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Morphology, development stages, and phylogeny of the Rhabditolaimus ulmi (Nematoda: Diplogastridae), a phoront of the bark beetle Scolytus multistriatus from the elm Ulmus glabra Huds. in Northwest Russia

Alexander Y. Ryss
Alexander Y. Ryss
, 
Kristina S. Polyanina
Kristina S. Polyanina
, 
Sergio Álvarez-Ortega
Sergio Álvarez-Ortega
 und 
Sergei A. Subbotin
Sergei A. Subbotin
   | 01. Jan. 2021
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Online veröffentlicht: 01. Jan. 2021
Seitenbereich: 1 - 25
DOI: https://doi.org/10.21307/jofnem-2021-025
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© 2021 Authors, published by Sciendo.
This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivatives 4.0 International License.

Figure 1:

Localization of the Rhabditolaimus ulmi dauer juveniles in pupae (A, B) and imago (C, D) of the beetle vector Scolytus multistriatus. A – on the elytra primordia; B – at the oviposititor opening; C, D – on the elytrae of imago. Scale: 1 mm for D, 2 mm for rest.
Localization of the Rhabditolaimus ulmi dauer juveniles in pupae (A, B) and imago (C, D) of the beetle vector Scolytus multistriatus. A – on the elytra primordia; B – at the oviposititor opening; C, D – on the elytrae of imago. Scale: 1 mm for D, 2 mm for rest.

Figure 2:

Rhabditolaimus ulmi. Morphology of adults. A – Male head; B – Female head; C – Head of molting juvenile (the molt of J4 into male); D – Male anterior part; E – Female tail; F – Male tail; G – Male genital system; H – Female genital system. Scale: 50 µm for D, G, H; 10 µm for rest.
Rhabditolaimus ulmi. Morphology of adults. A – Male head; B – Female head; C – Head of molting juvenile (the molt of J4 into male); D – Male anterior part; E – Female tail; F – Male tail; G – Male genital system; H – Female genital system. Scale: 50 µm for D, G, H; 10 µm for rest.

Figure 3:

Rhabditolaimus ulmi. Morphology of adults. A – Female anterior part; B – Male tail, lateral view; C – Male tail, ventral view; D – Female tail, lateral view; E, F – Female genital system sections; G – Female with J2 in eggshell (arrow, ovoviviparity). The location of the fragments (E: I-V) is shown in the diagram (F). Fragments: I – Mature oocyte before reverse movement along the oviduct to uterus; II – anterior spermatheca and oviduct; III – anterior uterus with maturating egg; IV – vulva, vagina and uteral chamber bordered by two sphincters; V – posterior spermatheca and oviduct. an – anus; bb – basal bulb; cd – cardia; ch – cheilostom; du – duct between spermatheca and uterus; e – egg; ex – excretory pore; gb – gubernaculum; gm – gymnostom; GR-I:1…GR-IV:4 – groups of ribs on the bursa alae: I…IV – number of the group, P+1… 4 – number of papillae in a group; mao – mature oocyte; mb – median bulb of pharynx; nr – nerve ring; oc – oocyte; ov – ovary; phm – phasmid, pr – head probolae; prc – procorpus of pharynx; s – spermatheca; sg – stegostom; sp – spicules, sph – sphincter; tt – tail tip; uch – uteral chamber; ut – uterus; v – vulva; va – vagina. Scale: 50 µm for G; 20 µm for rest.
Rhabditolaimus ulmi. Morphology of adults. A – Female anterior part; B – Male tail, lateral view; C – Male tail, ventral view; D – Female tail, lateral view; E, F – Female genital system sections; G – Female with J2 in eggshell (arrow, ovoviviparity). The location of the fragments (E: I-V) is shown in the diagram (F). Fragments: I – Mature oocyte before reverse movement along the oviduct to uterus; II – anterior spermatheca and oviduct; III – anterior uterus with maturating egg; IV – vulva, vagina and uteral chamber bordered by two sphincters; V – posterior spermatheca and oviduct. an – anus; bb – basal bulb; cd – cardia; ch – cheilostom; du – duct between spermatheca and uterus; e – egg; ex – excretory pore; gb – gubernaculum; gm – gymnostom; GR-I:1…GR-IV:4 – groups of ribs on the bursa alae: I…IV – number of the group, P+1… 4 – number of papillae in a group; mao – mature oocyte; mb – median bulb of pharynx; nr – nerve ring; oc – oocyte; ov – ovary; phm – phasmid, pr – head probolae; prc – procorpus of pharynx; s – spermatheca; sg – stegostom; sp – spicules, sph – sphincter; tt – tail tip; uch – uteral chamber; ut – uterus; v – vulva; va – vagina. Scale: 50 µm for G; 20 µm for rest.

