Table 1.
Morphometrics of C. tianzhuensis n. sp.
| Stage | Character | Holotype | Paratype |
|---|---|---|---|
| Cyst | |||
| n | 20 | ||
| L (excluding length) | 531 | 571.2 ± 79.1 (511.0–761.0) | |
| Diam. | 429 | 454.3 ± 56.8 (360.5–558.0) | |
| L/Diam. | 1.23 | 1.3 ± 0.1 (1.1–1.6) | |
| Fenestral diam. | 22.8 | 23.4 ± 3.8 (20.0–31.5) | |
| Second-stage juvenile | |||
| n | 20 | ||
| Body length | 538.5 ± 25.0 (494.5–591.5) | ||
| Body width at mid-body | 23.2 ± 1.4 (20.0–25.0) | ||
| a | 23.4 ± 2.0 (20.6–27.2) | ||
| b | 3.7 ± 0.3 (3.1–4.2) | ||
| c | 10.0 ± 0.8 (8.4–11.6) | ||
| c′ | 3.9 ± 0.3 (3.5–4.5) | ||
| Lip region height | 4.8 ± 0.4 (4.0–5.0) | ||
| Lip region diam. | 10.7 ± 0.8 (9.0–12.0) | ||
| Stylet length | 24.9 ± 0.8 (22.5–26.0) | ||
| Stylet base height | 2.5 ± 0.3 (2.0–3.0) | ||
| Stylet base width | 5.1 ± 0.54 (4.0–5.5) | ||
| Median bulb from anterior end (MB) | 77.8 ± 4.2 (69.5–85.0) | ||
| Opening of dorsal pharyngel gland from stylet base (DGO) | 4.9 ± 0.5 (4.0–6.5) | ||
| Excretory pore from anterior end (EP) | 111.2 ± 6.2 (97.5–124.0) | ||
| Median bulb width (MBW) | 11.8 ± 1.1 (9.5–13.5) | ||
| Diam. at anus | 13.8 ± 0.5 (11.5–15.0) | ||
| Tail length | 54.1 ± 3.6 (46.5–59.0) | ||
| Hyaline portion tail | 25.8 ± 2.1 (21.5 –28.5) | ||
| L/MB | 6.9 ± 0.5 (6.1–8.0) | ||
| TL/H | 2.1 ± 0.2 (1.8–2.4) | ||
| Egg | |||
| n | 20 | ||
| Length | 117.9 ± 11.6 (101.5–144.0) | ||
| Width | 51.3 ± 4.1 (41.0–61.5) | ||
| Length/Width | 2.3 ± 0.3 (1.9–2.9) | ||

Figure 1:
Line drawing of C. tianzhuensis n. sp. A: Anterior region of second-stage juvenile; B: Head of second-stage juvenile; C, H: Cyst and cysts. D: Lateral field; E: Fenestration in vulval cone; F, G: Tail of second-stage juvenile (Scale bar: H = 500 μm; C = 200 μm; A, F, E = 20 μm; B = 10 μm; D, G = 5 μm).

Figure 2:
Light micrographs of C. tianzhuensis n. sp. A: Cyst; B: Cysts; C: Crush cyst; C: Female attached on the root (Scale bar: A, D = 500 μm; B, C = 1 mm).

Figure 3:
Vulval cones of C. tianzhuensis n. sp. A, C: Fenestration in vulval cone (inside); B, D: Fenestration in vulval cone (outside); E, F: Cyst surface punctations. (Scale bar = 20 µm).

Figure 4:
Egg of C. tianzhuensis n. sp. A: Smooth eggshell; B: Embryo egg; C: Body of developed J2 in egg. (Scale bar = 20 µm).

