Figure 1:
Line drawings of Heterorhabditis ruandica n. sp. (A) A hermaphroditic female. (B) Cephalic region of a hermaphroditic female. (C) Pharyngeal region of a hermaphroditic female. (D) Anterior part of the reproductive system of a hermaphroditic female. (E) Posterior end of a hermaphroditic female. (F) An amphimictic female. (G) Pharyngeal region of an amphimictic female. (H) Posterior end of an amphimictic female. (I) A male adult. (J) Pharyngeal region of a male adult. (K) Posterior region of a male adult. (L) Pharyngeal region of an infective juvenile. (M) Posterior end of an infective juvenile. (N) An infective juvenile.
Figure 2:
Light microscope micrographs of Heterorhabditis ruandica n. sp. (A) An amphimictic female (black arrow pointing at the position of the vulva, white arrow pointing at the anus). (B) Pharyngeal region of an amphimictic female. (C) Posterior end of an amphimictic female. (D) Vulva of an amphimictic female. (E) A male adult. (F) Pharyngeal region of a male adult. (G) Posterior end of a male adult (arrows pointing at the genital papillae).
Figure 3:
Light microscope micrographs of Heterorhabditis ruandica n. sp. (A) A hermaphroditic female. (B) Anterior end of a hermaphroditic female. (C) Pharyngeal region of a hermaphroditic female. (D) Posterior end of a hermaphroditic female. (E) A sheathed third stage juvenile (J2). (F) Pharyngeal region of a sheathed third stage juvenile (J2). (G) Posterior end of a sheathed third stage juvenile (J2) (arrow pointing the anus). (H) A non-sheathed third stage juvenile (J3). (I) Pharyngeal region of a non-sheathed third stage juvenile (J3). (J) Posterior end of a non-sheathed third stage juvenile (J3).
Figure 4:
Scanning electron microscope (SEM) micrographs of Heterorhabditis ruandica n. sp. (A, B) Lip region in lateral and frontal views, respectively, of a hermaphroditic female. (C) Excretory pore of a hermaphroditic female (pointed by an arrow). (D) Vulva of a hermaphroditic female (pointed by an arrow). (E) Tail of a hermaphroditic female in lateral view. (F, G) Lip region of a female adult in lateral and frontal views, respectively. (H) Excretory pore (pointed by an arrow). (I) Vulva of a female adult (pointed by an arrow). (J) Tail of a female adult in ventral view. (K, L) Lip region of a male adult in sublateral and frontal views, respectively. (M, N) Posterior end of a male adult in ventral and lateral views, respectively (arrows pointing the genital papillae). (O) Lip region of a second-stage juvenile (J2) in lateral view. (P) Cuticle of a second-stage juvenile (J2). (Q) Tail of a second-stage juvenile (J2) in lateral and ventral views, respectively. (R) Lip region of a third-stage juvenile (J3) in ventral view (arrow pointing the frontal tooth). (S) Cuticle (arrow pointing the cuticle of a third-stage juvenile).
Table 1.
Morphometrics of infective juveniles and adult generations of Heterorhabditis ruandica n. sp.
| Male | |||||
|---|---|---|---|---|---|
| Characters | Holotype | Paratypes | Hermaphrodite (1st Gen) paratypes | Female (2nd Gen) paratypes | Infective juvenile paratypes |
| n | 1 | 20 | 20 | 20 | 25 |
| Body length (L) | 760 | 769 ± 60 (652-863) | 3295 ± 286 (2907-4123) | 1366 ± 123 (1131-1608) | 544 ± 29 (496-591) |
| a (L/BD) | 20.3 | 17 ± 1.5 (15-21) | 14.1 ± 1.1 (11.7-16.1) | 18 ± 1.4 (15-20) | 24 ± 3.0 (20-29) |
| b (L/NL) | 7.8 | 8.1 ± 1.0 (5.8-9.7) | 23 ± 1.8 (21-27) | 11.4 ± 1.2 (9.0-13.6) | 4.7 ± 0.4 (4.1-5.4) |
| c (L/T) | 26.2 | 31 ± 3.6 (23-36) | 42 ± 5.7 (34-51) | 20 ± 2.2 (16-24) | 8.2 ± 1.0 (7.6-8.6) |
| c’ (T/ABW) | 1.1 | 1.4 ± 0.2 (0.6-1.7) | 2.2 ± 0.3 (1.7-2.6) | 2.8 ± 0.5 (1.9-3.6) | 4.6 ± 0.8 (3.4-5.8) |
| V (VA/L × 100) | – | – | 48 ± 2.5 (45-55) | 48 ± 2.3 (41-51) | – |
| Max. Body Width (MBD) | 37.5 | 44 ± 3.0 (40-51) | 233 ± 17 (209-274) | 77 ± 4.0 (68-83) | 23 ± 2.7 (18-27) |