Figure 4:

Rhabditolaimus ulmi. SEM. A, B – Female cephalic region. C – Male cephalic region; anterior annulus (aa), cephalic papilla (cp), amphid (am), lip papillae (lp), probolae (pr), oral opening (oo). D – Female excretory pore (ex) and deirid (di). E – Male lateral field at mid-body, lateral field ridges (fr), outer body ridges (ou). F – Vulva (vu) with vulval pore-like papilla (vp), wrinkled zone (wz) and smooth zone (sz). G, H – Female anus (an): G – Ventral view; H – Lateral view with a minute phasmid (psm). I – Male tail with caudal papillae, ventral view. P1 – unpaired precloacal papilla, OP1…OP7 – papillae on bursal alae, P2…P3 – inner papillae on ventral side of tail; cloacal opening (cl), pore-like phasmid (psm), ventral cultcular ridge (vr). J – Cloacal opening with spicule tips (sc) and P1 unpaired papilla. K – Tail end with OP3, group of OP4-OP6 papillae and phasmid (psm). L, M – OP7 papillae: L – Ventral view, M – Dorsal view. N – Female tail.
Rhabditolaimus ulmi. SEM. A, B – Female cephalic region. C – Male cephalic region; anterior annulus (aa), cephalic papilla (cp), amphid (am), lip papillae (lp), probolae (pr), oral opening (oo). D – Female excretory pore (ex) and deirid (di). E – Male lateral field at mid-body, lateral field ridges (fr), outer body ridges (ou). F – Vulva (vu) with vulval pore-like papilla (vp), wrinkled zone (wz) and smooth zone (sz). G, H – Female anus (an): G – Ventral view; H – Lateral view with a minute phasmid (psm). I – Male tail with caudal papillae, ventral view. P1 – unpaired precloacal papilla, OP1…OP7 – papillae on bursal alae, P2…P3 – inner papillae on ventral side of tail; cloacal opening (cl), pore-like phasmid (psm), ventral cultcular ridge (vr). J – Cloacal opening with spicule tips (sc) and P1 unpaired papilla. K – Tail end with OP3, group of OP4-OP6 papillae and phasmid (psm). L, M – OP7 papillae: L – Ventral view, M – Dorsal view. N – Female tail.

Figure 5:

Rhabditolaimus ulmi. The development in egg. A to D – Embryo cleavage; E – Juvenile J1; F to I – Juvenile J2 at different focal levels; J – Egg shell with a longitudinal slit indicating the place from where the juvenile coming away. Scale 20 µm.
Rhabditolaimus ulmi. The development in egg. A to D – Embryo cleavage; E – Juvenile J1; F to I – Juvenile J2 at different focal levels; J – Egg shell with a longitudinal slit indicating the place from where the juvenile coming away. Scale 20 µm.

Figure 6:

The juvenile development stages of males (left) and females (right). J1 .. J4 – juvenile stage numbers; pairs J2 to J3 .. J3-J4 – the molt of the junior stage to the senior one indicated by the stages numbers. hd – head; ph – pharynx; tl –tail. Scale 50 µm.
The juvenile development stages of males (left) and females (right). J1 .. J4 – juvenile stage numbers; pairs J2 to J3 .. J3-J4 – the molt of the junior stage to the senior one indicated by the stages numbers. hd – head; ph – pharynx; tl –tail. Scale 50 µm.

