Nematodes of the order Dorylaimida from Andalucía Oriental, Spain. The genus Enchodelus Thorne, 1939. 2. Description of three known species with rounded tail and long odontostyle Author Guerrero, Pablo Departamento de Biología Animal, Biología Vegetal y Ecología, Universidad de Jaén, Campus ‘ Las Lagunillas’ s / n, Edificio B 3, 23071 Jaén, Spain Author Liébanas, Gracia Departamento de Biología Animal, Biología Vegetal y Ecología, Universidad de Jaén, Campus ‘ Las Lagunillas’ s / n, Edificio B 3, 23071 Jaén, Spain Author Santiago, Reyes Peña- Departamento de Biología Animal, Biología Vegetal y Ecología, Universidad de Jaén, Campus ‘ Las Lagunillas’ s / n, Edificio B 3, 23071 Jaén, Spain text Nematology 2008 2008-01-01 10 4 451 470 journal article 57140 10.1163/156854108784513815 2eb64730-71cc-4e01-8a33-9f9995fa41d6 8111856 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 3 , 4 ) MATERIAL EXAMINED Eleven females from Sierra Nevada (province of Granada ). MEASUREMENTS See Table 2 . SEM OBSERVATIONS Lip region considerably elevated. Oral aperture appearing as a dorso-ventral rhomboidal slit. Perioral region slightly elevated, with very subtle radial striae. Labial and cephalic papillae located on rather small elevations of lip region surface, comparatively less marked than in E. groenlandicus and E. saxifragae ; circular striae surrounding inner labial papillae, forming a ring-like structure; but outer labial and cephalic papillae located on less differentiated areas. One lateral pore observed at base of amphid fovea. Vulva an oval transverse aperture preceded by a slight depression in body contour. DISTRIBUTION The species was collected in three samples in Sierra Nevada (province of Granada ); two of them in wet meadows at 2925 and 1950 m a.s.l., and one from river bank soil around poplar at 1400 m a.s.l. It co-occurred with E. groenlandicus in one sample. REMARKS Iberian individuals are practically identical to American material collected by Thorne during the years 1924-1932 and redescribed by Guerrero and Peña-Santiago (2007), except for minor differences in morphometry such as the broadening of body length range (1.38-1.92 vs 1.49-1.70 mm ) and, consequently, the de Man indexes; slightly more posterior vulva (V = 42-46 vs 41-44); shorter uterus (56- 115 µ m long or 0.9-1.6 times corresponding body diam. vs 61-143 µ m long or 1.1-2.0 timesbody diam.); shorter prerectum (112-176 vs 162-212 µ m); and longer tail (22- 28 vs 18-24 µ m). These differences can be explained by the scarcity of available material (eight American and ten Spanish individuals). Fig. 4. Enchodelus macrodorus (de Man, 1880) Thorne, 1939 (female). A: Variability of anterior region; B: Neck showing dorsal gland nucleus (large arrow) and one of subventral gland nuclei (small arrow); C: Anterior genital branch showing sphincter between uterus and oviduct (arrow); D: Detail of uterus-oviduct junction, displaying sphincter (arrow) and weak development of uterus; E: Vulval region; F; Entire body; G: Variability of tail. (Scale bars: A, D, E, G = 20 µm; B = 100 µm; C = 50 µm; F = 250 µm.) Table 2. Morphometric data for female Enchodelus macrodorus (de Man, 1880) Thorne, 1939. All measurements are in µm (except L in mm) and in the form: mean ± standard deviation (range).

Parameter	Population	Total range
Sierra Nevada	Sierra Nevada	Sierra Nevada
1400 m a.s.l.	1950 m a.s.l.	2925 m a.s.l.
