
Figure 1:
Distribution map of soil sample sites and entomopathogenic nematodes in Syrian coastal regions. A: Soil sample sites. B: Sites positive for entomopathogenic nematodes. Those marked with green denote H. indica, yellow denotes H. bacteriophora, blue denotes H. pakistanense, and red denotes S. affine.
Table 1:
Locality, coordinates, altitude, sampling date, habitat, and soil characteristics of positive samples.
| Species | Site/Location | GenBank accession no. | Coordinates | Altitude | Sampling date | Habitats | Soil types | pH | Organic matter % |
|---|---|---|---|---|---|---|---|---|---|
| H. indica | Zeghreen / Latakia | OP221993 | 35°43′39″N 35°52′39″E | 40 m | Autumn | Olive field | Sandy | 7.3 | 3.39 |
| H. indica | Al-Hinadi / Latakia | OP221988 | 35°51′48″N 35°30′12″E | 36 m | Autumn | Citrus orchard | Silty loam | 7.5 | 1.4 |
| H. indica | Ein-Albida / Latakia | OP221987 | 35°39′26″N 35°35′18″E | 207 m | Autumn | Citrus orchard | Silty loam | 7.5 | 5.19 |
| H. indica | Al-Meherfha/Latakia | OP221992 | 35°36′12″N 35°51′20″E | 214 m | Autumn | Olive orchard | Sandy loam | 6.1 | 2.3 |
| H. indica | Al-Shamiea /Latakia | OP221989 | 35°38′22″N 35°48′30″E | 61 m | Autumn | Citrus orchard | Silty loam | 6.5 | 3.72 |
| H. indica | Fattiro /Latakia | OP221991 | 35°37′19″N 35°15′24″E | 138 m | Autumn | Olive orchard | Clay | 6.2 | 2 |
| H. indica | Kersana/Latakia | OP221983 | 35°37′18″N 35°49′40″E | 53 m | Autumn | Olive orchard | Sandy loam | 6.0 | 5.72 |
| H. indica | Al-Basa/Latakia | OP221984 | 35°30′14″N 35°52′15″E | 46 m | Autumn | Vegetables | Silty loam | 6.5 | 2.33 |
| H. indica | Al-Borjan /Latakia | OP221986 | 35°17′34″N 35°58′42″E | 47 m | Autumn | Vegetables | Sandy loam | 6.1 | 3.66 |
| H. indica | Al-Meghreet /Latakia | OP221990 | 35°63′17″N 35°49′53″E | 81 m | Spring | Citrus orchard | Silty loam | 6.4 | 2.9 |
| H. indica | Al-Hwiz/Latakia | OP221996 | 35°20′27″N 36°00′02″E | 108 m | Spring | Grassland | Sandy loam | 5.2 | 2.26 |
| H. indica | Hraeson / Tartus | OP221985 | 35°15′01″N 35°57′01″E | 15 m | Spring | Citrus orchard | Silty loam | 8.1 | 2.73 |
| H. indica | Fneiteq /Tartus | OP221994 | 35°06′51″N 36°07′17″E | 597 m | Autumn | Walnut orchard | Silty loam | 8.1 | 1.73 |
| H. indica | Karm-Alten/Tartus | OP221995 | 35°07′01″N 36°06′20″E | 580 m | Spring | Grassland | Sandy | 7.5 | 3.02 |
| H. bacteriophora | Al-Snobr /Latakia | OP222916 | 35°28′45″N 35°53′09″E | 26 m | Spring | Grassland | Clay loam | 5.8 | 6.2 |
| H. bacteriophora | Zeghreen /Latakia | OP222020 | 35°43′45″N 35°52′57″E | 46 m | Autumn | Citrus orchard | Clay loam | 6.7 | 4.72 |
| H. bacteriophora | Al-Shelfatea/Latakia | OP222019 | 35°32′54″N 35°53′53″E | 41 m | Autumn | Citrus orchard | Silty loam | 7.1 | 3.3 |
| H. bacteriophora | Al-Serskiea/Latakia | OP222018 | 35°42′33″N 35°55′36″E | 85 m | Autumn | Vegetables | Sandy loam | 6.7 | 5.39 |
| H. bacteriophora | Ras-Alein /Latakia | OP222015 | 35°17′38″N 35°57′53″E | 32 m | Spring | Tobacco field | Silty loam | 6.0 | 1.46 |
| H. bacteriophora | Al-Basa /Latakia | OP222017 | 35°29′34″N 35°50′43″E | 4 m | Autumn | Citrus orchard | Silty loam | 7.7 | 2.67 |
| H. bacteriophora | Dimsarkho /Latakia | OP222012 | 35°33′07″N 35°46′26″E | 15 m | Autumn | Tobacco field | Silty loam | 5.9 | 3.06 |
| H. bacteriophora | AL-Kawkae/Tartus | OP222014 | 35°09′24″N 36°06′47″E | 486 m | Spring | Grassland | Clay loam | 6.3 | 2.06 |
| H. bacteriophora | Bet-shofan /Tartus | OP222013 | 34°45′35″N 36°00′44″E | 70 m | Spring | Citrus orchard | Silty loam | 7.8 | 1.25 |
| H. pakistanense | Al-Zobar /Latakia | OP235514 | 35°37′57″N 36°00′12″E | 213 m | Autumn | Pear orchard | Sandy loam | 6.8 | 1.86 |
| H. pakistanense | Al-Bahloliea/Latakia | OP235513 | 35°38′09″N 35°57′40″E | 179 m | Summer | Vineyard | Sandy loam | 7.1 | 1.3 |
| S. affine | Amodeyha/Tartus | OM350119 | 35°06′10″N 36°07′57″E | 759 m | Spring | Walnut orchard | Silty loam | 6.1 | 6 |
| S. affine | Fneitiq/Tartus | OL437016 | 35°06′02″N 36°08′51″E | 604 m | Winter | Grass land | Silty | 6.4 | 4.1 |
Table 2:
Vegetation habitat and abiotic factors with their corresponding codes.
