Table 1.
Geocenamus brevidens female morphometrics.
| Characters | Present study | Allen (1955) a | Siddiqi (1961) | Alvani et al. (2017) | Tzortzakakis et al. (2018) | |
|---|---|---|---|---|---|---|
| Origin | Canada | USA | India | Iran | Greece | |
| Host | Potato field | Grass | Grass | Cauliflower, cabbage, mint, potato, pea | Jujube, saffron, barberry | Cultivated olives |
| n | 15 | 10 | 11 | 15 | 8 | 3 |
| Body length | 667.8 ± 64.2 (590.9-811.0) | 687.1 ± 53.3 (604.0-752.0) | 540-690 | 550-850 | 650 (600-718.5) | 564 (490-698) |
| a | 35.4 ± 2.8 (25.5-38.5) | 28.2 ± 1.8 (24.8-31.1) | 23-27 | 22-29 | 26.5 (23.9-29) | 26.3 (22.9-30.3) |
| b | 5.0 ± 0.4 (4.6-5.9) | 4.9 ± 0.3 (4.5-5.6) | 4.2-5.2 | 4.6-6.0 | 5.0 (4.5-5.2) | 4.4 (4.0-5.1) |
| c | 12.7 ± 0.6 (11.7-13.6) | 14.3 ± 1.1 (12.7-15.9) | 11-13 | 12-17 | 13.7 (13.7-15.7) | 14.3 (12.9-17.0) |
| c' | 4.2 ± 0.3 (3.8-4.9) | 3.2 ± 0.2 (3.0-3.5) | 2.5-4.4 | – | 2.7 (2.4-3.2) | 3.0 (2.9-3.2) |
| V | 56.6 ± 1.6 (53.8-61.0) | 56.2 ± 1.2 (54.5-58.1) | 52-58 | 54-61 | 55.6 (54.5-57) | 54.7 (52-58) |
| MB | 48.6 ± 2.0 (46.2-54.3) | 51.1 ± 1.5 (49.6-53.7) | 42-47 | – | (41.1-46.5) | – |
| Lip height | 3.6 ± 0.3 (3.1-4.0) | 3.6 ± 0.3 (3.0-4.0) | – | – | – | – |
| Lip width | 7.2 ± 0.5 (6.1-7.7) | 8.0 ± 0.2 (7.5-8.3) | 6.5–8.5 | – | – | – |
| Stylet length | 16.2 ± 0.7 (15.0-17.5) | 17.0 ± 0.5 (16.0-17.6) | 14-16 | 13-15 | 16.3 (16-16.5) | 14.3 (13-16) |
| Median bulb length | 14.2 ± 1.3 (12.3-16.4) | 15.1 ± 0.6 (14.3-15.8) | 13-16 | – | – | – |
| Median bulb width | 9.9 ± 0.9 (8.5-11.4) | 10.3 ± 0.9 (9.2-11.7) | 9-12 | – | – | – |
| Anterior end to excretory pore | 103.3 ± 6.7 (90.3-112.5) | 106.1 ± 1.9 (103.0-109.0) | – | – | – | – |
| Pharynx length | 133.5 ± 6.9 (120.3-144.0) | 139.6 ± 5.8 (130.0-148.0) | 110-118 | – | 129.9 (118.5-141) | 127.7 (123-136) |
| Maxim body width | 18.6 ± 1.7 (15.5-21.2) | 24.4 ± 1.2 (22.3-26.2) | 19-24 | – | 100.2 (90.5-110) | 21.3 (19.0-23.0) |
| Vulva body length | 19.4 ± 1.9 (16.4-22.3) | 23.5 ± 1.3 (21.0-25.3) | – | – | 24.7 (21.5-30) | – |
| Anal body width | 12.6 ± 1.0 (10.3-14.2) | 14.9 ± 0.5 (13.7-15.3) | – | – | 17.1 (15.5-19) | 13.0 (12.0-14.0) |
| Tail length | 52.5 ± 3.7 (46.0-59.8) | 48.2 ± 2.2 (46.0-52.0) | 34-58 | – | 47.5 (42-53) | 39.3 (38-41.0) |
| Phasmid position | Posterior to middle of tail | Posterior to middle of tail | Posterior to middle of tail | – | – | – |