(Granada)	(Granada)	(Granada)
n	1	7	2	10
L	1.38	1.47 ± 0.07	1.86, 1.92	1.54 ± 0.19
(1.39-1.57)	(1.38-1.92)
a	23.1	23.1 ± 1.9	22.6, 22.4	23.0 ± 1.6
(20.5-26.0)	(20.5-26.0)
b	4.7	4.5 ± 0.2	5.5, 6.1	4.8 ± 0.6
(4.3-4.8)	(4.3-6.2)
c	62.8	62.9 ± 4.9	72.9, 69.7	64.6 ± 5.4
(57.8-72.2)	(57.8-72.9)
cļ	0.6	0.6 ± 0.0	0.7, 0.6	0.6 ± 0.0
(0.6-0.7)	(0.6-0.7)
V	43.7	44.3 ± 1.4	43.5, 42.6	44.0 ± 1.3
(41.9-46.0)	(41.9-46.0)
Lip region diam.	18.0	18.9 ± 0.6	19.0, 19.5	18.9 ± 0.7
(18.0-19.5)	(18.0-19.5)
Odontostyle length	42	40.5 ± 1.6	44, 43	41.2 ± 1.9
(38-43)	(38-44)
Odontophore length	43	43.1 ± 1.5	47, 48	44.0 ± 2.2
(42-46)	(42-48)
Stylet total length	84	84 ± 1.6	91, 91	85 ± 3.3
(81-85)	(81-91)
Guiding ring from ant. end	26	26.8 ± 0.8	28, 29	27.1 ± 1.2
(25-28)	(25-29)
Neck length	293	325 ± 19	338, 312	322 ± 19
(288-344)	(288-344)
Pharyngeal expansion length	115	125 ± 9	131,?	124 ± 9
(113-138)	(113-138)
Diam. at neck base	53	59.0 ± 3.0	72, 77	61.0 ± 7.5
(55-63)	(53-77)
– at mid-body	60	63.6 ± 3.7	82, 86	67.2 ± 9.4
(60-69)	(60-86)
– at anus	35	36.7 ± 2.1	39, 42	37.3 ± 2.6
(34-41)	(34-42)
Prerectum length	176	137 ± 18	?,?	143 ± 22
(112-156)	(112-176)
Rectum length	38	35.7 ± 5.0	?,?	36.0 ± 4.7
(30-42)	(30-42)
Tail length	22	23.4 ± 1.3	26, 28	23.9 ± 1.9
(22-26)	(22-28)

THE IDENTITYOF ENCHODELUS MACRODORUS In spite of being the type species of the genus, identification of E. macrodorus is rather problematical. The original description by de Man (1880) was useful at the time to distinguish between a generalised Enchodelus species with rounded tail and the vast number of dorylaim species which were then classified within the genus Dorylaimus (including representatives of Dorylaimidae , Aporcelaimidae , Qudsianematidae , Nordiidae or even Nygolaimidae ). No illustrations were provided in 1880, but they were included in a subsequent work (de Man, 1884). Unfortunately, it seems that de Man’s slides containing individuals of E. macrodorus were lost before Loof (1961) revised the nematode collection of de Man, since this species is not redescribed in that work. This has been the most cited Enchodelus species , reported from all of the Holarctic region, from the USA (Thorne, 1939) to Korea (Choi et al. , 1997). However, the ambiguity of its diagnosis probably led to misidentifications and some of the records may correspond to other species. Asummary of E. macrodorus records, including morphometric data, is presented in Table 3 . Some remarks follow below. Table 3. Previous records for Enchodelus macrodorus (de Man, 1880) Thorne, 1939. All measurements are in µm (except Lin mm). Data marked with an asterisk (*) are calculated either from de Man’s indexes or from illustrations. Abbreviations: LRD = lip region diam., Odont. = odontostyle, Phar. exp. = pharyngeal expansion, abd = anal body diam., Ventr. sup. = number of ventromedian supplements, D = Denmark, No = Norway.

n	L	a	b	c	V	cļ LRD Odont. Odont./ Total					Neck Phar. abd Prerectum Tail Spicule Ventr. Country Reference
LRD	stylet	exp.		sup.
