m, pars distalis 2.5-3.5 µ m long, weakly sclerotised. Two cells (probably glands) clearly visible in several specimens on both sides of vulva, one anterior, one posterior. Vulva a transverse slit. Prerectum 4.1-6.6 and rectum 0.8-1.2 anal body diam. long. Tail short, rounded; inner cuticle layer bearing radial striations. Two pairs of subterminal caudal pores, one subdorsal, one practically lateral. ∗ Corresponding author, e-mail: icb@cluj.astral.ro ** The specific epithet refers to Carpates, - um, the Latin name of the Carpathian Mountains, or Carpathians, the geographical origin of this species. © Koninklijke Brill NV, Leiden, 2010 Also available online - www.brill.nl/nemy * According to Coldea (1991, 1993) and Pop et al. (1973). ** According to the Romanian System of Soil Classification (Conea et al., 1980). Male Not found, but certainly occurring because one fertilised female was observed in the material studied. TYPE LOCALITY AND HABITAT Mountainous grassland located at Padina Izvoarelor Valley-Piatra Clo&scedil;ani, Mehedin&tcedil;i Mountains (Southern Romanian Carpathians); site no. 4 in Table 1. TYPE MATERIAL Holotype female numbered “5” on slide 744 and six paratype females (749/1-4, 6-7) deposited inthe nematode collection of the Institute of Biological Research, Cluj-Napoca, Romania. Slide 749 containing seven paratype females numbered “1”-“3” and “5”-“8” deposited in the nematode collection of Departamento de Biología Animal, Biología Vegetal y Ecología, University of Jaén, Jaén, Spain. DIAGNOSIS AND RELATIONSHIPS The new species is distinguished by its body 1.59- 1.87 mm long, lip region offset by a marked depression and 17-20 µ m diam., odontostyle 39.5-47 µ m long and 2.1-2.5 lip region diam. or 2.3-2.8% of total body length, odontophore 42-51 µ m long and with distinct basal flanges, neck length 336-388 µ m long, pharyngeal expansion 136-167 µ m long or 39-45% of total neck length, presence of dorsal cell mass near cardia, female genital system amphidelphic, long tripartite uterus 144- 195 µ m or 2.2-2.9 corresponding body diam. long, pars refringens vaginae with two trapezoidal sclerotisations, V = 42-50, and female tail short and rounded (21-29 µ m long, c = 55-87, c &lcedil; = 0.5-0.7). In having an odontostyle 39.5-47 µ m long, very well developed flanges at the odontophore base and a tripartite uterus, the new species is very similar to E. macrodorus (de Man, 1880) Thorne, 1939 and E. saxifragae Popovici, 1995 . It differs from both in one subtle but reliable feature, the presence of a dorsal cell mass or chord between the cardia and the proximal end of the anterior ovary, a feature confirmed in all of the 14 specimens of the new species examined but never reported in E. macrodorus and E. saxifragae. Moreover, it can be distinguished from E. macrodorus by its longer pharyngeal expansion (136-167 µ m or 39-45% of total neck length vs 111-138 µ m or 33-38% of total neck length, although some variability is found in this character in E. macrodorus, see below), more posterior vulva with V = 42-50 (but less than 44 in only two specimens out of 14) vs 40-44 in both type and Romanian populations) and longer tripartite uterus (144-195 vs 56-143 µ m or 2.2-2.9 vs or 0.9-2.0 corresponding body diam. long) with well differentiated ( vs barely differentiated) intermediate region; and from E. saxifragae in its shorter female body (L = 1.59-1.87 vs 1.96-2.35 mm in type population, n = 6; L = 1.80- 2.38 in Iberian populations, n = 37) and lip region offset by depression vs constriction. It also resembles E. groenlandicus (Ditlevsen, 1927) Thorne, 1939 (see redescription by Guerrero et al., 2008), but can be separated in having the lip region offset by a well marked depression vs almost continuous or offset by weak depression, shorter odontostyle of 43.3 (39.5-47) vs 49.3 (44-53) µ m, shorter total stylet length (84-96 vs 94-108 µ m) and longer pharyngeal expansion (136-167 vs 112-142 µ m or 39-45 vs 31-39% of total neck length).

Enchodelus ponorensis Popovici, 1995 (Fig. 6) MATERIAL EXAMINED Six female paratypes, in not very good condition, collected from the Poiana Ponor area located within the Padi&scedil; karstic plateau, Bihor Mountains (Western Romanian Carpathians); sites 1 and 3 in Table 1. MEASUREMENTS See Table 3. Female (based on material examined) Slender nematodes of medium size, 1.63-2.04 mm long. Habitus after fixation curved ventrad, C-shaped. Body cylindrical, tapering towards both ends, but more so anteriorly. Cuticle 3.0 µm in anterior region, 3.0-4.0 µm at mid-body 7.0-8.0 µm on dorsal region of tail, and 11- 13 µm at tail terminus, its outer layer with very fine transverse striations and much thinner than inner one, especially on tail. Lateral chord 16-19 µm wide or occupying almost one-third (29-33%) of mid-body diam., lacking any particular differentiation. Body pores, if present, inconspicuous. Lip region more or less rounded, offset by a marked constriction, 2.5-3.0 times as broad as high and ca one-third (29-33%) of body diam. at neck base. Lips amalgamated, labial and cephalic papillae inconspicuous, barely protruding above cephalic contour. Amphid fovea cup-shaped, opening at level of cephalic depression, occupying 6.0-6.5 µm ca two-fifths (38-41%) of corresponding body diam. Cheilostom almost cylindrical. Odontostyle comparatively robust, 8.2-10.4 times as long as wide, 1.5-1.6 times longer than lip region diam. or 1.3-1.6% of total body length, aperture small, but precise extent not distinguishable in material examined. Odontophore 1.2-1.4 times as long as odontostyle, with small basal thickenings, no evidence of knobs or flanges observed. Guiding ring double, located at 14-16 µm or 0.8- 0.9 times lip region diam. from anterior end. Pharynx consisting of a slender muscular anterior portion expanding gradually; pharyngeal expansion occupying ca two-fifths (37-45%) of total neck length, 5.5-7.2 times as long as wide or 3.0-3.4 times corresponding body diam. Pharyngeal gland nuclei located as follows: DN = 62-67; S 1 N obscure; S 2 N = 80-85. Cardia rounded conoid, nearly as long as broad, (13-20) × (12-15) µm. Genital system didelphic-amphidelphic; both branches equally and well developed, anterior branch 155-225 µm, posterior branch 180-228 µm long. Ovaries 66-130 µm long, often extending to sphincter level, oocytes initially in two or more rows, then in one row. Oviduct 60-88 µm or 1.0-1.5 corresponding body diam. long, consisting of a slender portion with prismatic cells and a moderately developed pars dilatata with visible lumen. Sphincter prominent, located between oviduct and uterus. Uterus long, 70-90 µm or 1.2-1.6 times corresponding body diam., bipartite, i.e., consisting of a wider proximal region with distinct lumen followed by a narrower and shorter distal portion with narrow lumen and surrounded by a cluster of hyaline cells. Vagina extending inwards 30-33 µm or 54-59% of body diam., pars proximalis nearly as long as broad, (14-15) × (11-15) µm, with sigmoid to straight walls and enveloped by weak circular musculature; pars refringens with (in lateral view) two trapezoidal sclerotisations measuring (3.0- 5.0) × (8.0-9.0) µm and with a combined width of 16- 18 µm, pars distalis 4.0-5.0 µm long. Vulva a transverse slit. Prerectum 6.0-6.6 and rectum 1.1-1.3 anal body diam. long. Tail conoid, slightly straighter ventrally, most specimens lacking saccate bodies. Two pairs of caudal pores at middle of tail, one subdorsal, another practically lateral. The genus Enchodelus from Romania Male Unknown. DIAGNOSIS Based on the original description and the present study, E. ponorensis is distinguished by its 1.63-2.07 mm long body, lip region offset by a marked constriction and 15- 16.5 µm diam., odontostyle 23-26 µm or 1.5-1.6 lip region diam. long and comprising 1.3-1.6% of total body length, odontophore 31-38 µm long and with small thickenings, neck 355-520 µm long, pharyngeal expansion 150-245 µm long or 41-47% of total neck length, female genital system amphidelphic, uterus short and bipartite, pars refringens vaginae with two trapezoidal sclerotisations, V = 44-52, tail rounded conoid (35-45 µm long, c = 40-57, c&lcedil; = 0.9-1.3) and males unknown. REMARKS The description herein perfectly agrees with the original one (Popovici, 1995), although some additional morphological details have now been incorporated, especially those concerning the female genital system. Enchodelus ponorensis is an interesting taxon within the genus. It is an atypical round-tailed species due to its comparatively short, bipartite uterus, whereas all other similar species examined by the authors (Guerrero & Peña-Santiago, 2007; Guerrero et al., 2008a, b) are characterised by having a long, tripartite uterus, only E. macrodorus showing a shorter, but distinctly tripartite, uterus. Thus, somewhat surprisingly, in its short, bipartite uterus, E. ponorensis resembles the conical-tailed species of the genus (see Guerrero & Peña-Santiago, 2007; Guerrero et al., 2008b). On the other hand, E. ponorensis is peculiar in having a relatively short and robust (8-10 times as long as wide) odontostyle, long pharyngeal expansion and comparatively longer and more conoid female tail, similar to that found in some species of the genus Crassolabium Yeates, 1967 (= Thonus Thorne, 1974), for instance C. tenuistylum (Ciobanu, Popovici, Abolafia & Peña-Santiago, 2008) Peña-Santiago & Ciobanu, 2008, also known from Romania. Nevertheless, E. ponorensis conforms better to the general pattern of Enchodelus in the form of its lip region, odontophore, location of S 2 N and uterus., Published as part of Ciobanu, Marcel, Popovici, Iuliana, Guerrero, Pablo & Santiago, Reyes Peña-, 2010, Nematodes of the order Dorylaimida from Romania. The genus Enchodelus Thorne, 1939. 2. Species with rounded tail and medium-sized odontostyle, pp. 381-397 in Nematology 12 (3) on pages 394-396, DOI: 10.1163/138855409X12549869072329, http://zenodo.org/record/8111660, {"references":["POPOVICI, I. (1995). New species of Tubixaba and Enchodelus (Nematoda: Dorylaimida) from Romania. Nematologica 41, 435 - 448.","GUERRERO, P., LIEBANAS, G. & PENA- SANTIAGO, R. (2008 a). Nematodes of the order Dorylaimida from Andalucia Oriental, Spain. The genus Enchodelus Thorne, 1939. 2. Description of three known species with rounded tail and long odontostyle. Nematology 10, 451 - 470.","GUERRERO, P., LIEBANAS, G. & PENA- SANTIAGO, R. (2008 b). Nematodes of the order Dorylaimida from Andalucia Oriental, Spain. The genus Enchodelus Thorne, 1939. 3. Description of two new and one known species with rounded tail and medium-sized odontostyle. Nematology 10, 711 - 733.","CIOBANU, M., POPOVICI, I., ABOLAFIA, J. & PENA-SANTIAGO, R. (2008). Nematodes of the order Dorylaimida from Romania. The genus Thonus Thorne, 1974. Part II. Nematology 10, 167 - 188.","CIOBANU, M., POPOVICI, I., ABOLAFIA, J. & PENA- SANTIAGO, R. (2008). Nematodes of the order Dorylaimida from Romania. The genus Thonus Thorne, 1974. Part II. Nematology 10, 167 - 188."]}