| Lip region width | 6.5 | 7.2 ± 0.8 (5.7-8.4) | 12.4 ± 0.8 (11.0-14.0) | 10.3 ± 0.9 (8.8-12.2) | – |
| Stoma length | 9.5 | 11.1 ± 1.6 (8.7-13.9) | 14.9 ± 1.4 (13-18) | 13.6 ± 1.8 (10.4-16.0) | 13.8 ± 1.2 (12.1-16.0) |
| Bulb length (BL) | 18.5 | 20 ± 1.8 (18-25) | 35 ± 3.6 (29-42) | 27 ± 2 (23-30) | 13.8 ± 1.8 (11.0-19.0) |
| Pharynx length (PL) | 95.2 | 84 ± 7.1 (74-107) | 128 ± 6.3 (118-142) | 107 ± 6.9 (91-120) | 102 ± 7.0 (91-115) |
| Nerve ring – ant. end (NR) | 68 | 63 ± 5.2 (56-74) | 93 ± 7.5 (78-108) | 81 ± 6.4 (69-97) | 55 ± 3.6 (52-64) |
| Excretory pore– ant. end (EP) | 84.3 | 81 ± 10.1 (61-109) | 121 ± 11 (106-153) | 111 ± 10.8 (92-129) | 78 ± 3.4 (70-89) |
| Neck length (Stoma+Pharynx, NL) | 98 | 96 ± 7.3 (84-117) | 143 ± 6.3 (134-159) | 120 ± 6.0 (107-132) | 115 ± 7.3 (103-131) |
| Body width at neck base | 36 | 34 ± 1.9 (30-37) | 119 ± 8.9 (101-138) | 58 ± 4.3 (50-66) | 18 ± 3.0 (15-24) |
| Vagina length | – | – | 28 ± 4.0 (20-38) | 19.2 ± 2.9 (15-26) | – |
| Body width at vulva | – | – | 240 ± 21 (199-278) | 78 ± 3.8 (72-85) | – |
| Vulva – ant. end (VA) | – | – | 1581 ± 151 (1369-1882) | 655 ± 47 (572-706) | – |
| Vulva – post. End (PV) | – | – | 1713 ± 178 (1453-2241) | 710 ± 89 (559-949) | – |
| Rectum length | – | – | 36 ± 4.6 (29-49) | 30 ± 3.8 (24-35) | 8.5 ± 1.9 (6.1-13.7) |
| Anal body diam. (ABD) | 26.1 | 18 ± 2.4 (15-25) | 37 ± 5.5 (29-51) | 25 ± 4.5 (18-34) | 12.4 ± 1.8 (9.2-16.0) |
| Tail with sheath length (T) | – | – | – | – | 56 ± 4.9 (49-64) |
| Tail without sheath length | 29 | 25 ± 3.2 (21-29) | 80 ± 7.9 (63-98) | 68 ± 6.5 (62-88) | 30.4 ± 4.5 (22-39) |
| Spicule length (SL) | 49 | 43 ± 4.1 (34-50) | – | – | – |
| Gubernaculum length (GL) | 20.2 | 18 ± 1.5 (15-21) | – | – | – |
| Stoma length/lip region width | 1.5 | – | 1.2 ± 0.2 (1.0-1.6) | 1.3 ± 0.1 (1.1-1.6) | – |
| Nerve ring % (NR/NL × 100) | 69.4 | 67 ± 4.4 (58-75) | 65 ± 5.0 (56-78) | 67 ± 3.9 (61-75) | – |
| Excretory pore % (EP/NL × 100) | 86 | 85 ± 8.5 (61-97) | 85 ± 8.3 (67-103) | 92 ± 7.6 (74-104) | – |
| Rectum % (R/ABD × 100) | – | – | 90 ± 10 (80-130) | 128 ± 29 (90-181) | – |
| D % (EP /PL × 100) | 88.5 | 96 ± 9.2 (69-111) | 95 ± 9.3 (74-114) | 104 ± 9.3 (82-118) | 78 ± 7.6 (66-98) |
| E % (EP/T × 100) | 290 | 325 ± 49 (232-413) | 153 ± 24 (120-205) | 164 ± 23 (111-203) | 139 ± 13.4 (112-168) |
| SW % (SL/ABD × 100) | 242 | 243 ± 47 (150-306) | – | – | – |
| GS % (GL/SL × 100) | 41.2 | 42 ± 5.2 (35-57) | – | – | – |
| H % (H/T × 100 | – | – | – | – | 46 ± 4.0 (37–55) |
Table 2.
Morphometrics of infective juveniles and adult generations of Heterorhabditis zacatecana n. sp.
| Male | |||||
|---|---|---|---|---|---|
| Characters | Holotype | Paratypes | Hermaphrodite (1st Gen) paratypes | Female (2nd Gen) paratypes | Infective juvenile paratypes |
| n | 1 | 20 | 22 | 22 | 25 |
| Body length (L) | 808.1 | 861 ± 29 (811-914) | 5127 ± 494 (4408-6179) | 2244 ± 203 (1954-2798) | 539 ± 21 (493-578) |
| a (L/BD) | 19.0 | 18 ± 1.6 (15-22) | 16 ± 2.0 (13-20) | 12.3 ± 1.2 (10.5-15.0) | 22 ± 1.2 (19-24) |
| b (L/NL) | 8.1 | 9.1 ± 1.1 (7.6-12) | 26 ± 4.3 (20-34) | 18 ± 1.8 (16-21) | 5.0 ± 0.4 (4.4-5.9) |
| c (L/T) | 28.9 | 34 ± 4.2 (26-43) | 70 ± 10.4 (52-90) | 39 ± 7.4 (31-63) | 9.4 ± 0.6 (8.2-10.5) |
| c’ (T/ABW) | 1.4 | 1.6 ± 0.3 (1.2-2.5) | 1.6 ± 0.3 (1.2-2.4) | 1.7 ± 0.2 (1.3-2.0) | 5.3 ± 0.6 (4.3-6.7) |
| V (VA/L × 100) | – | – | 48 ± 4.3 (36-57) | 53 ± 4.2 (43-61) | – |
| Max. Body Width (MBD) | 42.5 | 48 ± 3.6 (41-56) | 319 ± 41 (235-385) | 183 ± 23 (160-228) | 24 ± 0.9 (23-27) |
| Lip region width | 6.2 | 7.4 ± 0.7 (6.2-8.8) | 11.7 ± 2.4 (9.2-19.2) | 10.1 ± 1.0 (7.7-11.4) | 4.0 ± 0.5 (3.2-5.2) |
| Stoma length | 10 | 9.3 ± 1.0 (6.3-11) | 19 ± 2.0 (14-23) | 11.5 ± 1.7 (8.0-15.2) | 13.5 ± 1.0 (12.0-15.3) |
| Bulb length (BL) | 20.2 | 22 ± 2.4 (19-28) | 40 ± 4.6 (28-49) | 30 ± 2.6 (28-38) | 20 ± 1.4 (17.1-23.0) |