Figure 1:
Light photomicrographs of Geocenamus brevidens. (A) Entire female, (B) Esophageal region, (C) Lip region, (D) Posterior esophageal region, (E) Deirids, (F) Posterior region with complete reproductive system, (G) Lateral lines, (H-J) Vulval region, (K-O) Female tails. Scale bars: (A) 50 μm; (B-D, E, H-O) 20 μm, (F) 50 μm, (G) 5 μm. Arrows point to (a) anus, (d) deirids, (exp) excretory pore, (ph) phamsid, and (v) vulva.
Table 2.
Quinisulcius capitatus female morphometrics.
| Characters | Present study | Allen (1955)a | Hopper (1959) | Loof (1959) | Siddiqi (1961) | Knobloch and Laughlin (1973) | Maqbool (1982) | Vovlas (1983) | Mekete et al. (2008) |
|---|---|---|---|---|---|---|---|---|---|
| Origin | Canada | USA | USA | Italy | India | Mexico | Pakistan | Italy | Ethiopia |
| Host | Grass | Pear | okra | Date palm | Grass Lily | Grass, pasture grass, wild poppy, barrel cactus, cotton | Potato | Maize | Coffee |
| n | 20 | 13 | 5 | 14 | 8 | – | 5 | 25 | 10 |
| Body length | 810.3 ± 44.6 (744.0-911.0) | 630-850 | 650-710 | 631-830 | 700-810 | 520-630 | 610-730 | 641-730 | 630-790 |
| a | 41.4 ± 1.8 (38.6-43.7) | 30-38 | 30.2-36.2 | 31-37.6 | 30-35 | 29-33 | 30-33 | 31-35 | 30.9-38.6 |
| b | 5.5 ± 0.3 (5.0-6.3) | 5.0-5.8 | 4.6-4.9 | 4.7-5.5 | 4.5-5.5 | 4.4-5.3 | 6.8-7.4 | 4.1-5.3 | – |
| c | 22.4 ± 1.1 (19.9-23.8) | 12-17 | 14.5-16.1 | 14.3-17.4 | 15-17 | 15-17 | 3.8-4.9 | 13-16 | 15.3-17.6 |
| c′ | 2.6 ± 0.2 (2.2-3.2) | – | – | – | – | – | – | 2.8-3.2 | |
| V | 57.4 ± 1.5 (53.4-59.8) | 51-58 | 55.2-56.9 | 53.5-58 | 54-56.5 | 52-59 | 53-56 | 54-57 | 54.7-63.6 |
| MB | 52.3 ± 1.7 (47.4-56.1) | – | – | – | – | – | – | 57-60 | – |
| Lip height | 4.0 ± 0.2 (3.7-4.4) | – | – | – | – | – | – | – | – |
| Lip width | 7.6 ± 0.4 (6.9-8.3) | – | – | – | – | – | – | – | – |
| Stylet length | 18.3 ± 1.0 (15.5-20.4) | 16-18 | 16.5-17.3 | 16-18 | 16-17 | 16-17 | 18.4-19.5 | 15-17 | 15-18 |
| Median bulb length | 14.0 ± 1.6 (11.3-16.9) | – | – | – | – | – | – | – | – |
| Median bulb width | 10.4 ± 1.4 (8.4-14.2) | – | – | – | – | – | – | – | – |
| Anterior end to excretory pore | 128.6 ± 5.3 (121.0-139.0) | – | – | – | – | – | – | 119-128 | 121 |
| Pharynx length | 147.8 ± 5.8 (140.2-159.0) | – | – | – | – | – | – | – | – |
| Maxim body width | 19.0 ± 1.5 (16.9-21.3) | – | – | – | – | – | – | – | – |
| Vulva body length | 18.4 ± 1.3 (15.7-20.8) | – | – | – | – | – | – | – | – |
| Anal body width | 13.8 ± 1.1 (11.2-15.2) | – | – | – | – | – | – | – | – |
| Tail length | 35.8 ± 2.4 (31.3-40.4) | – | – | – | – | – | – | 40-48 | – |
| Phasmid position | Middle of tail | Middle of tail | Middle of tail | Anterior to middle of tail | Anterior to middle of tail | Anterior to middle of tail | Anterior to middle of tail | Anterior to middle of tail | – |
Table 3.
Worldwide distribution and host plant association of Geocenamus brevidens and Quinisulcius capitatus.
| No. | Country | Host associations | References |
|---|---|---|---|
| G. brevidens | |||
| Americas | |||
| 1 | USA | WheatPotato | Mayol (1981), Smiley et al. (2006) Olthof et al. (1982) |
| Wheat grass | Griffin and Asay (1996) | ||
| Pasture filed, vegetable and horticultural crops | Hafez et al. (2010) | ||
| Asia | |||
| 2 | Iran | Wheat | Ghaderi et al. (2014) |
| Jujube, saffron, barberry | Alvani et al. (2017) | ||
| Africa | |||
| 3 | South Africa | Wheat | Jordaan et al. (1992) |
| 4 | Egypt | Soybean | Salem et al. (1994) |
| Europe | |||
| 5 | Spain | Cereals, sunflower, wheat | Tobar et al. (1995a, 1995b) |
| Chickpea | Castillo et al. (1996) | ||
| Grasslands | Talavera and Navas (2002) | ||
| Olives | Palomares-Rius et al. (2015) | ||
| 6 | Slovakia | Hop gardens | Lišková and Rencˇo (2007) |
| 7 | Czech Republic | Hop gardens | Cˇermák et al. (2011) |
| 8 | Turkey | Cultivated plants | Kasapog˘lu et al. (2014) |
| Cotton, barley, melons, tobacco, Watermelons, wheat, lentils | Kasapog˘lu Uludamar et al. (2018) | ||
| 9 | Greece | Olives | Tzortzakakis et al. (2018) |
| 10 | Poland | Jerusalem artichoke | Zapałowska and Skwiercz (2018) |
| Oceania | |||
| 11 | Australia | Cereal fields | Meagher (1970) |
| Wheat | Thompson et al. (2008, 2010) | ||
| Millet, soybean, grasses | Owen et al. (2014) | ||
| Q. capitatus | |||
| Americas | |||
| 1 | Ecuador | Avocado, barley, bean, carrot, cucumber, lettuce, pea, onion, tomato, soybean, sugarcane | Bridge (1976) |
| 2 | Argentina | Corn, sunflower | Doucet (1986) |
| 3 | USA | Okra | Hopper (1959) |
| Wild poppy, barrel cactus, cotton | Knobloch and Laughlin (1973) | ||
| Tobacco | Ponchillia (1975) | ||
| Red clover, Kentucky bluegrass | Malek (1980) | ||
| Sorghum | Cuarezma-Teran and Trevathan (1985) | ||
| Potato | Hafez et al. (2010) | ||
| Switchgrass | Cassida et al. (2005) | ||
| Asia | |||
| 4 | Pakistan | Potato | Maqbool (1982) |
| Maqbool and Hashmi (1986) | |||
| 5 | India | Lily | Siddiqi (1961) |
| Potato | Krishna Prasad (2008) | ||
| 6 | Iran | Cultivated crops | Kheiri et al. (2002) |
| Africa | |||
| 7 | Ethiopia | Coffee | Mekete et al. (2008) |
| 8 | South Africa | Potato | Marais et al. (2015) |
| Soybean | Mbatyoti et al. (2020) | ||
| Europe | |||
| 9 | Italy | Lily, maize | Loof (1959), Vovlas (1983) |
| 10 | Bulgaria | Apple | Braasch (1978) |
| 11 | Cypress | Grapes | Antoniou (1981) |
| 12 | Turkey | Tomato, tobacco | Kasapog˘lu Uludamar et al. (2018) |
| Oceania | |||
| 13 | New Zealand | Tomato, tobacco, squash | Knight et al. (1997) |

