Table 1
Numbers of species of Heterodera for which one or more gene sequences of one or more molecular markers (18S, ITS, 28S, and coxI) are available on GenBank.
| Species of Heterodera | 18S | ITS | 28S | coxI | All |
|---|---|---|---|---|---|
| Species with at least one sequence | 18 | 55 | 36 | 38 | 13 |
| Species with more than one sequence | 13 | 38 | 22 | 29 | 11 |
Table 2
Species pairs of Heterodera which are inadequately or poorly distinguished from one another using one or more of the standard molecular markers evaluated (18S, ITS, 28S, coxI), including minimum base pair differences observed between sequences of problematic species pairs.
| Species group | Species pair | Gene region | Minimum base pair difference | Delineation power | Notes |
|---|---|---|---|---|---|
| Avenae | H. avenae–H. filipjevi | 18S | 1 | Inadequate | |
| Avenae | H. avenae–H. hordecalis | 18S | 4 | Weak | H. hordecalis not monophyletic |
| in NJ/ML analyses; potentially | |||||
| distinguishable from closely | |||||
| related species in isolated | |||||
| analyses. | |||||
| Avenae | H. avenae–H. mani | 18S | 1 | Inadequate | |
| Avenae | H. filipjevi–H. hordecalis | 18S | 1 | Inadequate | |
| Avenae | H. filipjevi–H. mani | 18S | 2 | Inadequate | |
| Avenae | H. hordecalis–H. mani | 18S | 3 | Weak | See above note. |
| Avenae | H. arenaria–H. avenae | ITS | 0 | Inadequate | |
| Avenae | H. avenae–H. pratensis | ITS | 0 | Inadequate | |
| Avenae | H. avenae–H. australis | ITS | 2 | Weak | Although only two bp different, |
| sequences of H. australis form | |||||
| clade with some sequences | |||||
| of “H. avenae” which have | |||||
| been shown to be H. australis | |||||
| (see Subbotin et al., 2018). | |||||
| Avenae | H. avenae–H. mani | ITS | 2 | Weak | Despite only two bp difference, |
| sequences of H. mani form | |||||
| monophyletic clade to | |||||
| exclusion of other closely | |||||
| related sequences. | |||||
| Avenae | H. aucklandica–H. | 28S | 1 | Inadequate | Only one 28S sequence of |
| avenae | H. aucklandica available for | ||||
| comparison. | |||||
| Avenae | H. aucklandica–H. | 28S | 1 | Inadequate | Two 28S sequences of |
| hordecalis | H. hordecalis 9 bp different | ||||
| Avenae | H. aucklandica–H. | 28S | 1 | Inadequate | Only one 28S sequence of |
| pratensis | H. pratensis available for | ||||
| comparison. | |||||
| Avenae | H. avenae–H. pratensis | 28S | 0 | inadequate | See above note. |
| Avenae | H. avenae–H. hordecalis | 28S | 0 | Inadequate | Two 28S sequences of |
| H. hordecalis 9 bp different | |||||
| Avenae | H. arenaria–H. avenae | cox1 | 2 | Weak | Despite only two bp |
| difference, H. arenaria forms | |||||
| monophyletic clade within | |||||
| H. avenae group. | |||||
