The genus
One of the species recently transferred to the genus
A survey to obtain the nematodes was conducted in soils of the district Shamli (29.6189° N, 77.4329° E; 280 m altitude), Uttar Pradesh, India. This location has a semiarid and moderate-to-tropical monsoon (humid subtropical) predominant climate. The type of soil is sandy loam and loamy and the pH of soil samples ranged from 6.5 to 8.4. A total of eighty-nine soil samples were taken from meadows, pastures, agricultural fields, open fields, and orchards.
Each soil sample consisted of 1 kg of soil that was a mixture of five soil subsamples collected at 15 to 20 cm depth in five locations within each field (one sample from each corner of the field, and one from the center of the field). The soil was first made fine to remove any debris (i.e., rocks, pieces of wood or bark, leaves, etc.) and then moistened with distilled water using a spray bottle to facilitate the movement of nematodes. To recover insect-associated nematodes from these soil samples, the ‘
Emerged IJs were sterilized with 0.1% sodium hypochlorite and washed with ddH2O, and finally stored in tissue culture flasks at 15°C. Third-stage juveniles were used within seven days after emergence (Aasha et al., 2019; Bhat et al., 2020a).
Nematode third-stage juveniles were surface-sterilized with 1% NaOCl. Then, greater wax moth (
For the SEM, male and female specimens preserved in glycerin were selected for observation and processed according to the Abolafia’s (2015) protocol. Thus, they were hydrated in distilled water, dehydrated in a graded mixture of ethanol-acetone series, critical point-dried with liquid carbon dioxide, and coated with gold. The mounts were examined with a Zeiss Merlin microscope (5 kV).
Genomic DNA was isolated from approximately five hundred infective juvenile nematodes by using the Qiagen Blood and Tissue Analysis Kit following the manufacturer’s protocol. A fragment of the rDNA gene containing the ITS regions (ITS1, 5.8S, ITS2) was amplified using primers 18S: 5′-TTGATTACGTCCCTGCCCTTT-3′ (forward), and 26S: 5′-TTTCACTCGCCGTTACTAAGG-3′ (reverse) (Vrain et al., 1992). The fragment containing the D2/D3 regions of the 28S rDNA gene was amplified using primers D2F: 5′-CCTTAGTAACGGCGAGTGAAA-3′ (forward) and 536: 5′-CAGCTATCCTGAGGAAAC-3′ (reverse) (Nadler et al., 2006). The 18S rDNA was amplified using primers NEM18SF: 5′-CGCGAATRGCTCATTACAACAGC-3′ (forward) and NEM18SR: 5′-GGGCGGTATCTGATCGCC-3′ (reverse) (Floyd et al., 2005). The Polymerase Chain Reaction (PCR) protocol for ITS, 18S, and D2/D3 rDNA gene amplification followed was described by Bharti et al. (2020) and Suman et al. (2020). Briefly, PCR master mix consisted of ddH2O 16.8 μl, 10x PCR buffer 2.5 μl, dNTP mix (10 mM each) 0.5 μl, 1 μl of each forward and reverse primers, dream taq green DNA polymerase 0.2 μl, and 3 μl of DNA extract. The PCR profiles used were 1 cycle of 94°C for 3 min followed by 40 cycles of 94°C for 30 sec, 52°C for 30 sec for LSU (28S) rDNA or 55°C for 30 sec for ITS rDNA or 54°C for 30 sec for SSU (18S) rDNA, 72°C for 60 sec, and a final extension at 72°C for 10 min. PCR was followed by electrophoresis (45 min, 100 volts) of 5 μl of PCR product in a 1% TAE (Tris–acetic acid–EDTA) buffered agarose gel stained with ethidium bromide (Bhat et al., 2020c; Rana et al., 2020a). The amplified products were purified and Sanger sequenced in both directions by Bioserve Technologies Ltd. (Hyderabad, India). The obtained sequences were manually curated, trimmed, and submitted to the Center for Biotechnology Information (NCBI) under accession numbers, MT873043, MT872508, and MT872503 for ITS, 28S (D2/D3) and 18S rDNA regions, respectively for the isolate CJ6, and MT873044, MT872509, and MT872504 for the same respective genes for the isolate CJ13.