Enchodelus parahopedoroides * sp. n. (Figs 3-5) MATERIAL EXAMINED Three females from the Bucegi Mountains, three females from the Retezat Mountains, one male from Vl&abreve;deasa Mountain, and one male from Bihor Mountains, all in acceptable condition. MEASUREMENTS See Table 4. DESCRIPTION Female Moderately slender nematodes of medium size, 1.44- 1.64 mm long. Habitus after fixation curved ventrad, Cshaped. Body cylindrical, tapering towards both ends, but more so anteriorly. Cuticle 2.0-2.5 µm in anterior region, 2.0-3.0 µm at mid-body and 7.0-7.5 µm on tail; its outer layer with very fine transverse striations and much thinner than inner one, especially on tail. Lateral chord 4.5- 7.0 µm wide or occupying 8-12% of mid-body diam., lacking any particular differentiation. Lateral pores obscure, one dorsal and one ventral pore prominent at odontostyle level. Lip region more or less rounded, offset by a distinct depression, 2.8-3.1 times as broad as high and ca one-third (32-36%) of body diam. at neck base. Lips amalgamated, labial and cephalic papillae visible, protruding above cephalic contour. Amphid fovea cup-shaped, opening at level of cephalic depression, occupying 8-10 µm or ca one-half (42-51%) of corresponding body diam. Cheilostom curly-bracket-shaped. Odontostyle comparatively robust, 11-14 times as long as wide, 1.5-1.6 times longer than lip region diam. or 1.6-2.0% of total body length, aperture small, 4-5 µm long or 14-17% of total length. Odontophore 1.4-1.6 times as long as odontostyle, with basal thickenings or flanges. Guiding ring double, located at 17-20 µm or 1.0-1.1 times lip region diam. from anterior end. Pharynx consisting of a slender muscular anterior portion expanding gradually, pharyngeal expansion occupying ca two-fifths (39-41%) of total neck length and ca one-half of corresponding body diam. Pharyngeal gland nuclei located as follows: DN = 64-67; S 1 N 1 obscure; S 1 N 2 = 75-78; S 2 N = 85-87. Cardia rounded conoid, nearly as long as broad, (12-18) × (10-13) µm. Genital system didelphic-amphidelphic, both branches equally and well developed, anterior branch 240- 293 µm, posterior branch 215-280 µm long. Ovaries 53- 140 µm long, not reaching sphincter level; oocytes initially in two or more rows, then in one row. Oviduct 65- 105 µm or 1.2-1.9 corresponding body diam. long, consisting of a slender portion with prismatic cells and a moderately developed pars dilatata with visible lumen. Sphincter prominent, located between oviduct and uterus. Uterus long, 133-170 µm or 2.4-3.1 corresponding body diam., sometimes twisted, tripartite, i.e.: consisting of a wider proximal region with distinct lumen followed by a narrower and shorter intermediate portion with narrow lumen and surrounded by a cluster of hyaline cells, and ending with a well developed spheroid pars dilatata distalis. Vagina extending inwards 27-32 µm or 47-58% of body diam., pars proximalis longer than broad, (18-22) × (11-15) µm, with sigmoid to straight walls and enveloped by weak circular musculature; pars refringens with (in lateral view) two trapezoidal sclerotisations measuring (3.0- 4.5) × (6-8) µm and with a combined width of 12-15 µm; pars distalis 4.0-5.0 µm long. Vulva a transverse oval slit, preceded by a shallow depression in body surface. Prerectum 3.9-5.7 and rectum 0.9-1.4 anal body diam. long. Tail rounded conoid, in one specimen with a few small saccate bodies, inner cuticle layer marked by radial striation, its ventral margin slightly irregular, barely separated from outer layer. Two pairs of subterminal caudal pores, one subdorsal, another practically lateral. Male (see remarks) General morphology similar to that of female, but with more slender body, narrower lip region and slightly shorter odontophore. Genital system diorchic, with opposed testes. In addition to adcloacal pair at 11 or 10.5 µm from cloacal aperture, presence of series of nine ventromedian supplements 10-24 µm apart, posteriormost supplement located at 31 or 32.5 µm from adcloacal pair, barely in anterior to, or at level of, anterior end of spicules. Spicules well curved ventrad, rather robust, ca 5.1 or 6.0 times as long as wide or 1.6 or 1.9 anal body diam. long. Lateral guiding pieces 9.5 or 11.5 µm, ca 4.8 or 3.8 times as long as wide. Tail similar to that of female. * The specific epithet indicates the resemblance of this species to E. hopedoroides. DIAGNOSIS AND RELATIONSHIPS Enchodelus parahopedoroides sp. n. is distinguished by its 1.44-1.64 mm long body, lip region 17.0-18.5 µm diam. and offset by a marked depression, odontostyle 26-28 µm or 1.5-1.6 lip region diam. long and 1.6- 2.0% of total body length, odontophore 40-43 µm long and with distinct basal knobs and flanges, neck 300- 328 µm long, pharyngeal expansion 120-130 µm long or 39-41% of total neck length, female genital system amphidelphic, uterus long and tripartite, pars refringens vaginae with two trapezoidal sclerotisations, V = 45- 52, tail rounded conoid (20-24 µm long, c = 63-80, c&lcedil; = 0.6-0.8), spicules 59-67 µm long, and nine spaced ventromedian supplements starting just anterior to, or at level of, anterior end of spicules. Enchodelus parahopedoroides sp. n. is very similar to E. hopedoroides Altherr, 1963 (see description by Guerrero et al., 2008b), from which it differs in its narrower lateral chord (4.5-7.0 µm or 8-12% of body diam. vs 8-16 µm or 11-27% of body diam., respectively), shorter odontostyle (26-28 vs 31-34 µm, n = 23), pars refringens vaginae without vs with two massive and granular appearing pieces, vulva more posterior (V = 45-52 vs = 40-47), and female tail lacking abundant saccate bodies and any peculiar differentiation of the cuticle vs saccate bodies abundant and inner cuticle layer divided into two by a discontinuity, its ventral margin rather irregular and somewhat separated from the outer layer. In spite of the high similarity between both species, there are three features (odontostyle length, nature of pars refringens vaginae and female tail) that support the proposal of a new taxon for the Romanian material. In having an odontostyle less than 30 µm long and rounded tail, the new species is also close to E. altherri Vinciguerra & de Francisci, 1973, E. ameliae Guerrero, Liébanas & Peña-Santiago, 2008, E. analatus (Ditlevsen, 1927) Thorne, 1939, E. arcticus Nesterov, 1976, E. georgiensis Eliava, Tskitishvili & Bagathuria, 2006, E. hopedorus (Thorne, 1929) Thorne, 1939, E. longispiculus Guerrero, Liébanas & Peña-Santiago, 2008 and E. ponorensis Popovici, 1995. It differs from E. altherri in its longer body (vs 1.18-1.38 mm), comparatively longer odontostyle at 1.5-1.6 vs 1.8-2.0 lip region diam. long (i.e., 1.8 according to the original Figure 2A of E. altherri and 2.0 according to the original description), longer neck (vs 236-276 µm, as calculated from original description), vagina with well developed pars refringens (vs “vulva debolmente ispessita”), tail of both female and male more rounded vs conoid, shorter (vs c = 41- 59, c&lcedil; = 1.0, calculated from original illustrations) and lacking saccate bodies vs 3-8 saccate bodies present, and longer spicules (vs 50-52 µm). From E. ameliae in its less marked lip region (vs offset by constriction), shorter odontostyle (vs 30-33 µm, n = 11, including females and males), longer neck (vs 268-307 µm) and pharyngeal expansion (vs 95-107 µm), and fewer (vs 11-14) ventromedian supplements, the posteriormost located barely anterior to, or at level of, the anterior end of the spicules vs one or two within the range of the spicules. From E. analatus in its shorter odontostyle (vs 30-34 µm), pars refringens vaginae present vs absent and shorter female tail (vs c = 40-55, c&lcedil; = 0.8-1.1) which lacks vs presence of saccate bodies. From E. arcticus in its broader lip region (vs 12 µm, as inferred from original description), longer odontophore (vs up to 36 µm, as inferred from original description) and ventromedian supplements present vs reportedly absent (!). From E. georgiensis in its longer body (vs 0.88-1.10 mm), broader lip region (vs 12 µm), longer odontophore (vs 35-37 µm) and longer spicules (vs 36- 44 µm). From E. hopedorus in its broader female lip region (vs 14.5-16 µm), comparatively shorter odontostyle (vs 1.9-2.2 times the lip region diam.), longer pharyngeal expansion (vs 33-38% of total neck length) and shorter female tail (vs 27-39 µm, c = 41-53) lacking vs bearing abundant saccate bodies. From E. longispiculus in its more slender body (a = 26-30 vs 17-25; body diam. at mid-body = 50-57 vs 64-80 µm), longer pharyngeal expansion (vs 77-114 µm or 28-37% of total neck length, n = 40), somewhat more posterior vulva (vs V = 41- 48) and posteriormost ventromedian supplement located barely anterior to, or at level of, anterior end of spicules (vs two posteriormost, occasionally only one, within range of spicules). Finally, from E. ponorensis, the other Romanian species, in its shorter body (vs 1.6-2.1 mm), narrower lateral chord (vs one-fourth of mid-body diam.) broader female lip region (vs 15-16 µm), longer odontostyle (vs 23-25 µm), uterus tripartite vs bipartite, and shorter female tail (vs 35-45 µm, c = 41-56, c&lcedil; = 0.9- 1.3). TYPE HABITAT AND LOCALITY Cliff vegetation, Caraiman Peak, Bucegi Mountains (Southern Romanian Carpathians); site 5 in Table 1. OTHER HABITATS AND LOCALITIES Cliff vegetation on Albele Peak, Retezat Mountains (Southern Romanian Carpathians), mountainous grassland at Dealul Caprei, Vl&abreve;deasa Mountain (Western Romanian Carpathians) and spruce forest in a swamp area on the Padi&scedil; karstic plateau, Bihor Mountains (Western Romanian Carpathians); sites 9, 11 and 4 in Table 1. TYPE MATERIAL Holotype female on slide 1317 and one paratype female (1318/c) deposited in the nematode collection of the Institute of Biological Research, Cluj-Napoca, Romania. Slide 1316 containing one paratype female deposited in the nematode collection of Departamento de Biología Animal, Biología Vegetal y Ecología, University of Jaén, Spain. REMARKS Some doubts persist as to the conspecifity of the two males examined, which were collected from the Vl&abreve;deasa and Bihor Mountains whilst the females came from the Bucegi and Retezat Mountains, as there are some morphometrical differences, particularly their narrower lip region in addition to their different geographical origin. Nevertheless, some significant features, such as odontostyle and pharyngeal expansion lengths are similar to those found in females and serve to distinguish the males from E. longispiculus, the most similar species. In addition, as mentioned above, the male posteriormost ventromedian supplement is located barely anterior to, or at level of, the anterior end of the spicule, whereas in other species at least one ventromedian supplement is present within the range of the spicules. Thus, the males are provisionally considered to belong to the same species as the females from Bucegi and Retezat Mountains.

