Table 1.
Species and populations of Paratylenchus and Trophotylenchulus, used in the present study.
| Species | Locality | Associated plant | GenBank accession No. |
|---|---|---|---|
| Paratylenchus bukowinensis (Micoletzky, 1922) | Boldaji, Chaharmahal and Bakhtiari | Alfalfa (Medicago sativa L.) | MN088372 |
| P. coronatus (Colbran, 1965) | Dehdasht, Kohgiloyeh and Boyer-Ahmad | Alfalfa (Medicago sativa L.) | – |
| P. goodeyi (Oostenbrink, 1953) | Sureshjan, Chahrmahal and Bakhtiari | Rough bluegrass (Poa trivialis L.) | – |
| P. nawadus (Khan et al., 1967) | Behbahan, Khuzestan | Gundelia (Gundelia tournefortii L.) | MN088373 |
| P. paraperaticus (Kashi et al., 2009) | Andimeshk, Khuzestan | Howthorn (Crataegus aronia L.) | – |
| P. projectus Jenkins, 1956 | Dezful, Khuzestan | Jujube (Ziziphus spina-christi L.) | – |
| P. sheri (Raski, 1973; Siddiqi, 1986) | Dezful, Khuzestan | Ziziphus (Ziziphus nummularia (Burm. F.) Wight et Arn.) | MN088374 |
| P. similis (Khan et al., 1967) | Andimeshk, Khuzestan | Howthorn (Crataegus aronia L.) | MN088375 |
| P. teres (Raski, 1976; Siddiqi, 1986) | Dehdez, Khuzestan | Mount Atlas mastic or Baneh (Pistacia atlantica Desf.) | MN088376 |
| P. variabilis (Raski, 1975a, b) | Basht, Kohgiloyeh and Boyer-Ahmad | Unknown citrus | – |
| Trophotylenchulus arenarius (Raski, 1956; Siddiqi, 1999) | Baghmalek, Khuzestan | Weeping willow (Salix babylonica L.) | MK733978 |
| T. arenarius | Dehdasht, Kohgiloyeh and Boyer-Ahmad | Unknown grasses in a canebrake | MK733979 |
Table 2.
Morphometric characters of the two populations of Trophotylenchulus arenarius from Iran.
| Character\population | Dehdasht population | Baghmalek population | Total | ||||
|---|---|---|---|---|---|---|---|
| n | 10 females | 5 J2 | 1 male | 10 females | 5 J2 | 20 females | 10 J2 |
| L | 416 ± 36.6 (356–476) | 313 ± 9.8 (301–326) | 500 | 478 ± 42.5 (382–544) | 346 ± 21.8 (324–380) | 447 ± 50.1 (356–544) | 330 ± 23.7 (301–380) |
| a | 11.6 ± 0.7 (10.2–12.6) | 26.9 ± 1.9 (24.5–29.6) | 37.3 | 11.1 ± 0.8 (9.8–12.4) | 26.5 ± 1.5 (24.5–28.5) | 11.3 ± 0.8 (9.8–12.6) | 26.7 ± 1.6 (24.5–29.6) |
| b | 3.6 ± 0.3 (3.1–4.2) | 2.9 ± 0.1 (2.7–3.2) | 5.1 | 3.7 ± 0.1 (3.3–4) | 3.1 ± 0.1 (2.9–3.3) | 3.6 ± 0.2 (3.1–4.2) | 3 ± 0.2 (2.7–3.3) |