The sequences were edited and compared with those already present in GenBank using the Basic Local Alignment Search Tool (BLASTN) of the National Center for Biotechnology Information (NCBI) (Altschul et al., 1990). The newly obtained ribosomal LSU (D2/D3 rDNA), SSU (18S rDNA), and ITS (ITS1, 5.8S, ITS2) rDNA sequences were manually edited using BioEdit 7.2.6 (Hall, 1999) and aligned with other relevant segments of same rDNA gene sequences available in GenBank using Clustal W alignment in the program MEGA7 (Kumar et al., 2016). Poorly aligned regions were removed from the alignments using MEGA7. The base substitution model was evaluated using jModeltest2.1.10 (Darriba et al., 2012). Phylogenetic trees were elaborated using the Bayesian inference method as implemented in the program MrBayes 3.2.7 (Ronquist et al., 2012). For analysis in jModeltest, the HKY + I + G model was selected for the ITS tree, the GTR + I + G model was selected for the 18S tree, and the GTR + G was selected for the 28S tree. The selected models were initiated with a random starting tree and ran with the Markov chain Monte Carlo (MCMC) for 1 × 106 generations. The Bayesian tree was ultimately visualized using the FigTree program 1.4.4 (Rambaut, 2018).
The details of all the nematode species used in the molecular and phylogenetic study, including their updated nomenclature, accession numbers of rDNA genes, isolation source, and origin of the sequences are given in Table 4.
Nematodes of four genera:
(Ali, Pervez, Andrabi, Sharma and Verma, 2011) Sudhaus, 2011.
Morphometrics of
Characters | Female | Male | L3 |
---|---|---|---|
n | 20 | 20 | 20 |
Body length ( |
939 ± 119 (718-1135) | 800 ± 91 (653-999) | 383 ± 55 (305-475) |
|
16.4 ± 2.7 (10.9-21.1) | 16.7 ± 2.0 (12.7-20.7) | 18.9 ± 2.0 (15.7-22.9) |
|
5.4 ± 0.6 (4.5-6.6) | 4.9 ± 0.5 (3.8-5.9) | 3.4 ± 0.5 (2.7-4.3) |
|
10.9 ± 1.7 (8.7-14.0) | 13.7 ± 2.7 (9.8-20.9) | 7.5 ± 1.2 (6.0-10.2) |
|
4.1 ± 0.6 (2.9-5.2) | 2.9 ± 0.5 (1.5-3.8) | 4.8 ± 1.0 (3.0-6.0) |
|
51.2 ± 2.1 (46-56) | – | – |
Lip region width | 7.9 ± 1.4 (5-11) | 7.1 ± 1.1 (6-10) | 3.3 ± 0.5 (2-5) |
Stoma length | 18.4 ± 2.3 (14-22) | 17.1 ± 2.3 (13-21) | 12.8 ± 1.1 (11-15) |
Stomatal tube width | 5.0 ± 0.4 (2.5-3.5) | 3.5 ± 0.7 (2.5-5.5) | ? |
Pharyngeal corpus length | 74 ± 5.6 (68-98) | 68 ± 4.5 (58-76) | 35 ± 2.9 (30-42) |
Metacorpus length | 31 ± 3.9 (24-36) | 30 ± 2.6 (24-33) | 22 ± 2.4 (19-27) |
Isthmus length | 40 ± 5.5 (40-48) | 39 ± 2.5 (35-41) | 27 ± 2.6 (23-31) |
Bulb length | 29 ± 2.3 (26-34) | 29 ± 2.6 (24-35) | 16.8 ± 2.8 (12-24) |
Pharynx length | 175 ± 13.3 (156-195) | 166 ± 7.3 (141-175) | 101 ± 6.5 (88-112) |
Nerve ring – anterior end | 123 ± 15.0 (98-153) | 112 ± 9.1 (92-125) | 69 ± 10.9 (53-92) |
Excretory pore–anterior end (EP) | 137 ± 18.2 (110-166) | 130 ± 9.1 (113-144) | 78 ± 11.7 (56-103) |
Deirid–anterior end | 133 ± 16.0 (110-167) | 125 ± 11.0 (107-147) | ? |
Neck length (stoma + pharynx, NL) | 173 ± 15.0 (148-195) | 163 ± 8.3 (144-176) | 114 ± 6.1 (101-124) |
Body width at neck base | 43 ± 5.2 (32-50) | 38 ± 4.2 (32-50) | 18.7 ± 2.9 (14-25) |
Mid-body width (MBW) | 58 ± 10.4 (43-81) | 48 ± 7.2 (40-66) | 20.5 ± 3.3 (16-29) |