Figure 7:

The development of juveniles of the propagative generation. A – Juvenile J1 in egg; B – Juvenile J2 in egg; genital primordia and anal regions of juveniles: C, D – Juvenile J2; E, F – Molt J2-J3; G, H – Juvenile J3 (female); I, J – Juvenile J4 (female); K, L – Juvenile J3 (male); M, N – Molt J3-J4 (male); O, P – Juvenile J4 (male, early molt phase); Q – Molt J4 – male; R, S – Molt J4 – female: R – Only anterior branch of genital primordium; S – Tail of molting female J4 (arrows – shed cuticule). gp – genital primordium; mb – median bulb of pharynx; me – meiosis; mi – mitosis; sd – spermatids. Scale: 50 µm for Q to S; 20 µm for rest.
The development of juveniles of the propagative generation. A – Juvenile J1 in egg; B – Juvenile J2 in egg; genital primordia and anal regions of juveniles: C, D – Juvenile J2; E, F – Molt J2-J3; G, H – Juvenile J3 (female); I, J – Juvenile J4 (female); K, L – Juvenile J3 (male); M, N – Molt J3-J4 (male); O, P – Juvenile J4 (male, early molt phase); Q – Molt J4 – male; R, S – Molt J4 – female: R – Only anterior branch of genital primordium; S – Tail of molting female J4 (arrows – shed cuticule). gp – genital primordium; mb – median bulb of pharynx; me – meiosis; mi – mitosis; sd – spermatids. Scale: 50 µm for Q to S; 20 µm for rest.

Figure 8:

Molt of pre-adult juveniles J4 to adult male and female. Male tail (A) and its anterior part (B); Female: tail (C), Vulval region (D), Anterior part (E). an – anus; bs – bursa; gb – gubernaculum; hd – head with shed cuticle; sp – spicule; tl – tail. Scale: 20 µm.
Molt of pre-adult juveniles J4 to adult male and female. Male tail (A) and its anterior part (B); Female: tail (C), Vulval region (D), Anterior part (E). an – anus; bs – bursa; gb – gubernaculum; hd – head with shed cuticle; sp – spicule; tl – tail. Scale: 20 µm.

Figure 9:

Dauer juveniles J3D of the transmissive generation and the juveniles J3 of the propagative generation (males). Total outline: A, B – Dauer juvenile; C – Juvenile J3 of propagative generation; anterior part and pharynx: D – Dauer, E – Juvenile J3; head: F – Dauer, G – Juvenile J3; Genital primordium (male): H – Dauer, I – Juvenile J3. Scale: 50 µm for A, B, C; 20 µm for D, E; 10 µm for F to I.
Dauer juveniles J3D of the transmissive generation and the juveniles J3 of the propagative generation (males). Total outline: A, B – Dauer juvenile; C – Juvenile J3 of propagative generation; anterior part and pharynx: D – Dauer, E – Juvenile J3; head: F – Dauer, G – Juvenile J3; Genital primordium (male): H – Dauer, I – Juvenile J3. Scale: 50 µm for A, B, C; 20 µm for D, E; 10 µm for F to I.

Figure 10:

Phylogenetic relationships within Rhabditolaimus spp. and other related taxa as inferred from Bayesian analysis of the D2-D3 expansion segments of 28S rRNA gene sequences. Posterior probability values more than 70% are given on appropriate clades. New sequence indicated in bold.
Phylogenetic relationships within Rhabditolaimus spp. and other related taxa as inferred from Bayesian analysis of the D2-D3 expansion segments of 28S rRNA gene sequences. Posterior probability values more than 70% are given on appropriate clades. New sequence indicated in bold.

Measurements (in µm) and ratios of Rhabditolaimus ulmi juveniles.