| c | 12.7 ± 1.1 (11.1–15.1) | 8.7 ± 0.4 (8.2–9.3) | 9.4 | 12.7 ± 1.3 (11.1–15.1) | 9.9 ± 0.8 (8.5–10.6) | 12.7 ± 1.2 (11.1–15.1) | 9.3 ± 0.9 (8.2–10.6) |
| c′ | 2.7 ± 0.2 (2.4–3.1) | 4.8 ± 0.2 (4.5–5) | 4.1 | 2.9 ± 0.3 (2.5–3.6) | 4.2 ± 0.3 (3.6–4.7) | 2.8 ± 0.3 (2.4–3.6) | 4.5 ± 0.4 (3.6–5) |
| V | 75.9 ± 2.2 (73.2–78.5) | – | – | 77.3 ± 1.5 (74.5–79.9) | – | 76.6 ± 1.9 (73.2–79.9) | – |
| Stylet | 13.5 ± 0.5 (12.3–14.5) | 12.5 ± 0.3 (12–13) | 9.6 | 13.5 ± 0.7 (12.4–14.3) | 13.2 ± 0.6 (12.5–13.8) | 13.5 ± 0.6 (12.3–14.5) | 12.9 ± 0.5 (12–13.8) |
| Conus | 6.7 ± 0.4 (6–7.5) | 6.2 ± 0.2 (6–6.6) | – | 7.1 ± 0.8 (6–8.3) | 6.9 ± 0.3 (6.5–7.3) | 6.9 ± 0.6 (6–8.3) | 6.5 ± 0.4 (6–7.3) |
| m (conus/stylet %) | 49.8 ± 2.2 (46.6–52.9) | 49.4 ± 0.9 (48.4–50.7) | – | 52.8 ± 5.6 (44.4–62.9) | 52.5 ± 1.3 (51–54.4) | 51.3 ± 4.4 (44.4–62.9) | 50.9 ± 1.9 (48.4–54.4) |
| Pharynx | 114 ± 12 (99–129) | 108 ± 8.7 (92–113) | 97 | 129 ± 9.2 (106–140) | 109 ± 5.7 (102–117) | 122 ± 12.8 (99–140) | 108 ± 7 (92–117) |
| Median bulb | 68.3 ± 5.6 (60–78) | 60.4 ± 5.1 (56–69) | 45 | 70.9 ± 3.7 (62.7–76) | 55.6 ± 2.9 (52–60) | 69.6 ± 4.8 (60–78) | 58 ± 4.7 (52–69) |
| MB | 59.9 ± 3.9 (55.2–66.6) | 56.2 ± 4.6 (51.8–61.6) | 46 | 55.1 ± 2.6 (51–59) | 50.9 ± 0.5 (50–51.3) | 57.5 ± 4.1 (51–66.6) | 53.6 ± 4.2 (50–61.6) |
| Excretory pore | 88.9 ± 9.8 (77–104) | 77.2 ± 4.4 (71–83) | 70 | 96.3 ± 9 (72.4–103) | 78.2 ± 3.1 (75–82) | 92.6 ± 9.9 (72.4–104) | 77.7 ± 3.6 (71–83) |
| Head-vulva | 316 ± 35.2 (261–373) | – | – | 370 ± 39.4 (284–435) | – | 343 ± 45.6 (261–435) | – |
| Head-anus | 383 ± 34.9 (327–441) | 277.4 ± 10 (265–289) | 447 | 440 ± 41.7 (349–506) | 312 ± 21.7 (291–344) | 412 ± 47.6 (327–506) | 294 ± 24 (265–344) |
| Tail length | 32.9 ± 3.3 (28–37) | 35.8 ± 1.3 (34–37) | 53 | 37.6 ± 3.4 (32.9–43) | 34.8 ± 2.7 (32–39) | 35.2 ± 4.1 (28–43) | 35.3 ± 2.1 (32–39) |
| Body width | 35.8 ± 3.7 (30–41.5) | 11.6 ± 0.5 (11–12.5) | 13.4 | 43.1 ± 5.4 (35.2–55) | 13 ± 1.3 (12–15.5) | 39.4 ± 5.9 (30–55) | 12.3 ± 1.2 (11–15.5) |
| Anal body width | 11.8 ± 0.8 (10.5–13.5) | 7.4 ± 0 (7.3–7.5) | 12.8 | 12.9 ± 0.9 (11.5–14.5) | 8.3 ± 0.5 (7.5–9) | 12.3 ± 1 (10.5–14.5) | 7.8 ± 0.5 (7.3–9) |
| Annulus width | 1.2 ± 0.1 (1.0–1.4) | – | 1.4 | 1.0 ± 0.1 (0.9–1.3) | – | 1.1 ± 0.1 (0.9–1.4) | – |
| Spicules | – | – | 21 | – | – | – | – |
| Gubernaculum | – | – | 4.5 | – | – | – | – |
Table 3.