| Cyperi | H. elachista–H. oryzae | ITS | 1 | Inadequate | |
| Cyperi | H. elachista–H. oryzae | 28S | 0 | Inadequate | |
| Goettingiana | H. carotae–H. cruciferae | ITS | 1 | Inadequate | |
| Goettingiana | H. goettingiana–H. | 28S | 0 | Inadequate | Two 28S sequences of |
| microulae | H. goettingiana are 19 bp | ||||
| different from one another; one | |||||
| potentially misidentified | |||||
| Goettingiana | H. carotae–H. urticae | 28S | 2 | Weak | Only one 28S sequence |
| of H. urticae available for | |||||
| comparison. | |||||
| Goettingiana | H. carotae–H. cruciferae | 28S | 2 | Weak | Only one 28S sequence of |
| H. cruciferae available for | |||||
| comparison (two on GenBank | |||||
| but one appears to actually be | |||||
| H. schachtii). | |||||
| Goettingiana | H. carotae–H. cruciferae | cox1 | 1 | Inadequate | Three cox1 sequences |
| available for H. cruciferae, one | |||||
| 15 bp different from others. | |||||
| Schachtii | H. betae–H. glycines | 18S | 2 | Weak | H. glycines forms |
| monophyletic clade to | |||||
| exclusion of H. betae, | |||||
| H. schachtii & H. trifolii in ML, | |||||
| but not NJ, analyses. | |||||
| Schachtii | H. betae–H. schachtii | 18S | 0 | Inadequate | |
| Schachtii | H. betae–H. trifolii | 18S | 0 | Inadequate | |
| Schachtii | H. schachtii–H. glycines | 18S | 2 | Weak | See above note. |
| Schachtii | H. schachtii–H. trifolii | 18S | 0 | Inadequate | |
| Schachtii | H. trifolii–H. glycines | 18S | 2 | Weak | See above note. |
| Schachtii | H. betae–H. daverti | ITS | 0 | Inadequate | |
| Schachtii | H. betae–H. schachtii | ITS | 0 | Inadequate | |
| Schachtii | H. betae–H. trifolii | ITS | 0 | Inadequate | |
| Schachtii | H. ciceri–H. schachtii | ITS | 2 | Inadequate | |
| Schachtii | H. ciceri–H. trifolii | ITS | 1 | Inadequate | |
| Schachtii | H. daverti–H. schacthii | ITS | 0 | Inadequate | |
| Schachtii | H. daverti–H. trifolii | ITS | 0 | Inadequate | |
| Schachtii | H. glycines–H. | ITS | 1 | Inadequate | |
| medicaginis | |||||
| Schachtii | H. schachtii–H. trifolii | ITS | 0 | Inadequate | |
| Schachtii | H. betae–H. daverti | 28S | 1 | Inadequate | |
| Schachtii | H. betae–H. schachtii | 28S | 2 | Inadequate | |
| Schachtii | H. betae–H. trifolii | 28S | 1 | Inadequate | |
| Schachtii | H. daverti–H. schachtii | 28S | 3 | Weak | |
| Schachtii | H. daverti–H. trifolii | 28S | 1 | Inadequate | |
| Schachtii | H. schachtii–H. trifolii | 28S | 1 | Inadequate |

Figure 1
Geographic origin of 18S, ITS, 28S, and coxI gene sequences analyzed in the present study. The size of the circle reflects the total number of sequences of all markers centered on the country of origin.

Figure 2
Neighbor-joining tree based on coxI sequences of Heterodera avenae from the NCBI database GenBank showing a clade of mislabeled sequences.