Enchodelus longispiculus Guerrero, Liébanas & Peña-Santiago, 2008 (Fig. 1) MATERIAL EXAMINED Eighteen females and 11 males from five localities: Gil&abreve;u Mountains (one female); Retezat Mountains (two populations, one consisting of four females and five males and the other including seven females and two males, respectively); Trasc&abreve;u Mountains (two females and three males); and Vrancei Mountains (four females and one male). MEASUREMENTS See Table 2. DISTRIBUTION Beech forest growing in the Ierii Valley, Gil&abreve;u Mountains (Western Romanian Carpathians); cliff vegetation on Albele Peak, Retezat Mountains (Southern Romanian Carpathians); cliff vegetation in the Piatra Iorgovanului area, Retezat Mountains; grassland on a mild slope at Tureni Gorges, Trasc&abreve;u Mountains (Western Romanian Carpathians); and cliff vegetation at Cheile Ti&scedil;i&tcedil;ei, Vrancei Mountains (Eastern Romanian Carpathians); sites 6, 9, 8, 10 and 12 in Table 1. * Corresponding author, e-mail: icb@cluj.astral.ro © Koninklijke Brill NV, Leiden, 2010 Also available online - www.brill.nl/nemy * According to Coldea (1991), Pop et al. (1973) and Coldea (1993). ** According to the Romanian System of Soil Classification (Conea et al., 1980). REMARKS The description of the material herein studied fits that of the Iberian populations well, although the Romanian specimens display a wider variability in several morphological or morphometrical features, such as odontostyle length (24-34 vs 27-31 µm) and vulva position (42.2-51.5 vs 41.0-47.8). The morphology of the vagina in the Romanian females deserves a special discussion. A few (four out of 18) specimens examined had the pars refringens well developed and nearly identical to that described in the type and other Iberian populations. In the remaining females, however, the pars refringens is poorly developed, even lacking or impossible to distinguish from the pars distalis. Such intraspecific variation was observed among females collected from the same locality, and represents a very unusual feature in dorylaims in which the pars refringens is either present or absent, such a condition being often assumed to represent a good diagnostic character to separate species belonging to the same genus. This is the first record of the species in Romania., Published as part of Ciobanu, Marcel, Popovici, Iuliana, Guerrero, Pablo & Santiago, Reyes Peña-, 2010, Nematodes of the order Dorylaimida from Romania. The genus Enchodelus Thorne, 1939. 2. Species with rounded tail and medium-sized odontostyle, pp. 381-397 in Nematology 12 (3) on pages 381-382, DOI: 10.1163/138855409X12549869072329, http://zenodo.org/record/8111660, {"references":["GUERRERO, P., LIEBANAS, G. & PENA- SANTIAGO, R. (2008 a). Nematodes of the order Dorylaimida from Andalucia Oriental, Spain. The genus Enchodelus Thorne, 1939. 2. Description of three known species with rounded tail and long odontostyle. Nematology 10, 451 - 470.","COLDEA, G. (1991). Prodrome des associations vegetales des Carpates du Sud-Est (Carpates Roumaines). Documents Phytosociologiques, Camerino 13, 317 - 539.","POP, E., BOSCAIU, N. & RADULESCU, D. (1973). L'efficacite de la protection de la tourbiere de Lucina (dep. Suceava, Roumanie). Studii si Comunicari de Ocrotirea Naturii 3, 21 - 28.","COLDEA, G. (1993). Cormofite. Sintaxonomia si descrierea asocia ¸ tiilor vegetale. In: Popovici, I. (Ed.). Parcul national Retezat-Studii ecologice. Brasov, Romania, Editura West Side, pp. 31 - 48.","CONEA, A., FLOREA, N. & PUIU, S. (EDS) (1980). Sistemul roman de clasificare a solurilor. ASAS, Institutul de Cercetari Pedologice si Agronomice. Metode, rapoarte, indrumari 12, 178 pp."]}

Enchodelus lucinensis Popovici, 1978 (Fig. 2) MATERIAL EXAMINED Four female paratypes, in acceptable condition, and two males (but see remarks) from Bihor Mountains, in a good state of conservation. MEASUREMENTS See Table 3. DESCRIPTION Female Moderately slender nematodes of medium size, 1.24- 1.46 mm long. Habitus after fixation curved ventrad, Cshaped. Body cylindrical, tapering towards both ends, but more so anteriorly. Cuticle 2.0 µm in anterior region, 3.0-4.5 µm at mid-body and 6.0-8.0 µm on dorsal side of tail, its outer layer with fine, but distinct transverse striations and much thinner than inner one, especially on tail. Lateral chord 11-15 µm wide or occupying one-fifth to one-fourth (21-26%) of mid-body diam., lacking any particular differentiation. Body pores obscure in material examined. Lip region somewhat angular, offset by a weak depression, 2.2-2.6 times as broad as high, ca one-fourth (22-28%) of body diam. at neck base. Lips amalgamated, labial and cephalic papillae slightly protruding above cephalic contour. Amphid fovea funnel-shaped, opening at level of cephalic depression, occupying 5-6 µm or ca one-half of corresponding body diam. Cheilostom almost cylindrical, with thick walls. Odontostyle comparatively robust, 10-14 times as long as wide and 1.6-1.9 times longer than lip region diam. or 1.4-1.7% of total body length, aperture small, 4-5 µm long or 14-17% of total length. Odontophore 1.4-1.5 times as long as odontostyle, lacking distinct basal thickenings or flanges (but see remarks). Guiding ring double, located at 10-12 µm or 1.1-1.3 lip region diam. from anterior end. Pharynx consisting of a slender muscular anterior portion gradually expanding, pharyngeal expansion occupying more than two-fifths (44-46%) of total neck length and ca one-half of corresponding body diam. Pharyngeal gland nuclei located as follows: DN = 61-65; S 1 N obscure; S 2 N = 83-86. Nerve ring at 85-112 µm from anterior end or 36-38% of total neck length. Cardia rounded conoid, nearly as long as broad, (10-15) × (11-14) µm in size. Genital system didelphic-amphidelphic; both branches equally and well developed: anterior branch 262-293 µm, posterior branch 310-353 µm long. Ovaries 80-109 µm long, usually reaching or slightly surpassing sphincter level; oocytes initially in two or more rows, then one. Oviduct 70-87 µm, or 1.3-1.4 corresponding body diam. long, consisting of a slender portion with prismatic cells and a moderately developed pars dilatata with visible lumen. Sphincter prominent, located between oviduct and uterus. Uterus long, 153-203 µm or 3.1-3.9 corresponding body diam., more or less twisted in females examined, tripartite, i.e., consisting of a wider proximal region with distinct lumen followed by a narrower and shorter intermediate portion with narrow lumen and surrounded by a cluster of hyaline cells, and ending with a well developed spheroid pars dilatata distalis. Sperm often abundant in proximal region of uteri. Vagina extending inwards 25-32 µm for one-half to three-fifths (51-61%) of body diam., pars proximalis longer than broad, (14-15) × (10-14) µm, usually with straight walls and enveloped by weak circular musculature, pars refringens with (in lateral view) two massive, granular, square sclerotisations measuring (5.0-6.0) × (6.0-7.0) µm and with a combined width of 13-14 µm; pars distalis 4.0 µm long. Vulva a transverse oval slit, preceded by a pronounced depression in body surface. Prerectum 2.2-2.9 and rectum 0.7-1.1 anal body diam. long. Tail short, rounded conoid, with abundant and distinct saccate bodies along ventral region, inner cuticle layer marked by radial striation, its ventral margin slightly irregular, barely separated from outer layer, cuticle 8-10 µm thick or 40-60% of tail length at terminus. Two pairs of subterminal caudal pores, one subdorsal, another practically lateral. Male (see remarks) General morphology similar to that of female, but with somewhat shorter body, 1.13 mm long. Genital system diorchic, with opposed testes. In addition to adcloacal pair, located at 10 µm from cloacal aperture, a series of seven or nine spaced ventromedian supplements, 8- 25 µm apart, extending anteriorly, posteriormost located at 17 or 22 µm from adcloacal pair, well within range of spicules. Other supplement located at level of anterior end of spicules. Spicules conspicuously robust, 3.8 or 4.2 times as long as wide and 2.1 or 2.3 anal body diam. long. Lateral guiding pieces 12 or 12.5 µm long, ca four times as long as wide. Tail somewhat more conoid than in female, saccate bodies present, but not so distinct (certainly due to fixation). DIAGNOSIS AND RELATIONSHIPS This diagnosis is based on the original description and a re-examination of the paratypes. Enchodelus lucinensis is distinguished by its 1.24-1.62 mm long body, lip region offset by weak depression and 11-14 µm diam., odontostyle 19-23 µm or 1.6-1.9 lip region diam. long and 1.4-1.7% of total body length, odontophore rod-like (but see remarks) and 29-32 µm long, neck length 238- 286 µm long, pharyngeal expansion 111-132 µm long or 42-46% of total neck length, female genital system amphidelphic, uterus long and tripartite, pars refringens vaginae with two massive and granular sclerotisations, V = 45-53, tail rounded conoid (18-21 µm long, c = 61- 76, c&lcedil; = 0.5-0.7), spicules 50-56 µm long and 7-9 spaced ventromedian supplements, the posteriormost one or two within the range of the spicules. In having a comparatively small body (E. lucinensis resembles E. teres Thorne, 1939 and E. parateres Baqri & Jairajpuri, 1974. It differs from E. teres, a nearly identical species (see remarks) in its longer pharyngeal expansion (vs ca two-fifths of total neck length) and shorter tail with less saccate bodies (vs 30 µm, c = 50, c&lcedil; = 1.0 as calculated from Thorne’s original drawing). Moreover, Thorne mentioned that the ovaries were “reflexed halfway to vulva”, an indication that the female genital tract is relatively short and the uterus (tentatively) not very long. It can be distinguished from E. parateres by its lip region offset by depression (vs deep constriction), longer pharyngeal expansion (vs 34- 40% of total neck length) and uterus tripartite (vs bipartite, lacking a spherical distal section). DISTRIBUTION Peat bog at Lucina nature reserve (Suceava District, Northern Moldavia; type locality) and mezohygrophilous meadow at Ghe&tcedil;ar-Sc&abreve;ri&scedil;oara, Bihor Mountains (Western Romanian Carpathians); sites 7 and 2 in Table 1. REMARKS As expected, the above description perfectly fits the original although new morphological details are added herein, particular for the female genital system. Some doubts persist as to the true identity and/or taxonomy of E. lucinensis. The precise nature of the odontophore base could not be assessed with accuracy in the available specimens, but the observations suggest that it is simple, rod-like, at most with weak basal thickening and without evidence of distinct knobs or flanges, i.e., in agreement with the original description. Its separation from E. teres is very problematic and based (see above) on minor differences. Unfortunately, available information about this species is very poor and many important morphological details are lacking, therefore preventing a detailed comparison. Zullini (1970) recorded E. teres from Italy, but only provided its basic ratios which, however, fit well with those of both E. lucinensis and E. teres. Until additional information about E. teres is available, separate status for these two species should be maintained. Some doubts persist as to the conspecificity of the two males from the Bihor Mountains with the four female paratypes due to their shorter body and, mainly, because of their separate geographical origin. However, the general morphology and morphometry are very similar, including lip region diam., odontostyle length, etc., and the morphology of male sexual structures is practically identical with the only male paratype known. Thus, they are provisionally considered to belong to E. lucinensis., Published as part of Ciobanu, Marcel, Popovici, Iuliana, Guerrero, Pablo & Santiago, Reyes Peña-, 2010, Nematodes of the order Dorylaimida from Romania. The genus Enchodelus Thorne, 1939. 2. Species with rounded tail and medium-sized odontostyle, pp. 381-397 in Nematology 12 (3) on pages 382-387, DOI: 10.1163/138855409X12549869072329, http://zenodo.org/record/8111660, {"references":["POPOVICI, I. (1978). New nematode species (Nematoda, Dorylaimida) from Romania. Nematologica 24, 404 - 411.","GUERRERO, P., LIEBANAS, G. & PENA- SANTIAGO, R. (2008 a). Nematodes of the order Dorylaimida from Andalucia Oriental, Spain. The genus Enchodelus Thorne, 1939. 2. Description of three known species with rounded tail and long odontostyle. Nematology 10, 451 - 470.","POPOVICI, I. (1995). New species of Tubixaba and Enchodelus (Nematoda: Dorylaimida) from Romania. Nematologica 41, 435 - 448.","THORNE, G. (1939). A monograph of the nematodes of the superfamily Dorylaimoidea. Capita Zoologica 8, 1 - 261.","BAQRI, Q. H. & JAIRAJPURI, M. S. (1974). Nematodes of high altitudes in India. V. Five new species of the genus Enchodelus Thorne, 1939 (Dorylaimida). Nematologica 20, 131 - 146.","ZULLINI, A. (1970). I nematodo muscicoli della Val Zebru' (Parco Nazionale dello Stelvio). Istituto Lombardo (Rendiconti Scienze) B 104, 88 - 137."]}