Anterior genital branch or Testis | 260 ± 38.2 (192-372) | 198 ± 15 (188-222) | – |
Posterior genital branch | 278 ± 27.5 (229-321) | – | – |
Vagina length | 24.5 ± 4.1 (17-30) | – | – |
Vulva–anterior end (VA) | 480 ± 64 (380-579) | – | – |
Rectum length | 31 ± 6.4 (22-42) | – | 15.0 ± 4.0 (9-23) |
Anal body width (ABW) | 22 ± 2.7 (16-28) | 20.9 ± 2.5 (17-27) | 11.2 ± 2.2 (9-17) |
Tail length (T) | 87 ± 10.5 (68-101) | 63 ± 8.2 (49-62) | 52 ± 5.9 (48-58) |
Spicules length | – | 41 ± 7.5 (34-49) | – |
Gubernaculum length | – | 19.6 ± 3.5 (16-28) | – |
Female: Reproductive system didelphic-amphidelphic, the anterior and the posterior branches in sinistral and dextral position to intestine, respectively. Ovaries moderately developed, dorsally reflexed but with distal end not reaching to vulval level, anterior ovary slightly larger. Usually one or two small rounded pseudocoelomocytes observed in close proximity to the proximal end of ovaries. Oviducts proximally enlarged, connected to ovoid spermatheca frequently filled with sperm. Uteri well developed, differentiated into long glandular and muscular regions, filled with sperm and one to ten intrauterine eggs, 40−49 × 22−23 µm, in different stages of embryonation. Vagina thick-walled, often cuticularized, at right angle to the longitudinal body axis, with length equal to about one-third of the vulval body diameter. Vulva a wide transverse slit, with protruding lips, unremarkable or weak epiptygma but distinct cuticular flap. Rectum short, 1.0 to 1.8 times anal body diameter, allied with rectal glands at its junction with prerectum. Prerectum distinguishable from intestine in lacking prominent cell nuclei. Tail elongate conoid, gradually tapering to a fine terminus. Phasmids short tubular, located posterior to anus, about 38 to 42% of tail length.
Male: Similar to female in general morphology except for smaller size, posterior body curvature prominent and cuticular striations relatively fine. Reproductive system monorchic, with single testis reflexed ventrad anteriorly on the right side of the intestine. Vas deferens a broad tube, packed with sperm without delineation of seminal vesicles. Ejaculatory glands not observed. Spicules paired and symmetrical, ventrally arcuate, free with slightly bent ventrad manubrium, ventrally hooked, calamus short conoid and slightly ventrally curved lamina with ventrally bent finely rounded tip in lateral view. Gubernaculum well-developed, slightly ventrad curved with long manubrium and narrow corpus, 50 to 60% of spicule length. Three small gland-like cells are distinguishable around the anterior end of the cloaca. Tail conoid with posterior two-thirds abruptly tapering and reduced. Bursa anteriorly open, narrow, leptoderan, not enclosing large tail spike, having smooth margins and eight pairs (1 + 1 + 1/3 + 2 + ph) of genital papillae, with GP1 and GP2 spaced, precloacal, GP3 slightly posterior to cloaca in most specimens, pairs GP4 to GP6 located at conoid part of tail and GP7 to GP8 located at posterior part of the bursa, dorsally directed. Phasmid small, tubular, located posterior to the GP8, at 45 to 50% of tail length.