? ♀♀	1.8	25	4.5-5	70-80	< 50	< 1?	?	?	?	?	360-400*?		?	?	24*	–	–	Netherlands	1
? ♀♀	1.8	25	4.5-5	70-80	44*	0.6*	?	?	2.07*	?	400* 135*		?	?	24*	–	–	Netherlands	2
2 ♂♂	1.56-1.69	35	5.3-5.5 80-100 –			?	?	?	?	?	300*	?	?	?	18*	?	6	Netherlands	3
1 ♀	1.55	23	5.5	60	45*	?	?	?	?	?	280	?	?	?	26	–	–	Germany	4
? ♀♀	1-1.2	25-30	4-6	50-90	< 50	?	?	?	?	?	<300*	?	?	?	16*	–	–	Switzerland	5
? ♂♂	1.3-2.16	27-35	4-6	60-80	–	?	?	?	?	?	> 215*	?	?	?	25*	?	6-10
4 ♀♀	1.44-1.53	27-31	4.65	53-64 45-49?			?	?	?	?	310-330*?		?	?	25*	–	–	Germany	6
5	41-45 0.5-0.6 17-20			42-43	2.2*	94* 340*		?	36-45 175-256 19-28			–	–
3 ♂♂	1.36-1.41	28-31	4.6	47-64	–	?	?	?	?	?	295-310*?		?	?	25*	?	5-7
10 ♀♀	0.82-1.01	26-34	3.6-5.2	35-46	< 50	?	?	?	?	?	< 280*	?	?	?	23*	–	–	Switzerland	7
21 ♀♀	1.23-1.93	18-35	4.5-6	53-71	< 50	?	?	< 35*	?	69-70	300*	?	?	?	25*	–	–	Poland	8
15 ♂♂ 1.12-1.8		19-27	4.5-5.5	41-57	–	?	?	”	?	”	290*	?	?	?	30*	?	9-14
? ♀♀	1.64	18-35	5.5	53-71	?	?	?	?	?	?	300*	?	?	?	23-31*	–	–	Romania	9

Previous records considered to represent the true E. macrodorus Taking into account that holotype and paratypes were so briefly described and that they are probably lost, the diagnosis of the species must be inferred from other sources. The only reports from the original country of description, i.e. , The Netherlands , are those by de Man (1880, 1884 and 1912, the latter cited by Menzel, 1914) and Bongers (1988), so they are considered to correspond with the true E. macrodorus. In a recent work, two of the authors of the present paper had the opportunity to redescribe American individuals of Enchodelus species collected by Thorne (Guerrero & Peña-Santiago, 2007), and, among them, eight females originally identified as E. macrodorus , which apparently fit well with specimens from The Netherlands . From this starting point, the most relevant characters for E. macrodorus are body length of ca 1.5-1.8 mm , odontostyle ca 40 µ m long, and a particularly anterior located vulva (V = ca 40-44). The following records agree well with this short diagnosis and can be considered to be conspecific with E. macrodorus : allpopulations reported by Thorne (1939), Jairajpuri and Loof (1968), Ahmad and Jairajpuri (1980), Nasira et al. (1992), and Ahmad et al. (2002). In addition, three of the populations reported by Popovici (1995), specifically those composed of nine and five females , and that by four females measuring 1.55-1.73 mm offer no reason to reject this attribution. Table 3. (Continued).

n	L a b c V	cļ LRD Odont. Odont./ Total					Neck	Phar.	abd Prerectum Tail Spicule Ventr. Country Reference
LRD	stylet	exp.	sup.
LRD	exp.
1 ♂	???? –	?	?	?	?	?	?	?	?	?	?	?	11
1 ♀	2 19 4.3 4.5 [sic] 45	?	?	?	?	?	465*	?	?	?	?	–	–	Faroe Isl. (D)	10
8 ♀♀	1.49-1.70 20-28 4.5-5.4 62-91 41-44 0.5-0.6 17-20 38-43				1.9-2.4 85		303-345 111-130 36-41 162-212				18-24	–	–	USA	11, 12
1 ♂	1.5 28 5 67 –	0.6*	?	?	ļļ	?	300*	?	?	?	22*	?	12	11
12 ♀♀	1.17-1.36 23-27 4.5-5.3 52-65 44-50	?	?	20-29	?	?	260*	?	?	?	22*	–	–	Hungary	13
7 ♂♂	1.42 27 5.9 57.5 –	?	?	24-35	?	?	240*	?	?	?	25* 46-60		9-13
? ♀♀	1.92 27 4.8 69 47	0.7	?	?	?	?	400*	?	40*	?	28*	–	–	Jan Mayen (No)	14
1 ♀	1.96 32 4.3 54 45	?	?	?	?	91*	455*	?	?	?	36*	–	–	Switzerland	15
? ♀♀	1.32-1.93 22-27 4-5 67-91 46-49	?	?	40-50	2.5 100-110* 265-483* 132-241*?					? 15-29* –			–	Uzbekistan	16
? ♂♂	???? –	?	?	?	?	?	?	?	?	?	? 50-60		6-13
? ♀♀	???? 47-51	?	?	?	?	?	?	?	?	?	?	–	–	Sweden	17
1 ♀	1.47 25 4.8 67 37	?	?	42	2.5	88*	305*	?	?	?	22*	–	–	India	18
2 ♀♀	1.64-1.65 24-25 4.8-5.4 79-98 42-44	?	?	?	?	? 305-345*?			?	? 17-21* –			–	Italy	19