Character/stage J2 in egg J2 free J3 male J3 female J4 male J4 female Dauer
n 5 10 7 11 7 10 10
L 190 ± 11.6 (176-200) 290.2 ± 31.4 (246-352) 358 ± 48 (292.6-427) 354.4 ± 38.6 (298-423) 475.6 ± 51.4 (373-558) 526.4 ± 66.8 (409-666) 338.8 ± 13.7 (326-354)
Body width 6.4 ± 0.6 (5.5-7.0) 7.8 ± 1.2 (5.5-10.5) 9.7 ± 1.5 (7.5-12.5) 9.6 ± 1 (7.5-12.0) 11.6 ± 1.1 (10-14) 12.6 ± 1.4 (9.5-16.5) 8.3 ± 0.3 (8.0-8.5)
Stoma 18 ± 3 (14-20) 23 ± 2 (21-26) 25 ± 2 (23-29) 25 ± 2 (23-29) 29 ± 2 (26-33) 30 ± 2 (25-35) 10 ± 1 (9-11)
Corpus (pro- & metacorpus) 22 ± 2 (20-24) 25 ± 2 (22-28) 27 ± 3 (21-31) 26 ± 3 (23-32) 33 ± 4 (28-41) 36 ± 5 (30-46) 33 ± 3 (30-37)
Pharynx 74.9 ± 12.9 (59-90) 91.7 ± 3.3 (86-97) 97 ± 7.4 (83-109) 92.6 ± 7.1 (87-111) 107.4 ± 7.8 (92-117) 119.4 ± 6.5 (106-132) 86.5 ± 3.9 (83-92)
Lip region width 1.9 ± 0.2 (1.5-2.0) 3.7 ± 0.3 (3.0-4.5) 3.9 ± 0.3 (3.5-4.5) 3.9 ± 0.7 (2.5-5.0) 4.5 ± 0.7 (3.5-6.0) 5.0 ± 0.8 (3.5-6.5) 4.2 ± 0.1 (4.0-5.5)
Nerve ring from anterior – 66.2 ± 3.6 (61-76) 73.1 ± 5 (64-81) 70.7 ± 5.8 (62-80) 83 ± 9.2 (65-96) 88.6 ± 5.4 (81-99) 61.5 ± 1.7 (60-64)
Excretory pore – 66.8 ± 4 (62-72) 79 ± 5 (73-86) 77 ± 5 (73-83) 84 ± 2 (82-86) 85 ± 2 (83-87) 65.4 ± 1.3 (64-671)
Tail 15 ± 3.9 (11-19) 46.3 ± 6.1 (35-60) 56.8 ± 9.2 (42-68) 55 ± 7.6 (38-65) 67.4 ± 6.6 (50-78) 71 ± 11 (41-88) 49.1 ± 0.5 (48-50)
Anal body width 3.5 ± 0.7 (2.5-4.0) 5.6 ± 0.8 (4.0-7.0) 7.3 ± 0.9 (5.5-8.5) 6.9 ± 0.9 (5.0-8.5) 8.9 ± 1.1 (7.5-11.5) 11 ± 1.6 (9.0-13.5) 5.3 ± 0.3 (5.0-6.0)
Genital primordium length (GPL) – 7.7 ± 1.7 (5.5-10.5) 12.5 ± 3.6 (6.5-20.5) 13.4 ± 6.8 (6.0-29.0) 77.7 ± 48.3 (12.5-149.5) 100.8 ± 63 (11.0-170.5) 11.5 ± 1.5 (10.0-13.5)
Genital primordium width (GPW) – 3.5 ± 0.8 (2.0-4.5) 4.1 ± 0.9 (2.5-5.5) 5.1 ± 1.5 (3.0-7.0) 6.4 ± 2.3 (3.5-12.0) 6.9 ± 1.3 (4.0-10.5) 3.8 ± 0.9 (2.6-4.7)