Table 3
Intraspecific differences observed in the 18S, ITS, 28S, and coxl gene regions for those species of Heterodera for which more than one sequence of one or more of the respective gene regions are available, excluding problematic sequences.
| 18S | ITS | 28S | cox1 | |||||
|---|---|---|---|---|---|---|---|---|
| Species | n | bp (%) | n | bp (%) | n | bp (%) | n | bp (%) |
| Heterodera arenaria | 4 | 0–1; 0.50 (0–0.29; 0.15) | 6 | 0–2; 2 (0–0.48; 0.48) | ||||
| Heterodera aucklandica | 3 | 0(0) | 5 | 0–1; 0 (0–0.24; 0) | ||||
| Heterodera australis | 18 | 0–9; 3 (0–0.93; 0.31) | 5 | 0(0) | ||||
| Heterodera avenae | 17 | 0–11; 1 (0–1.09; 0.06) | 542 | 0–80; 8 (0–11.89; 0.89) | 76 | 0–5; 0 (0–0.67; 0) | 91 | 0–34; 9 (0–8.17; 2.40) |
| Heterodera betae | 2 | 1 (0.06) | 7 | 0–12; 3 (0–1.29; 0.37) | 2 | 0(0) | 2 | 0(0) |
| Heterodera bifenestra | 3 | 2–10; 8 (0.20–1.02; 0.81) | ||||||
| Heterodera cajani | 12 | 0–12; 4 (0–1.18; 0.39) | 13 | 0–11; 2 (0–1.46; 0.26) | ||||
| Heterodera carotae | 13 | 0–13; 6 (0–1.35; 0.62) | 8 | 0–6; 3 (0–0.83; 0.41) | 17 | 0–20; 3 (0–4.77; 0.82) | ||
| Heterodera ciceri | 3 | 7–14; 11 (0.74–1.49; 1.18) | ||||||
| Heterodera cruciferae | 11 | 0–10; 3 (0–1.21; 0.36) | 2 | 52 (9.49) | 3 | 1–16; 16 (0.24–4.01; 3.77) | ||
| Heterodera cyperi | 2 | 0(0) | ||||||
| Heterodera daverti | 5 | 0–2; 1 (0–0.24; 0.11) | 3 | 0–2; 2 (0–0.51; 0.51) | ||||
| Heterodera dunensis | 3 | 0(0) | 3 | 0(0) | 3 | 0(0) | 8 | 0–1; 0 (0–0.28; 0) |
| Heterodera elachista | 3 | 3–4; 3 (0.25–0.36; 0.36) | 43 | 0–48; 11 (0–4.52; 1.05) | 18 | 0–4; 1 (0–0.54; 0.15) | 2 | 0(0) |
| Heterodera fici | 2 | 2 (0.34) | 8 | 0–3; 1 (0–0.31; 0.10) | 9 | 0–2; 1 (0–0.47; 0.24) | ||
| Heterodera filipjevi | 3 | 1–2; 1 (0.12–0.17; 0.17) | 256 | 0–43; 2 (0–4.46; 0.21) | 10 | 0–2; 1 (0–0.30; 0.13) | 75 | 0–44; 10 (0–10.58; 2.42) |
| Heterodera glycines | 6 | 0–1; 0 (0–0.16; 0) | 319 | 0–112; 3 (0–11.89; 0.32) | 56 | 0–43; 2 (0–5.79; 0.26) | 81 | 0–9; 0 (0–1.04; 0) |
| Heterodera goettingiana | 4 | 0–10; 5 (0–1.21; 0.59) | 15 | 0–81; 37 (0–11.34; 5.08) | 2 | 19 (2.92) | 3 | 0–17; 17 (0–4.84; 4.84) |
| Heterodera goldeni | 5 | 0–6; 2 (0–0.59; 0.23) | ||||||
| Heterodera guangdongensis | 3 | 0–3; 3 (0–0.26; 0.26) | 7 | 1–6; 4 (0.15–0.91; 0.58) | ||||
| Heterodera hordecalis | 2 | 5 (0.31) | 37 | 0–68; 7 (0–8.89; 0.84) | 2 | 9 (1.33) | 10 | 0–47; 14 (0–11.4; 3.37) |
| Heterodera humuli | 13 | 0–4; 1 (0–0.57; 0.17) | 19 | 0–9; 2 (0–5.92; 0.59) | ||||
| Heterodera koreana | 3 | 1–22; 2 (0.17–1.27; 0.34) | 29 | 0–28; 7 (0–3.08; 0.73) | 48 | 0–21; 2 (0–2.88; 0.27) | 44 | 0–18; 5 (0–4.57; 1.27) |
| Heterodera latipons | 154 | 0–122; 13 (0–13.32; 1.46) | 2 | 1 (0.15) | 16 | 0–50; 27 (0–12.05; 6.52) | ||