Figure 5:
Light micrographs of second stage juvenile of C. tianzhuensis n. sp. A: Entire body; B, C: Head region; D: Anterior region of J2; E: Lateral field; F: Genital primordium; G: Annulation of tail; H-I: Tail; J: Lateral field of tail. (Scale bar: A = 50 μm, B, C, D, E, F = 20 μm, H, I = 10 μm, G, J = 5 μm).
Table 2.
Morphological and morphometrics of characters of cysts, eggs and J2s, useful for identification of Cactodera species.
| Stage | ||||||||||||
|---|---|---|---|---|---|---|---|---|---|---|---|---|
| Cyst | Second-stage juvenile | Egg | ||||||||||
| Species | Length | Width | L/W ratio | Vulval denticles | Fenestral diam. | Length | DGO | Stylet Length | Tail Length | Hyaline tail length | Eggshell surface | Original des. |
| C. acnidae | 504–857 | 319–493 | 1.2–2.4 | – | 30 × 33 | 361–448 | 2.5–3.0 | 19–25 | 43–48 | 17–26 | Smooth | Schuster and Brezina (1979) |
| C. amaranthi | 525–774 | 370–550 | 1.1–1.7 | Present | 25–38 | 340–460 | 3.9–5.1 | 20–21 | 32–40 | 12–16 | Smooth | Golden and Raski (1977) |
| C. cacti | 389–658 | 323–598 | 1.0–1.4 | Present | 16–30 | 456–540 | 2.9–4.4 | 24–26 | 49–60 | 14–21 | Punctate | Graney and Bird (1990) |
| C. chenopodiae | 423–585 | 283–398 | 1.2–1.7 | Absent | 20–26 | 438–539 | 3.1–4.3 | 22–26 | 39–51 | 17–28 | Punctate | Feng et al. (2018) |
| C. eremica | 530–810 | 290–590 | 1.2–1.9 | Absent | 14–25 | 440–510 | 3.5–6.0 | 25–28 | 36–47 | 17–23 | Punctate | Baldwin and Bell (1985) |
| C. estonica | 686–1014 | 312–468 | 2.0–2.4 | Present | 18–30 | 426–465 | 3.4–4.3 | 22–24 | 36–44 | 14–21 | Smooth | Golden and Raski (1977) |
| C. evansi | 416–528 | 284–384 | 1.2–1.7 | – | 18–23 | 358–420 | 2.8–4.0 | 20–24 | 34–44 | 16–23 | Punctate | Cid Del Prado and Rowe (2000) |
| C. galinsogae | 453–675 | 284–508 | 1.1–1.7 | Absent | 33–56 | 358–443 | – | 19–31 | 26–45 | 10–24 | Punctate | Soto et al. (2003) |
| C. milleri | 550–849 | 419–598 | 1.2–1.6 | Present | 14–18 | 370–479 | 3.2–5.1 | 21–23 | 37–49 | 15–21 | Punctate | Graney and Bird (1990) |
| C. radicale | 553–986 | 220–626 | 1.3–2.6 | Absent | 17–28 | 467–520 | 3.6–5.8 | 20–27 | 46–60 | 15–28 | Smooth | Subbotin et al. (2010) |
| C. rosae | 460–840 | 280–560 | 1.2–2.1 | Present | 10–21 | 348–472 | – | 16–26 | 31–68 | 4–8 | Punctate | Cid Del Prado and Miranda (2008) |
| C. salina | 415–742 | 193–475 | 1.4–2.2 | Absent | 20–28 | 410–514 | 2.5–4.0 | 23–25 | 31–48 | 10–31 | Smooth | Baldwin et al. (1997) |
| C. solani | 291–581 | 204–505 | 1.2–1.4 | Present | 20–36 | 379–511 | 3.7–6.9 | 24–27 | 28–49 | 12–23 | Punctate | Escobar-Avila et al. (2020) |
| C. thornei | 485–806 | 286–581 | 1.2–1.9 | Present | 31–36 | 446–620 | 5–7 | 25–28 | 49–64 | 23–28 | Punctate | Golden and Raski (1977) |
| C. tianzhuensis n. sp. | 511–761 | 361–558 | 1.1–1.6 | Present | 20–32 | 495–592 | 4.0–6.5 | 23–26 | 47–59 | 22–29 | Smooth | In this study |
| C. torreyanae | 364–712 | 92–432 | 1.4–2.9 | Absent | 20×26 | 390–550 | 2–4 | 21–23 | 32–45 | 16–25 | Smooth | Cid Del Prado and Subbotin (2014) |
| C. weissi | 524–598 | 350–394 | 1.2–2.3 | Present | 29–38 | 407–489 | 4.5–5.6 | 20–22 | 43–50 | 17–24 | Smooth | Golden and Raski (1977) |

Figure 6:
Molecular phylogenetic tree of C. tianzhuensis n. sp. (highlighted in bold) inferred from 28 S D2/D3 extension region under GTR + I + G model. The posterior probability values exceeding 50% are given on appropriate clades. *Originally identified as C. estonica in the GenBank. **Originally identified as C. rosae in the GenBank. ***Originally identified as C. estonica in the GenBank.

Figure 7:
Molecular phylogenetic tree of C. tianzhuensis n. sp. (highlighted in bold) inferred from ITS region under GTR + I + G model. The posterior probability values exceeding 50% are given on appropriate clades. *Originally identified as C. estonica in the GenBank. **Originally identified as C. eremica in the GenBank. ***Originally identified as C. estonica in the GenBank. ****Originally identified as C. estonica in the GenBank.