| Pharynx length (PL) | 95.2 | 86 ± 9.8 (57-100) | 182 ± 23 (155-211) | 113 ± 9.5 (101-133) | 95 ± 7.2 (82-111) |
| Nerve ring – ant. end (NR) | 65.4 | 66 ± 5.3 (60-78) | 131 ± 22 (96-169) | 83 ± 7.3 (71-96) | 81 ± 6.3 (69-72) |
| Excretory pore– ant. end (EP) | 96.2 | 93 ± 9.6 (77-109) | 150 ± 24 (108-190) | 113 ± 11 (100-133) | 89 ± 6.8 (72-99) |
| Neck length (Stoma+Pharynx, NL) | 99.3 | 96 ± 9.6 (71-108) | 201 ± 21 (174-231) | 124 ± 10 (112-148) | 109 ± 6.9 (96-124) |
| Body width at neck base | 34.5 | 36 ± 2.3 (31-40) | 167 ± 13 (133-188) | 95 ± 13.9 (74-121) | 23 ± 1.3 (19-26) |
| Vagina length | – | – | 31 ± 4.0 (24-36) | 25 ± 6.4 (17-42) | – |
| Body width at vulva | – | – | 331 ± 33 (257-379) | 185 ± 27 (153-230) | – |
| Vulva – ant. end (VA) | – | – | 2470 ± 279 (1959-3038) | 1182 ± 129 (910-1397) | – |
| Vulva – post. end (PV) | – | – | 2657 ± 279 (1990-3938) | 1062 ± 147 (860-1455) | – |
| Rectum length | – | – | 36 ± 4.6 (30-41) | 27 ± 4.1 (19-39) | – |
| Anal body diam. (ABD) | 19.6 | 17 ± 2.3 (13-22) | 47 ± 8.1 (34-58) | 35 ± 3.2 (31-41) | 11.1 ± 1.3 (8.6-14.1) |
| Tail with sheath length (T) | – | – | – | – | 58 ± 3.1 (52-63) |
| Tail without sheath length | 28 | 26 ± 3.3 (21-33) | 74 ± 8.3 (63-87) | 58 ± 8.2 (45-75) | 29.4 ± 2.5 (25-34) |
| Spicule length (SL) | 54.1 | 45 ± 3.7 (38-55) | – | – | – |
| Gubernaculum length (GL) | 18.7 | 20 ± 2.1 (15-25) | – | – | – |
| Stoma length/lip region width | 1.6 | – | 1.6 ± 0.3 (1.1-2.1) | 1.2 ± 0.2 (0.8-1.7) | – |
| Nerve ring % (NR/NL × 100) | 65.9 | 69 ± 9.9 (61-96) | 65 ± 9.4 (49-86) | 67 ± 4.8 (60-82) | – |
| Excretory pore % (EP/NL × 100) | 96.9 | 98 ± 17 (78-134) | 75 ± 11 (51-95) | 67 ± 4.8 (60-82) | – |
| Rectum % (R/ABD × 100) | – | – | 79 ± 17 (54-112) | 76 ± 13 (52-106) | – |
| D % (EP /NL × 100) | 101.05 | 109 ± 21 (83-156) | 75 ± 11 (55-95) | 92 ± 7.9 (80-111) | 94 ± 12 (68-120) |
| E % (EP/T × 100) | 343.6 | 365 ± 68 (236-503) | 206 ± 46 (145-303) | 197 ± 27 (145-246) | 154 ± 14 (128-184) |
| SW % (SL/ABD × 100) | 276 | 270 ± 50 (170-320) | – | – | – |
| GS % (GL/SL × 100) | 34.56 | 40 ± 10 (40-60) | – | – | – |
| H % (H/T × 100 | – | – | – | – | 47 ± 5.6 (35-56) |
Table 3.
Comparative morphometrics of adult males of Heterorhabditis ruandica n. sp., H. zacatecana n. sp., and of different closely related Heterorhabditis species. All measurements are in µm (except ratios and percentages)
| Species | L | BD | EP | NR | NL | T | SL | GL | a | b | c | c´ | SW% | GS% | D% | Country | Reference |
|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
| H. amazonensis | 692–826 | 36–43 | 96–116 | 71–88 | 97–114 | 29–41 | 35–45 | 19–23 | 18.7* | 7.7** | 27.5** | 1.3** | 120–187 | 44–56 | 95–109 | Brazil | Andaló et al. (2006) |
| H. atacamensis | 842–1025 | 42–55 | 116–149 | 69–93 | 99–119 | 24–36 | 40–49 | 17–22 | 19.7* | 9.6** | 29.3** | 1.5** | 179–249 | 38–51 | 108–126 | Chile | Edgington et al. (2011) |
| H. bacteriophora | 780–960 | 38–46 | 114–130 | 65–81 | 99–105 | 22–36 | 36–44 | 18–25 | 20.8* | 9.1* | 34.3* | 1.8* | 174 | 50 | 117 | Australia | Poinar (1976) |
| 700–940 | 37–50 | 113–140 | 70–85 | 95–110 | 20–27 | 39–47 | 18–24 | – | – | – | – | – | – | – | Argentina | Agüera de Doucet and Doucet (1986) | |
| 689–880 | 38–46 | 78–123 | 55–90 | 92–124 | 21–32 | 34–48 | 17–26 | – | – | – | 1.2* | 147–256 | 41–49 | 68–106 | Australia | Sagun et al. (2015) | |
| 782–927 | 92–120 | 103–139 | 58–76 | 84–105 | 28–37 | 51–53 | 17–26 | 6.6–8.5 | 8.5–10 | 23–32 | 1.4–2.2 | 194–282 | 37–57 | 108–157 | India | Bhat et al. (2019a) | |
| 805–1075 | 42–57 | 84–111 | 84–75 | 80–119 | 24–39 | 39–51 | 17–27 | 16–22 | 7.1–12 | 22–41 | 1.0–1.7 | 170–225 | 40–62 | 77–136 | India | Rana et al. (2020) | |
| as H. argentinensis # | 1000–2000 | 42–70 | 145–170 | 64–82 | 103–120 | 28–49 | 42–49 | 20–26 | 16.7* | 8.3* | 14.3* | 1.4* | 198* | 62* | 92* | Argentina | Stock (1993) |