| Heterodera litoralis | 4 | 0(0) | ||||||
| Heterodera mani | 6 | 1–9; 3 (0.10–0.93; 0.31) | 6 | 0–1; 1 (0–0.27; 0.24) | ||||
| Heterodera medicaginis | 14 | 0–9; 3 (0–0.95; 0.42) | 14 | 0–2; 0 (0–0.49; 0) | ||||
| Heterodera mediterranea | 2 | 1 (0.17) | 6 | 1–9; 7 (0.11–0.95; 0.74) | 2 | 3 (0.38) | ||
| Heterodera mothi | 2 | 3 (0.30) | 2 | 2 (0.44) | ||||
| Heterodera orientalis | 5 | 6–18; 12 (0.66–1.94; 1.28) | ||||||
| Heterodera pratensis | 13 | 0–10; 6 (0–1.08; 0.62) | 20 | 0–8; 2 (0–1.93; 0.53) | ||||
| Heterodera ripae | 5 | 0–4; 2 (0–0.46; 0.22) | 19 | 0–11; 1 (0–2.59; 0.24) | ||||
| Heterodera sacchari | 3 | 0–1; 1 (0–0.09; 0.09) | ||||||
| Heterodera salixophila | 5 | 0–20; 15 (0–2.10; 1.58) | 2 | 1 (0.15) | 16 | 0–18; 2 (0–4.57; 0.51) | ||
| Heterodera schachtii | 9 | 1–9; 4 (0.06–0.99; 0.31) | 64 | 0–31; 12 (0–3.04; 1.29) | 20 | 0–4; 1 (0–0.58; 0.13) | 38 | 0–7; 1 (0–1.88; 0.12) |
| Heterodera sojae | 5 | 0–12; 4.5 (0–1.36; 1.03) | 10 | 0–8; 4 (0–1.06; 0.53) | 12 | 0–6; 1 (0–1.59; 0.24) | ||
| Heterodera sturhani | 7 | 0–3; 0 (0–0.72; 0) | ||||||
| Heterodera trifolii | 5 | 0–8; 3 (0–0.47; 0.29) | 23 | 0–14; 3 (0–1.48; 0.43) | 19 | 0–1; 0 (0–0.15; 0) | 26 | 0–5; 0 (0–1.24; 0) |
| Heterodera ustinovi | 5 | 1–8; 3.5 (0.10–0.88; 0.36) | 5 | 0–1; 1 (0–0.24; 0.24) | ||||
| Heterodera zeae | 16 | 0–39; 14 (0–5.65; 2.19) | 4 | 0–2; 1 (0–0.31; 0.15) | ||||
[i] Intraspecific differences are presented in the format “base pair difference range; median of base pair difference range (percent difference range; median of percent difference range).” Medians indicated only in cases of three or more comparisons; n = the number of sequences compared.

Figure 3
Neighbor joining (A) and maximum likelihood (B) trees derived from analyses of ITS gene sequences for the Schachtii species group showing multiple instances where species cannot be differentiated.
Table S1
Total number of sequences of species of Heterodera per country available on the NCBI database GenBank, up to 10 August 2021.
| ITS | 18S | 28S | coxI | Total | |
|---|---|---|---|---|---|
| Afghanistan | 1 | 1 | |||
| Algeria | 62 | 7 | 8 | 77 | |
| Australia | 11 | 1 | 6 | 18 | |
| Azerbaijan | 36 | 1 | 37 | ||
| Belgium | 21 | 1 | 2 | 14 | 38 |
| Canada | 19 | 1 | 7 | 5 | 32 |
| Chile | 1 | 1 | |||
| China | 761 | 16 | 131 | 21 | 937 |
| Costa Rica | 1 | 1 | 1 | 3 | |
| Cyprus | 21 | 21 | |||
| Czech Republic | 7 | 7 | 7 | 21 | |
| Egypt | 7 | 3 | 10 | ||
| Estonia | 2 | 2 | |||
| France | 11 | 9 | 20 | ||
| Georgia | 1 | 1 | 2 | ||
| Germany | 35 | 1 | 28 | 64 | |
| Ghana | 2 | 2 | |||
| Greece | 6 | 1 | 1 | 7 | 15 |
| Guatemala | 2 | 2 | |||
| India | 26 | 15 | 1 | 42 | |
| Iran | 41 | 16 | 51 | 108 | |
| Iraq | 3 | 8 | 11 | ||