(V): GP center from anterior/ L, % – 62 ± 4 (56-66) 54 ± 4 (49-57) 55 ± 4 (50-59) 51 ± 4 (46-55) 57 ± 4 (50-60) 57 ± 1 (56-59)
a 30.2 ± 3.5 (25.7-33.0) 37.5 ± 2.7 (33.4-43.1) 37.1 ± 4.3 (29.5-43.7) 37.3 ± 3.8 (27.9-43.0) 41.2 ± 2.1 (38.1-44.2) 41.7 ± 2.9 (36.4-44.5) 40.6 ± 1.4 (38.7-42.0)
b 2.6 ± 0.4 (2.2-3.2) 3.2 ± 0.4 (2.6-3.8) 3.7 ± 0.4 (2.9-4.2) 3.7 ± 0.2 (3.4-3.9) 4.4 ± 0.4 (3.6-4.9) 4.4 ± 0.4 (3.6-5.2) 3.9 ± 0.2 (3.7-4.1)
c 12.9 ± 2.6 (10.5-15.7) 6.3 ± 0.9 (5.3-8.3) 6.4 ± 0.7 (5.0-7.8) 6.5 ± 0.7 (5.7-7.9) 7.1 ± 0.7 (6.0-8.3) 7.6 ± 1.7 (6.2-13.5) 6.9 ± 0.3 (6.6-7.2)
c′ 4.3 ± 0.8 (3.7-5.2) 8.3 ± 0.9 (6.2-9.6) 7.8 ± 0.9 (5.9-9.2) 8.0 ± 0.8 (6.1-9.2) 7.6 ± 1 (6.0-9.2) 6.7 ± 1.7 (3.0-8.7) 9.2 ± 0.4 (8.7-9.7)
Stoma/corpus 0.8 ± 0.2 (0.6-1.0) 0.9 ± 0.1 (0.8-1.1) 1 ± 0.1 (0.9-1.1) 0.9 ± 0.1 (0.8-1.1) 0.9 ± 0.1 (0.7-1.1) 0.8 ± 0.1 (0.6-1.1) 0.32 ± 0.03 (0.29-0.34)
Stoma/pharynx 0.23 ± 0.01 (0.22-0.24) 0.24 ± 0.02 (0.22-0.26) 0.25 ± 0.02 (0.23-0.27) 0.28 ± 0.02 (0.26-0.30) 0.27 ± 0.02 (0.24-0.29) 0.27 ± 0.02 (0.24-0.29) 0.12 ± 0.01 (0.11-0.12)
Corpus L/diam. 4.0 ± 0.4 (3.6-4.5) 3.6 ± 0.3 (3.2-4.4) 3.3 ± 0.4 (2.6-4.1) 3.5 ± 0.4 (2.7-4.2) 3.6 ± 0.4 (3.1-4.7) 3.6 ± 0.3 (3.0-4.1) 5.4 ± 0.5 (5.0-6.2)
Corpus/pharynx 0.30 ± 0.03 (0.26-0.34) 0.27 ± 0.02 (0.22-0.31) 0.27 ± 0.01 (0.25-0.29) 0.27 ± 0.01 (0.25-0.29) 0.3 ± 0.03 (0.2-0.3) 0.30 ± 0.04 (0.24-0.39) 0.37 ± 0.01 0.36-0.39)
Pharynx anter. part/pharynx L 0.53 ± 0.04 (0.48-0.57) 0.52 ± 0.03 (0.43-0.57) 0.54 ± 0.04 (0.48-0.57) 0.56 ± 0.04 (0.50-0.59) 0.55 ± 0.04 (0.50-0.59) 0.58 ± 0.04 (0.52-0.61) 0.48 ± 0.02 (0.47-0.51)
GPL/GPW – 2.3 ± 0.5 (1.4-3.1) 3.2 ± 1.1 (1.7-6.2) 2.5 ± 0.7 (1.7-4.4) 11.4 ± 4.9 (2.6-17.2) 14.4 ± 8.9 (2.7-30.0) 3.1 ± 0.6 (2.6-3.9)
GPL/L, % – 3 ± 1 (2-4) 4 ± 1 (2-7) 4 ± 2 (2-7) 16 ± 9 (3-27) 19 ± 12 (3-37) 4 ± 1 (3-5)

