Enchodelus macrodorus ( de Man, 1880 ) Thorne, 1939

Elshishka, Milka, Mladenov, Aleksandar, Altash, Stela, Álvarez-Ortega, Sergio, Peña-Santiago, Reyes & Peneva, Vlada, 2024, Cyber catalogue and revision of the nematode genus Enchodelus (Dorylaimida, Nordiidae), Biodiversity Data Journal 12, pp. e 126315-e 126315 : e126315-

Enchodelus macrodorus ( de Man, 1880) Thorne, 1939

Dorylaimus macrodorus de Man, 1880 ; de Man (1880) BLR: https://doi.org/10.5281/zenodo.10852798

Dorylaimus macrodorus ; de Man (1884) BLR: https://doi.org/10.5281/zenodo.10867883

Dorylaimus macrodorus ; de Man (1912): 454 -456 BLR: https://doi.org/10.5281/zenodo.11003008

Dorylaimus macrodorus ; Menzel (1913) BLR: https://doi.org/10.5281/zenodo.10797350

Dorylaimus macrodorus ; Brakenhoff (1914) BLR: https://doi.org/10.5281/zenodo.10822895

Dorylaimus macrodorus ; Steiner (1914): 262 [list] BLR: https://doi.org/10.5281/zenodo.11003080

Dorylaimus macrodorus ; Hofmänner and Menzel (1915): 186 -187 BLR: https://doi.org/10.5281/zenodo.11003145

Dorylaimus macrodorus ; Steiner (1916 a): 69-70 BLR: https://doi.org/10.5281/zenodo.11003157

Dorylaimus macrodorus ; Steiner (1916 b): 345 BLR: https://doi.org/10.5281/zenodo.11003151

Dorylaimus macrodorus ; Micoletzky (1921) BLR: https://doi.org/10.5281/zenodo.10932729

Dorylaimus macrodorus ; Schneider (1923) BLR: https://doi.org/10.5281/zenodo.10795142

Dorylaimus macrodorus ; Kreis (1924) BLR: https://doi.org/10.5281/zenodo.10892094

Dorylaimus macrodorus ; Stefanski (1924): 55-57

Dorylaimus macrodorus ; Micoletzky (1925): 181-182

Dorylaimus macrodorus ; Rahm (1925): 182

Dorylaimus macrodorus ; Stefanski (1927) BLR: https://doi.org/10.5281/zenodo.10941762

Dorylaimus macrodorus ; Soós (1936): 61 [list] BLR: https://doi.org/10.5281/zenodo.10944890

Dorylaimus macrodorus ; Allgén (1953) BLR: https://doi.org/10.5281/zenodo.10809558

Dorylaimus (Doryllium) macrodorus de Man, 1880; Seidenschwarz (1923): 42

Dorylaimus (Doryllium) macrodorus de Man, 1880 ; Allgén (1925) BLR: https://doi.org/10.5281/zenodo.10998093

Dorylaimus (Doryllium) macrodorus de Man, 1880 ( Ditlevsen, 1928); Ditlevsen (1928): 9

Dorylaimellus macrodorus ( de Man, 1880) Thorne & Swanger, 1936 ; Thorne and Swanger (1936) BLR: https://doi.org/10.5281/zenodo.10845809

Dorylaimellus macrodorus ; Altherr (1950) BLR: https://doi.org/10.5281/zenodo.10810136

Dorylaimellus macrodorus ; Altherr (1952): 342

Dorylaimellus macrodorus ; Andrássy (1952) BLR: https://doi.org/10.5281/zenodo. 11012991

Dorylaimus (Dorylaimellus) macrodorus de Man, 1880 ( Thorne & Swanger, 1936): Thorne and Swanger (1936) BLR: https://doi.org/10.5281/zenodo.10845809

Dorylaimus (Dorylaimellus) macrodorus ; Schneider (1939) BLR: https://doi.org/10.5281/zenodo.11028113

Enchodelus (Enchodelus) macrodorus ( de Man, 1880) Thorne, 1939

Enchodelus (Enchodelus) macrodorus ; Ahmad and Jairajpuri (1980) BLR: https://doi.org/10.5281/zenodo.8144942

Enchodelus (Enchodelus) macrodorus ; Jairajpuri and Ahmad (1992): 180 [figure] BLR: https://doi.org/10.5281/zenodo.10850166

Enchodelus (Enchodelus) macrodorus ; Choi et al. (1997) BLR: https: // doi. org / 10.5281/zenodo.10822472