Enchodelus ameliae * sp. n. (Figs 4B; 5, 6) MATERIAL EXAMINED Three females and eight males from only one sample in Sierra Mágina, province of Jaén. MEASUREMENTS See Table 1. DESCRIPTION Female Moderately slender, medium-sized nematodes, 1.37- 1.55 mm long. Habitus after fixation curved ventrad, adopting a rather open C-shape. Body cylindrical, tapering towards both ends but more so anteriorly. Cuticle 2.5- 3.0 µ m in anterior region, 3.0 µ m at mid-body and 5.0-6.5 µ m on tail; its outer layer marked with very fine transverse striations, and thinner than inner one, especially at tail level. Lateral chord very narrow, ca 5 µ m wide or occupying 9% of mid-body diam., lacking any particular differentiation. Lateral pores obscure. Lip region moderately angular, offset by more or less marked constriction, 2.6-2.7 times as broad as high and ca one-third to two-fifths of body diam. at neck base. Lips amalgamated; labial and cephalic papillae distinct, visibly protruding above cephalic contour. SEM pictures showing a rhomboid oral aperture surrounded by elevated perioral area; inner labial papillae offset from adjacent surface by a circular striation, outer labial and cephalic papillae lacking this differentiation. Amphid fovea cup-shaped, opening at level of cephalic constriction, occupying ca one-half of corresponding body diam; under SEM, one pore visible a little posterior to amphid fovea surface. Cheilostom almost cylindrical but with slight discontinuity at anterior third. Odontostyle long and slender, 13- 17 times as long as wide or 1.8-1.9 times longer than lip region diam.; aperture small, occupying 8-9% of total length. Odontophore 1.2-1.4 times longer than odontostyle, provided with distinct basal flanges along posterior half, consisting of six sclerotised pieces with distinct basal knobs. Guiding ring double, situated at 18- 19 µ m or 1.1 times lip region diam. from anterior end. Pharynx consisting of slender muscular anterior part expanding gradually into basal expansion at 61-62% of total neck length, reaching full diam. at 68-70%; pharyngeal expansion occupying 34-35% of total neck length, ca one-half to two-thirds of corresponding body diam. Pharyngeal gland nuclei located as follows: DN = 68-71; S 1 N 1 and S 1 N 2 not observed; S 2 N 1 = 51-53; S 2 N 2 = 52- 57. Delicate membrane-like structure surrounding base of pharyngeal expansion. Cardia rounded conoid, practically as wide as long, 11-13 × 12-14 µ m. No greenish material inside intestine. Genital system didelphic-amphidelphic, both branches equally and well developed. Ovaries 92- 139 µ m long, reaching sphincter level; oocytes first in two or more rows, then one. Oviduct 82-123 µ m or 1.6- 2.3 corresponding body diam. long, consisting of slen- der portion with prismatic cells and well developed pars dilatata with distinct lumen and containing sperm. Distinct sphincter located between oviduct and uterus. Uterus long, 159-188 µ m or 2.9-3.6 times corresponding body diam., occasionally twisted; tripartite, i.e., consisting of: i) wide proximal region with distinct lumen; ii) followed by a narrower intermediate section with very narrow lumen and surrounded by long cluster of hyaline cells; and iii) ending with well developed spheroid pars dilatata distalis. No eggs observed. Two females with sperm inside proximal region of uterus and pars dilatata uteri. Vagina extending inwards for 48-62% of body diam.; pars proximalis longer than broad, 16-21 × 12-14 µ m, with sigmoid walls and surrounded by weak circular musculature; pars refringens consisting of (in lateral view) two arcuate drop-shaped sclerotisations with combined width of 17-19 µ m; pars distalis 3.0-5.0 µ m long. Vulva a transverse oval slit preceded by slight depression in body contour. Prerectum 3-4 and rectum 1.0-1.1 anal body diam. long. Tail rounded conoid; terminal cuticle 4.0-6.5 µ mthick or 28-31% of total tail length, its inner layer with barely visible radial striation, neither irregularities nor saccate bodies observed. Two pairs of subterminal caudal pores in posterior half of tail, one lateral, another subdorsal. * Named after the late Dr Amelia Ocaña, a zoologist, for her valuable contributions to nematology, especially on the taxonomy and ecology of freshwater nematodes. Male General morphology similar to female, albeit posterior region more curved, adopting a J-shape. Lateral chord very narrow, ca 2 µ m wide. Pharyngeal gland nuclei located as follows (n = 4): DN = 70-72; S 1 N 1 and S 1 N 2 not observed; S 2 N 1 = 51-57; S 2 N 2 = 55-61. Genital system diorchic, testes opposed. Sperm spindle shaped, 8.0-10.0 µ m long. In addition to acloacal pair, a series of 11-14 irregularly spaced ventromedian supplements present, posteriormost two being within spicule range; in two cases only posteriormost supplement located inside spicule range. Posterior supplement located at 18-25 µ m from adcloacal pair with latter at 9-11 µ m from cloaca aperture. Spicules dorylaimoid, 3.9-4.8 times as long as wide or 1.5-1.9 times longer than anal body diam. Lateral guiding pieces 11.5-14.5 × 3.0-3.5 µ m, with bifurcate tip. Prerectum longer than in female, 4-6 times anal body diam. Tail more conoid than in female, with straight ventral contour, terminal cuticle 4.0-6.0 µ m thick or 16- 26% of total tail length; cuticle lacking differentiations as in female. TYPE HABITAT AND LOCALITY Hedgehog heath at 1950 m a.s.l., in Sierra Mágina (province of Jaén, Spain). TYPE MATERIAL Two females (holotype and paratype) and six paratype males on slides 308-310, deposited in the Departamento de Biología Animal, Biología Vegetal y Ecología, Universidad de Jaén, Spain. One paratype female and two paratype males deposited in the Nematode Collection of the United States Department of Agriculture, Beltsville, MD, USA. DIAGNOSIS AND RELATIONSHIPS Enchodelus ameliae sp. n. is characterised by its body length of 1.37-1.55 mm, verynarrow lateral chord, slightly angular lip region offset by a constriction and 16-17 µ m broad, odontostyle 30-32 µ m long or 1.8-1.9 times lip region diam., odontophore with distinct basal flanges and 1.2-1.4 times longer than odontostyle, neck 282-307 µ m long, pharyngeal expansion 100-107 µ m long or 34-35% of total neck length, female genital system amphidelphic, uterus long and tripartite, pars refringens vaginae with two arcuate drop-shaped sclerotisations, vulva a transverse slit and almost equatorial (V = 47-50), tail rounded conoid (15-21 µ m long, c = 71-91, c &lcedil; = 0.6- 0.7 in females and 23-30 µ m long, c = 46-68, c &lcedil; = 0.7-1.0 in males), spicules 49-60 µ m long and presence of 11-14 irregularly spaced ventromedian supplements, the one or two posteriormost ones being located within the range of the spicules. Because of its short rounded conoid tail and medium sized odontostyle (ca 30 µ m) this species resembles E. altherri, E. analatus, E. arcticus, E. georgiensis, E. hopedoroides, E. hopedorus and E. ponorensis. However, it can be distinguished from them by the following characteristics: from E. altherri by its slightly longer body (1.37-1.55 vs 1.18-1.38 mm) and longer odontostyle (30- 32 vs 25-27 µ m), lip region offset by constriction vs slight depression, absence of saccate bodies on tail vs presence of small bodies and a greater number of ventromedian supplements (11-14 vs 7-9); from E. analatus by its lip region offset by constriction vs slight depression, shorter odontostyle, odontophore and total stylet (30- 31, 37-44 and 67-76, respectively, vs 34, 47 and 81 µ m, as estimated from original drawing), guiding ring located further posterior (18-19 µ m from anterior end vs 15 µ m, as estimated from original drawing) and males more frequent than females vs males not known; from E. arcticus by its broader lip region (16-17 vs 12 µ m, as estimated from original drawing) offset by constriction vs scarcely differentiated by a shallow depression, longer odontostyle, odontophore and total stylet (30-31, 37-44 and 67-76, respectively, vs 24-29, 29-33 and 56-64 µ m, as maximum range estimates from data on stylet length and original drawing) and presence of 11-14 ventromedian supplements vs absence [sic] of any supplements apart from the adanal pair; from E. georgiensis by its longer body (1.37-1.55 vs 0.88-1.10 mm), broader lip region (16- 17 vs ca 12 µ m), and longer spicules (49-60 vs 36-44 µ m); from E. hopedoroides by its narrower lip region (16-17 vs 18-20 µ m) offset by a constriction vs a slight depression, shorter neck (282-307 vs 317-442 µ m), shape of pars refringens vaginae (drop-shaped vs semicircular), shorter prerectum (95-111 vs 140-369 µ m) and males more frequent than females vs males not known; from E. hopedorus by its slightly broader lip region (16-17 vs 15-16 µ m) offset by a constriction vs depression, slightly shorter neck (282-307 vs 305-374 µ m), shorter tail (15-21 vs 27-39 µ m) lacking saccate bodies (vs present) and males frequent vs not known; and, finally, from E. ponorensis by its shorter body (1.37-1.55 vs 1.65-2.07 mm), longer odontostyle, odontophore and total stylet (30-32, 37-44 and 67-76, respectively, vs 23-25, 33- 38 and 55-63 µ m), shorter pharyngeal expansion (100- 107 vs 150-245 µ m), uterus tripartite vs a simple tube, as illustrated in the original description, although this feature must be interpreted with caution), shorter prerectum (95- 111 vs 107-281 µ m), shorter tail (15-21 µ m long and c &lcedil; = 0.6-0.7 vs 35-45 µ m long and c &lcedil; = 0.9-1.3) and frequent males vs absence.

Enchodelus hopedoroides Altherr, 1963 (Figs 1-3; 4A, D) MATERIAL EXAMINED Twenty-four females collected from the province of Granada. In addition, the holotype female deposited at the Museo Cantonale di Storia Naturale di Lugano (MCSN, Switzerland) was made available for study by courtesy of Ms Lucia Pollini. MEASUREMENTS See Table 1. DESCRIPTION Female Based on Andalusian individuals. Moderately slender to slender nematodes of medium size, 1.69-1.99 mm long. Habitus after fixation curved ventrad, adopting a C-shape. Body cylindrical, tapering towards both ends but more so anteriorly. Cuticle 4.0-6.0 µ m in anterior region, 3.0-5.0 µ m at mid-body and 7.0-9.5 µ m on tail; its outer layer with very fine transverse striations and much thinner than inner one, especially at tail level. Lateral chord 8-16 µ m wide or occupying 11-27% of mid-body diam., lacking any particular differentiation. Lateral pores obscure. Lip region more or less rounded, offset by shallow depression, 2.2-3.5 times as broad as high and ca one-third to two-fifths of body diam. at neck base. Lips amalgamated; labial and cephalic papillae visible, scarcely protruding above cephalic contour. Under SEM, oral aperture appearing as circular to square opening surrounded by a slight elevation of perioral area; inner labial papillae located on ring-like protuberances offset from adjacent surface by circular striation; outer labial papillae on similar but less differentiated areas; cephalic papillae lacking such differentiation. Amphid fovea cup-shaped, opening at level of cephalic depression, occupying two- to three-fifths of corresponding body diam; under SEM, one pore visible on amphid fovea surface. Cheilostom curly bracket-shaped. Odontostyle long and slender, 12-17 times as long as wide or 1.6-1.9 times longer than lip region diam.; aperture small, occupying 7-15% of total length. Odontophore 1.1- 1.4 times as long as odontostyle, bearing distinct basal flanges extending over posterior half of its total length and consisting of three sets of two sclerotised pieces with basal knobs. Guiding ring double, located at 19-23 µ m or 1.0-1.3 times lip region diam. from anterior end. Pharynx consisting of slender muscular anterior portion expanding gradually into basal expansion at 54-64% of total neck length, reaching full diam. at 61-70%; pharyngeal expansion occupying 34-43% of total neck length, ca twoto three-fifths of corresponding body diam. Pharyngeal gland nuclei located as follows (n = 6): DN = 61-67; S 1 N 1 = 32 (n = 1); S 1 N 2 = 43 (n = 1); S 2 N 1 = 50-57; S 2 N 2 = 50-58. Weak membrane-like structure surrounding base of pharyngeal expansion. Cardia rounded conoid, wider than long, 12-18 × 8-16 µ m. Some individuals with a cell mass located on dorsal surface, ca one body diam. posterior to cardia. Intestine not containing any greenish material. Genital system didelphic-amphidelphic, both branches equally and well developed. Ovaries 86-128 µ m long, normally not reaching sphincter level; oocytes first in two or more rows, then one. Oviduct 104-190 µ m or 1.6-2.7 corresponding body diam. long, consisting of slen- der portion with prismatic cells and well developed pars dilatata with