Measurements (in µm) and ratios of Rhabditolaimus ulmi adults.

Character/sex Adult male Adult female
n 20 20
L 715 ± 148 (557-1007) 804.6 ± 139.2 (633-1012)
Body width 20.7 ± 5.2 (15.5-28.0) 27.0 ± 7.0 (19.0-36.0)
Stoma 35 ± 4 (29-40) 34.2 ± 3.6 (31-41)
Corpus (pro- and metacorpus) 41 ± 3 (34-47) 43 ± 4 (36-47)
Pharynx 130.6 ± 12.3 (112-149) 132.5 ± 11.7 (116-146)
Lip region width 6.4 ± 0.9 (5.5-8.0) 6.9 ± 1.0 (5.5-8.0)
Nerve ring from anterior end 101.7 ± 12.7 (84-127) 102.6 ± 8.6 (90-112)
Excretory pore from anterior 107.5 ± 13.1 (92-134) 111.6 ± 9.8 (97-127)
Tail 28 ± 7 (19-39) 83 ± 8 (68-92)
Anal body width 12.2 ± 1.7 (10-15) 12.4 ± 2.2 (10-16)
Genital tube length (GTL) 411 ± 100 (295-586) 403 ± 172.7 (204-760)
Genital tube width (GTW) 19 ± 8.8 (9.5-35.5) 27.6 ± 9.0 (15.5-39.0)
Spicule length along arc 45.9 ± 6 (34.5-53.0) –
V % – 56 ± 2 (55-60)
a 35.1 ± 4.4 (27.8-39.9) 30.6 ± 4.6 (26.1-38.0)
b 5.4 ± 0.7 (4.8-6.8) 6.1 ± 0.6 (5.3-6.9)
c 25.8 ± 4.0 (20.5-33.4) 9.7 ± 1.0 (8.5-11.5)
c′ 2.3 ± 0.4 (1.7-3) 6.8 ± 1.0 (5.5-8.2)
Stoma/corpus 0.81 ± 0.02 (0.79-0.85) 0.82 ± 0.06 (0.74-0.87)
Stoma/pharynx 0.28 ± 0.01 (0.26-0.27) 0.27 ± 0.02 (0.25-0.29)
Corpus L/diam. 3.4 ± 0.3 (2.7-3.8) 3.2 ± 0.3 (2.7-3.6))
Corpus/pharynx 0.31 ± 0.03 (0.29-0.37) 0.32 ± 0.01 (0.30-0.36)
Pharynx anter./post. 0.60 ± 0.01 (0.59-0.61) 0.61 ± 0.04 (0.55-0.65)
GTL/GW 24.8 ± 10.8 (13.5-41.4) 14.7 ± 4.4 (10.8-23.3)
GTL/L, % 60 ± 10 (40-70) 50 ± 20 (30-90)

Measurements (in µm) and ratios of molting juveniles of Rhabditolaimus ulmi.