| Habitat | Code-Hab | Soil characteristics | Code | Altitude and Season | Code |
|---|---|---|---|---|---|
| Citrus orchard | Hab1 | Soil type |
| Altitude |
|
| Olive orchard | Hab2 |
|
| ||
| Grassland | Hab3 | ||||
| Vegetables | Hab4 | Season | |||
| Walnut orchard | Hab5 |
|
|
|
|
| Tobacco field | Hab6 | ||||
| Pear orchard | Hab7 | ||||
|
| ||||
| Vineyard | Hab8 |
Table 3:
Distribution of EPNs in Syrian coast region according to the vegetation habitat, altitude and sampling seasons variables.
| Habitat | Frequency | Abundance | No. of identified samples to species |
|---|---|---|---|
| Coniferous forest | 0 c | 0 b | 0 |
| Broadleaf forest | 0 c | 0 b | 0 |
| Citrus orchards | 27.27 a | 2.4 a | 9 |
| Olive orchards | 10.81b | 1.06 a | 4 |
| Walnut orchards | 8 b | 0.53 a | 2 |
| Pear orchards | 4.34 b,c | 0.26 a | 1 |
| vineyard | 4 bc | 0.26 a | 1 |
| Fruit orchards (e.g., cherry, apple) | 0 c | 0 b | 0 |
| Vegetable fields | 7.5 b | 0.8 a | 3 |
| Oil crop | 0 c | 0 b | 0 |
| Leguminous crop | 0 c | 0 b | 0 |
| Leafy vegetables | 0 c | 0 b | 0 |
| Tobacco fields | 8.33b | 0.53 a | 2 |
| Riverbanks | 0 c | 0 b | 0 |
| Greenhouses | 0 c | 0 b | 0 |
| Fodder crop | 0 c | 0 b | 0 |
| Grassland | 11.11b | 1.33 a | 5 |
| Medicinal and aromatic plants | 0 c | 0 b | 0 |
| Sea coast | 0 c | 0 b | 0 |
| Public parks | 0 c | 0 b | 0 |
| P = 0.01491 | P = 0.01892 | ||
| Altitude | |||
| 0–200 | 8.37a | 19.31 | 18 |
| 200–400 | 1.79b | 3.79 | 3 |
| 400–600 | 1.93b | 3.94 | 3 |
| 600–800 | 1.39b | 2.7 | 2 |
| >800 | 0c | 0 | 0 |
| P = 0.0138 | P = 0.1565 | ||
| Season | |||
| Autumn | 5.01 a | 10.37 | 16 |
| Spring | 2.4a,b | 6.33 | 9 |
| Summer | 1.4b | 1.96 | 1 |
| Winter | 1.78 b | 2.12 | 1 |
| P = 0.0474 | P = 0.08053 | ||

Figure 2:
Heterorhabditis indica male. A: Male entire body. B–H: Male tail with bursal papillae. B, C: Ventral view of bursa showing papillae (3–2) in the terminal group with some variation. E, F, G, H: Lateral and ventral view of bursa showing papillae (2–2) in the terminal group with some modifications (namely, being short and swollen). I: Lateral view of spicule and gubernaculum. J: Ventral view of spicule and gubernaculum. A = 705 μm. I, J: SL = 40 μm, GL = 19 μm.

Figure 3:
Heterorhabditis bacteriophora male. A: Male entire body. B: Male tail with bursal lateral view of spicules and gubernaculum. C: Lateral view of bursa showing the arrangement of papillae, arrow refers to curved outward fourth pair. D: Lateral view of bursa showing the terminal group of bursal papillae pairs (7, 8, and 9). E: Anterior portion; the left arrows refer to the ring position and excretory pore, while the right arrow refers to the end of the esophagus. A = 861 μm. B: SL = 42 μm, GL = 20 μm. E: NR = 72 μm, ES = 101 μm.