Juveniles: Third-stage juveniles ensheathed in a cuticle of second stage juveniles. Sheath free anteriorly in third-stage juveniles, firmly bound to the posterior region of the body. Body lean, from anus to tail terminus. Cuticle with transverse striae. Lip region smooth; stoma opening closed. Stoma tubular. Pharynx with pharyngeal corpus and isthmus both long and narrow, and basal bulb spheroid, valvate. Nerve ring and excretory pore located at isthmus level. Tail conoid with pointed terminus.
From the two populations of
Comparing with other sequences (unpublished) of the species available from GenBank, the 18S rDNA fragment, from a common aligned fragments with 794 bp, the present populations from India shows one (0.1%) change from the other sequence available from India (NM453373), 1 (0.1%) and 16 (2.0%) changes from the sequences submitted from Philippines (MT012150 and MT043860), respectively. For the 28S rDNA fragment, there are no other available sequences to compare. The ITS rDNA sequences, from a common aligned fragments with 645 bp, show 3 (0.5%) or 6 (0.9%) changes from other sequences submitted from India (KP834432/KP834433/KY083045 and MH392568), respectively; 1 (0.2%) or 2 (0.3%) changes with respect to two sequences from Philippines (MT422254 and MT576957), while other two sequences (MT452472 and MT576957) deposited from Philippines show too much changes (51 and 64, respectively); the sequence submitted from Pakistan (MK973071) show 24 (3.7%) changes, the most of them consistent in two long contiguous deletions in the Pakistani sequence (10 and 12 gaps, respectively, after aligning sequences), which must be considered as
Twenty females and twenty males of each isolate were deposited at the museum of the Department of Zoology, Chaudhary Charan Singh University Meerut, India. Ten females and ten males were deposited at the nematode collection of the Department of Animal Biology, Plant Biology and Ecology of the University of Jaén, Spain.
The material examined in this study agrees well with the type material described by Ali et al. (2011) and the redescription of Asif et al. (2013) as
Compendium of females of
Species |
|
|
|
|
|
|
|
|
|
---|---|---|---|---|---|---|---|---|---|
Reference | Present study | Ali et al. (2011) | Shaheen et al. (2011) | Shaheen et al. (2011) | Pervez et al. (2012) | Asif et al. (2013) | Tabassum et al. (2019) | Tabassum et al. (2019) | Tabassum et al. (2019) |
Country | India | India | India | India | India | India | Pakistan | Pakistan | Pakistan |
Habitat | Rhizosphere of sugarcane and groundnut | Rhizosphere of mungbean | Rhizosphere of chickpea | Rhizosphere of pigeonpea | Rhizosphere of ginger | Decaying matter | Rhizosphere of mango tree | Decomposed guava fruit | Rhizosphere of chicko |