distinct lumen. Sphincter prominent, located between oviduct and uterus. Uterus long, 136-255 µ m or 2.3-4.1 corresponding body diam., sometimes twisted, tripartite, i.e., consisting of: i) a wider proximal region with distinct lumen; ii) followed by a narrower and shorter intermediate portion with narrow lumen and surrounded by a cluster of hyaline cells; and iii) ending with a well developed, spheroid, pars dilatata distalis. Two females bearing one egg each, eggs measuring 113-116 × 34-36 µ m. Sperm not observed in genital tract. Vagina extending inwards for 42-67% of body diam.; pars proximalis longer than broad, 15-27 × 13-21 µ m, with sigmoid to straight walls and enveloped by weak circular musculature; pars refringens with (in lateral view) two massive trapezoidal sclerotisations with combined width of 15-21 µ m and an apparently granular texture; pars distalis 4.0-6.5 µ m long. Vulva a transverse oval slit, preceded by shallow depression in body surface. Prerectum 4-12 and rectum 0.8-1.7 anal body diam. long. Tail rounded conoid, with abundant saccate bodies, especially on ventral surface but sometimes extending to subdorsal region; terminal cuticle 8.0- 11.5 µ m thick or 25-54% of total tail length, its inner layer marked by radial striation and divided into two by a discontinuity along its middle; ventral margin of inner layer rather irregular, somewhat separated from outer. Two pairs of subterminal caudal pores, one subdorsal, another practically lateral. Male Unknown. Based on the original description and data of Andalusian individuals, E. hopedoroides is distinguished by its 1.42-1.99 mm long body, lip region offset by a shallow depression and 18-20 µ m diam., odontostyle 31-34 µ m long or 1.6-1.9 lip region diam. long, odontophore with distinct basal flanges and 1.1-1.4 times longer than odontostyle, neck length 317-442 µ m long, pharyngeal expansion 114- 161 µ m long or 34-43% of total neck length, female genital system amphidelphic, uterus long and tripartite, pars refringens vaginae with two trapezoidal sclerotisations, vulva a transverse pre-equatorial slit (V = 40-48), tail rounded conoid (16-33 µ m long, c = 47-117, c &lcedil; = 0.5- 0.9) with ventral margin of inner cuticle layer distinctly irregular and separated from outer layer and males unknown. Because of its short rounded conoid tail and medium sized odontostyle (ca 30 µ m), E. hopedoroides resembles E. altherri Vinciguerra & De Francisci, 1973, E. analatus (Ditlevsen, 1927) Thorne, 1939, E. arcticus Nesterov, 1976, E. distinctus Ahmad & Jairajpuri, 1980, E. georgiensis Eliava, Tskitishvili & Bagathuria, 2006, E. hopedorus (Thorne, 1929) Thorne, 1939 and E. ponorensis Popovici, 1995. However, it can be distinguished from these species by the following characters: from E. altherri by its longer body (1.42-1.99 vs 1.18-1.38 mm) and longer odontostyle (31-34 vs 25-27 µ m), broader lip region (18- 20 vs 14.5 µ m as calculated from original drawing), vulva located more anterior (V = 40-48 vs 50-52), longer prerectum (3.8-12.3 vs 1.5-2.5 times longer than anal body diam.) and males absent vs present; from E. analatus, a little known species described on the basis of only two females, by its guiding ring located further posterior (19- 23 vs 15 µ m) from the anterior end (as estimated from the original drawing), presence of flanges in odontophore vs absence – a striking feature which should be viewed with caution, vulva pre-equatorial (V = 40-48 vs 51), and longer prerectum (4-12 vs 1.5 times anal body diam., as calculated from original drawing); from E. arcticus by its longer body (1.42-1.99 vs 1.2-1.5 mm), longer odontostyle, odontophore and total stylet (31-34, 38-45 and 70- 79, respectively, vs 27-31, 29-33 and 56-64 µ m as maximum range estimates from data on stylet length and original drawing), vulva pre-equatorial (V = 40-48 vs 49-51), and absence of males vs presence; from E. distinctus, a species described on the basis of only a single female, by its lip region offset by a shallow depression (vs a clear constriction as depicted in the original drawing), slightly shorter odontostyle, odontophore and total stylet (31-34, 38-45 and 70-79, respectively, vs 36, 46 and 82 µ m), and vulva pre-equatorial (V = 40-48 vs 53); from E. georgiensis by its rather longer body (1.42-1.99 vs 0.88-1.10 mm), broader lip region (18-20 vs ca 12 µ m), longer odontostyle (31-34 vs 27-29 µ m) and absence of males vs presence; from E. hopedorus, the closest species and one recently redescribed by Guerrero and Peña-Santiago (2007), by its more rounded (vs moderately angular) and broader lip region (18-20 vs 15-16 µ m), odontostyle slightly but consistently longer (33 (31-34) vs 31 (29-33) µ m), guiding ring located posterior (20 (19-23) vs 18 (17-21) µ m), vulva located anteriorly (V = 40-48 vs 48-55), pars refringens vaginae differently shaped (trapezoidal and with granular texture vs drop-shaped and smooth), longer eggs (113-120 vs < 90 µ m long), longer prerectum (140-369 µ m long or 4-12 times anal diam. vs 62-162 µ m or 1.5- 4.5 times anal diam.) and tail more rounded and relatively shorter (c &lcedil; = 0.5-0.9 vs 0.7-1.0 and more conoid); and, finally, from E. ponorensis by its broader lip region (18- 20 vs 15-16 µ m) offset by a depression vs a constriction, rather longer odontostyle, odontophore and total stylet (31-34, 38-43 and 70-77, respectively, vs 23-25, 33-38 and 55-63 µ m), guiding ring located further posterior (19-23 vs 15-17 µ m from anterior end), uterus tripartite (vs a simple tube as illustrated in original description, although this feature should be considered with caution) and shorter tail (16-33 µ m long and c &lcedil; = 0.5-0.9 vs 35-45 µ m long and c &lcedil; = 0.9-1.3). DISTRIBUTION This species was collected in only one sampling point, a wet meadow on a glacial valley in Sierra Nevada, province of Granada, at 2800 ma.s.l. * Probably due to flattening. REMARKS To date, the only available information concerning E. hopedoroides was the original description by Altherr (1963) which was based on three females from Dischma valley, Switzerland, and a detailed drawing of the anterior region of a type specimen by Siddiqi (1969). After the death of Dr Altherr in 1976, his nematode collection was inherited by the Station Fédérale de Recherches Agronomiques de Changins, Nyon (Switzerland). It is now deposited in the Museo Cantonale di Storia Naturale di Lugano (Switzerland), under the charge of Ms Lucia Pollini, who kindly provided the holotype of E. hopedoroides for study. The original material is still in acceptable condition, although the pharynx is twisted, an obvious flattening of the body distorts some of the measurements and the low contrast prevents accurate observations on the genital system. General measurements and morphology, especially those relative to lip region, odontostyle, odontophore and guiding ring, fit well with the individuals collected in Sierra Nevada. Only minor differences have been observed, i.e., the body seems to be slightly longer in the Andalusian individuals (1.69-1.99 vs 1.42-1.85 mm) whereas the odontophore is somewhat shorter (38-43 vs 43-45 µ m)., Published as part of Guerrero, Pablo, Liébanas, Gracia & Santiago, Reyes Peña-, 2008, Nematodes of the order Dorylaimida from Andalucía Oriental, Spain. The genus Enchodelus Thorne, 1939. 3. Description of two new and one known species with rounded tail and medium-sized odontostyle, pp. 711-733 in Nematology 10 (5) on pages 712-718, DOI: 10.1163/156854108785787208, http://zenodo.org/record/8114910, {"references":["ALTHERR, E. (1963). Nematodes des sols forestiers subalpins du Val Dischma (Grisons). Bulletin de la Societe Vaudoise des Sciences Naturelles 68, 333 - 349.","VINCIGUERRA, M. T. & DE FRANCISCI, M. (1973). Nematodi muscicoli delle Alpi Apuane. Bollettino delle Sedute dell'Accademia Gioenia di Scienze Naturali in Catania. Serie IV 9, 1 - 24."]}

Enchodelus longispiculus * sp. n. (Figs 4C, E, F; 7, 8) MATERIAL EXAMINED Forty-eight females and 29 males collected in Sierra Nevada, province of Granada. MEASUREMENTS See Table 2. DESCRIPTION Female Moderately slender to stout, medium-sized nematodes, 1.32-1.87 mm long. Habitus after fixation slightly curved ventrad, adopting an open C- or J-shape. Body cylindrical, tapering towards both ends but more so anteriorly. Cuticle 3.5-5.5 µ m in anterior region, 3.0-4.5 µ m at mid-body and 5.5-9.5 µ m on tail; its outer layer with very fine transverse striations and much thinner than inner, especially at tail level. Lateral chord relatively narrow, 7.0-12.0 µ m wide or occupying 9-17% of mid-body diam., granular, lacking any particular differentiation. Lateral pores obscure. Lip region moderately angular, offset by shallow depression, 2.4-3.7 times as broad as high and ca oneto two-fifths of body diam. at neck base. Lips amalgamated; labial and cephalic papillae distinct, moderately protruding above cephalic contour. Under SEM, oral aperture rhomboid, located on moderate elevation of perioral area; inner and outer labial papillae surrounded by ring-like differentiation of body surface offset by circular striation; cephalic papillae on less marked areas. Amphid fovea cup-shaped, opening at level of cephalic depression, occupying one-half to three-fifths of corresponding body diam. Under SEM, pore visible slightly posterior to amphidial fovea surface. Cheilostom cylindrical but with marked constriction at its middle. Odontostyle long and slender, 12-16 times as long as wide or 1.6-1.9 times longer than lip region diam.; aperture small, occupying 5-13% of total length. Odontophore 1.2-1.5 times longer than odontostyle, with distinct basal flanges over posterior half of its total length, consisting of six sclerotised pieces with basal knobs. Guiding ring double, located at 16-19 µ m or 0.9-1.2 times lip region diam. from anterior end. Pharynx consisting of slender muscular anterior portion expanding gradually into basal expansion at 61-70% of total neck length, reaching full diam. at 66-74%; pharyngeal expansion occupying 28-37% of total neck length and ca two- to three-fifths of corresponding body diam. Pharyngeal gland nuclei located as follows (n = 22): DN = 69-75; S 1 N 1 = 26-31 (n = 3); S 1 N 2 = 33- 37 (n = 3); S 2 N 1 = 51-60; S 2 N 2 = 53-60. Weak membrane-like structure surrounding base of pharyngeal expansion. Cardia rounded conoid, wider than long, 11- 19 × 10-17 µ m. Intestine containing olive green or brownish material. Genital system didelphic-amphidelphic, both branches equally and well developed. Ovaries 78-293 µ m long, usually surpassing sphincter level; oocytes first in two or more rows, then in one. Oviduct 95-231 µ m or 1.3-3.1 times corresponding body diam. long, consisting of slender portion with prismatic cells and well developed pars dilatata with distinct lumen. Distinct sphincter between oviduct and uterus. Uterus long, 160-283 µ m or 2.2-3.9 times corresponding body diam., sometimes twisted; tripartite, i.e., consisting of: i) a wider proximal region with distinct lumen; ii) followed by a narrower and shorter intermediate portion with narrow lumen and surrounded by a cluster of hyaline cells; and iii) ending with a well developed spheroid pars dilatata distalis. Fourteen females bearing one to three eggs measuring 75-87 × 32- 46 µ m. Thirty out of 40 females containing sperm inside genital system. Vagina extending inwards for 41-64% of body diam.; pars proximalis longer than broad, 20- 34 × 11-26 µ m, with sigmoid to straight walls, surrounded by weak circular musculature; pars refringens with (in lateral view) two arcuate drop-shaped sclerotisations with combined width of 16-24 µ m and divided into proximal and distal area by change in refringence; pars distalis 1.5- 6.0 µ m long. Vulva a transverse oval slit. Prerectum 2-5 and rectum 0.8-1.6 anal body diam. long. Tail rounded conoid; terminal cuticle 6.5-14.0 µ m thick or 28-76% of total tail length, its inner layer marked by radial striation and divided into two by a discontinuity along its middle; aspect of cuticle rather variable, ranging from presence of saccate bodies or small irregularities to no differentiation at all; some individuals showing very faint discontinuity dividing inner layer of cuticle along its middle. Two pairs subterminal caudal pores, one lateral, another subdorsal. * The specific epithet refers to the relatively long male spicules. Male General morphology similar to female, although posterior region more curved, adopting a J- or G-shape. Lateral chord narrower than in female, 3.0-8.0 µ m wide or occupying 5-9% of body diam. Pharyngeal gland nuclei located as follows (n = 20): DN = 70-75; S 1 N 1 = 21- 27 (n = 2); S 1 N 2 = 32-37 (n = 2); S 2 N 1 = 48- 61; S 2 N 2 = 52-63. Genital system diorchic, testes opposed. Sperm spindle shaped, 8.5-11.0 µ m long. In addition to acloacal pair, a series of 8-14 more or less regularly