| as H. heliothidis £ | 1000–1200 | 32–60 | 125* | 125* | 113–131 | 29–36 | 42–52 | 22–27 | 19–35 | 8–11 | 28–38 | 1.3* | 185* | 51* | 95* | USA | Khan et al. (1976) |
| H. baujardi | 818–970 | 45–53 | 71–93 | 54–77 | 105–132 | 28–38 | 33–45 | 18–22 | 16–22 | 6.4–8.8 | 24–33 | 1.5** | 138–208 | 44–61 | 79** | Vietnam | Phan et al. (2003) |
| 710–903 | 40–50 | 83–93 | 53–68 | 98–110 | 33–40 | 43–48 | 20–28 | 16–20 | 6.7–9.3 | 18–28 | – | 154–200 | 47–61 | 80–90 | India | Vanlalhlimpuia et al. (2018) | |
| as H. somsookae # | 737–870 | 37–44 | 68–93 | 72–83 | 90–120 | 20–30 | 32–45 | 17–23 | 20.7** | 8.3** | 32.3** | 1.2** | 133–198 | 42–59 | 74–99 | Thailand | Maneesakorn et al. (2015) |
| H. beicherriana | 889–1192 | 51–73 | 130–157 | 81–108 | 116–143 | 32–45 | 40–49 | 22–27 | 15–23 | 7.2–10 | 22–34 | 1.3–2.3 | 153–208 | 48–59 | 102–120 | China | Li et al. (2012) |
| H. downesi | 699–876 | 33–40 | 86–91 | 62–78 | 97–106 | 29–34 | 41–47 | 17–19 | 26.6* | 8.8** | 27.4** | 1.4** | 170–220 | 36–47 | 90 | Ireland | Stock et al., 2002 |
| H. egyptii + | 594–848 | 31–56 | 80–97 | 56–84 | 96–109 | 23–34 | 25–50 | 16–22 | 17.1* | 6.6** | 19.5** | 1.5* | 120–220 | 40–65 | 84–91 | Egypt | Abd–Elgawad and Ameen (2005) |
| H. floridensis | 785–294 | 43–50 | 104–128 | 73–90 | 97–111 | 29–40 | 36–46 | 17–30 | 19.9* | 7.9** | 24.1** | 1.4** | 133–209 | 47–65 | 112 | USA | Nguyen et al. (2006) |
| H. georgiana | 721–913 | 43–55 | 101–145 | 72–93 | 100–122 | 29–41 | 41–49 | 20–28 | 16.5* | 7.7** | 26.1** | 1.4** | 150–200 | 51–64 | 100–122 | USA | Nguyen et al. (2008) |
| H. hambletoni + | 510–800 | 38–60 | 80–100 | 80–90 | – | – | – | – | – | – | – | – | – | – | – | Brazil | Pereira (1937) |
| H. hoptha $ | 554–837 | – | – | – | – | 30.9* | 43–60 | 26–30 | 18–22 | 5.9–8.2 | 18–37 | 1.1* | 167** | 55** | – | USA | Turco (1970) |
| H. indica | 573–788 | 35–46 | 109–138 | 72–85 | 93–109 | 24–32 | 35–48 | 18–23 | 17.6* | 6.7** | 23.0** | 1.1** | 187 | 49 | 121 | India | Poinar et al. (1992) |
| 724–864 | 41–48 | 96–113 | 63–80 | 89–109 | 29–36 | 30–40 | 21–31 | 17–20 | 7.6–8.6 | 22–27 | 1.4–1.8 | 155–210 | 49–68 | 101–111 | India | Kajol et al. (2020) | |
| 609–916 | 26–50 | 78–109 | 62–83 | 90–116 | 18–33 | 37–48 | 19–26 | 16–28 | 6.5–8.2 | 25–37 | 1.0–1.5 | 116–225 | 49–64 | 86–106 | India | Bhat et al. (2021b) | |
| as H. brevicaudis # | 840–950 | 40–48 | 92–100 | 80–88 | 104–112 | 28–36 | 44–48 | 20–24 | – | – | – | 2.9* | 170* | 47* | 84* | China | Liu (1994) |
| as H. gerrardi # | 508–916 | 34–48 | 93–141 | 54–87 | 78–115 | 28–46 | 34–48 | 16–27 | – | – | – | – | 138–274 | 40–69 | 100–172 | Australia | Plichta et al. (2009) |
| as H. hawaiiensis # | 864–1130 | 49–84 | 71–146 | 67–112 | 100–149 | 26–40 | 40–51 | 18–26 | – | – | – | – | – | – | – | USA | Gardner et al. (1994) |
| as H. pakistanense # | 720–1013 | 38–43 | 112–133 | 80–110 | 100–105 | 30–42 | 35–42 | 20–22 | 18–24 | 7.2–9.8 | 19–25 | 1.4** | 144–191 | 48–65 | 110–126 | Pakistan | Shahina et al. (2017) |
| H. marelatus | 805–1046 | 48–56 | 110–168 | 61–95 | 99–123 | 24–38 | 41–49 | 18–22 | 15.5* | 7.8** | 30.0** | 1.1** | 196 | 36–50 | 113** | USA | Liu and Berry (1996) |
| 960–1010 | 48–80 | 107–116 | 89–95 | 115–130 | 37–47 | 48–52 | 21–24 | – | – | – | – | – | – | – | USA | Stock (1997) | |
| as H. hepialius # | 8000–1000 | 65–98 | 102–131 | 84–114 | 113–139 | 37–49 | 42–52 | 17–24 | – | – | – | – | – | – | – | USA | Stock et al. (1996) |
| H. megidis | 800–1100 | 44–50 | 139–176 | 96–112 | 122–134 | 35–43 | 46–54 | 17–24 | 18–22 | 7–9 | 23–31 | 1.6* | 188 | 43 | 122 | USA | Poinar et al. (1987) |
| H. mexicana | 614–801 | 38–47 | 108–145 | 61–83 | 89–108 | 21–36 | 30–47 | 18–32 | 21.7* | 6.8** | 27.6** | 1.1** | 130–196 | 43–70 | 114–149 | Mexico | Nguyen et al. (2004) |
| H. noenieputensis | 530–775 | 34–46 | 75–102 | 64–75 | 88–106 | 21–32 | 37–49 | 17–24 | 14–18 | 5.6–7.9 | 21–33 | 1.1–1.7 | 202–301 | 38–56 | 81–108 | S. Africa | Malan et al. (2014) |