Enchodelus macrodorus ( de Man, 1880) Thorne, 1939 ; Thorne (1939) BLR: https://doi.org/10.5281/zenodo.11003121

Enchodelus macrodorus ; Goodey (1951) BLR: https://doi.org/10.5281/zenodo.11013374

Enchodelus macrodorus ; Altherr (1953) BLR: https://doi.org/10.5281/zenodo. 11012949

Enchodelus macrodorus ; Andrássy (1958 b) BLR: https://doi.org/10.5281/zenodo.10842858

Enchodelus macrodorus ; Meyl (1961) BLR: https://doi.org/10.5281/zenodo.11015551

Enchodelus macrodorus ; van Rossen and Loof (1962) BLR: https://doi.org/10.5281/zenodo.10845171

Enchodelus macrodorus ; Goodey (1963) BLR: https://doi.org/10.5281/zenodo.10891552

Enchodelus macrodorus ; Jairajpuri and Loof (1967) BLR: https://doi.org/10.5281/zenodo.8122568

Enchodelus macrodorus ; Loof and Coomans (1970) BLR: https://doi.org/10.5281/zenodo.11014428

Enchodelus macrodorus ; Zullini (1970) BLR: https://doi.org/10.5281/zenodo.10850512

Enchodelus macrodorus ; Loof (1971) BRL: h https://doi.org/10.5281/zenodo.8152932

Enchodelus macrodorus ; Thorne (1974) BLR: https://doi.org/10.5281/zenodo.10819174

Enchodelus macrodorus ; Andrássy (1978): 116 [list]

Enchodelus macrodorus ; Nesterov (1979) BLR: https://doi.org/10.5281/zenodo.11015718

Enchodelus macrodorus ; Vinciguerra (1984) BLR: https://doi.org/10.5281/zenodo.10818036

Enchodelus macrodorus ; Winiszewska-Slipinska (1987) BLR: https://doi.org/10.5281/zenodo.10854940

Enchodelus macrodorus ; Bongers (1988) BLR: https://doi.org/10.5281/zenodo.10822198

Enchodelus macrodorus ; Eliava and Eliashvily (1990): 50 -51, 96 BLR: https://doi.org/10.5281/zenodo.11003031

Enchodelus macrodorus ; Gerber (1991) BLR: https://doi.org/10.5281/zenodo.10883357

Enchodelus macrodorus ; Nasira et al. (1992) BLR: https://doi.org/10.5281/zenodo.8122525

Enchodelus macrodorus ; Popovici (1995 a) BLR: https://doi.org/10.5281/zenodo.8125537

Enchodelus macrodorus ; Loof (1999) BLR: https://doi.org/10.5281/zenodo.11003046

Enchodelus macrodorus ; Ahmad et al. (2002) BLR: https://doi.org/10.5281/zenodo.10683060

Enchodelus macrodorus ; Poiras (2006): 93 [list] BLR: https://doi.org/10.5281/zenodo.10717726

Enchodelus macrodorus ; Guerrero and Peňa-Santiago (2007) BLR: https://doi.org/10.5281/zenodo.8111782

Enchodelus macrodorus ; Guerrero et al. (2008 b) BLR: https://doi.org/10.5281/zenodo.8111849

Enchodelus macrodorus ; Pedram et al. (2009) BLR: https://doi.org/10.5281/zenodo.8114781

Enchodelus macrodorus ; Andrássy (2009 b) BLR: https://doi.org/10.5281/zenodo.10821949

Enchodelus macrodorus ; Ciobanu et al. (2010 a) BLR: https://doi.org/10.5281/zenodo.8111705;

Enchodelus macrodorus ; Holovachov (2014): 22 [list] Plazi treatment

Enchodelus macrodorus ; Shahina et al. (2019): 223 [list] BLR: https://doi.org/10.5281/zenodo.10710707

Enchodelus macrodorus ; Peña-Santiago (2021 b): 208-210 BLR: https://doi.org/10.5281/zenodo. 11191905