| Ireland | 1 | 2 | 3 | ||
| Israel | 7 | 7 | |||
| Italy | 19 | 6 | 8 | 12 | 45 |
| Ivory Coast | 1 | 1 | |||
| Japan | 47 | 64 | 50 | 161 | |
| Jordan | 2 | 3 | 5 | ||
| Kazakhstan | 53 | 53 | |||
| Kyrgyzstan | 32 | 1 | 33 | ||
| Madagascar | 2 | 2 | |||
| Mexico | 3 | 3 | 6 | ||
| Morocco | 34 | 5 | 39 | ||
| Myanmar | 2 | 2 | |||
| Netherlands | 16 | 2 | 3 | 7 | 28 |
| New Zealand | 2 | 2 | 7 | 11 | |
| Norway | 4 | 4 | |||
| Pakistan | 2 | 1 | 3 | ||
| Poland | 1 | 2 | 3 | ||
| Portugal | 1 | 1 | 2 | ||
| Qatar | 1 | 1 | |||
| Russia | 12 | 23 | 35 | ||
| Saudi Arabia | 23 | 1 | 1 | 25 | |
| Serbia | 7 | 3 | 10 | ||
| Slovak Republic | 3 | 4 | 7 | ||
| South Africa | 3 | 1 | 1 | 5 | |
| South Korea | 30 | 38 | 36 | 104 | |
| Spain | 10 | 3 | 3 | 13 | 29 |
| Sweden | 4 | 2 | 2 | 8 | |
| Switzerland | 3 | 3 | |||
| Syria | 15 | 1 | 21 | 37 | |
| Tajikistan | 1 | 3 | 4 | ||
| Tunisia | 4 | 1 | 2 | 6 | 13 |
| Turkey | 219 | 53 | 272 | ||
| Ukraine | 3 | 1 | 7 | 11 | |
| United Kingdom | 11 | 1 | 9 | 21 | |
| USA | 71 | 7 | 18 | 158 | 254 |
| Vietnam | 2 | 1 | 3 | ||
| Unknown | 2 | 21 | 1 | 1 | 25 |
| Total | 1724 | 77 | 346 | 592 | 2739 |
Table S2
Species of Heterodera, and associated number of sequences, recorded from 59 countries based on data from the NCBI database GenBank, up to 10 August 2021.
| Species | Country | 18S | ITS | 28S | coxI |
|---|---|---|---|---|---|
| Heterodera arenaria | Italy | 1 | 3 | ||
| Netherlands | 2 | ||||
| United Kingdom | 3 | 1 | |||
| Heterodera aucklandica | Belgium | 1 | |||
| New Zealand | 1 | 1 | 3 | ||
| United Kingdom | 1 | 2 | |||
| Heterodera australis | Australia | 7 | 5 | ||
| China | 11 | ||||
| Heterodera avenae | Algeria | 4 | 6 | ||
| Australia | 1 | ||||
| Azerbaijan | 9 | ||||
| China | 4 | 359 | 57 | 4 | |
| Czech Republic | 7 | 7 | 7 | ||
| Egypt | 5 | 3 | |||
| France | 9 | 7 | |||
| Germany | 3 | 4 | |||
| India | 4 | 1 | |||
| Iran | 2 | 1 | 4 | ||
| Iraq | 3 | ||||
| Israel | 3 | 4 | |||
| Morocco | 27 | 5 | |||
| Norway | 4 | ||||
| Pakistan | 1 | ||||
| Saudi Arabia | 23 | 1 | 1 | ||
| Serbia | 1 | 1 | |||
| South Korea | 1 | 2 | 1 | ||
| Spain | 3 | 3 | |||
| Switzerland | 1 | ||||
| Syria | 3 | 6 | |||
| Tunisia | 1 | 6 | |||
| Turkey | 39 | 26 | |||
| United Kingdom | 1 | ||||
| USA | 1 | 10 | 3 | 14 | |
| Unknown | 4 | ||||
| Heterodera betae | Belgium | 1 | |||
| Germany | 3 | ||||
| Netherlands | 3 | 2 | 2 | ||
| Unknown | 2 | ||||
| Heterodera bifenestra | Belgium | 2 | |||
| Sweden | 1 | 1 | |||
| Heterodera cajani | India | 12 | 13 | ||
| Heterodera cardiolata | Pakistan | 1 | 1 | ||
| United Kingdom | 1 | ||||
| Heterodera carotae | Belgium | 1 | |||
| Algeria | 2 | ||||
| Canada | 5 | 6 | 5 | ||
| France | 1 | 2 | |||
| Italy | 3 | 1 | 5 | ||