| H. poinari $ | 970–1100 | 43–70 | – | – | 150–150 | 36–65 | 43–55 | 24–32 | 95–100 | 51–95 | 11–97 | – | – | – | – | USA | Kakulia and Mikaia (1997) |
| H. ruandica Rw14_NC4a | 652–863 | 40–51 | 61–109 | 56–74 | 84–117 | 21–29 | 34–50 | 16–23 | 15–21 | 5.8–9.7 | 23–36 | 0.6–1.7 | 150–306 | 35–57 | 61–97 | Rwanda | This study |
| H. safricana | 777–1009 | 40–58 | 104–147 | 52–61 | 105–126 | 27–49 | 35–54 | 19–27 | 20.1* | 7.9** | 43.0** | 1.5* | 130–259 | 43–62 | 92–133 | S. Africa | Malan et al. (2008) |
| H. taysearae | 648–736 | 38–48 | 78–120 | 54–88 | 85–123 | 20–29 | 30–42 | 12–21 | 15.1* | 6.5** | 14.0** | 1.3** | 156 | 46 | 88 | Egypt | Shamseldean et al. (1996) |
| as H. sonorensis # | 500–750 | 32–42 | 60–84 | 60–80 | 80–100 | 25–45 | 31–45 | 20–31 | – | – | – | – | 110–180 | 40–75 | 72–91 | Mexico | Stock et al. (2009) |
| H. zealandica | 848–1044 | 36–45 | 130–150 | – | 110–128 | 30–41 | 48–55 | 19–25 | – | – | – | 1.7* | 246 | 44 | 118 | N. Zealand | Poinar (1990) |
| as H. heliothidis # | 848–1044 | 36–45 | 130–150 | – | 110–128 | 30–41 | 48–55 | 19–25 | – | – | – | 1.7* | 246 | 44 | 118 | N. Zealand | Wouts (1979) |
| H. zacatecana MEX–39 | 811–914 | 41–56 | 77–109 | 60–78 | 71–108 | 21–33 | 38–55 | 15–25 | 15–25 | 7.6–12 | 26–43 | 1.2–2.5 | 170–320 | 40–60 | 78–134 | Mexico | This study |
Table 4.
Comparative morphometrics of hermaphrodite females of Heterorhabditis ruandica n. sp., H. zacatecana n. sp., and of different closely related Heterorhabditis species.
| Species | L | BD | EP | NR | NL | T | a | b | c | c´ | V | ABD | D% | Country | Reference |
|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
| H. amazonensis | 3517-5587 | 220-316 | 184-238 | 128-171 | 180-225 | 104-154 | – | – | – | 2.3* | 42-47 | 59-83 | 103* | Brazil | Andaló et al. (2006) |
| H. atacamensis | 1791-2904 | 88-122 | 165-206 | 101-132 | 174-200 | 72–112 | – | – | – | 2.7* | 39-48 | 30-46 | 90-114 | Chile | Edgington et al. (2011) |
| H. bacteriophora | 3630-4390 | 160-180 | 189-217 | 121-130 | 189-205 | 81-93 | – | – | – | – | 41-47 | 40-53 | 106 | Australia | Poinar (1976) |
| 4200-5600 | 175-242 | 163-225 | 125-152 | 180-220 | 50-87 | – | – | – | – | 35-45 | 45-62 | – | Argentina | Agüera de Doucet and Doucet (1986) | |
| 2686-4893 | 131-241 | 150-379 | 80–196 | 162–302 | 70-120 | – | – | – | 1.8* | 36-52 | 43-76 | 76-126 | Australia | Sagun et al. (2015) | |
| 3086-5492 | 221-352 | 127-260 | 79-162 | 101–200 | 71-123 | 9.2-28 | 23-37 | 25-75 | 1.2-3.7 | 37-52 | 34-75 | 112-155 | India | Bhat et al. (2019a) | |
| 3916-5155 | 205-206 | 153-198 | 94-127 | 158-207 | 70-98 | 17-21 | 21-28 | 46-69 | 1.7-2.2 | 37-46 | 39-51 | 75-103 | India | Rana et al. (2020) | |
| as H. argentinensis # | 5000-7500 | 250-575 | 250-340 | 132-196 | 235-300 | 100–140 | – | – | – | 1.8* | 40-50 | 70-120 | 102* | Argentina | Stock (1993) |
| as H. heliothidis £ | 3000-5100 | 200-344 | 250* | 250* | 163-286 | 76-100 | 11-18 | 11-25 | 30-63 | 2.2* | 45-52 | 62.5* | 80* | USA | Khan et al. (1976) |
| H. baujardi | 3135-4170 | 180-240 | 156-192 | 119-147 | 186-206 | 66-114 | 15-19 | 16-21 | 36-50 | 2.0* | 43-48 | 47-63 | 88* | Vietnam | Phan et al. (2003) |
| 3250-3970 | 190-250 | 98-115 | 120-135 | 180-205 | 80-105 | 13-19 | 16-20 | 31-45 | – | 41-49 | 50-65 | 73-92 | India | Vanlalhlimpuia et al. (2018) | |
| as H. somsookae # | 2275-3952 | 108-183 | 156-214 | 118-144 | 158-193 | 56-87 | – | – | – | 2.3* | 41-56 | 30-53 | 86-113 | Thailand | Maneesakorn et al. (2015) |
| H. beicherriana | 3671-5543 | 198-374 | 165-297 | 135-243 | 192-343 | 68-130 | 13-20 | 13-25 | 34-62 | 1.0-2.3 | 41-49 | 51-92 | 76-94 | China | Li et al. (2012) |
| H. downesi | 3030-5051 | 183-291 | 200-254 | 175-230 | 230-244 | 60-70 | – | – | – | 1.1* | 50-55 | 57-65 | 117* | Ireland | Stock et al. (2002) |