Figure 2:
Light photomicrographs Quinisulcius capitatus. (A) Entire female, (B) Esophageal region, (C) Lip region, (D) Basal esophageal bulb, (E) Posterior region with complete reproductive system, (F) Posterior region with eggs, (G-J) Vulval region, (K) Lateral lines, (L-P) Female tails. Scale bars: (A) 100 μm, (B, C, F-J; L-P) 20 μm, (E) 50 μm, (D, K) 5 μm. Arrows point to (a) anus, (d) deirids, (exp) excretory pore, (ph) phasmid, and (v) vulva.

Figure 3:
Phylogenetic relationships within selected genera of subfamily Telotylenchinae and subfamily Merliniinae as inferred from Bayesian analysis using the 18S of the rRNA gene sequence dataset with the GTR + I + G model (lnL = 1,910.5101; AIC = 4,017.0201; freq A = 0.2500; freq C = 0.2500; freq G = 0.2500; freq T = 0.2500; R(a) = 1.0000; R(b) = 3.9248; R(c) = 1.0000; R(d) = 1.0000; R(e) = 4.6930; R(f) = 1.0000). Posterior probability of more than 70% is given for appropriate clades. Newly obtained sequences are indicated in bold. *** need to be revised by integrative taxonomy.

Figure 4:
Phylogenetic relationships within selected genera of subfamily Telotylenchinae and subfamily Merliniinae as inferred from Bayesian analysis using the 28S of the rRNA gene sequence dataset with the GTR + I + G model (lnL = 6,015.1425; AIC = 12,526.2851; freq A = 0.1987; freq C = 0.2072; freq G = 0.3206; freq T = 0.2736; R(a) = 0.4322; R(b) = 2.5823; R(c) = 1.2662; R(d) = 0.2497; R(e) = 5.4146; R(f) = 1.0000). Posterior probability of more than 70% is given for appropriate clades. Newly obtained sequences are indicated in bold.

Figure 5:
Phylogenetic relationships within selected genera of subfamily Telotylenchinae and subfamily Merliniinae as inferred from Bayesian analysis using the ITS of the rRNA gene sequence dataset with the GTR + I + G model (lnL = 10,413.7629; AIC = 21119.5049; freq A = 0.1932; freq C = 0.2202; freq G = 0.2725; freq T = 0.3141; R(a) = 0.8338; R(b) = 3.3701; R(c) = 1.6297; R(d) = 0.6490; R(e) = 3.3701; R(f) = 1.0000). Posterior probability of more than 70% is given for appropriate clades. Newly obtained sequences are indicated in bold. **previously unidentified.