1 ♀	2.00 28 5.3 69 42	?	?	48	?	103*	375*	?	?	279	29*	–	–	Spitzbergen (No)	20
1 ♀	1.5 26 4.3 68 44	?	?	?	?	87*	350*	?	?	?	22*	–	–	USA	21
? ♀♀	1.1-1.93 23-37 4.5-6.0 50-71 43-50	?	27*	52*	1.9*	110*	211*	83*	51*	?	33*	–	–	Moldova	22
? ♂♂	1.2-1.8 19-35 5.3-5.9 41-100 –	?	?	?	?	?	?	?	?	?	?	?	11
15 ♀♀	1.42-1.79 24-32 4.5-5.2 55-75 40-45 0.65*		41-42 1.7-1.8* 84-88*			330*	125*	38*	?	25*	–	–	India	23
47 ♀♀	0.90-1.32 17-25 3.8-5.5 32-64 45-52	?	?	27-37	?	?	240*	?	?	?	23*	–	–	Italy	24
19 ♂♂ 0.94-1.22 20-32 3.6-5.3 32-66 –		?	?	ļļ	?	?	240*	?	?	?	22*	?	7-11
2 ♀♀	1.6-1.8 21-22 5.0-5.3 42-52 46 0.7-0.9		?	46	?	?	330*	?	?	?	36*	–	–	Poland	25
? ♀♀	1.6-1.8 24-28 4.5-5.0 62-80 42	0.6	?	40	1.9* 90-100* 360*			?	?	?	?	–	–	Netherlands	26
1? ♂	1.5 28 5 67 –	0.6	?	ļļ	ļļ	?	300*	?	37*	?	22*	?	11
9 ♀♀	1.47-1.65 19-25 4.4-5.2 59-81 42-45 0.5-0.6		?	33-43	2.2*	78-92	325* 110-120* 180-249				18-22	–	–	Pakistan	27
9 ♀♀	1.52-1.87 24-28 4-5 64-83 41-47 0.5-0.7 15-19 37-48				2.5*	90*	360*	?	36-40 200-331		21-25	–	–	Romania	28
4 ♀♀	1.24-1.48 20-24 4.5-4.8 52-63 47-50 0.6-0.8 15-18 29-35				2.1*	75*	290*	?	33-36	87-194	23-25	–	–
4 ♂♂	1.29-1.40 23-28 4.4-4.7 48-62 – 0.6-0.8 15-16 32.5				2.1*	76*	300*	?	33-36 150-205		22-28 52-55		8-9
2 ♀♀	1.48-1.61 26-27 5.2-5.4 64-66 42-48 0.6-0.7		15	32.5	2.2*	74*	290*	?	34-38	106	22-25	–	–
6 ♂♂	1.44-1.67 24-29 5.4-6.0 58-66 – 0.7-0.8 15-18 25-27				1.6*	65*	290*	?	32-38 109-194		24-28 55-70		7-11
5 ♀♀	1.58-1.67 24-27 4.5-4.7 56-8
1 ♀	1.32 26 5 59 48	0.7	15	25	1.7*	63*	260*	?	33	93	23	–	–
1 ♀	1.75 29 4.9 80 45	0.7	18	33	1.9*	70*	360*	?	32	125	22	–	–
1 ♂	1.70 39 5.4 62 –	0.9	13	28	2.2*	65*	315*	?	31	172	28	53	9
4 ♀♀	1.55-1.73 22-26 4.2-4.5 59-67 41-47 0.6-0.7		20	42-46	2.2*	96*	380*	?	42-44 234-288		25-30	–	–
1 ♂	1.62 33 5 52 –	0.9	15	33	2.2*	68*	324*	?	35	163	31	63	9
13 ♀♀	1.47-1.77 25-32 4.3-5.9 47-63 48-55	0.7* 16-18* 35-40			2.1*	78*	318-370 150-170* 36-47			?	25-34	–	–	Korea	29
15 ♀♀	1.38-1.54 21-26 4.0-4.9 57-73 41-43 0.5-0.7 17-19 42-47				2.5*	90*	304-361 116-144 38-43 114-222				20-29	–	–	China	30

References: (1) de Man (1880); (2) de Man (1884); (3) de Man (1912) (in Menzel, 1914); (4) Brakenhoff (1914); (5) Menzel (1914); (6) Schneider (1923); (7) Kreis (1924); (8) Stefański (1924); (9) Stefański (1927); (10) Ditlevsen (1928); (11) Thorne (1939); (12) Guerrero and Peña-Santiago (2007); (13) Andrássy (1952); (14) Allgén (1953); (15) Altherr (1953); (16) Tulaganov (1958) (in Tulaganov and Usmanova, 1978); (17) van Rossen and Loof (1961); (18) Jairajpuri and Loof (1968); (19) Zullini (1970); (20) Loof (1971); (21) Thorne (1974); (22) Nesterov (1979); (23) Ahmad and Jairajpuri (1980); (24) Vinciguerra (1984); (25) Winiszewska-Slipinska (1987); (26) Bongers (1988); (27) Nasira et al . (1992); (28) Popovici (1995); (29) Choi et al . (1997); (30) Ahmad et al . (2002). Previous records which probably do not correspond to E. macrodorus Menzel (1914): body too short in females (L = 1-1.2 mm ) and excessively variable in males (L = 1.3-2.16 mm ). May correspond to more than one species. Schneider (1923): vulva located not especially anterior (V = 45.5-48.5). Kreis (1924): body exceedingly short (L = 0.82-1.01 mm ). Stefański (1924): odontostyle and total stylet length too short (total stylet length is reported to be 69-70 µ m. Since odontophore is normally as long as or longer than odontostyle, the latter will barely reach 35 µ m long). Andrássy (1952): body length somewhat short (L = 1.15-1.36 mm in females, 1.42 mm in males); vulva located not particularly