| H. egyptii + | 2100-3100 | 107-164 | 154-205 | 101-147 | 144-192 | 83-115 | – | – | – | 2.7* | 46-59 | 33-51 | 104* | Egypt | Abd–Elgawad and Ameen (2005) |
| H. floridensis | 3731-5865 | 217-331 | 211-301 | 169-271 | 271-391 | 84-126 | – | – | – | 2.5* | 44-49 | 42-78 | 104* | USA | Nguyen et al. (2006) |
| H. georgiana | 3232-4928 | 157-267 | 200-277 | 143-217 | 132-271 | 65-96 | – | – | – | 1.2* | 44-55 | 42.6* | – | USA | Nguyen et al. (2008) |
| H. hambletoni + | – | – | – | – | – | – | – | – | – | – | – | – | – | Brazil | Pereira (1937) |
| H. hoptha $ | – | – | – | – | – | – | – | – | – | – | – | – | – | USA | Turco (1970) |
| H. indica | 2300-3100 | 107-145 | 163-187 | 104-123 | 163-179 | 72-110 | – | – | – | – | 45-50 | 38-51 | – | India | Poinar et al. (1992) |
| 2751-4481 | 168-273 | 184-238 | 115-157 | 167-204 | 67-108 | 16-18 | 15-25 | 35-55 | 1.4-2.5 | 37-48 | 30-71 | 103-132 | India | Kajol et al. (2020) | |
| 2861-4227 | 152-208 | 140-179 | 119-146 | 165-186 | 79-114 | 16-23 | 17-24 | 30-47 | 1.5-2.4 | 39-55 | 37-56 | 81-100 | India | Bhat et al. (2021b) | |
| as H. brevicaudis # | 3550-5040 | 200-312 | 160-200 | 144-176 | 192-240 | 72-128 | – | – | – | – | 37-50 | 56-88 | 91* | China | Liu (1994) |
| as H. gerrardi # | 2049-4288 | 93-209 | 103–288 | 82-210 | 146-317 | 90-196 | – | – | – | 2.4* | 40-48 | 40-80 | 90-147 | Australia | Plichta et al. (2009) |
| as H. hawaiiensis # | 4000-7000 | 270-376 | 219-318 | 102-212 | 187-283 | 67–98 | – | – | – | – | – | 38-79 | – | USA | Gardner et al. (1994) |
| as H. pakistanense # | 1939-4625 | 102-240 | 145-186 | 130-180 | 155-220 | 64–95 | 16–23 | 11–24 | 23–58 | 1.7* | 41-49 | 37-55 | 68-106 | Pakistan | Shahina et al. (2017) |
| H. marelatus | 3000-4500 | 161–233 | 212-287 | 133-182 | 190-244 | 75-101 | – | – | – | 1.3* | 45-50 | 20-28 | 109* | USA | Liu and Berry (1996) |
| – | – | – | – | – | – | – | – | – | – | – | – | – | USA | Stock (1997) | |
| as H. hepialius # | 4000-5200 | 205-335 | 175-258 | 117-161 | 190-223 | 60-126 | – | – | – | 1.9* | 45-50 | 34-60 | – | USA | Stock et al. (1996) |
| H. megidis | 2400-4900 | 120-133 | 193-270 | 139-178 | 106-269 | 95-124 | 14-24 | 12-21 | 23-49 | – | 45-50 | 36-86 | – | USA | Poinar et al. (1987) |
| H. mexicana | 2440-4606 | 135-267 | 103-201 | 114-171 | 168-221 | 94-170 | – | – | – | 2.6* | 30-58 | 40-46 | 90* | Mexico | Nguyen et al. (2004) |
| H. noenieputensis | 2987-5498 | 168-289 | 152-209 | 112-152 | 166-220 | 79-120 | 14-23 | 18-28 | 37-58 | 1.7-3.4 | 39-47 | 26-56 | 77-112 | S. Africa | Malan et al. (2014) |
| H. poinari $ | 1350-2800 | 54-105 | – | – | – | 108-112 | – | – | – | – | – | – | – | USA | Kakulia and Mikaia (1997) |
| H. ruandica Rw14_NC4a | 2907-4123 | 209-274 | 106-153 | 78-108 | 134-159 | 63-98 | 12-16 | 21-27 | 34-51 | 1.7-2.6 | 45-55 | 29-51 | 67–103 | Rwanda | This study |
| H. safricana | 3373-4073 | 127-188 | 210-267 | 121-163 | 199-236 | 64-91 | – | – | – | – | 43-46 | 40-54 | 98-119 | S. Africa | Malan et al. (2008) |
| H. taysearae | 2200-2800 | 116-170 | 137-182 | 83–120 | 161-200 | 72-100 | – | – | – | – | 40-64 | 41-67 | – | Egypt | Shamseldean et al. (1996) |
| as H. sonorensis # | 2856-5799 | 150-200 | 115-203 | 105-180 | 133-215 | 122-178 | – | – | – | 3.0* | 50-58 | 40-75 | - | Mexico | Stock et al. (2009) |
| H. zealandica | – | – | – | – | – | – | – | – | – | – | – | – | – | N. Zealand | Poinar (1990) |
| as H. heliothidis # | 4000* | 247* | – | 181* | 236 | 90* | 16* | 17* | 44* | 1.7* | 46* | 53* | – | N. Zealand | Wouts (1979) |
| H. zacatecana MEX-39 | 4408-6179 | 235-385 | 108-190 | 96-169 | 174-231 | 63-87 | 13-20 | 20-34 | 52-90 | 1.2-2.4 | 36-57 | 34-58 | 55-95 | Mexico | This study |
Table 5.
Comparative morphometrics of adult females of Heterorhabditis ruandica n. sp., H. zacatecana n. sp., and of different closely related Heterorhabditis species.