Figure 4:
Heterorhabditis pakistanense male. A: Male entire body. B: The anterior portion; the left arrow refers to the nerve ring position, while the right arrow refers to the excretory pore. C, D: Male tail with bursal papilae. B, C: Ventral view of bursa showing papillae (3–2) in the terminal group with some variation in tall. D: Lateral view of bursa; the arrow refers to curved outward seventh pair. E, F, G, H: Lateral and ventral view of bursa showing (2–2) papillae in the terminal group, with some modifications (namely, being short and swollen). I: Lateral view of spicule and gubernaculm. J: Ventral view of spicule and gubernaculum. A = 705 μm. I, J: SL = 40 μm, GL = 19 μm.
Table 4:
Morphometrics of infective juvenile of Syrian Heterorhabditis indica isolates. All measurements are in μm (except n, ratio, and percentage), and in the form: mean ± s.d. (range).
| Character | SY-BS | SY-E | SY-AS | SY-K | SY-BO | SY-FA | SY-SH | SY-HR | SY-KA | SY-Z | SY-MS | SY-ME | SY-HW | SY-AF | H. indica Type strain Poinar et al. (1992) |
|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
| n | 20 | 20 | 20 | 20 | 20 | 20 | 20 | 20 | 20 | 20 | 20 | 20 | 20 | 20 | 25 |
| L | 532±31 (503–589) | 551±25 (532–591) | 547±45 (523–598) | 559±37 (518–589) | 553±37 (510–595) | 561±41 (529–602) | 569±27 (520–596) | 578±46 (545–610) | 570±21 (531–604) | 589±69 (520–610) | 597±51 (532–614) | 602±62 (541–610) | 607±66 (539–615) | 604±59 (551–618) | 528 (479–573) |
| MBD | 20.5±0.7 (18–22) | 20±1.1 (19–23) | 19±0.8 (17–21) | 19.5±0.9 (18–22) | 20 ±1 (18–21) | 21±1.3 (17–24) | 21±1.2 (18–23) | 22±1.4 (18–24) | 21.5±1.5 (19–24) | 21±1.2 (18–25) | 22±1.4 (17–24) | 21±1.8 (18–26) | 22±1.2 (19–25) | 22±1 (18–25) | 20 (19–22) |
| EP | 97.2±6 (85–103) | 95±5.5 (83–101) | 94.5±5.3 (85–100) | 96.5±5 (87–102) | 94±4.9 (82–102) | 96±5.2 (83–102) | 101±6.1 (85–107) | 98.5±4.7 (87–103) | 101±4.5 (81–105) | 102±6.7 (89–107) | 102±6.4 (84–106) | 101.5±58 (89–110) | 103±6.5 (87–112) | 102±6.3 (89–109) | 98 (88–107) |
| NR | 81.4±3 (70–86) | 78.6±2.4 (74–87) | 83±1.9 (78–88) | 82.5±2.1 (73–86) | 82±2.3 (75–84) | 81±2.8 (70–85) | 83±2.7 (74–88) | 80±2.3 (75–86) | 82.5±2.2 (73–85) | 84±2.6 (74–88) | 83.5±2.9 (71–87) | 84±2.5 (75–88) | 81±2.7 (72–86) | 80.5±2.9 (75–84) | 82 (72–85) |
| ES | 118±4 (112–122) | 117±5.1 (104–123) | 118±6 (109–121) | 117.5±5 (110–120) | 115±5.6 (109–121) | 121±5.4 (110–123) | 121±5.8 (112–124) | 122±7.4 (105–125) | 118±6.1 (107–120) | 119.5±6 (107–122) | 120±6.1 (114–125) | 119±5.4 (108–121) | 118±5.7 (110–123) | 120±5.5 (109–122) | 117 (109–123) |