anterior (V = 44-50); and odontostyle exceedingly short (20-29 µ m in females, 24-35 µ m in males). Van Rossen and Loof (1961): vulva located not particularly anterior (V = 47-51). Nesterov (1979): vulva not always located particularly anterior (V = 43-50), odontostyle too long (52 µ m as estimated from original illustration), lip region expanded and rather broad (27 µ m as calculated from drawing). Vinciguerra (1984): body too short (L = 0.90-1.32 mm in females and 0.94-1.22 mm in males); vulva located practically equatorial (V = 45-52); and odontostyle exceedingly short (27-37 µ m). Popovici ( 1995 in part): all populations with odontostyle shorter than 36 µ m, including all described males. Some of the females have, in addition, the body rather short and vulva located quite posterior. Choi et al. (1997): vulva located practically equatorial (V = 48-55); odontostyle somewhat short (35-40 µ m). Previous records doubtful or with insufficient data to assess identity Tulaganov (1958, cited in Tulaganov and Usmanova, 1978): vulva located only slightly pre-equatorial (V = 46- 49); and odontostyle rather too long (40-50 µ m). Loof (1971): odontostyle length seems to be somewhat long (48 µ m, but compare with odontostyle length 42-47 µ m in individuals from China described by Ahmad et al. , 2002). Remaining measurements fit well with diagnosis of E. macrodorus . Winiszewska-Slipinska (1987): odontostyle length somewhat long (46 µ m, see above) and c index apparently too low (c = 42-52 vs 70-80 in original description, which leads to a rather long tail, i.e ., 36 µ m as estimated from L and c index vs 24 µ m as estimated from the original description and 18-24 µ m in American individuals reported by Guerrero and Peña-Santiago (2007). Identityof populations describedby Brakenhoff (1914), Stefański (1927), Ditlevsen (1928), Allgén (1953), Altherr (1953), Zullini (1970) and Thorne (1974) cannot be confirmed due to lack of information. As a curiosity, Coomans (1985) beautifully illustrated the anterior region of E. macrodorus in his study on longidorid phylogeny. The specimens used for the drawings were provided by Loof, but nothing is said about their origin and no morphometric data are available. EMENDED DIAGNOSIS OF E. MACRODORUS Taking into account all the information collected from the most reliable bibliography, the redescription of the species on the basis of American individuals by Guerrero and Peña-Santiago (2007), and observations on Iberian specimens, the following emended diagnosis for the species is provided: body 1.38-1.92 mm long, lip region offset by more or less marked depression and 15-20 µ m diam., odontostyle 37-44 µ m long or 1.7-2.5 times lip diam. and rarely somewhat shorter or longer (ranging from 33-48 µ m), odontophore with distinct basal flanges consisting of six sclerotised pieces and 1.0-1.5 times longer than odontostyle, total length of stylet 78-100 µ m, neck length 288-400 µ m, pharyngeal expansion 110- 145 µ m and occupying 35-40% of total neck length, female genital system amphidelphic, uterus rather short, tripartite but with poor development of each region, 0.9-2.0 times body diam. long, pars refringens vaginae formed by two trapezoidal sclerotisations, vulva a transverse slit and uniquely anterior (V = 37-47), tail short, rounded to hemispherical (18-30 µ m long, c = 55-91, cļ = 0.5-0.7 in females, and 18-22 µ m long, c = 67-100, cļ = 0.6 in males), males very rare, bearing 6-12 spaced ventromedian supplements with posterior one in range of the spicules.