spaced ventromedian supplements present, posteriormost two being within spicule range; in few cases ventromedian supplements irregularly spaced forming two or three groups; in three specimens, only one supplement located within spicule range. Posterior supplement located at 15- 35 µ m from adcloacal pair, latter being 10-14 µ m from cloaca aperture. Spicules dorylaimoid, relatively long due to extension of dorsal surface of head (manubrium), 4.3- 5.4 times as long as wide and 1.9-2.3 times longer than anal body diam. Lateral guiding pieces 13.5-17.5 × 3.5- 4.5 µ m, with bifurcate tip. Prerectum considerably longer than in female, 4-8 times anal body diam. Tail rounded conoid, practically straight ventrally, terminal cuticle 5.5- 11.5 µ m thick or 20-47% of total tail length; aspect of cuticle variable as in female, although saccate bodies, if present, generally less pronounced. TYPE HABITAT AND LOCALITY Wet meadow at 2925 ma.s.l. in Sierra Nevada (province of Granada, Spain). Enchodelus macrodorus also occurred in this site. OTHER HABITATS AND LOCALITIES The species has been collected from three other samples, all in Sierra Nevada, province of Granada, viz. a gorse scrubland at 2700 m a.s.l. and two wet meadows at 2590 and 2600 ma.s.l. TYPE MATERIAL Twenty females (holotype and paratypes) and five paratype males on slides 311-317, deposited in the Departamento de Biología Animal, Biología Vegetal y Ecología, Universidad de Jaén, Spain. Four paratype females and three paratype males deposited in the Nematode Collection of the United States Department of Agriculture, Beltsville, MD, USA. DIAGNOSIS AND RELATIONSHIPS Enchodelus longispiculus sp. n. is characterised by its 1.32-1.87 mm long body, slightly angular lip region offset by a shallow depression and 15-18 µ m broad, odontostyle 27-31 µ m long or 1.6-1.9 times lip region diam., odontophore with distinct basal flanges and 1.2- 1.5 times longer than odontostyle, neck 251-315 µ m long, pharyngeal expansion 77-114 µ m long or 28-37% of total neck length, female genital system amphidelphic, uterus long and tripartite, pars refringens vaginae with two arcuate drop-shaped sclerotisations, vulva a transverse slit and pre-equatorial (V = 41-48), tail rounded conoid (17- 29 µ mlong, c = 49-90, c &lcedil; = 0.6- 0.8 infemalesand 20-29 µ m long, c = 54-71, c &lcedil; = 0.6- 0.8 in males), spicules 63-72 µ m long and presence of 8-14 regularly spaced ventromedian supplements, the two posteriormost being within the range of the spicules. Its short rounded conoid tail and medium sized odontostyle (ca 30 µ m) make this species similar to E. altherri, E. ameliae sp. n., E. analatus, E. arcticus, E. georgiensis, E. hopedoroides, E. hopedorus and E. ponorensis. Nevertheless, it differs from these species by the following features: from E. altherri by its slightly longer body (1.32- 1.87 vs 1.18-1.38 mm) and longer odontostyle (27-31 vs 25-27 µ m), vulva located more anterior (V = 41-48 vs 50-52) and longer spicules (63-72 vs 50-52 µ m); from E. ameliae sp. n., certainly a very similar species, by its more robust body (a = 17-25 and mid-body diam. 64-80 µ m vs a = 26-29 and mid-body diam. 52-54 µ m), lip region offset by a slight depression vs constriction, vulva located anterior (V = 41-48 vs 47-50) and longer spicules (63-72 vs 49-60); from E. analatus by its shorter odontostyle, odontophore and total stylet (27-31, 36-44 and 64-74, respectively, vs 34, 47 and 81 µ m, as estimated from original drawing), vulva pre-equatorial (V = 41- 48 vs 51) and frequent presence of males vs not known; from E. arcticus, by its longer odontostyle, odontophore and total stylet (27-31, 36-44 and 64-74, respectively, vs 24-29, 29-33 and 56-64 µ m, as maximum range estimates from data on stylet length and original drawing), vulva pre-equatorial (V = 41-48 vs 49-51), longer spicules (63- 72 vs 58 µ m), and presence of 8-14 ventromedian supplements vs absence [sic] of any supplements apart from the adanal pair; from E. georgiensis by its longer body (1.32- 1.87 vs 0.88-1.10 mm), broader lip region (15-18 vs ca 12 µ m) and much longer spicules (63-72 vs 36-44 µ m); from E. hopedoroides by its slightly narrower lip region (16-18 vs 18-20 µ m), shorter odontostyle and total stylet (27-31 and 64-74, respectively vs 31-34 and 70-77 µ m), guiding ring located further anterior at 16-19 vs 19-23 µ m from anterior end, shorter neck (251-315 vs 317-442 µ m), shape of pars refringens vaginae (drop-shaped vs semicircular), slightly shorter prerectum (65-162 vs 140-369 µ m) and frequent presence of males vs not known; from E. hopedorus by its more anterior vulva (V = 41-48 vs 48-55), slightly broader lip region (16-18 vs 15-16 µ m), relatively shorter odontostyle (1.6-1.9 vs 1.9-2.2 times lip region diam.), guiding ring located further anterior at 16- 19 vs 18-21 µ m from anterior end, slightly shorter neck (251-315 vs 305-374 µ m), shorter tail (17-29 vs 27-39 µ m) and males frequent vs not known; and, finally, from E. ponorensis by its stouter body (a = 17-25 vs 30-39 and mid-body diam. 64-80 vs 50-61 µ m), lip region offset by a depression vs constriction, odontostyle, odontophore and total stylet slightly longer (27-31, 36-44 and 64-74, respectively vs 23-25, 33-38 and 55-63 µ m), shorter pharyngeal expansion (77-114 vs 150-245 µ m), uterus tripartite vs a simple tube, as illustrated in original description – a rather striking condition and to be interpreted cautiously, shorter tail (17-29 µ m long and c &lcedil; = 0.6-0.8 vs 35-45 µ m long and c &lcedil; = 0.9-1.3) and frequent presence of males vs absence.

Enchodelus striatus Thorne, 1939 ( Figs 12, 13 ) MATERIAL EXAMINED Fourteen females, 15 males and a juvenile. Of them, three females and eight males on slide labelled Enchodelus striatus 8, and ten females, six males and one juvenile on slide labelled Enchodelus striatus 8b, both from Broads Fork and Twin Peaks, Utah, USA, collected in August 1924; one female and one male on slide labelled Enchodelus macrodoroides 2e, collected from summit soil in Lone Peak, Utah, USA, on 4 July 1926. Several specimens are in quite good condition, but most are damaged to some extent. MEASUREMENTS See Table 5. Female Moderately slender to slender nematodes of medium size, 1.5-2.0 mm long. Body cylindrical, tapering towards both ends but more so towards posterior. Cuticle 2.5- 3.5 µ m thick at anterior region, 2.5-4.5 µ m at mid-body and 4.0-6.0 µ m at tail; its outer layer provided with coarse transverse striations. Lateral chord 9-13 µ m wide or one-sixth to one-fifth of mid-body diam., lacking any particular differentiation. Lateral pores obscure. Lip region offset by marked constriction, sometimes appearing narrower than adjacent body, 2.1-2.8 times as wide as high and ca one-third of body diam. at neck base; perioral area weakly differentiated; lips practically amalgamated; labial and cephalic papillae rounded but protruding. Amphid fovea cup-shaped, opening at level of cephalic constriction and occupying ca half of lip region diam. Cheilostom a truncate cone, with no particular differentiation. Odontostyle slender, 14-21 times as long as wide or 2.5-3.0 times as long as lip region diam.; aperture small, less than one-tenth of total length. Odontophore rod-like, lacking any specialisation, 1.0-1.2 times the odontostyle length. Guiding ring apparently double, at 19.5-21.5 µ m or 1.4- 1.6 times lip region diam. from anterior end. Pharynx consisting of a slender but well muscled anterior portion expanding gradually into basal expansion at 60-65% of total neck length, reaching full diam. at 67-72%; pharyngeal expansion occupying 32-37% of total neck length, and ca half of corresponding body diam. Pharyngeal gland nucleilocatedas follows: DN = 71-75; S 1 N 1 =?; S 1 N 2 =?; S 2 N 1 = 47-53; S 2 N 2 = 49-55. Base of pharyngeal expansion surrounded by faint membrane-like structure. Cardia rounded conoid, practically as long as wide (11- 16 × 8-14 µ m). Genital system didelphic-amphidelphic, with both branches equally and well developed. Ovaries relatively large, 105-218 µ m long, usually not reaching sphincter level; oocytes first in two or more rows, then in one. Oviduct 123-218 µ m long or 2.0-4.4 times corresponding body diam., consisting of slender portion with prismatic cells and well developed pars dilatata bearing distinct lumen and containing sperm. Distinct sphincter present between oviduct and uterus. Uterus bipartite, i.e., with a wider proximal portion having distinct lumen and containing sperm, and a more slender distal part with narrow lumen, refractive inner lining and completely surrounded by a group of hyaline cells; length 114-223 µ m or 2.1-3.3 times the body diam. at its level. Vagina extending inwards one-third to one-half body diam.; pars proximalis vaginae 13-22 × 12-22 µ m, with sigmoid walls and enveloped by moderately developed circular musculature; pars refringens vaginae with (in lateral view) two triangular to trapezoidal sclerotisations with a combined width of 12-16 µ m; pars distalis vaginae well developed, 4.0-8.0 µ m long. Vulva a transverse slit, preceded by a depression in body contour; vulval lips distinctly raised. Prerectum 3.1-5.9, rectum 0.9-1.2 anal body diam. long. Tail conical, regularly ventrad curved, with rounded tip; hyaline terminal portion 12-19 µ m or 24-36% of total length. Two pairs of caudal pores at the middle of tail; one subdorsal, another practically lateral. ). Male General morphology similar to female, but with posterior body region more ventrally curved, G-shaped. Genital system diorchic, with opposite testes. In addition to adanal pair a series of 6-9 regularly spaced ventromedian supplements present outside of spicule range. Spicules dorylaimoid, relatively slender, ventrally curved, 5.1-7.7 times as long as wide and 1.3-1.9 times anal body diam. long. Lateral guiding pieces 12-14 µ m long and ca four times as long as wide, with bifurcate tip. Tail regularly ventrad curved, with rounded tip; hyaline portion occupying 12- 14 µ m or 19-25% of total length. DIAGNOSIS Enchodelus striatus is distinguished by its body 1.5- 2.0 mm long, outer cuticle with coarse transverse striations along the entire body, lip region offset by constriction and 12.5-14.5 µ m diam., odontostyle 35-38 µ m long or 2.5-3.0 times the lip region diam., odontophore lacking distinct flanges and equal in length to odontostyle, neck length 267-316 µ m long, pharyngeal expansion occupying 93-109 µ m or ca one-third (32-37%) of total neck length, female genital system amphidelphic, uterus bipartite, pars refringens vaginae with distinct trapezoidal sclerotisations, vulva in form of a transverse slit (V = 46-52), tail conical and regularly ventrally curved (46-59 µ m, c = 28-36, c&lcedil; = 1.6-1.9 in females; 51-63 µ m, c = 25-32, c&lcedil; = 1.5-1.8 in males), spicules 49-58 µ m long and presence of 6-9 regularly spaced ventromedian supplements. REMARKS Thorne (1939) originally described this species and provided the only report. The above description fits well with the original, although of course many new morphological details have been added. Concerning the odontophore, Thorne wrote “often appearing almost flanged” (see also his Fig. 80); however, our observations do not confirm any major specialisation of this structure. Other minor morphometric differences have also been observed. Enchodelus striatus might be endemic to the Neartic region., Published as part of Guerrero, Pablo & Santiago, Reyes Peña-, 2007, Redescription of Enchodelus species studied by Thorne in 1939 (Dorylaimida: Nordiidae), pp. 93-121 in Nematology 9 (1) on pages 116-120, DOI: 10.1163/156854107779969646, http://zenodo.org/record/8111793, {"references":["THORNE, G. (1939). A monograph of the nematodes of the superfamily Dorylaimoidea. Capita Zoologica 8, 1 - 261."]}