Character/stage J2-J3 J3-J4 male J3-J4 female J4-Adult male J4-Adult female
n 2 5 5 5 2
L 283, 321 461 ± 22.9 (435-482) 445.5 ± 67.3 (378-525) 582 ± 81 (472-669) 643, 646
Body width 6.0, 7.0 12.8 ± 0.9 (11.5-14.0) 12.0 ± 1.6 (10.5-14.5) 14.2 ± 1.6 (12-16) 15.5, 16.0
Stoma 21, 22 31 ± 3 (26-34) 29 ± 3 (24-32) 33 ± 3 (29-37) 33, 36
Corpus (pro- and metacorpus) 25, 26 34 ± 2 (31-35) 30 ± 5 (23-37) 39 ± 4 (35-45) 44, 48
Pharynx 88, 94 122 ± 10 (109-130) 118 ± 12 (103-131) 129 ± 9 (118-141) 139, 145
Lip region width 3.5, 4.0 5.1 ± 0.8 (4.0-6.0) 5 ± 1 (4.5-6.5) 6.2 ± 1.2 (5.0-7.5) 6.5, 7.0
Nerve ring from anterior end 65, 66 82 ± 5 (77-90) 81 ± 9 (69-93) 95 ± 6 (88-101) 97, 101
Tail 47, 53 64 ± 4 (59-69) 64 ± 6 (59-73) 71 ± 4 (67-75) 74, 76
Anal body width 4.5, 5.0 9.4 ± 1.1 (8.0-11.0) 7.6 ± 0.6 (7.0-8.5) 12.4 ± 2.3 (9.5-15.0) 10.0, 11.0
Genital primordium length (GPL) 7, 10 31.9 ± 13.2 (13-49) 21.2 ± 17.6 (9-52) 226.6 ± 111.2 (95-357) 145, 262
Genital primordium width (GPW) 2.0, 3.0 6.6 ± 0.9 (6.0-8.0) 6.1 ± 1.1 (5.5-8.0) 10.7 ± 4.6 (5.5-15.5) 9.5, 12.0
Spicule length along arc – – – 34.5-42.0 –
(V): GP center from anterior/L, % 47, 55 55 ± 4 (49-58) 55 ± 3 (49-57) 88 ± 6 (79-93) 50, 59
a 40.0, 54.3 36.1 ± 1.2 (34.3-37.4) 37.5 ± 7 (26.4-45.5) 40.9 ± 2.2 (38.5-43.4) 40.0, 41.8
b 3.0, 3.6 3.8 ± 0.1 (3.6-3.9) 3.7 ± 0.2 (3.4-4) 4.5 ± 0. 5 (3.8-4.9) 4.5, 4.6
c 6.0, 6.1 7.0 ± 0.5 (6.6-7.8) 7.0 ± 0.6 (6.2-7.6) 8.2 ± 0.8 (7.0-8.9) 8.5, 8.7
c′ 10.3, 11.7 6.9 ± 1 (5.6-8.4) 8.3 ± 0.8 (7.2-9.4) 5.8 ± 0.8 (4.9-7) 7.2, 7.6
Stoma/corpus 0.8, 1.0 0.9 ± 0.1 (0.8-1.0) 1.0 ± 0.2 (0.9-1.2) 0.8 ± 0.1 (0.7-0.9) 0.7, 0.8
Stoma/pharynx 0.23, 0.25 0.25 ± 0.01 (0.24-0.26) 0.24 ± 0.01 (0.23-0.25) 0.25 ± 0.01 (0.24-0.26) 0.24, 0.25
Corpus L/diam. 4.0, 4.5 3.7 ± 0.2 (3.5-4.0) 4.2 ± 0.6 (3.7-5.3) 3.8 ± 0.6 (2.9-4.3) 4.0, 5.2
Corpus/pharynx 0.27, 0.3 0.28 ± 0.01 (0.26-0.30) 0.25 ± 0.03 (0.21-0.28) 0.31 ± 0.02 (0.28-0.34) 0.30, 0.35
Pharynx anter. part/ pharynx L. 0.51, 0.52 0.53 ± 0.01 (0.52-0.54) 0.50 ± 0.02 (0.46-0.53) 0.56 ± 0.02 (0.54-0.59) 0.55, 0.58
GPL/GPW 3.2, 3.4 4.8 ± 1.6 (2.1-6.1) 3.6 ± 3.3 (1.7-9.4) 20.8 ± 3.5 (17.0-24.7) 15.1, 21.7
GPL/L, % 2, 3 7 ± 3 (3 - 11) 5 ± 3 (2-12) 37 ± 14 (20-55) 20, 40

Tabular key for identification of the Rhabditolaimus species.

Species/characters C1 C2 C3 C4 C5 C6 C7 C8 C9 C10 C11
R. neolongistoma 1 1 2 1 2 1 3 1 2 3 2
R. platypi 1 1 2 4 3 3 2 1 23 2 1
R. rifflei 1 1 3 1 2 2 1 1 3 2 3
R. curzii 1 1 3 1 12 3 2 2 1 2 3
R. erectus 1 1 3 2 2 3 1 2 1 2 2
R. anoplophorae 1 1 3 2 3 3 2 1 1 3 3
R. kishtwarensis 1 1 3 2 12 3 2 1 1 3 2
R. pellucidus 1 1 3 3 4 3 2 1 1 2 3
R. macrolaimus 1 1 3 12 1 3 2 2 1 2 3
R. pini 1 2 1 1 4 23 2 2 1 2 1
R. carolinensis 1 2 1 2 3 2 3 1 2 3 1
R. leuckarti 1 2 1 2 3 3 2 2 1 3 1
R. picei 1 2 1 2 4 3 2 1 3 3 1
R. nacogdochensis 1 2 1 3 3 3 1 1 3 3 1
R. vitautasi 1 2 2 23 3 23 2 2 1 2 2
R. inevectus 1 2 3 1 3 3 1 1 1 1 3
R. zamithi 1 3 3 2 2 2 2 1 1 1 2
R. robiniae 1 3 3 2 3 3 2 1 1 2 2
R. walkeri 1 3 3 2 3 3 2 1 1 2 3
R. ulmi 1 3 3 3 2 23 2 2 1 3 2
R. goodeyi 2 1 2 5 4 3 2 1 1 1 1
R. dendrophilus 2 1 3 4 4 3 3 1 1 1 2
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