| Mexico | 2 | 1 | |||
| South Africa | 1 | 1 | 1 | ||
| Switzerland | 1 | ||||
| Heterodera ciceri | Syria | 3 | 1 | ||
| Heterodera circeae | Germany | 1 | |||
| Heterodera cruciferae | Algeria | 5 | |||
| Iran | 1 | 1 | |||
| Italy | 1 | 1 | |||
| Netherlands | 1 | ||||
| Russia | 1 | 1 | |||
| Serbia | 1 | ||||
| USA | 1 | 2 | |||
| Heterodera cyperi | Spain | 1 | |||
| USA | 1 | 1 | |||
| Heterodera daverti | Germany | 3 | 2 | ||
| Italy | 1 | 1 | 1 | ||
| Netherlands | 1 | 1 | |||
| Heterodera dunensis | Spain | 3 | 3 | 3 | 8 |
| Heterodera elachista | China | 1 | 27 | 10 | |
| Iran | 2 | 1 | 1 | ||
| Italy | 2 | 4 | 4 | 2 | |
| Japan | 11 | 3 | |||
| Heterodera fici | Canada | 2 | |||
| Georgia | 1 | 1 | |||
| Greece | 4 | ||||
| Iran | 1 | 3 | |||
| Italy | 2 | 1 | |||
| Qatar | 1 | ||||
| USA | 2 | 1 | 1 | ||
| Heterodera filipjevi | Algeria | 2 | |||
| Azerbaijan | 20 | ||||
| China | 58 | 7 | 2 | ||
| Germany | 2 | 1 | |||
| Greece | 1 | ||||
| India | 1 | ||||
| Iran | 5 | 19 | |||
| Italy | 1 | ||||
| Kazakhstan | 25 | ||||
| Kyrgyzstan | 31 | ||||
| Russia | 3 | 7 | |||
| Serbia | 1 | 2 | |||
| Slovakia | 1 | ||||
| Spain | 1 | 2 | |||
| Sweden | 1 | 1 | |||
| Syria | 6 | ||||
| Tajikistan | 1 | 3 | |||
| Turkey | 80 | 20 | |||
| Ukraine | 3 | ||||
| United Kingdom | 1 | 3 | |||
| USA | 3 | 1 | 5 | ||
| Unknown | 2 | ||||
| Heterodera glycines | Canada | 1 | 12 | 1 | |
| Chile | 1 | ||||
| China | 5 | 275 | 29 | ||
| Iran | 2 | 1 | |||
| Japan | 6 | 10 | |||
| South Korea USA | 1 | 4 21 | 7 7 | 81 | |
| Unknown | 1 | 1 | |||
| Heterodera goettingiana | China | 1 | 10 | 1 | 3 |
| Germany | 2 | 1 | |||
| Iran | 1 | ||||
| Ireland | 1 | ||||
| Italy | 2 | ||||
| Netherlands | 4 | ||||
| USA | 1 | ||||
| Heterodera goldeni | Egypt | 2 | |||
| Iran | 2 | ||||
| Israel | 1 | ||||
| Heterodera graminis | India | 1 | |||
| Heterodera graminophila | USA | 1 | |||
| Heterodera guangdongensis | China | 3 | 3 | ||
| Myanmar | 2 | ||||
| Vietnam | 2 | 1 | |||
| Heterodera hainanensis | China | 1 | 1 | ||
| Heterodera hordecalis | Algeria | 2 | 21 | 2 | |
| Estonia | 1 | ||||
| Germany | 1 | 2 | |||
| Iran | 4 | 3 | |||
| Israel | 3 | 2 | |||
| Italy | 2 | ||||
| Netherlands | 1 | 2 | |||
| Slovakia | 1 | 2 | |||
| Sweden | 1 | 1 | |||
| Tunisia | 1 | ||||
| United Kingdom | 1 | ||||
| Heterodera humuli | Belgium | 1 | |||
| Germany | 1 | 2 | |||
| Iran | 3 | 1 | 8 | ||
| Kyrgyzstan | 1 | 1 | |||
| Portugal | 1 | ||||
| Russia | 1 | 2 | |||
| USA | 6 | 5 | |||
| Heterodera koreana | China | 1 | 2 | 2 | |
| Iran | 1 | 1 | |||
| Japan | 20 | 41 | 41 | ||
| South Korea | 3 | 3 | 3 | ||
| USA | 2 | 3 | 1 | ||
| Heterodera latipons | Cyprus | 21 | |||
| Iran | 3 | 3 | |||
| Israel | 2 | ||||
| Jordan | 2 | 3 | |||