Enchodelus arcuatus Thorne, 1939 ( Figs 2, 3 ) MATERIAL EXAMINED Two females, one mounted onslide labelled Enchodelus arcus, and collected from Arlington Farm, Virginia (USA) on April 8, 1931, in quite good condition. Another female mounted on slide labelled Enchodelus arcus, and collected from Granddaddy Lake, Utah (USA) in August, 1924, in poor condition. MEASUREMENTS See Table 2. DESCRIPTION Female Moderately slender nematodes of medium size, 1.5, 2.0 mm long. Body cylindrical, tapering towards both ends but more so towards posterior. Cuticle 2.5, 3.5 µ m thick at anterior region, 3.0, 4.0 µ m at mid-body and 4.0, 6.0 µ m at tail; its outer layer thinner than inner, with very fine transverse striations. Lateral chord 13 µ m wide or ca one-fifth of mid-body diam., lacking any particular differentiation. Lateral pores obscure. Lip region rounded, offset by shallow constriction, 2.8, 3.0 times as broad as high and ca one-third of body diam. at neck base; lips practically amalgamated; perioral area weakly differentiated; labial and cephalic papillae hardly perceptible. Amphid fovea cup-shaped, opening at level of cephalic constriction and occupying almost two-thirds of corresponding body diam. Cheilostom a truncate cone, with no particular differentiation. Odontostyle slender, slightly dorsad curved; its length 10.9, 12.9 times as long as wide or 1.5, 1.6 times as long as lip region diam.; aperture small, ca one-tenth of total length. Odontophore rod-like, lacking any specialisation, 1.3, 1.4 times the odontostyle length long. Guiding ring apparently double, at 10.5-12.0 µ m, or 0.9 times lip region diam., from anterior end. Pharynx consisting of a slender, but well muscled portion expanding gradually into basal expansion at 54, 58% of total neck length, reaching full diam. at 63, 67%; pharyngeal expansion occupying ca two-fifths (38, 42%) of total neck length, and ca three-fifths of corresponding body diam. Pharyngeal gland nuclei located as follows: DN = 70, 65; S 1 N 1 = 23; S 1 N 2 = 27; S 2 N 1 = 51, 48; S 2 N 2 = 53, 49. Base of pharyngeal expansion surrounded by a weakly refractive membrane-like structure. Cardia rounded conoid, practically as long as wide (8 µ m). Intestine containing abundant green-coloured material, mainly in posterior portion. Genital system didelphic-amphidelphic, with both branches equally and well developed. Ovaries relatively large, 87-139 µ m long, but not reaching sphincter level; oocytes first in two or more rows, then in one row. Oviduct 108-147 µ m long or 1.6-2.4 times corresponding body diam., consisting of a slender portion with prismatic cells and a well developed pars dilatata with distinct lumen. Sphincter clearly visible between oviduct and uterus; a group of hyaline cells surrounding sphincter and distal part of uterus. Uterus bipartite, i.e., consisting of a wider proximal portion with distinct lumen, and a narrower distal part with narrow lumen and refractive inner lining, portion close to sphincter bearing very small refractive granules; uterus length 133-161 µ m or 2.0-2.7 times body diam. at its level. Sperm not observed in genital tract. Uterine egg measuring 35, 38 × 80, 81 µ m (n = 2). Vagina extending inwards to ca two-fifths of body diam.; pars proximalis vaginae 11.5, 13.5 × 16.5, 15.0 µ m, with almost straight walls and enveloped by moderately developed circular musculature; pars refringens vaginae with (in lateral view) two, rather close, trapezoidal sclerotisations that have a combined width of 12.0, 18.0 µ m; pars distalis vaginae short, 2.5, 5.0 µ m long. Vulva a transverse slit. Prerectum 2.1, 2.9; rectum 1.1, 1.2 anal body diam. long. Tail conical, regularly ventrad curved, with rounded tip; hyaline terminal portion 16, 23 µ m or 26, 29% of total length. Two pairs of caudal pores at the middle of tail: one subdorsal, another practically lateral. Male Unknown. DIAGNOSIS Enchodelus arcuatus is distinguished by its body 1.5- 2.0 mm long, lip region 12-14 µ m diam., odontostyle 18.5-22 µ m long or 1.5-1.6 times the lip region diam., odontophore rod-like and 1.3-1.4 times the odontostyle, neck length 287-340 µ m long, pharyngeal expansion 108- 142 µ m or occupying ca two-fifths (38-42%) of total neck length, female genital system amphidelphic, uterus bipartite, pars refringens vaginae with two well developed trapezoid sclerotisations, vulva in form of a transverse slit and slightly posterior (V = 52-54), tail conical and regularly ventrad curved (62-80 µ m, c = 24-25, c&lcedil; = 2.0- 2.2), and males unknown. REMARKS Thorne (1939) provided measurements, description and illustrations of the female collected from Arlington Farm, and mentioned that he had studied another two females from Kailab Forest, Arizona, and three from Tryal Lake, Uinta Mountains, Utah. One of the two females examined here is, without doubt, that from Arlington Farm; another might be one of the three collected from Uinta Mountains. After its original description, E. arcuatus was later reported from several habitats and localities. Andrássy (1958) studied one Bulgarian female having longer prerectum (4-5 ABD) and shorter tail (c = 35.9). Also Andrássy (1959) described one female from Romania having longer body (L = 2.2), more anterior vulva (V = 47) and shorter tail (c = 32.7, c&lcedil; = 1.8). Zullini (1970) reported fourteen females from Italy having a more anterior vulva (V = 49-52, n = 3) and shorter tail (c = 27-35, n = 3). Taking into account that the differences observed are morphometric and with no large gap among them, and that only a few specimens have hitherto been studied, it is tentatively assumed that these four records belong to the same species, although European material can be distinguished from that from America by a shorter tail. On the other hand, Vinciguerra and De Francisci (1973) and Vinciguerra and La Fauci (1978) described two Italian populations characterised by a smaller body (L = 1.1-1.5, n = 18), shorter odontostyle (13-16 µ m), pars refringens vaginae apparently absent (“la vagina... e la sua estremità prossimale è cuticolarizzata, ma non sclerificata”), and male as frequent as the female. This material is certainly not conspecific with those referred above., Published as part of Guerrero, Pablo & Santiago, Reyes Peña-, 2007, Redescription of Enchodelus species studied by Thorne in 1939 (Dorylaimida: Nordiidae), pp. 93-121 in Nematology 9 (1) on pages 94-100, DOI: 10.1163/156854107779969646, http://zenodo.org/record/8111793, {"references":["THORNE, G. (1939). A monograph of the nematodes of the superfamily Dorylaimoidea. Capita Zoologica 8, 1 - 261.","ANDRASSY, I. (1958). Erd- und Susswassernematoden aus Bulgarien. Acta Zoologica Academiae Scientiarum Hungaricae 4, 1 - 88.","ANDRASSY, I. (1959). Freilebende Nematoden aus Rumanien. Annales Universitatis Scientiarum Budapestinensis Rolando Eotvos, Sectio Biologia, pp. 3 - 27.","ZULLINI, A. (1970). I nematodi muscicoli della ' Val Zebru' (Parco Nazionale dello Stelvio). Istituto Lombardo (Rediconti Scienze) B 104, 88 - 137.","VINCIGUERRA, M. T. & DE FRANCISCI, M. (1973). Nematodi muscicoli delle Alpi Apuane. Bolletino dell'Accademia Gioenia di Scienze Naturali in Catania 9, 1 - 24.","VINCIGUERRA, M. T. & LA FAUCI, G. (1978). Nematodi muscicoli dell'Isola di Lampedusa (Sicilia). Animalia 5, 13 - 37."]}

Enchodelus hopedorus (Thorne, 1929) Thorne, 1939 = Dorylaimellus hopedorus Thorne, 1929 ( Figs 8, 9 ) MATERIAL EXAMINED Nineteen females, mounted on slide labelled Enchodelus hopedorus 5c, and collected from Longs Peak, Colorado, USA, in summit soil, ont 19 July 1924; in acceptable condition although the specimens have become somewhat flattened. MEASUREMENTS See Table 4. DESCRIPTION Female Moderately stout nematodes of medium size, 1.4- 1.8 mm long. Habitus after fixation slightly ventrad curved. Body cylindrical, tapering towards both ends but more so towards anterior. Cuticle 2.5-3.0 µ m thick at anterior region, 2.5-3.5 µ m at midbody and 4.5-6.5 µ m at tail; its outer layer practically smooth or bearing very fine transverse striations, and much thinner than inner, especially at level of tail. Lateral chord relatively narrow, occupying 13-18% of midbody diam., lacking any particular differentiation. Lateral pores obscure. Lip region practically continuous or barely offset by a shallow depression, 2.4-3.1 times as broad as high and ca one-third of body diam. at neck base. Lips amalgamated, with their inner portions visibly projecting; labial and cephalic papillae distinct. Amphid fovea cup-shaped, opening at level of cephalic depression and occupying one-half to three-fifths of corresponding body diam. Cheilostom a truncate cone, with thickened walls. Odontostyle long and slen- der, with distinct walls and narrow lumen, slightly curved; 11-16 times as long as wide or 1.9-2.2 times as long as lip region diam.; aperture small, one-twelfth to one-tenth of total length. Odontophore lacking basal flanges but with basal portion slightly thickened, clearly separated from pharyngeal lining; 1.3-1.6 times longer than odontostyle. Guiding ring double, at 17-21 µ m or 1.1-1.3 times lip region diam. from anterior end. Pharynx consisting of a slender but well muscular anterior portion expanding gradually into basal expansion at 59-64% of total neck length, reaching full diam. at 65-70%; pharyngeal expansion occupying 33-38% of total neck length, and one-half to four-fifths of the corresponding body diam. Pharyngeal gland nuclei located as follows: DN = 66-72; S 1 N 1 =?; S 1 N 2 =?; S 2 N 1 = 52-63; S 2 N 2 = 53-64. Weak membrane-like structure surrounding base of pharyngeal expansion. Cardia very small, rounded conoid, as wide as long. In some specimens, intestine containing greenish material. Genital system didelphic-amphidelphic, with both branches equally and well developed. Ovaries relatively large, 69-187 µ m long, but usually not reaching sphincter level; oocytes first in two or more rows, then in one row. Oviduct 79-156 µ m long and consisting of a slender portion with prismatic cells and a well developed pars dilatata bearing distinct lumen. Sphincter distinct, located between oviduct and uterus. Uterus long, 189-270 µ m or 2.9-4.2 times corresponding body diam.; tripartite with wider proximal portion with distinct lumen followed by narrower and often twisted intermediate length region with narrow lumen joining a well developed spheroid pars dilatata distalis. Sperm not observed in genital tract. Vagina extending inwards 39-55% of body diam.; pars proximalis 12-17 × 16-25 µ m, with almost straight or somewhat sigmoid walls, enveloped by weak circular musculature; pars refringens with (in lateral view) two close together drop-shaped sclerotisations which have a combined width of 13-19 µ m; pars distalis short, 3.0-6.0 µ m long. Vulva a transverse slit. Prerectum 1.5-4.6, rectum 0.8-1.2 anal body diam. long. Tail rounded conoid, slightly more straight ventrally; numerous saccate bodies distinct in tail, most in ventral side. Two pairs of caudal pores, both subdorsal, one at middle of tail, other subterminal. Male Unknown. DIAGNOSIS Enchodelus hopedorus is distinguished by its body 1.4- 1.8 mm long, lip region practically continuous and 14.5- 16 µ m diam., odontostyle 29-33 µ m long or 1.9-2.2 times the lip region diam., odontophore lacking distinct basal flanges and longer (1.3-1.6 times) than the odontostyle, neck length 305-374 µ m long, pharyngeal expansion 106- 136 µ m and occupying 33-38% of total neck length, female genital system amphidelphic, uterus long and tripartite, pars refringens vaginae with two drop-shaped sclerotisations, vulva in form of a transverse slit and equatorial (V = 48-55), tail rounded conoid (27-39 µ m, c = 41-53, c&lcedil; = 0.7-1.0) with numerous saccate bodies, and males unknown. REMARKS There is no doubt concerning the identity of these specimens with those Thorne (1939) refers to in his description. The re-examination of this material has provided some remarkable differences from Thorne’s original description. The lip region is practically continuous, not “set off by slight constriction”; in fact, Thorne’s Figure 70 shows a constriction, but the specimens are viewed in dorsal or ventral position, not laterally. The presence of basal flanges in the odontophore is not confirmed, because the nature of odontophore base resembles that observed in E. macrodoroides (see under E. geraldi) rather than the corresponding to E. macrodorus, as can be seen if we compare his Figures 70a, 68c and 76b, respectively. After its original description, E. hopedorus was reported from Poland (Brzeski, 1963; Winiszewska-Slipinska, 1987) and Nepal (Zullini, 1973), but some doubts persist concerning the identity of non-American material. Brzeski (1963) studied two females having shorter odontostyle (20-23 µ m) and neck (b = 5.9-7.1), and slightly anterior vulva (V = 46); he also described two males. Zullini (1973) recorded one female, comparatively smaller (L = 1.15), and having a shorter odontostyle (21 µ m) and more anterior vulva (V = 43). Also, Winiszewska-Slipinska (1987) studied three females having a shorter neck (b = 5.0-5.7), relatively longer tail (c&lcedil; = 1.1-1.2) and more anterior vulva (V = 45-46)., Published as part of Guerrero, Pablo & Santiago, Reyes Peña-, 2007, Redescription of Enchodelus species studied by Thorne in 1939 (Dorylaimida: Nordiidae), pp. 93-121 in Nematology 9 (1) on pages 108-112, DOI: 10.1163/156854107779969646, http://zenodo.org/record/8111793, {"references":["THORNE, G. (1929). Nematodes from the summit of Long's Peak, Colorado. Transactions of the American Microscopical Society 48, 181 - 195.","THORNE, G. (1939). A monograph of the nematodes of the superfamily Dorylaimoidea. Capita Zoologica 8, 1 - 261.","DE MAN, J. G. (1880). Die einheimischen, frei in der reinen Erde und im sussen Wasser lebenden Nematoden. Tijdschrift Nederlandsche Dierkundige Vereeniging 5, 1 - 104.","BRZESKI, M. (1963). [Further studies on nematodes (Nematoda) of the Sphagnaceae of the Tatra Mountains.] Fragmenta Faunistica 10, 309 - 315.","ZULLINI, A. (1973). Su alcuni nematodi di alta quota del Nepal. Khumbu Himal 4, 401 - 413."]}