Distribution

A typical member of Palearctic nematode fauna, recorded in a myriad of countries and habitats: Netherlands (type habitat: moist soil in meadows and marshes - de Man (1880), de Man (1884) / sandy soil with moss - de Man (1912), Loof and Oostenbrink (1962), Loof and Coomans (1970) Loof and Coomans (1970), Bongers (1988)), Alps (moss - Franz (1942), Franz and Gunhold (1954)), Austria (moss Hypnum cupressiforme Hedw - Steiner (1916 b), Micoletzky (1921) / moss - Seidenschwarz (1923)), Bulgaria (moss - Andrássy (1958 a), Katalan-Gateva (1968)), China (grassland and shrubs - Ahmad et al. (2002)), Czech Republic (meadow, grassland - Háněl (1998)), Denmark (moor - Micoletzky (1925)), Faroe Islands ( Ditlevsen (1928)), Germany (river bank - Brakenhoff (1914) / moss - Schneider (1923), Rahm (1925), Schneider (1939) / grassland - Diedrich et al. (1998) / lake, 1136 m a. s. l. - Michiels and Traunspurger (2005)), Hungary (moss - Soós (1936), Soós (1940 a), Soós (1940 b), Andrássy (1952), Andrássy (1958 b) / moss, grass, pine - Andrássy (1996), Andrássy (2009 b)), Iran (grasslands - Pedram et al. (2009), Jabbari et al. (2019)), Italy (moss - Zullini (1970) / moss - Vinciguerra (1984)), Korea ( Betula platyphylla var. japonica Hara - Choi et al. (1997)), Moldova (deciduous forests - Poiras (2006)), Norway (Jan Mayen Island - Allgén (1953) / Spitzbergen, grasses - Loof (1971)), Pakistan (soil around roots of weeds and grasses - Nasira et al. (1992) / freshwater - Nasira et al. (2016) / freshwater - Nasira and Shamim (2018)), Poland (moss - Stefanski (1924), Brzeski (1963 b) / Arrhenatherum sp. - Winiszewska-Slipinska (1987) / primeval forest - Brzeski and Winiszewska-Slipinska (1996)), Romania (freshwater - Stefanski (1926), Stefanski (1927), Popovici (1993), Popovici (1995 a) / hornbeam-beech forest, 400 m a. s. l - Popovici (1995 b), Popovici and Ciobanu (1997) / grasslands - Popovici (1998), Ciobanu and Popovici (2001) / hornbeam-beech forest, grasslands, cliff vegetation, 400-1250 m a. s. l. - Ciobanu et al. (2010 a) / grass - Ciobanu and Popovici (2017)), Russia (Arctic - Novaya Zemlya Archipelago - Steiner (1916 a) / Arctic tundra - Kuzmin (1973), Kuzmin and Gagarin (1990) / Novaya Zemlya Archipelago - Gagarin (1997), Gagarin (2001) / polar desert - cape Cheluyskin - Chernov et al. (1979), Soloveva et al. (1976)), Slovakia (moss - Koniar (1957) / corn - Sabova et al. (1979), Šály (1979) / vineyard - Liskova (1980) / Alnus glutinosa Gaertn. - Šály (1980) / Fagus sylvatica L., Abies alba Mill, Picea abies H. Karst. , meadow, moss, riverbank, brook, waterfall - Šály (1985) / Pinus mugo Turra , P. abies - Liskova et al. (1996) / meadow, river bank - Liskova and Čerevkova (2005) / grassland - Čerevková (2006) / grasslands - Čerevková (2008) / Lariceto - Picetum - Čerevková and Renčo (2009) / moss, forests, grasslands, potato, cereal, vineyard, riverbank - Liskova and Čerevkova (2011) / spruce forests - Čerevková et al. (2013) / P. abies , Pinus sylvestris L., Quercus robur L., Acer pseudoplatanus L., F. sylvatica - Renčo (2013) / maize - Čerevková and Cagan (2015) / alpine meadows, 1763-1994 m a. s. l. - Háněl 2017 / P. abies , Larix decidua - Renčo and Čerevková (2017) / deciduous forest, wetland, Fallopia japonica (Houtt.) , 386-455 m a. s. l. - Čerevková et al. (2019) / Asclepias syriaca L., grassland - Jurová et al. (2020) / deciduous forests, grassland, Solidago gigantea Ait. - Čerevková et al. (2020)), Spain ( Palomo (1979) / wet meadows, soil around poplar, 1400-2925 m a. s. l. - Guerrero et al. (2008 b)), Sweden (moss - Allgén (1925), van Rossen and Loof (1962)), Switzerland (Alps, 2000-4000 m a. s. l. - Menzel (1913), Steiner (1914), Hofmänner and Menzel (1915) / lake with the melting snow water - Kreis (1924) / alpine meadow with Alchemilla sp. , Nardus stricta L., Trifolium sp. , pine forest, 1900-2200 m a. s. l. - Altherr (1950), Altherr (1952), Altherr (1953)), UK (moor, grassland - Banage (1962), Loof and Coomans (1970) / grasslands, 110-160 m a. s. l. - Hodda and Wanless (1994) / spruce forests, 200 m a. s. l. - Ruess et al. (1996)), Ukraine (grasses - Nesterov (1979)) and Uzbekistan (rice, bean, cotton - Tulaganov (1958) / Cucumis sativus L. - Tukhtasinov (2023)).