|
718-1,135 | 658-786 | 964-1,018 | 902-989 | 1,418-1,813 | 786-902 | 1,366-1,684 | 1,769-2,078 | 1,546-1,694 |
|
10.9-21.1 | 19.7-22.9 | 20.1-22.5 | 24.4-25.5 | 18.5-21.2 | 19.2-23.5 | 14.0-18.0 | 11.7-20.0 | 15.0-18.0 |
|
4.5-6.6 | 4.1-4.8 | 5.7-5.9 | 3.8-4.3 | 5.1-5.3 | 4.1-5.0 | 5.5-7.8 | 7.0-8.0 | 6.0-8.0 |
|
8.7-14.0 | 8.9-12.1 | 10.3-12.9 | 10.2-12.7 | 12.1-13.2 | 9.6-11.2 | 12.0-16.0 | 12.0-18.0 | 13.0-19.0 |
|
2.9-5.2 | 4.1-4.6 | 3.6-4.3 | 3.3-4.8 | 4.6 | 3.8-4.5 | 2.5.-4.4 | 4.0-8.0 | 3.0-4.0 |
|
46-56 | 50-58 | 43a | 42a | 51-60 | 51-55 | 48-56 | 49-58 | 50-54 |
Lip region width | 5-11 | 7-8 | 8-11 | 6-8 | 8-12 | 9-10 | 11-15 | 12a | 12a |
Stoma length | 14-22 | 16-18 | 18-19 | 22-23 | 19-21 | 20-25 | 22-28 | 26-30 | 22-26 |
Corpus length | 68-98 | 96-115 | 92-100 | 133 | 95-170 | 50a | 62-106 | 57a | 42a |
Isthmus length | 40-48 | 32-44 | 30-40 | 44-56 | 28-57 | 35-45 | 30a | 27a | 33a |
Bulb length | 26-34 | 24-35 | 26-31 | 33-45 | 44a | 25-35 | 19a | 16a | 17a |
Nerve ring-ant. end | 98-153 | 99-112 | 111-134 | 170-179 | 178-203 | 110-143 | 154-190 | 76a | 95a |
Excretory pore-ant. end | 110-166 | 109-130 | 134-136 | 166-172 | 187-223 | 121-160 | 148-250 | 98a | 112a |
Pharynx length | 156-195 | 159-178 | 167-176 | 220-225 | 189-284 | 177-218 | 206-265 | 250-279 | 218-247 |
Mid-body width | 43-81 | 32-39 | 39-46 | 32-40 | 75-89 | 26-40 | 78-108 | 92-111 | 88-105 |
Anal body width | 16-28 | 16-17 | 20-28 | 19-23 | 25-28 | 19-22 | 25-33 | 12-30 | 30-36 |
Tail length | 68-101 | 65-80 | 75-96 | 77-87 | 115-129 | 81-100 | 94-112 | 100-148 | 84-132 |
Compendium of males of
Species |
|
|
|
|
|
|
|
|
|
---|---|---|---|---|---|---|---|---|---|
Reference | Present study | Ali et al. (2011) | Shaheen et al. (2011) | Shaheen et al. (2011) | Pervez et al. (2012) | Asif et al. (2013) | Tabassum et al. (2019) | Tabassum et al. (2019) | Tabassum et al. (2019) |
Country | India | India | Pakistan | Pakistan | India | India | Pakistan | Pakistan | Pakistan |
Habitat | Rhizosphere of sugarcane and groundnut | Rhizosphere of mungbean | Rhizosphere of chickpea | Rhizosphere of pigeonpea | Rhizosphere of ginger | Decaying matter | Rhizosphere of mango tree | Descomposed guava fruit | Rhizosphere of chicko |
|
653-999 | 594-804 | 754-973 | 855-889 | 673-821 | 683-868 | 1,154-1,325 | 1,100-1,392 | 1,234-1,498 |
|
12.7-20.7 | 16.6-19.3 | 19.4-20.8 | 25.0.-28.0 | 18.3-24.0 | 18.1-21.7 | 14.4-19.4 | 15.0-24.0 | 14.0-20.0 |
|
3.8-5.9 | 4.0-5.0 | 5.0-5.6 | 3.83-3.89 | 4.32-5.3 | 4.3-4.5 | 5.3-6.6 | 4.0-6.0 | 6.0-8.0 |
|
9.8-20.9 | 10.7-17.8 | 13.6-16.7 | 13.9-13.6 | 11.5-16.7 | 11.5-13.7 | 14.9-19.0 | 23.0-37.0 | 16.0-20.0 |
|
1.5-3.8 | 2.6-2.8 | 2.1-2.7 | 3.2 | 2.8-3.1 | 2.5-3.0 | 2.1-3.7 | 1.0-2.0 | 2.0-3.0 |