Morphometric characters of Paratylenchus goodeyi, P. nawadus and P. sheri from Iran.
| Characters\species | P. goodeyi | P. nawadus | P. sheri | ||
|---|---|---|---|---|---|
| n | 8 females | 10 females | 5 males | 12 females | 6 males |
| L | 335 ± 38.8 (263–368) | 358 ± 24.1 (330–400) | 353 ± 10.1 (340–367) | 424 ± 12.5 (406–446) | 449 ± 24.7 (412–488) |
| a | 20.7 ± 2.2 (15.4–22.5) | 25.2 ± 2.5 (21.1–29.3) | 28 ± 1.6 (26.1–30.3) | 25.1 ± 1.6 (22–27) | 33.2 ± 2 (30.8–36.1) |
| b | 4.1 ± 0.7 (2.8–5.3) | 3.5 ± 0.2 (3.1–3.8) | 3.5 ± 0.2 (3.3–3.9) | 4 ± 0.1 (3.8–4.2) | 4.5 ± 0.2 (4.1–4.8) |
| c | 12 ± 1.3 (9.3–13.2) | 12.7 ± 1.6 (10.1–15.6) | 11.1 ± 1 (10–12.5) | 11.4 ± 0.5 (10.7–12.5) | 10.5 ± 1.4 (8.5–12.8) |
| c′ | 2.9 ± 0 (2.8–3) | 3 ± 0.3 (2.5–3.6) | 3.1 ± 0.3 (2.5–3.6) | 3.6 ± 0.1 (3.3–3.9) | 3.9 ± 0.5 (3.2–4.6) |
| V | 81.8 ± 2.4 (79.4–87.4) | 83.1 ± 0.7 (81.8–84.2) | – | 80.2 ± 1 (78.5–82.3) | – |
| Stylet | 48.8 ± 1.4 (46.9–51.2) | 20.5 ± 1 (18.7–22) | – | 24.1 ± 0.5 (23–25) | – |
| m (conus/stylet %) | 76.7 ± 1.2 (74.8–78.1) | 65 ± 2.4 (61.1–69.9) | – | 64.5 ± 1.3 (62.5–66.5) | – |
| Pharynx | 82.5 ± 9.9 (67–93.5) | 99.8 ± 6.9 (91.7–112) | 100 ± 4.8 (93–105.6) | 104.6 ± 4.6 (98–112) | 99.1 ± 8.3 (85–109) |
| MB | 73 ± 6.6 (65.8–80.5) | 56.4 ± 1 (55.1–58) | – | 56.3 ± 1 (54.7–58) | – |
| Excretory pore | 77 ± 9 (65–89) | 82.5 ± 5.1 (77–89) | 74.6 ± 8.4 (67–88.7) | 91.6 ± 3.2 (87–96) | 86 ± 10.3 (66–95) |
| Head-vulva | 274 ± 31.2 (209–300) | 298 ± 20.3 (274.6–332) | – | 340 ± 12.4 (319–360) | – |
| Head-anus | 307 ± 38.8 (235–340) | 329 ± 23 (305–369) | 321 ± 9.5 (313–336) | 389 ± 11.4 (372–409) | 406 ± 20.7 (376–441) |
| Tail length | 27.8 ± 0.5 (27–28.5) | 28.4 ± 3.9 (22.1–34.4) | 31.8 ± 3 (27–34.9) | 37 ± 1.9 (33–40) | 43.1 ± 7 (34–53) |
| Body width | 16.2 ± 1.3 (13.9–17.4) | 14.2 ± 1.2 (12.5–16.3) | 12.5 ± 0.5 (11.8–13) | 16.9 ± 0.9 (15.5–18.5) | 2.8 ± 0.2 (2.5–3.2) |
| Annulus width | 1.3 ± 0.1 (1.2–1.5) | 1.3 ± 0.1 (1.1–1.5) | 1.3 ± 0.1 (1.2–1.4) | 1.3 ± 0.1 (1.1–1.5) | 1.3 ± 0.1 (1.2–1.5) |
| st/L × 100 | 14.7 ± 1.7 (13.1–18.2) | 5.7 ± 0.4 (5.1–6.4) | – | 5.7 ± 0.1 (5.5–5.8) | – |
| Spicules | – | – | 23.4 ± 1.4 (22–25.2) | – | 24.8 ± 0.6 (24–25.5) |
| Gubernaculum | – | – | 4.5 ± 0.4 (4.1–5.2) | – | 4.8 ± 0.2 (4.5–5) |
Table 4.