Very sporadically, this species was also recorded in Nearctic ( USA, Sporobolus compositus (Poir.) Merr. - Orr and Dickerson (1967) / mountain soil - Thorne (1974)) and Indomalayan ( India, soil near roots of apple - Jairajpuri and Loof (1967) / Pinus pinea L. - Ahmad and Jairajpuri (1980)) enclaves.

Taxon discussion

Enchodelus macrodorus was originally described by de Man (1880) from the Netherlands. It is the most widely spread Enchodelus species. Guerrero and Peňa-Santiago (2007) and Guerrero et al. (2008 b) re-described American specimens of E. macrodorus studied by Thorne (1939), noted that these specimens apparently fitted very well with those described by de Man (1880), considered that many subsequent records of the species were doubtful or might correspond to other species, provided an emended diagnosis of this species and regarded the populations reported by Thorne (1939), Jairajpuri and Loof (1967), Ahmad and Jairajpuri (1980), Nasira et al. (1992) and Ahmad et al. (2002), also three of the populations reported by Popovici (1995 a) to be conspecific with E. macrodorus . Later Pedram et al. (2009) reported the species for the first time from Iran and presented the first integrative study of the species.

Notes

Morphological characterisation

Material examined. One female in good condition, mounted on one slide and collected from Rasquilla, Navalsauz, Avila Province, Spain, in grassland riparian zone close to the Alberche River (N 40 º 24.350 ' W 05 º 01.817 ', elevation 1247 m a. s. l.) in May 2021.

Measurements: Table 1 , Fig. 2

Diagnosis (based in examined specimen)

Enchodelus macrodorus is characterised by its 1.44 mm long body, lip region 18.5 μm wide and offset by weak depression, odontostyle 39.0 μm long or 2.1 times lip region diameter, odontophore flanged and 1.1 times as long as odontostyle, neck 326 μm long, pharyngeal expansion occupying two-fifths of the total neck length, female genital system diovarian, uterus short (63, 68 μm long), pars refringens vaginae with two rectangular sclerotisations, vulva ( V = 44 %), tail short and rounded (20 μm, c = 71, c’ = 0.5) bearing saccate bodies.

Remarks

The studied female perfectly fits, morphologically and morphometrically, with the previous descriptions of this species, especially with the re-description of the Thorne material provided by Guerrero and Peňa-Santiago (2007).

Molecular characterisation

One partial 18 S rRNA and one D 2 - D 3 of 28 S rRNA gene sequences were obtained for this species. In the partial 18 S rRNA (Fig. 3 ) gene tree, the sequence of the Spanish E. macrodorus clustered together to another sequence of the same species ( FJ 042953 ) and inside of a highly-supported clade ( PP = 100) with other Enchodelus spp. In the D 2 - D 3 of 28 S rRNA (Fig. 4 ) gene tree, the sequence of the Spanish E. macrodorus formed a highly-supported clade ( PP = 100) with E. enguriensis sp. nov. and another three Enchodelus sequences from GenBank.

Regarding the molecular data for this species, it appears that the sequence AY 593054 available in GenBank was incorrectly identified or labelled. This sequence is herein identified as belonging to Pungentus silvestris (Figs 3 , 4 ). Besides, the sequence AY 284791 , available in GenBank, was incorrectly identified too and most probably belongs to a Pungentus species.

Figures

Figure 2. Enchodelus macrodorus (de Man, 1880) Thorne, 1939, female. A Entire body; B-D Anterior region; E Neck; F Genital system; G Pharyngeal dorsal nuclei; H Pharyngeal intestinal junction; I Uterus and oviduct junction; J Vagina; K Posterior body region; L-N Tail ends (M detail of the lateral chord ending; N detail of the saccate bodies of tail). Scale bars: A 200 µm; B-D, G, H, J, L-N 10 µm; E, F, K 50 µm; I 20 µm.

Figure 3. Bayesian 50 % majority rule consensus tree as inferred from the partial 18 S rRNA gene sequence alignment under the GTR + I + G model. Posterior probability values more than 70 % are given on appropriate clades. New sequences are indicated by bold letters.

Figure 4. Bayesian 50 % majority rule consensus tree as inferred from the D 2 - D 3 expansion segments of 28 S rRNA gene sequence alignment under the GTR + I + G model. Posterior probability values more than 70 % are given on appropriate clades. New sequences are indicated by bold letters.