| ABW | 9.6±1.1 (8–11) | 7±0.7 (6.5–8) | 7.1±0.8 (6.5–7.8) | 7±0.7 (6.4–7.5) | 6.7±1 (6.2–7.5) | 7.2±0.8 (6.5–7.9) | 7.5±0.9 (6.7–7.9) | 7.2±0.4 (6.8–7.5) | 7±0.6 (6.4–7.6) | 7.8±0.9 (6.9–8) | 6.8±0.8 (6.4–7.8) | 6.7±0.9 (6.5–7) | 7.1±0.5 (6.7–7.4) | 7±0.5 (6.5–7.6) | - |
| T | 103±5.1 (88–105) | 94±2.3 (90–97) | 96±4.1 (92–106) | 97.5±2.3 (93–102) | 97±1.4 (92–99) | 98.5±2.9 (89–103) | 95±2.1 (90–101) | 102±5 (89–107) | 98±4.9 (88–105) | 101±4.1 (90–110) | 98.5±4.5 (93–104) | 102±4 (94–107) | 104±4.3 (93–105) | 101±4.9 (95–107) | 101 (93–109) |
| a | 25.9 | 27.5 | 28.7 | 28.6 | 27.6 | 26.7 | 27 | 26.2 | 26.5 | 28 | 27.1 | 27.3 | 27.5 | 27.4 | 26 (25–27) |
| b | 4.50 | 4.7 | 4.63 | 4.75 | 4.82 | 4.63 | 4.70 | 4.81 | 4.83 | 4.90 | 4.97 | 5 | 5.14 | 5 | 4.5 (4.3–4.8) |
| c | 5.1 | 5.86 | 5.69 | 5.73 | 5.70 | 5.69 | 5.98 | 5.66 | 5.81 | 5.83 | 6 | 5.90 | 5.83 | 5.98 | 5.3 (4.5–5.6) |
| D | 0.82 | 0.81 | 0.80 | 0.82 | 0.82 | 0.79 | 0.83 | 0.80 | 0.85 | 0.85 | 0.85 | 0.85 | 0.87 | 0.85 | 0.84 (0.79–0.90) |
| E | 0.94 | 1.01 | 0.98 | 0.98 | 0.96 | 0.97 | 1.06 | 0.96 | 1.03 | 1 | 1.03 | 0.99 | 0.99 | 1 | 0.90 (0.83–1.03) |
Table 5:
Morphometrics of male of the Syrian Heterorhabditis indica isolates. All measurements are in μm (except n, ratio, and percentage), and in the form mean ± S.D. (range).
| Character | SY-BS | SY-E | SY-AS | SY-K | SY-BO | SY-FA | SY-SH | SY-HR | SY-KA | SY-Z | SY-MS | SY-ME | SY-HW | SY-AF | H. indica Type strain Poinar et al. (1992) |
|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
| n | 20 | 20 | 20 | 20 | 20 | 20 | 20 | 20 | 20 | 20 | 20 | 20 | 20 | 20 | 12 |
| L | 685±89 (569–719) | 697±80 (598–731) | 689±75 (604–740) | 715±69 (591–730) | 703±81 (590–741) | 718±87 (610–773) | 726±89 (598–761) | 741±91 (589–767) | 735±85 (602–759) | 749±80 (584–773) | 738±90 (609–770) | 745±77 (614–780) | 754±94 (598–783) | 762±79 (590–790) | 721 (573–788) |
| MBD | 37±2.3 (35–41) | 38.5±2.4 (35–43) | 37±2.1 (33–40) | 41±1.5 (38–47) | 40.5±1.8 (37–43) | 41±1.3 (37–42) | 41.5±1 (35–44) | 42±1.2 (37–45) | 42±1.9 (35–44) | 43.5±1.6 (38–47) | 43±0.9 (37–44) | 44±1.4 (38–46) | 43.5±1.5 (36–45) | 43±1.9 (38–45) | 42 (35–46) |
| EP | 113±6.7 (105–125) | 114.5±5.9 (107–121) | 114±6.1 (107–127) | 119±8.1 (107–127) | 115.5±5.7 (109–125) | 119±5.9 (105–130) | 124±5.1 (109–131) | 126±5.5 (110–137) | 124.5±5.9 (105–134) | 126±5.8 (114–136) | 124.5±6.1 (107–133) | 126.5±5.4 (112–136) | 129±5.8 (110–139) | 132±6 (109–140) | 123 (109–138) |