| Kazakhstan | 28 | ||||
| Morocco | 3 | ||||
| Russia | 1 | 1 | |||
| Syria | 9 | 1 | 7 | ||
| Turkey | 87 | 3 | |||
| Heterodera litoralis | New Zealand | 1 | 1 | 4 | |
| Heterodera mani | Germany | 4 | 1 | ||
| United Kingdom | 1 | ||||
| USA | 1 | 4 | |||
| Unknown | 1 | ||||
| Heterodera medicaginis | Russia | 1 | |||
| USA | 13 | 1 | 14 | ||
| Heterodera mediterranea | Italy | 2 | 2 | ||
| Spain | 2 | ||||
| Tunisia | 1 | 2 | 2 | ||
| Heterodera microulae | China | 1 | 1 | 1 | |
| Heterodera mothi | Azerbaijan | 1 | 1 | ||
| Iran | 1 | 1 | |||
| Heterodera orientalis | Guatemala | 2 | |||
| Russia | 1 | ||||
| USA | 2 | 1 | |||
| Heterodera oryzae | South Korea | 1 | 1 | 1 | |
| Heterodera oryzicola | India | 1 | 1 | ||
| Heterodera persica | Iran | 1 | |||
| Heterodera pratensis | Belgium | 1 | |||
| Germany | 8 | 8 | |||
| Iran | 5 | ||||
| Netherlands | 1 | 1 | |||
| Russia | 1 | 2 | |||
| South Korea | 2 | 1 | 2 | ||
| USA | 1 | 1 | |||
| Heterodera ripae | Belgium Germany | 1 | 5 4 | ||
| Greece | 1 | 2 | |||
| Russia Serbia | 1 1 | 4 | |||
| Slovakia | 2 | ||||
| Sweden | 1 | ||||
| United Kingdom | 2 | ||||
| Heterodera sacchari | Ghana | 2 | |||
| Ivory Coast | 1 | ||||
| Heterodera salixophila | Belgium | 1 | 1 | 3 | |
| Germany | 4 | ||||
| Iran | 1 | 1 | |||
| Russia | 5 | ||||
| Ukraine | 3 | 4 | |||
| Heterodera schachtii | Algeria | 1 | |||
| Australia | 4 | ||||
| Belgium | 1 | 15 | |||
| China | 1 | 1 | 8 | 3 | |
| France | 1 | 3 | |||
| Germany | 4 | ||||
| Iran | 2 | 1 | |||
| Ireland | 2 | ||||
| Japan | 1 | 1 | 1 | ||
| Mexico | 1 | 2 | |||
| Morocco | 3 | ||||
| Netherlands | 8 | 1 | 1 | ||
| Poland | 1 | 1 | |||
| Serbia | 3 | ||||
| South Africa | 1 | ||||
| South Korea | 3 | 4 | 7 | ||
| Turkey | 12 | ||||
| USA | 2 | 3 | 22 | ||
| Unknown | 6 | 1 | |||
| Heterodera scutellariae | Germany | 1 | |||
| Heterodera sinensis | China | 1 | |||
| Heterodera skohensis | India | 1 | |||
| Heterodera sojae | China | 1 | 1 | ||
| Japan | 1 | 1 | |||
| South Korea | 3 | 8 | 12 | ||
| Heterodera sorghi | India | 1 | |||
| Heterodera sturhani | China | 7 | |||
| Heterodera trifolii | Costa Rica | 1 | 1 | 1 | |
| Estonia | 1 | ||||
| Iran | 1 | 1 | 2 | ||
| Italy | 1 | ||||
| Japan | 8 | 8 | 8 | ||
| South Korea | 8 | 8 | 9 | ||
| United Kingdom | 1 | 2 | 1 | ||
| USA | 2 | 5 | |||
| Unknown | 4 | ||||
| Heterodera turcomanica | Iran | 1 | 1 | ||
| Heterodera urticae | Belgium | 1 | 1 | 3 | |
| Heterodera ustinovi | Belgium | 1 | |||
| Germany | 1 | ||||
| Slovakia | 1 | ||||
| United Kingdom | 1 | ||||
| USA | 1 | 2 | |||
| Heterodera vallicola | Russia | 1 | 1 | ||
| Heterodera zeae | Afghanistan | 1 | |||
| China | 5 | ||||
| Greece | 1 | 5 | 1 | ||
| India | 4 | ||||
| Portugal | 1 | ||||
| USA | 2 | 1 |