Morphometric characters of Paratylenchus teres, P. bukowinensis and P. coronatus from Iran.
| Characters\species | P. teres | P. bukowinensis | P. coronatus | |
|---|---|---|---|---|
| n | 8 females | 7 females | 5 males | 6 females |
| L | 337 ± 26.7 (285–365) | 382 ± 41 (304–425) | 367 ± 25.3 (325–390) | 308 ± 24.1 (273–347) |
| a | 22.5 ± 1.4 (19.5–23.9) | 24.6 ± 2.1 (22.1–28.6) | 30.1 ± 2.5 (27–32.8) | 23.7 ± 2.2 (20.5–26.5) |
| b | 2.4 ± 0.1 (2.2–2.6) | 3.9 ± 0.3 (3.3–4.5) | 4.1 ± 0.4 (3.4–4.6) | 3.7 ± 0.3 (3.4–4.2) |
| c | 15.5 ± 2.5 (12.3–19.1) | 15.5 ± 1.7 (12.1–17.3) | 11.2 ± 0.4 (10.7–11.8) | 15.2 ± 1.5 (13–17.7) |
| c′ | 2.8 ± 0.2 (2.5–3.3) | 2.7 ± 0.1 (2.6–2.8) | 3 ± 0.2 (2.8–3.3) | 2.4 ± 0 (2.3–2.5) |
| V | 75.7 ± 1.2 (74.6–78) | 85.6 ± 4.7 (83.4–96.4) | – | 80.5 ± 0.9 (79.1–82.1) |
| Stylet | 76.7 ± 6.4 (61.3–82) | 25.4 ± 1.4 (22.7–26.7) | 11.7 ± 1 (10–13) | 30.2 ± 1 (28.9–32) |
| m (conus/stylet %) | 89.5 ± 0.7 (88.2–90.7) | 63.3 ± 1.2 (61.2–64.9) | – | 68.9 ± 1.8 (66.7–71.1) |
| Pharynx | 139 ± 10.7 (117–149) | 97.6 ± 6 (90.3–105.7) | 90 ± 12.2 (79–110) | 82.5 ± 5.1 (79–92.8) |
| MB | 70.5 ± 1.4 (68–72.3) | 58.2 ± 1.4 (56.7–60.6) | – | 57.7 ± 3.2 (51.5–60.5) |
| Excretory pore | 81.5 ± 5.5 (68–85) | 97.6 ± 6 (90.3–105.7) | 91 ± 7.3 (79–98) | 82.5 ± 5.1 (79–92.8) |
| Head-vulva | 255.6 ± 20.8 (219.7–285) | 327 ± 39.4 (255–369) | – | 248 ± 19.5 (220–280) |
| Head-anus | 315 ± 27.4 (264–345) | 357 ± 41 (279–401) | 335 ± 23 (297–357) | 287 ± 24.5 (252–327) |
| Tail length | 22.1 ± 2.9 (19–27) | 24.5 ± 0.6 (23.5–25.1) | 32.6 ± 2.6 (28–34) | 20.2 ± 0.4 (19.5–21) |
| Body width | 14.9 ± 0.5 (14.2–15.8) | 15.5 ± 1.9 (12.9–19.2) | 12.2 ± 0.5 (11.5–13) | 12.9 ± 1 (11.7–14.5) |
| Annulus width | 1.2 ± 0.1 (0.9–1.3) | 1.4 ± 0.2 (1.2–1.7) | 1.3 ± 0.1 (1.2–1.4) | 1.2 ± 0.1 (1.1–1.3) |
| st/L × 100 | 22.8 ± 1.2 (21.1–24.6) | 6.7 ± 0.5 (6.3–7.5) | 3.2 ± 0.3 (2.7–3.6) | 9.9 ± 0.9 (8.5–11.1) |
| Spicules | – | – | 20.1 ± 0.8 (19.5–21.5) | – |
| Gubernaculum | – | – | 4.5 ± 0.4 (4.2–5) | – |
Table 5.