Lip region width | 6-10 | 7-8 | 8-11 | 6-8 | 7a | 9-10 | 11-14 | ? | ? |
Stoma length | 13-21 | 15-17 | 19 | 22-23 | 17-19 | 18-20 | 20-28 | 24-28 | 20-24 |
Corpus length | 58-76 | 81-109 | ? | 134 | 71-112 | 72a | ? | ? | ? |
Isthmus length | 35-41 | 27-42 | ? | 44-56 | 21-38 | 27a | ? | ? | ? |
Bulb length | 24-35 | 20-36 | ? | 33-45 | ? | 23a | ? | ? | ? |
Nerve ring-ant. end | 92-125 | 79-108 | 116-136 | 149-179 | 90-114 | 100-127 | 143-185 | ? | ? |
Excretory pore-ant. end | 113-144 | 87-114 | 127-138 | 155-168 | 110-142 | 119-141 | 137-179 | ? | ? |
Pharynx length | 141-175 | 134-169 | 149-172 | 223-228 | 142-187 | 155-190 | 184-256 | 211-256 | 204-236 |
Midbody width | 40-66 | 31-45 | 39-46 | 30-35 | 32-39 | 26-40 | 64-69 | 54-88 | 70-98 |
Anal body width | 17-27 | 16-20 | 26-30 | 19-21 | 16-19 | 20-24 | 21-40 | 26-34 | 27-32 |
Tail length | 49-62 | 41-55 | 55-58 | 61-65 | 43-59 | 58-67 | 62-76 | 32-56 | 66-78 |
Spicules length (SL) | 34-49 | 31-36 | 35-44 | 41-44 | 24-27 | 33-39 | 40-46 | 32-60 | 42-60 |
Gubernaculum length (GL) | 16-28 | 13-17 | 19-20 | 14-18 | 9-10 | 14-20 | 20-34 | 13-23 | 16-22 |
GL/SL × 100 | 50-60 | 43-46 | 45-54 | 34-40 | 36 | 42-51 | 50-74 | 40 | 37-38 |
Nematode species, GenBank accession number, and origin of the sequences used for phylogenetic study.
GenBank accession number | ||||||
---|---|---|---|---|---|---|
Speciesa | 18S rDNA | 28S rDNA | ITS rDNA | Country | Isolation source | Reference |
|
EU196013 | EU195976 | USA | Unknown | Kiontke et al. (2007) | |
|
AF083012 | USA | Unknown | Fitch (unpublished) | ||
|
EU196004 | USA | Unknown | Kiontke et al. (2007) | ||
|
EU196020 | USA | Unknown | Kiontke et al. (2007) | ||
|
MK359049 | India | Soil | Palanisamy et al. (unpublished) | ||
|
EF990722 | USA | Unknown | Kiontke et al. (2007) | ||
|
JN636068 | USA | Rotting fruits | Kiontke et al. (2011) | ||
|
Z92784 | UK | Unknown | Sulston and Waterston (1998) | ||
|
X03680 | USA | Unknown | Ellis et al. (1986) | ||
|
JN636142 | USA | Rotting fruits | Kiontke et al. (2011) | ||
|
EU196005 | USA | Unknown | Kiontke et al. (2007) | ||
|
EU195995 | USA | Unknown | Kiontke et al. (2007) | ||
|
U73449 | Unknown | Baldwin et al. (1997) | |||
|
AY593921 | Netherlands | Soil | Helder et al. (2006) | ||
|
EU195959 | USA | Unknown | Kiontke et al. (2007) | ||
|
EF990725 | USA | Unknown | Kiontke et al. (2007) | ||
|
EU040125 | China | Soil | Li et al. (unpublished) | ||
|
AF083017 | USA | Unknown | Fitch (unpublished) | ||
|
EF990727 | USA | Unknown | Kiontke et al. (2007) | ||
|
KJ636408 | Netherlands | Unknown | van Megen (unpublished) | ||
|
KU573061 | Ireland | Soil | Maher et al. (2016) | ||
|
KJ636320 | Netherlands | Unknown | van Megen (unpublished) | ||
|
AF083021 | USA | Unknown | Fitch (unpublished) | ||
|
AM399053 | Belgium |