| NR | 71±4.5 (69–76) | 72±5.8 (69–79) | 71.5±4.3 (67–75) | 74±4.7 (70–80) | 72±6.1 (68–83) | 73±5.7 (69–84) | 73±4.1 (70–80) | 78±5.5 (69–80) | 77.5±4.3 (70–81) | 78.5±4.9 (70–85) | 77±3.9 (73–83) | 78±4.5 (72–84) | 79±5.1 (74–86) | 78.5±5.2 (71–85) | 75 (72–85) |
| ES | 95±5.7 (94–104) | 97.5±5.1 (92–103) | 95.5±5.5 (93–105) | 98±5.9 (92–107) | 98.5±6.1 (94–107) | 99±6.3 (95–108) | 101±5.8 (95–107) | 103±5.9 (94–106) | 101.5±6 (96–109) | 104±5.8 (94–107) | 103±5.2 (96–105) | 103.5±4.9 (95–106) | 104.5±5.6 (94–109) | 106±6 (97–109) | 101 (93–109) |
| ABW | 20±1.2 (18–22) | 20±1.3 (17–21) | 20.5±1.2 (18–23) | 20±1.4 (17–23) | 21±0.9 (18–24) | 22±1.3 (19–24) | 22.5±1.4 (18–24) | 23±0.8 (19–24) | 22±1.4 (19–25) | 23±1.1 (17–25) | 21.5±0.9 (19–24) | 22.5±1.4 (18–25) | 23.5±1.2 (18–25) | 23±1.1 (18–25) | 23 (19–24) |
| T | 23±2.1 (21–26) | 25±1.9 (22–29) | 24.5±1.3 (23–30) | 26±1.7 (21–30) | 25±1.8 (22–30) | 26.5±2.3 (24–33) | 27.5±2.1 (24–31) | 29±2.6 (21–32) | 29±2 (24–31) | 30±2.5 (22–32) | 28±2.1 (24–31) | 29.5±1.9 (24–32) | 30±1.6 (24–31) | 30±1.5 (25–33) | 28 (24–32) |
| a | 18.51 | 17.9 | 18.61 | 17.43 | 17.35 | 17.51 | 17.49 | 17.64 | 17.5 | 17.21 | 16.9 | 16.93 | 17.33 | 17.7 | 17.16 |
| b | 7.21 | 7.14 | 7.21 | 7.29 | 7.13 | 7.25 | 7.18 | 7.19 | 7.24 | 7.20 | 7.16 | 7.19 | 7.21 | 7.18 | 7.13 |
| c | 29.7 | 27.88 | 28.1 | 27.5 | 28.12 | 27 | 27.39 | 25.5 | 25.34 | 25.82 | 26.3 | 25.25 | 25.13 | 25.4 | 25.75 |
| D | 1.18 | 1.17 | 1.19 | 1.21 | 1.16 | 1.20 | 1.22 | 1.22 | 1.22 | 1.21 | 1.20 | 1.22 | 1.23 | 1.24 | 1.21 |
| E | 4.91 | 4.58 | 4.63 | 4.6 | 4.5 | 4.49 | 4.50 | 4.34 | 4.29 | 4.2 | 4.44 | 4.28 | 4.3 | 4.4 | 4.39 |
| SL | 37±3.9 (34–43) | 38.5±3.7 (35–44) | 37±3.5 (35–43) | 41±4 (37–42) | 39.5±3.2 (38–44) | 42±2.1 (39–46) | 43±3.1 (37–46) | 44.5±3.4 (38–47) | 43.5±3.2 (39–45) | 45±3.9 (38–47) | 44±3.1 (40–48) | 44.5±3.2 (40–47) | 46±3.8 (39–48) | 47.5±4.1 (39–50) | 43 (35–48) |
| GL | 17.5±1.9 (17–22) | 19±1.5 (17–22) | 18±1.3 (17–21) | 21±1.7 (19–22) | 19.5±1.2 (18–22) | 20.5±1.3 (19–23) | 21±1.4 (17–22) | 21.5±1.3 (19–22) | 22±1.2 (19–23) | 23±1.6 (19–25) | 21±1.2 (20–24) | 22±1.6 (19–23) | 22.5±1.4 (19–24) | 23±1.2 (18–24) | 21 (18–22) |
| SW | 1.85 | 1.92 | 1.80 | 2 | 1.88 | 1.90 | 1.91 | 1.93 | 1.89 | 1.95 | 2 | 1.97 | 1.95 | 2 | 1.86 |
| GS | 0.47 | 0.49 | 0.48 | 0.51 | 0.49 | 0.48 | 0.48 | 0.48 | 0.50 | 0.51 | 0.47 | 0.49 | 0.48 | 0.48 | 0.50 (0.40–0.60) |
Table 6:
Morphometrics of infective juvenile of the Syrian Heterorhabditis bacteriophora isolates. All measurements are in μm (except n, ratio, and percentage), and in the form mean ± S.D. (range).