| Species | L | BD | EP | NR | NL | T | a | b | c | c´ | V | ABD | D% | Country | Reference |
|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
| H. amazonensis | 1279-2070 | 70-122 | 103-126 | 68-100 | 119-142 | 25-38 | – | – | – | 2.4* | 46-50 | 25-38 | – | Brazil | Andaló et al. (2006) |
| H. atacamensis | 1754-2628 | 86-129 | 154-182 | 79-119 | 129-167 | 80-108 | – | – | – | 3.8* | 43-49 | 24-33 | 100-113 | Chile | Edgington et al. (2011) |
| H. bacteriophora | 3180-3850 | 160-220 | 174-214 | 93-118 | 155-183 | 71-93 | 21.4* | 18.8 | 41.5* | 3.1* | 42-53 | 22-31 | 114 | Australia | Poinar (1976) |
| 1800-2400 | 100-162 | 122-162 | 83-102 | 108–145 | 40-65 | – | – | – | – | 41-50 | 23-40 | – | Argentina | Agüera de Doucet and Doucet (1986) | |
| 1690-3214 | 100-224 | 101-212 | 67-103 | 120-163 | 54-101 | – | – | – | 2.4* | 44-50 | 21-24 | 72-137 | Australia | Sagun et al. (2015) | |
| 1513-2290 | 84-150 | 128–181 | 71-99 | 113–135 | 41-79 | 11-22 | 11-19 | 26-42 | 1.6–2.5 | 38-51 | 24-39 | 108–150 | India | Bhat et al. (2019a) | |
| 1226-1819 | 58-115 | 108-157 | 68-91 | 101-127 | 29-94 | 16-25 | 7.8-16 | 16-48 | 1.0-3.4 | 44-58 | 24-31 | 83-116 | India | Rana et al. (2020) | |
| as H. argentinensis # | 2000-3500 | 90-180 | 105-240 | 88-140 | 162-200 | 75-108 | 12.5* | 7.8* | 31.2* | 2.0* | 42-48 | 33-35 | 100* | Argentina | Stock (1993) |
| as H. heliothidis £ | 2000-3300 | 184-240 | 146* | 126* | 148-177 | 71-93 | 11-15 | 14-21 | 26-46 | 2.8* | 48-53 | 33* | 95* | USA | Khan et al. (1976) |
| H. baujardi | 1335-2130 | 90-150 | 104-149 | 75-122 | 131-185 | 68-89 | 12-16 | 10-12 | 19-32 | – | 46-51 | 27-41 | – | Vietnam | Phan et al. (2003) |
| 2060-2290 | 120-150 | 98-115 | 80-95 | 123-148 | 78-108 | 15-17 | 16-18 | 20-27 | - | 41–48 | 30-38 | 63-78 | India | Vanlalhlimpuia et al. (2018) | |
| as H. somsookae # | 2159-2666 | 117-194 | 143-156 | 90-112 | 128-144 | 41-80 | – | – | – | 2.9* | 36-51 | 21-35 | 104-111 | Thailand | Maneesakorn et al. (2015) |
| H. beicherriana | 1581-3026 | 125-218 | 95-165 | 59-138 | 105-186 | 68-105 | 10-18 | 10–23 | 19–34 | 1.6-2.4 | 41-49 | 35–81 | 88-98 | China | Li et al. (2012) |
| H. downesi | 1231-2728 | 74–131 | 99-126 | 117-151 | 111–155 | 70-122 | – | – | – | 2.5* | 47-60 | 25-38 | – | Ireland | Stock et al. (2002) |
| H. egyptii + | 1050-1420 | 56-84 | 69-106 | 69-94 | 106-125 | 56-78 | 17.5** | 14.4** | 22.2** | 3.1** | 44-51 | 19-27 | 78** | Egypt | Abd–Elgawad and Ameen (2005) |
| H. floridensis | 2054-2548 | 120-156 | 110-168 | 86-122 | 126-178 | 69-87 | – | – | – | – | 44-50 | 32-42 | – | USA | Nguyen et al. (2006) |
| H. georgiana | 1640-2779 | 101-188 | 111-177 | 96–162 | 136-219 | 62-88 | – | – | – | 1.5* | 46-53 | 42* | – | USA | Nguyen et al. (2008) |
| H. hambletoni + | 600-1200 | 70-100 | 80-90 | 70-80 | – | – | – | – | – | – | 50-58** | – | – | Brazil | Pereira (1937) |
| H. hoptha $ | 2826-3983 | – | 148* | 161* | 219* | 28* | 13-19 | 12-21 | 47-67 | 0.8* | 43–49 | 33* | 92* | New Jersey | Turco (1970) |
| H. indica | 1200-1800 | 76-113 | 118-138 | 88–96 | 120-139 | 66-88 | – | – | – | – | 40-53 | 22-32 | – | India | Poinar et al. (1992) |
| 1713-2242 | 110-156 | 135-172 | 77-92 | 120-138 | 61-83 | 13-17 | 11-18 | 22-36 | 1.9-2.9 | 44-50 | 27-33 | 102-128 | India | Kajol et al. (2020) | |
| 1274-1993 | 70-135 | 105-129 | 84-111 | 124-155 | 64-83 | 12-18 | 10-13 | 16-31 | 2.6-4.9 | 45-52 | 22-30 | 77-99 | India | Bhat et al. (2021b) | |
| as H. brevicaudis # | 2100-2500 | 128-168 | 124-160 | 100-108 | 144-160 | 76-92 | – | – | – | – | 45-53 | 36-48 | – | China | Liu (1994) |
| as H. gerrardi # | 1428-2533 | 71-161 | 108-157 | 73-141 | 120-182 | 66-95 | – | – | – | 3.3* | 43-55 | 22-38 | 74-112 | Australia | Plichta et al. (2009) |
| as H. hawaiiensis # | 1300-2300 | 104-171 | 116-175 | 78-116 | 110-153 | 49-87 | – | – | – | – | 49-56 | 20-35 | – | USA | Gardner et al. (1994) |
| as H. pakistanense # | 1413-1785 | 71-86 | 130-150 | 80-100 | 130-145 | 65-95 | 19-21 | 11-12 | 16-22 | 3.1* | 44-53 | 24-27 | 95* | Pakistan | Shahina et al. (2017) |
| H. marelatus | 1600-2600 | 113-177 | 139-178 | 79-119 | 129–164 | 55-81 | – | – | – | 1.3* | 45-50 | 29-48 | 110* | USA | Liu and Berry (1996) |
| – | – | – | – | – | – | – | – | – | – | – | – | – | USA | Stock (1997) | |