|
Derycke et al. (2008a) | ||
|
AM937053 | Greece | Decaying seaweeds | Derycke et al. (2008b) | ||
|
AF083020 | USA | Unknown | Fitch (unpublished) | ||
|
AM399068 | New Zealand | Unknown | Derycke et al. (2008a) | ||
|
AF083013 | USA | Unknown | Fitch (unpublished) | ||
|
EF990723 | USA | Unknown | Kiontke et al. (2007) | ||
|
EU196014 | EU195980 | USA | Unknown | Kiontke et al. (2007) | |
|
MT872503, MT872504 | MT872508, MT872509 | MT873043, MT873044 | India | Soil | Present study |
|
MT422254 | Philippines | Soil | Dichusa et al. (unpublished) | ||
|
MT452472, MT452471 | Philippines | Soil | Sangcopan and Sumaya (unpublished) | ||
|
MH392568, KY083045 KP834433, KP834432 | India | Soil | Chavan et al. (unpublished) | ||
|
MK973071 | Pakistan | Soil | Tabassum et al. (unpublished) | ||
|
U13935 | Germany | Unknown | Fitch et al. (1995) | ||
|
MT043860 | Philippines | Soil | Andalan et al. (unpublished) | ||
|
MF989442 | Colombia |
|
Chaves et al. (unpublished) | ||
|
MT012150 | Philippines | Soil compost | Guadalquiver (unpublished) | ||
|
MN453373 | India | Soil | Kandhasamy and Muthugounder (unpublished) | ||
|
EU195965 | USA | Unknown | Kiontke et al. (2007) | ||
|
KM233152, KM233153 | Brazil | Cow | Bossi et al. (2015) | ||
|
DQ121436 | South Africa | Unknown | Jumba and Gray (unpublished) | ||
|
MT012135, MT012133 | Philippines | Soil | Kabalu et al. (unpublished) | ||
|
MT012153 | Philippines | Soil | Buldiman et al. (unpublished) | ||
|
JQ237848 | USA | Citrus groves | Campos-Herrera et al. (2012) | ||
|
AF083008 | USA | Unknown | Fitch (unpublished) | ||
|
EU195966 | USA | Unknown | Kiontke et al. (2007) | ||
|
KR011843, KR011846 | Brazil | Soil | de Brida et al. (2017) | ||
|
MK973071 | Pakistan | Soil | Tabassum et al. (unplublished) | ||
|
KU180665 | KU180676 | Sweden | Unknown | Holovachov et al. (2015) | |
|
FJ547240 | FJ547239 | USA | Vermicompost | Ye et al. (2010) | |
|
EF503692 | China | Soil | Zhang et al. (2008) | ||
|
EU273599 | China | Soil | Liu et al. (2012) | ||
|
MT548593 | China |
|
Li et al. (unpublished) | ||
|
AY751546 | Colombia |
|
Stock et al. (2005) | ||
|
EU196010 | USA | Unknown | Kiontke et al. (2007) | ||
|
AF082998 | USA | Unknown | Fitch (unpublished) | ||
|
EU195970 | USA | Unknown | Kiontke et al. (2007) | ||
|
EU196022 | USA | Unknown | Kiontke et al. (2007) | ||
|
MF441252 | India | Soil | Kumar et al. (2019) | ||
|
AF083019 | USA | Unknown | Fitch (unpublished) | ||
|
EU195968 | USA | Unknown | Kiontke et al. (2007) | ||
|
AY602176 | USA | Unknown | Kiontke et al. (2004) | ||
|
MG742117 | Thailand | Soil | Nitjarunkul et al. (unpublished) | ||
|
U13936 | USA | Soil | Fitch et al. (1995) | ||
|
KT884894 | Australia |
|
Carta et al. (2018) | ||
|
MG551687 | Portugal | Unknown | Campos-Herrera (unpublished) | ||