| Character | SY-ALS | SY-SR | SY-ALB | SY-AL | SY-K | SY-S | SY-D | SY-Z | SY-B | H. bacteriophora Type strain Poinar (1976) |
|---|---|---|---|---|---|---|---|---|---|---|
| n | 20 | 20 | 20 | 20 | 20 | 20 | 20 | 20 | 20 | 15 |
| L | 603±80 (532–613) | 600±67 (521–611) | 597±71 (527–608) | 589±67 (529–603) | 593±78 (522–610) | 578±79 (524–602) | 553±51 (519–597) | 564±63 (520–602) | 575±59 (518–590) | 588 (512–617) |
| MBD | 26±3.2 (18–33) | 26±3 (17–31) | 24.5±2.8 (18–29) | 24±2.9 (18–31) | 24±2.9 (18–33) | 23.5±2.8 (18–31) | 21.5±3.1 (18–29) | 23±3.3 (18–31) | 23±2.7 (18–31) | 23 (18–31) |
| EP | 106.5±5.4 (91–112) | 106±5.6 (86–114) | 105.5±6 (89–109) | 102±6.3 (85–105) | 104±6.8 (89–110) | 98.5±5.7 (87–102) | 93.5±5.1 (88–100) | 94.5±5 (89–101) | 97±5.5 (88–100) | 103 (87–110) |
| NR | 90±4.8 (75–95) | 88.5±4.5 (72–92) | 88.5±4.6 (72–90) | 86.5±4.9 (74–90) | 88±4.7 (71–94) | 84±5 (72–90) | 80.5±4.3 (70–91) | 84±4.6 (71–92) | 83±4.2 (70–89) | 85 (72–93) |
| ES | 132±6.2 (113–138) | 131.5±5.7 (103–135) | 129.5±5.9 (102–135) | 126.5±6.1 (105–130) | 128±6.3 (104–138) | 124±6 (102–132) | 114±5.9 (104–130) | 119±5.6 (107–130) | 122±6.2 (106–129) | 125 (100–139) |
| ABW | 8.2±1.2 (6–8) | 8±1.1 (6.3–8.4) | 7.8±0.9 (6.5–8) | 7.4±0.7 (6.5–7.9) | 7.5±1 (6.5–8) | 7.2±0.7 (6.3–7.9) | 6.5±0.9 (6.2–7.4) | 6.8±0.9 (6.5–7.8) | 7±1 (6–7.4) | - |
| T | 108.5±6.7 (89–115) | 107±6.4 (81–114) | 104±6.1 (83–109) | 100±5.8 (85–104) | 102±6.2 (81–112) | 95±6.3 (82–101) | 89±6 (81–100) | 92.5±5.6 (82–102) | 93.5±5.1 (81–98) | 98 (83–112) |
| a | 23.1 | 23 | 24.36 | 24.5 | 24.7 | 24.59 | 25.72 | 24.52 | 25 | 25 (17–30) |
| b | 4.56 | 4.56 | 4.61 | 4.65 | 4.63 | 4.66 | 4.85 | 4.73 | 4.71 | 4.5 (4–5.1) |
| c | 5.55 | 5.60 | 5.74 | 5.89 | 5.81 | 6 | 6.21 | 6 | 6.14 | 6.2 (5.5–7) |
| D | 0.80 | 0.80 | 0.81 | 0.80 | 0.83 | 0.79 | 0.82 | 0.79 | 0.79 | 0.84 (0.76–0.92) |
| E | 0.98 | 0.99 | 1.01 | 1.02 | 1 | 1.03 | 1.05 | 1.02 | 1.03 | 1.12 (1.03–1.30) |
Table 7:
Morphometrics of male of the Syrian Heterorhabditis bacteriophora isolates. All measurements are in μm (except n, ratio, and percentage), and in the form: mean ± s.d. (range).
| Character | SY-ALS | SY-SR | SY-ALB | SY-AL | SY-K | SY-S | SY-D | SY-Z | SY-B | H. bacteriophora Type strain Poinar (1976) |
|---|---|---|---|---|---|---|---|---|---|---|
| n | 20 | 20 | 20 | 20 | 20 | 20 | 20 | 20 | 20 | 15 |
| L | 845±91 (789–910) | 829±85 (778–903) | 810±65 (785–895) | 819±69 (791–901) | 824±84 (775–915) | 831±95 (793–942) | 804±79 (773–891) | 820±83 (786–900) | 815±74 (781–906) | 820 (780–960) |
| MBD | 46.5±4.8 (37–48) | 44.5±5.1 (39–47) | 41±4.1 (36–43) | 41±4.5 (38–46) | 42.5±4.6 (37–47) | 44.5±4.1 (39–46) | 40±4.2 (35–45) | 42±4 (36–45) | 41±4.3 (36–45) | 43 (38–46) |
| EP | 126±5.5 (115–129) | 124±5.7 (114–126) | 119.5±5.1 (115–125) | 121.5±5.5 (115–125) | 123±5.9 (115–128) | 124.5±5.8 (115–130) | 120±5.9 (112—124) | 122.5±5.1 (116–128) | 122±5.4 (114–129) | 121 (114–130) |
| NR | 76±4.7 (67–79) | 73±4.6 (64–77) | 69±4.1 (67–74) | 71±4.5 (67–77) | 72±5.1 (65–80) | 73.5±5.2 (66–83) | 68.5±4 (65–75) | 71.5±4.3 (67–76) | 71±4.7 (66–78) | 72 (65–81) |
| ES | 103.5±2.1 (100–104) | 103±2 (99–104.5) | 102±1.7 (99–104) | 101±1.1 (100–103) | 102.5±1.2 (98–104) | 103±1.2 (99–104) | 100±0.8 (98–102) | 101±0.9 (99–103) | 100.5±1 (97–104) | 103 (99–105) |