Morphometric characters of Paratylenchus paraperaticus, P. projectus, P. similis and P. variabilis from Iran.
| Characters\species | P. paraperaticus | P. projectus | P. similis | P. variabilis |
|---|---|---|---|---|
| n | 5 females | 5 females | 22 females | 17 females |
| L | 303 ± 17.2 (283–325) | 405 ± 28.1 (363–437) | 358 ± 44.4 (281–451) | 325 ± 18.1 (291–355) |
| a | 25.6 ± 0.6 (25–26.4) | 20.9 ± 1.6 (18.7–23.4) | 22.7 ± 1.9 (17.2–26.5) | 23.2 ± 1.5 (20.7–27.1) |
| b | 2.4 ± 0.1 (2.2–2.5) | 4.3 ± 0.3 (4–4.7) | 3.8 ± 0.3 (3–4.6) | 3.9 ± 0.2 (3.5–4.3) |
| c | 8.6 ± 0.8 (7.4–9.8) | 13.2 ± 1 (11.5–14.5) | 10.9 ± 0.6 (10–12.6) | 16.1 ± 1 (14.2–17.7) |
| c′ | 3.2 ± 0.1 (3–3.4) | 3 ± 0.1 (2.7–3.2) | 3.4 ± 0.2 (2.9–4.1) | 2.4 ± 0.1 (2.2–2.5) |
| V | 75.4 ± 1.9 (72.2–77.3) | 81.8 ± 0.8 (80.4–83) | 81.2 ± 2 (75.3–84.3) | 85.2 ± 1 (83.4–87.4) |
| Stylet | 80.1 ± 2.2 (76.7–82.6) | 25.4 ± 0.5 (24.3–25.9) | 16.9 ± 0.8 (15.5–18.9) | 14.1 ± 0.6 (12.8–14.9) |
| m (conus/stylet %) | 90.5 ± 5.3 (87.6–100) | 62.2 ± 2.8 (58.4–66.5) | 61.8 ± 2 (57.6–65.8) | 62.5 ± 2.6 (57.3–69.9) |
| Pharynx | 124.4 ± 6.5 (113.2–128.9) | 92.6 ± 5.1 (88.1–102.1) | 92.5 ± 6.2 (79.6–114.6) | 82.1 ± 3.1 (76.4–87.8) |
| MB | 76.6 ± 3.3 (74.1–82.3) | 58.6 ± 2.6 (56.5–63.6) | 54.1 ± 2.1 (45.6–56.4) | 50.5 ± 1.9 (45.5–53.9) |
| Excretory pore | 83.4 ± 2.3 (81.7–87.4) | 92.6 ± 5.1 (88.1–102.1) | 86.2 ± 4.8 (76–102) | 72.7 ± 2.6 (69–78) |
| Head-vulva | 229 ± 16.5 (208–247) | 331 ± 25.1 (292–358) | 291 ± 32.5 (220–346) | 277 ± 17.2 (246–304) |
| Head-anus | 268 ± 18.6 (245–292) | 374 ± 28 (332–406) | 325 ± 41.8 (254–413) | 305 ± 18.1 (270–335) |
| Tail length | 35 ± 1.8 (33–38) | 30.6 ± 1.5 (29.1–33.1) | 32.5 ± 2.8 (27–38) | 20.1 ± 0.5 (19–21.5) |
| Body width | 11.8 ± 0.6 (10.9–12.5) | 19.3 ± 2.1 (16.8–22.6) | 15.7±1.7 (13.5–21.2) | 14 ± 0.9 (11.4–15.4) |
| Annulus width | 0.9 ± 0.1 (0.8–1.1) | 1.5 ± 0.1 (1.4–1.6) | 1.1 ± 0.1 (0.9–1.4) | 1.3 ± 0.1 (1.1–1.5) |
| st/L × 100 | 26.5 ± 0.8 (25.4–27.4) | 6.3 ± 0.5 (5.6–7.0) | 4.8 ± 0.5 (3.9–5.8) | 4.4 ± 0.3 (3.8–4.7) |
Figure 1:
Trophotylenchulus arenarius from Iran: Dehdasht population with narrow tail terminus in females (B, E–H, J, O) and Baghmalek population with blunt tail terminus in females (A, C, D, I, K–N). Female (A, B, K–O), male (F, J) and second-stage juvenile (C–E, G–I). A, B: entire body; C–E: head and stylet; F: anterior end and pharyngeal region; G–O: posterior end.