|
LN613263 | Italy | Soil | Torrini et al. (2015) | ||
|
JQ002566 | China | Soil | Zhang et al. (2012) | ||
|
KT884891 | Japan |
|
Carta et al. (2018) | ||
|
MK959600 | MK959601 | Spain | Decaying wood | Abolafia and Pena-Santiago (2019b) | |
|
EU195969 | USA | Unknown | Kiontke et al. (2007) | ||
|
KP792649 | Brazil | Soil | Campos-Herrera and Pu˚ža (unpublished) | ||
|
EF990724 | Germany | Bark beetles | Kiontke et al. (2007) | ||
|
EU003189 | USA | Unknown | Kiontke et al. (2007) | ||
|
AF036946 | Kenya | Feces of an antelope | Adams et al. (1998) | ||
|
EU196023 | EU195997 | USA | Unknown | Kiontke et al. (2007) | |
|
AF083002 | USA | Unknown | Fitch (unpublished) | ||
|
EU195979 | USA | Unknown | Kiontke et al. (2007) | ||
|
DQ639980 | Scotland | Nemaslug | MacMillan et al. (2006) | ||
|
DQ639982 | Scotland |
|
MacMillan et al. (2006) | ||
|
KX267675 | South Africa |
|
Pieterse et al. (2017) | ||
|
AB370214 | USA | Termites | Kanzaki et al. (2009) | ||
|
AF083023 | USA | Unknown | Fitch (unpublished) | ||
|
EU195984 | USA | Unknown | Kiontke et al. (2007) | ||
|
DQ059060 | Germany | Unknown | Hong et al. (2005) | ||
|
AF083022 | USA | Unknown | Fitch (unpublished) | ||
|
DQ059057 | Germany | Unknown | Hong et al. (2005) | ||
|
AF083001 | USA | Unknown | Fitch (unpublished) | ||
|
AY602168 | USA | Unknown | Kiontke et al. (2007) | ||
|
U13934 | Germany | Unknown | Fitch et al. (1995) | ||
|
AY602177 | USA | Unknown | Kiontke et al. (2004) | ||
|
EU196006 | USA | Unknown | Kiontke et al. (2007) | ||
|
MN082353, MN082355 | Taiwan | Soil | Yang et al. (2020) | ||
|
AF082996 | Unknown | Fitch (unpublished) | |||
|
EF990726 | USA | Unknown | Kiontke et al. (2007) | ||
|
KX385908 | Mexico | White grub | Jiménez-Cortés et al. (2016) | ||
|
EF990716 | EF990721 | USA | Soil compost | Kanzaki et al. (2008) | |
|
U13937 | USA | Unknown | Fitch et al. (1995) |
Recently, Tabassum et al. (2019) described a new species,
Bayesian Inference tree from previously and the newly sequenced
According to this, the updated list of synonyms of
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The phylogenetic relationships as inferred from the Bayesian Inference analysis between
Bayesian Inference tree from the newly sequenced
Bayesian Inference tree from the newly sequenced
The phylogenetic tree inferred using 18S rDNA gene sequences, shows three clusters that contain sequences of nematodes that have been suggested to belong to
The phylogenetic tree inferred using 28S rDNA gene sequences show also three clusters containing
Finally, analyzing ITS rRNA gene sequences, we arrive to the same conclusions derived from the analysis of 28S- and 18S rDNA gene sequences. ITS-based phylogenetic tree show a clear cluster that separates