| ABW | 24.5±1.7 (22–26) | 23±1.3 (21–24) | 21±1.1 (20–23) | 21.5±1.5 (20–24) | 23±1.2 (21–24) | 23.5±1 (21–25) | 21±0.9 (20–23) | 23±1.4 (20–24) | 21±1.1 (20–24) | 23 (22–25) |
| T | 32.5±3.9 (24–34) | 29±4.3 (22–32) | 25±3.8 (24–30) | 26.5±3.1 (24–31) | 27±4.1 (22–34) | 29.5±3.5 (24–35) | 23.5±4.5 (21–30) | 27±3.4 (24–30) | 26±3.1 (24–31) | 28 |
| a | 18.1 | 18.6 | 19.7 | 19.9 | 19.3 | 18.6 | 20 | 19.5 | 19.8 | 19 |
| b | 8.16 | 8.04 | 7.94 | 8.10 | 8.03 | 8.06 | 8.04 | 8.11 | 8.01 | 7.9 |
| c | 25.22 | 28.58 | 32.5 | 30.9 | 30.5 | 28.16 | 24.73 | 30.3 | 31.34 | 29.2 |
| D | 1.21 | 1.20 | 1.17 | 1.20 | 1.2 | 1.20 | 1.2 | 1.21 | 1.21 | 1.17 |
| E | 3.76 | 4.27 | 4.78 | 4.58 | 4.55 | 4.22 | 5.1 | 4.53 | 4.69 | 4.32 |
| SL | 42.5±2.5 (37–44) | 40±2.6 (35–43) | 37.5±2 (36–41) | 39±1.7 (37–44) | 39.5±1.8 (36–44) | 41±2.9 (36–46) | 37±2 (35–42) | 39±1.7 (36–42) | 39±1.6 (35–42) | 40 (36–44) |
| GL | 21±0.8 (19–23) | 19±0.7 (18–23) | 18±0.7 (17–22) | 19±0.9 (18–23) | 20±1 (17–23) | 21±1.1 (18–24) | 19±1 (17–22) | 20±1.2 (17–23) | 19±1.1 (17–23) | 20 (18–25) |
| SW | 1.73 | 1.73 | 1.78 | 1.81 | 1.71 | 1.74 | 1.76 | 1.69 | 1.85 | 1.74 |
| GS | 0.49 | 0.47 | 0.48 | 0.48 | 0.48 | 0.51 | 0.51 | 0.51 | 0.48 | 0.50 |
Table 8:
Morphometrics of infective juvenile and male of the Syrian Heterorhabditis pakistanense isolates. All measurements are in μm (except n, ratio, and percentage), and in the form mean ± S.D. (range).
| Character | Heterorhabditis pakistanense SY-BH | Heterorhabditis pakistanense SY-ZO | Heterorhabditis pakistanense n. sp. (Shahina et al., 2016) | |||
|---|---|---|---|---|---|---|
| Stage | Infective Juvenile | Male | Infective Juvenile | Male | Infective Juvenile | Male |
| n | 20 | 20 | 20 | 20 | 25 | 25 |
| L | 575±23 (536–597) | 839.5±91 (710–945) | 583±31 (542–603) | 810±85 (721–991) | 581 (558–624) | 819 (720–1013) |
| MBD | 20±2.7 (17–21) | 39.5±2.4 (36–41) | 19±0.8 (17–22) | 38±2.1 (35–40) | 21 (19–23) | 39.8 (38–43) |
| EP | 95.5±3.6 (85–98) | 120.4±3.2 (113–129) | 92±2.7 (87–98) | 119±3.4 (114–125) | 99.3 (95–106) | 122.8 (112–133) |
| NR | 83±2.5 (70–89) | 87±3.8 (81–97) | 81±1.5 (73–86) | 85.6±4.2 (79–95) | 82 (73–90) | 89 (80–110) |
| ES | 116.5±2.9 (109–121) | 100.5±1.1 (97–103) | 114±1.6 (107–119) | 98.5±0.9 (97–101) | 117 (113–125) | 102 (100–105) |
| ABW | 12.5±1.5 (9.4–15) | 23.3±0.9 (21–25) | 12±1.2 (10–15) | 22.5±1.4 (21–26) | 13.7 (12–16) | 24.5 (22–26) |
| T | 96.5±2.3 (92–103) | 38±1.8 (32–41) | 94±2.1 (90–105) | 35.5±2.3 (31–40) | 99 (95–110) | 37 (30–42) |
| a | 28.75 | 21.2 | 30.6 | 21.3 | 27.4 (25–29) | 20.5 (18–23.5) |
| b | 4.9 | 8.35 | 5.11 | 8.22 | 4.8 (4.7–5.3) | 7.9 (7.2–9.8) |
| c | 5.95 | 22 | 6.2 | 22.8 | 5.78 (5.4–6.2) | 22 (19.3–25) |
| D | 0.81 | 1.19 | 0.80 | 1.20 | 0.84 (0.78–0.97) | 1.19 (1.1–1.26) |
| E | 0.98 | 3.16 | 0.97 | 3.35 | 1 (0.95–1.07) | 3.31 (2.67–4) |
| SL | - | 39±2.4 (35–41) | - | 37±2.7 (34–40) | - | 38.5 (35–42) |
| GL | - | 21.4±1.9 (19–23) | - | 20.5±1.5 (19–22) | - | 21 (20–22) |
| SW | - | 1.67 | - | 1.64 | - | 1.56 (1.44–1.91) |
| GS | - | 0.54 | - | 0.55 | - | 0.58 (0.48–0.65) |

Figure 5:
Evolutionary relationship of Syrian Heterorhabditis isolates using ITS rDNA sequences. The evolutionary history was inferred by using the maximum likelihood method. The bootstrap consensus tree inferred from 1,000 replicates is taken to present the evolutionary history of the analyzed taxa. Caenorhabditis elegans was used as an outgroup.