Figure 2:
Trophotylenchulus arenarius from Iran: Dehdasht population (A, C, E, F, H, J–L) and Baghmalek population (B, D, G, I, M, N). Female (A–D, M, N), male (E, L) and second-stage juvenile (F–K). A, B: entire body; C–G: head and stylet; H–J: anterior end and pharyngeal region; K: lateral field at mid-body; L: posterior end; M, N: pharyngeal median bulb and excretory pore (scale bars: A = 50 µm; B = 100 µm; C–N = 10 µm).
Figure 3:
Posterior end of females of Trophotylenchulus arenarius from Iran: Dehdasht population (A–E) and Baghmalek population (F–O) (scale bars = 10 µm).
Figure 4:
Posterior end of second-stage juveniles of Trophotylenchulus arenarius from Iran. Dehdasht population (A–H) and Baghmalek population (I–P) (scale bars = 10 µm).
Figure 5:
Diagnostic characters of Paratylenchus goodeyi (A–E), P. nawadus (F–J) and P. teres (K–M) from Iran. Female (A–F, H, I, K–M) and male (G, J). A, B, F, G, K: anterior end and pharyngeal region; C, H, J: reproductive system; D, E, I, L, M: posterior end.
Figure 6:
Paratylenchus goodeyi from Iran. Female (A–G). A: entire body; B: anterior end; C: part of reproductive system; D–G: posterior end (scale bars: A = 20 µm; B–G = 5 µm).
Figure 7:
Paratylenchus nawadus from Iran. Female (A, C–G) and male (B, H–J). A, J: entire body; B, C, F: anterior end; D, E: vulval region; G, I: posterior end; H: spicules (scale bars: A = 20 µm; B–I = 5 µm, J = 50 µm).
Figure 8:
Diagnostic drawings of Paratylenchus sheri from Iran. Female (A, C, D, F–H, J–O) and male (B, E, I). A, B: entire body; C: head and stylet; D, E: anterior end and pharyngeal region; F–H: reproductive system; I–M: posterior end; N, O: cross sections at mid-body.
Figure 9:
Paratylenchus sheri from Iran. Female (A, B, E–M) and male (C, D, N). A, D: entire body; B, C: anterior end; E: cross sections from near mid-body; F–I: vulval region and part of reproductive system; J–M: posterior end; N: cloaca and genital organs (scale bars: A, D = 50 µm; B, C, E–I = 5 µm).
Figure 10:
Paratylenchus teres from Iran. Female (A–D, F–H, J–L) and fourth-stage juvenile (E, I). A: entire body; B–E: head and stylet; F, J: vulval region; G, H: lateral field; I–L: posterior end (scale bars: A = 20 µm; B–I = 5 µm).
Figure 11:
The 50% majority rule consensus tree from Bayesian analysis generated from the D2–D3 expansion segments of 28 S rRNA gene dataset under the GTR + I + G model. Posterior probabilities for BI analysis more than 50% are given for appropriate clades. The new sequencesare indicated in bold.