Enchodelus brevidentatus Thorne, 1939 ( Figs 4, 5 ) MATERIAL EXAMINED Nineteen females, 18 of them mounted on one slide labelled Enchodelus brevidens, and collected from foothill soil in Lehi, Utah (USA) on March 8, 1926; a few specimens in acceptable condition, but most of them becoming flattened. The remaining female, mounted on slide labelled Enchodelus macrodoroides 2a, and collected from Wellsville Hill, Utah (USA) on March 28, 1925, is in quite bad condition. MEASUREMENTS See Table 2. DESCRIPTION Female Moderately slender nematodes of medium size, 1.6- 1.9 mm long. Body cylindrical, tapering towards both ends but more so towards posterior. Cuticle 2.0-3.0 µ m thick at anterior region, 2.0-4.0 µ m at midbody and 3.5- 6.0 µ m at tail; its outer layer thinner than inner, bearing fine but distinct transverse striations that are more visible at level of anterior body region and caudal region. Lateral chord occupying 18-30% of midbody diam, with granular appearance. Lateral pores obscure. Lip region rather angular, offset by distinct depression, 2.4-2.9 times as broad as high and ca one-third (some specimens ca one-fifth, but obviously flattened) of body diam. at neck base; lips mostly amalgamated; labial and cephalic papillae protruding, in particular the inner labial ones. Amphid fovea cup-shaped, opening at level of cephalic depression and occupying one-third to one-half of corresponding body diam. Cheilostom a truncate cone to practically cylindrical, with no particular differentiation. Odontostyle relatively slen- der, practically straight; 7.7-11.3 times as long as wide or 1.1-1.4 times as long as lip region diam.; aperture small, one-tenth to one-fifth of total length. Odontophore rod-like, lacking any specialisation, 1.2-1.5 times odontostyle length long. Guiding ring simple, at 8.5-10.5 µ m or 0.7- 0.8 lip region diam. from anterior end. Pharynx consisting of a slender but well muscular portion expanding gradually into basal expansion at 54-65% of total neck length, reaching full diam. at 62-70%; pharyngeal expansion occupying 33-42% of total neck length, and ca three-fifths of corresponding body diam. Pharyngeal gland nucleilocatedas follows: DN = 67-72 (n = 10); S 1 N 1 = 22; S 1 N 2 = 31 (n = 1); S 2 N 1 = 49-55; S 2 N 2 = 50- 57 (n = 7). Base of pharyngeal expansion surrounded by faint membrane-like structure. Cardia rounded conoid, wider than long, 16 × 11 µ m. Genital system didelphic-amphidelphic, with both branches equally and well developed. Ovaries relatively large, 109-320 µ m long, usually reaching and surpassing sphincter level; oocytes first in two or more rows, then in one row. Oviduct 120-273 µ m long or 2-3 times corresponding body diam., consisting of a slender portion with prismatic cells and a well developed pars dilatata with distinct lumen. Sphincter present between oviduct and uterus. Uterus bipartite, i.e., with a wider proximal portion with distinct lumen, and a narrower distal part with narrow lumen and refractive inner lining; length 65-126 µ m or 0.9-2.0 times body diam. at its level; a group of hyaline cells observed surrounding distal part of uterus. Sperm not observed in genital tract. Only one uterine egg observed in female from Wellsville, measuring 91 × 35 µ m. Vagina extending inwards one-third (flattened specimens) to two-fifths of body diam.; pars proximalis vaginae 11-16 × 13-25 µ m, with almost straight or slightly convergent walls and enveloped by moderately developed circular musculature; pars refringens vaginae with (in lateral view) two trapezoidal sclerotisations which have a combined width of 9.5-15.5 µ m and are somewhat separated by a weakly sclerotised intermediate area; pars distalis vaginae short, 2.5-5.0 µ m long. Vulva a transverse slit. Prerectum 1.8-3.2, rectum 0.8-1.2 anal body diam. long. Tail conical, regularly ventrad curved, with rounded tip; hyaline terminal portion 10-15 µ m or 15-21% of total length. Two pairs of caudal pores at the middle of the tail: one subdorsal, another practically lateral. Male Unknown. Enchodelus brevidentatus is distinguished by its body 1.6-1.9 mm long, lip region 12-13.5 µ m diam., odontostyle 14.5-17 µ m long or 1.1-1.4 times the lip region diam., odontophore rod-like, neck length 264-308 µ m long, pharyngeal expansion 91-125 µ m long or occupying 33-42% of total neck length, female genital system amphidelphic, uterus bipartite, pars refringens vaginae with two trapezoidal sclerotisations, vulva in form of a transverse slit (V = 47-53), tail conical and regularly ventrad curved (55-71 µ m, c = 24-33, c&lcedil; = 1.3-2.1), and males unknown. REMARKS The material examined is certainly the original described by Thorne (1939) and, as a consequence, the above description fits that by the American author, although ratio a is lower (vs a = 30-35) due to flattening of the specimens. Since its original description, E. brevidentatus has been recorded from Italy (Vinciguerra, 1972), Hungary (Andrássy, 1996) and Spain (Liébanas et al., 2004). The material described by Vinciguerra differs from type population in several relevant features: shorter body (L = 1.2-1.5) and odontostyle (10-12 µ m) and males as frequent as females. Some doubts therefore exist as to the true identity of the Italian material., Published as part of Guerrero, Pablo & Santiago, Reyes Peña-, 2007, Redescription of Enchodelus species studied by Thorne in 1939 (Dorylaimida: Nordiidae), pp. 93-121 in Nematology 9 (1) on pages 100-103, DOI: 10.1163/156854107779969646, http://zenodo.org/record/8111793, {"references":["THORNE, G. (1939). A monograph of the nematodes of the superfamily Dorylaimoidea. Capita Zoologica 8, 1 - 261.","VINCIGUERRA, M. T. (1972). Nematodi di Sicilia. Nota I. Bolletino delle Setude dell'Accademia Gioenia di Scienze Naturali in Catania 11, 3 - 35.","ANDRASSY, I. (1996). Free-living nematodes in the Bukk Mountains, Hungary. The Fauna of the Bukk National Park, 1996, pp. 33 - 63.","LIEBANAS, G., GUERRERO, P., MARTIN- GARCIA, J. - M. & PENA- SANTIAGO, R. (2004). Spatial distribution of dorylaimid and mononchid nematodes from the southeast Iberian peninsula: environmental characterization of chorotypes. Journal of Nematology 36, 114 - 122."]}

Enchodelus macrodorus (de Man, 1880) Thorne, 1939 = Dorylaimus macrodorus de Man, 1880 = Dorylaimus (Doryllium) macrodorus ( de Man, 1880 ) Ditlevsen, 1928 = Dorylaimellus macrodorus ( de Man, 1880 ) Thorne & Swanger, 1936 ( Figs 10, 11 ) Nine females, five of them mounted on a slide labelled Enchodelus macrodorus 1h, and collected from Aspen Creek, Utah, USA, on 22 June 1932, in very good condition. The other four females mounted on slide labelled Enchodelus macrodorus 1d, and collected from Strawberry Reservoir, Utah, USA, in sage brush soil, in September, 1924, three of them also in good condition. MEASUREMENTS See Table 4. DESCRIPTION Female Moderately stout nematodes of medium size, 1.5- 1.7 mm long. Habitus after fixation slightly ventrad curved. Body cylindrical, tapering towards both ends but more so towards anterior. Cuticle 3.0-4.5 µ m thick at anterior region, 3.5-4.5 µ m at mid-body and 8.5-11.0 µ m at tail; its outer layer practically smooth or bearing very fine transverse striations, and much thinner than inner, specially at level of tail. Lateral chord very narrow, less than one-tenth of mid-body diam., lacking any particular differentiation. Lateral pores obscure. Lip region rounded, offset by a marked depression, 2.5-3.5 times as broad as high and ca one-third of body diam. at neck base. Lips amalgamated, perioral area hardly differentiated; labial and cephalic papillae visibly protruding from lip region contour. Amphid fovea cup-shaped, opening at level of cephalic depression and occupying one-half to two-thirds of corresponding body diam. Cheilostom a truncate cone, with thickened walls. Odontostyle long and slender, with distinct walls and narrow lumen; 11- 18 times as long as wide or 1.9-2.4 times as long as lip region diam.; aperture small, ca one-twelfth of total length. Odontophore bearing distinct flanges occupying ca half of total length; scarcely longer (1.0-1.2 times) than odontostyle. Guiding ring double, at 25-28 µ m or 1.3-1.5 times lip region diam. from anterior end. Pharynx consisting of slender but well muscled anterior portion expanding gradually into basal expansion at 59-62% of total neck length, reaching maximum diam. at 64- 68%; pharyngeal expansion occupying 35-38% of total neck length, and one-half to two-thirds of corresponding body diam. Pharyngeal gland nuclei located as follows: DN = 65-68; S 1 N 1 = 24-36; S 1 N 2 = 27-37; S 2 N 1 = 54-59; S 2 N 2 = 57-61. Base of pharyngeal expansion surrounded by weak membrane-like structure. Cardia very small, rounded conoid, and wider than long (12- 19 × 9-16 µ m). Genital system didelphic-amphidelphic, with both branches equally and well developed. Ovaries relatively large, 83-188 µ m long, but usually not reaching sphincter level; oocytes first in two or more rows, then one. Oviduct 97-159 µ m long, consisting of slender portion with prismatic cells and well developed pars dilatata with distinct lumen. Sphincter distinct, located between oviduct and uterus. Uterus short, tripartite: wider proximal portion having distinct lumen followed by a narrower intermediate region with narrow lumen joining a small spheroid pars dilatata distalis; length 61-143 µ m or 1.1-2.0 times body diam. at its level. Sperm not observed in genital tract. Two eggs observed inside both genital branches of one female, measuring 57 × 92 and 55 × 87 µ m. Vagina extending inwards to ca half (47-57%) of body diam.; pars proximalis 17-21 × 21-27 µ m, with almost straight or somewhat sigmoid walls, and enveloped by moderately developed circular musculature; pars refringens with (in lateral view) two close trapezoidal sclerotisations with a combined width of 19-20 µ m; pars distalis short, 4.0-6.0 µ m long. Vulva a transverse slit. Prerectum long, 3.9-5.8 anal body diam. Rectum almost equal to anal body diam. Tail short rounded to hemispheroid. Two pairs of caudal pores, both subterminal, one subdorsal, another subventral; most specimens bearing some small irregularities between inner and outer layers of cuticle, mainly at ventral side of the tail, sometimes appearing as vesicles in dorsal view. Male Not seen. DIAGNOSIS Enchodelus macrodorus is distinguished by its body 1.5-1.7 mm long, lip region offset by marked depression and 17.5-20 µ m diam., odontostyle 38-43 µ m long or 1.9-2.4 times the lip region diam., odontophore provided with distinct basal flanges and slightly longer (1.0-1.2 times) than odontostyle, neck length 303-345 µ m long, pharyngeal expansion 111-130 µ m long and occupying 35-38% of total neck length, female genital system amphidelphic, uterus tripartite, pars refringens vaginae with two sclerotisations, vulva in form of a transverse slit and uniquely anterior (V = 41-44), and tail short rounded to hemispherical (18-24 µ m, c = 68-91, c&lcedil; = 0.5-0.6). REMARKS Thorne (1939) wrote that this species was “common at elevations of 8,000 to 12,000 feet in Wasatch and Uinta Mountains, Utah, U.S.A.”, and, although he did not mention the two localities from where the specimens studied herein were collected, it seems more than likely that this material belongs to that studied by him since his morphometric and morphological details fit perfectly with those given above. Thorne (1939) described the male of the species but unfortunately nomale was found in the two available slides from the USDA Nematode Collection. The present description and that of Thorne (1939) of American specimens of E. macrodorus apparently fit very well that of de Man (1880), although, of course, the latter is not very detailed. Enchodelus macrodorus seems to be very widely distributed in Holartic areas., Published as part of Guerrero, Pablo & Santiago, Reyes Peña-, 2007, Redescription of Enchodelus species studied by Thorne in 1939 (Dorylaimida: Nordiidae), pp. 93-121 in Nematology 9 (1) on pages 112-116, DOI: 10.1163/156854107779969646, http://zenodo.org/record/8111793, {"references":["DE MAN, J. G. (1880). Die einheimischen, frei in der reinen Erde und im sussen Wasser lebenden Nematoden. Tijdschrift Nederlandsche Dierkundige Vereeniging 5, 1 - 104.","THORNE, G. (1939). A monograph of the nematodes of the superfamily Dorylaimoidea. Capita Zoologica 8, 1 - 261.","DITLEVSEN, H. J. (1928). Land- and freshwater nematodes. In: Jensen, A. S., Lundbeck, W. & Mortensen, Th. (Eds). Zoology of the Faroes 13, pp. 1 - 28.","THORNE, G. & SWANGER, H. H. (1936). A monograph of the nematode genera Dorylaimus Dujardin, Aporcelaimus n. g., Dorylaimoides n. g., and Pungentus n. g. Capita Zoologica 6, 1 - 220."]}