Figure 6:
Cluster dendrogram of Syrian Heterorhabditis species isolates based on male morphological traits.

Figure 7:
Co-inertia loading plot for EPN species and habitat and abiotic factors data in the Syrian coastal ecosystems surveyed.
Table 9:
Mortality of third- and fourth-instar larvae of Tuta absoluta on outside leaves at each dose rate for three different entomopathogenic nematodes (mean ± S.D.).
| EPNs species | Dose 1IJ/ L | Dose 5Ij/ L | Dose 10Ij/L | Dose 15Ij/ L | Dose 25Ij/ L | Dose 50Ij/ L | |
|---|---|---|---|---|---|---|---|
| Third-instar larvae of T. absoluta | S. affine. 313 | 25.00a±1.62 | 37.50a±2.16 | 45.83a±1.33 | 54.17a±1.45 | 66.67a±0 | 79.17a±0.95 |
| H. indica. Fn | 16.67b±0 | 26.17ab±2.27 | 37.50a±1.33 | 45.83a±1.45 | 62.50a±1.43 | 70.83a±0.84 | |
| H. bacteriophora. H | 8.33bc±0.32 | 16.67bc±0.15 | 25.00b±2.62 | 33.33b±0 | 37.50b±1.43 | 45.83b±0.84 | |
| Control | 0c±0 | 0c±0 | 0c±0 | 0c±0 | 0c±0 | 0c±0 | |
| P=0.00139** | P=0.0000657*** | P=0.0000114*** | P=0.0000105*** | P=0.0000559*** | P=0.0000011*** | ||
| df= 3 | df= 3 | df=3 | df=3 | df=3 | df=3 | ||
| F=10 | F=30.7 | F=27.6 | F= 65.33 | F=108.5 | F=97.22 | ||
| Fourth-instar larvae of T. absoluta | S. affine. 313 | 29.17a±1.33 | 41.67a±1.64 | 50.0a ±0 | 59.52a±1.16 | 71.43a±1.59 | 91.67a±1.62 |
| H. indica. Fn | 20.83ab±1.33 | 36.50ab±1.33 | 41.67ab±1.62 | 50.0ab±0 | 50.00a±1.45 | 87.50a±1.33 | |
| H. bacteriophora. H | 12.50ab±2.17 | 25.00b±1.64 | 33.33b±0 | 41.67b±1.62 | 45.83a±1.33 | 54.17b±1.95 | |
| Control | 4.17b±1.62 | 4.17c±1.34 | 4.17c±1.33 | 4.17c±1.33 | 4.17b±1.33 | 4.17c±1.33 | |
| P=0.00671** | P=0.000318*** | P=0.0000175*** | P=0.0000462*** | P=0.00295** | P=0.0000311*** | ||
| df= 3 | df= 3 | df= 3 | df= 3 | df= 3 | df= 3 | ||
| F=6.67 | F=14 | F=39.29 | F=32.78 | F=8.29 | F=53.86 |
[i] The mortalities including the same letter are not statistically different (P < 0.01).

Figure 8:
Larvae and pupae of T.absoluta infected by S.affine M.313. A: S. affine after completing its development within the fourth-instar larva. B: Emergence of the IJs from the infected fourth instar larva. C: Infected third instar larva. D, E: S.affine female inside pupae observed with the naked eye and 4× magnification.

Figure 9:
Larvae and pupae of T.absoluta infected by H.indica H. A, B: infected fouth-instar larvae outside leaf and inside mine. C, D: Infected third-instar larva outside leaf, and hermaphroditic inside it. E: Infected pupae. F: Hermaphroditics inside pupa.

Figure 10:
Mortality rates (mean ± SD) of third- and fourth-instar larvae inside the galleries and pupae of T.absoluta treated by LD50 inoculation rates of S.affine and H.indica (P = 0.01).