| as H. hepialius # | 3500-4500 | 99-161 | 133-177 | 99-135 | 150-183 | 76–113 | 18* | 13* | 27* | 1.3* | 49-51 | 24-60 | 88* | USA | Stock et al. (1996) |
| H. megidis | 1500-2500 | 95-140 | 158-206 | 105-120 | 155-168 | 70-101 | 15-19 | 10-16 | 18-32 | 2.6* | 47-51 | 25-38 | 119* | USA | Poinar et al. (1987) |
| H. mexicana | 1144-2108 | 65-123 | 114-148 | 76-103 | 121-150 | 76-106 | – | – | – | – | 44-51 | 21-36 | – | Mexico | Nguyen et al. (2004) |
| H. noenieputensis | 1075-1697 | 76-129 | 102-125 | 73-90 | 115-132 | 63-75 | 13-17 | 9-14 | 17-24 | 2.3-3.1 | 40-53 | 22-32 | 83-104 | S. Africa | Malan et al. (2014) |
| H. poinari $ | 910-1520 | 62-80 | – | – | 152-172 | 86-105 | 11-14 | 50-51 | 10-11 | – | 38-50 | – | – | USA | Kakulia and Mikaia (1997) |
| H. ruandica Rw14_NC4a | 1131-1608 | 68-83 | 92-129 | 69-97 | 107-132 | 62-88 | 15-20 | 9.0-14 | 16-24 | 1.9–3.6 | 41-51 | 18-34 | 74-104 | Rwanda | This study |
| H. safricana | 1679-2937 | 102-229 | 151-196 | 87–139 | 148-180 | 55-111 | – | – | – | 1.3* | 45-50 | 25-72 | 97–120 | S. Africa | Malan et al. (2008) |
| H. taysearae | 830-1400 | 42-96 | 120-166 | 76-109 | 129-179 | 62-80 | – | – | – | 4.0* | 44-73 | 19-28 | 82* | Egypt | Shamseldean et al. (1996) |
| as H. sonorensis # | 1500-2500 | 85-210 | 95-140 | 85-105 | 129-215 | 75-99 | – | – | – | 3.5* | 49-53 | 36-46 | 93* | Mexico | Stock et al. (2009) |
| H. zealandica | – | – | – | – | – | – | – | – | – | – | – | – | – | N. Zealand | Poinar (1990) |
| as H. heliothidis # | – | – | – | – | – | – | – | – | – | – | – | – | – | N. Zealand | Wouts (1979) |
| H. zacatecana MEX-39 | 1954-2798 | 160-228 | 100-133 | 71-96 | 112-148 | 45-75 | 11–15 | 16–21 | 31-63 | 1.3–2.0 | 43-61 | 31–41 | 80-111 | Mexico | This study |
Table 6.
Comparative morphometrics of infective juveniles of Heterorhabditis ruandica n. sp., H. zacatecana n. sp., and of different closely related Heterorhabditis species.
| Species | L | BD | EP | NR | NL | T | a | b | C | c´ | D% | E% | Country | Reference |
|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
| H. amazonensis | 567-612 | 20-24 | 89-115 | 76-93 | 107-132 | 98-115 | 24-29 | 4.4-5.5 | 5.1-6.1 | 7.3* | 83-92 | 89–109 | Brazil | Andaló et al. (2006) |
| H. atacamensis | 578-666 | 19-26 | 101-126 | 79-101 | 124-144 | 94-107 | 25-31 | 4.8-5.7 | 5.7-7.1 | 5.7* | 79-94 | 149-182 | Chile | Edgington et al. (2011) |
| H. bacteriophora | 512-671 | 18-31 | 87-110 | 72-93 | 100-139 | 83–112 | 17-30 | 4.0-5.1 | 5.7-7.0 | 6.0* | 76-92 | 103-130 | Australia | Poinar (1976) |
| 530-660 | 22-30 | 93-108 | 80-90 | 110-130 | 84-105 | 23* | 4.4* | 5.7* | 4.8* | 81* | 106* | Argentina | Agüera de Doucet and Doucet (1986) | |
| 537-587 | 19-22 | 87-104 | 67-83 | 112-121 | 94-111 | 25-31 | 3.3-3.6 | 3.6-4.6 | 7.0* | 73-88 | 87-105 | Australia | Sagun et al. (2015) | |
| 474-568 | 22-28 | 110-127 | 61-90 | 90-115 | 57-90 | 19-25 | 4.7-6.1 | 5.5-9.3 | 3.4-7.5 | 105-139 | 131-211 | India | Bhat et al. (2019a) | |
| 453-167 | 19-27 | 72-102 | 50-74 | 83-106 | 47-89 | 19-29 | 4.9-7.4 | 6.0-12 | 3.7-6.5 | 78–107 | 105-189 | India | Rana et al. (2020) | |
| as H. argentinensis # | 610-710 | 24-38 | 68-112 | 82-116 | 101-150 | 70-105 | 18.3* | 3.7* | 6.5* | 4.3* | 80* | 141* | Argentina | Stock (1993) |
| as H. heliothidis £ | 619-671 | 23-29 | 112* | 108* | 130-139 | 104-112 | 22-28 | 4.6-5.4 | 5.8-6.3 | 6.0* | 83* | 97* | USA | Khan et al. (1976) |
| H. baujardi | 497-595 | 18-22 | 91–103 | 75-86 | 107-120 | 83-97 | 26-30 | 4.5-5.1 | 6-6.7 | 7.2* | 78-88 | 98–114 | Vietnam | Phan et al. (2003) |
| 525-615 | 18-25 | 88-96 | 68-85 | 98-120 | 95-108 | 24-32 | 4.6-5.9 | 5.2-6.1 | – | 74-86 | 89-92 | India | Vanlalhlimpuia et al. (2018) | |
| as H. somsookae # | 502-565 | 19-23 | 81-95 | 78-94 | 106-117 | 91-131 | 23-27 | 5-5 | 4-6 | 8.0* | 76-87 | 64–95 | Thailand | Maneesakorn et al. (2015) |
| H. beicherriana | 566-687 | 21-25 | 100-122 | 85-106 | 118-146 | 86-111 | 24–29 | 4.2-4.9 | 5.9-6.8 | 6.0-7.4 | 80-93 | 103–121 | China | Li et al. (2012) |
| H. downesi | 588-692 | 15-22 | 96-128 | 96-105 | 126-141 | 62-74 | 29-42 | 4.4-5.3 | 8.5-10.5 | 4.4* | 76-96 | 160-180 | Ireland | Stock et al., 2002 |
| H. egyptii + | 484-515 | 18-23 | 81-94 | 78-100 | 100-119 | 53-75 | 20-27 | 4.2-5.2 | 6.8-9.1 | 6.9* | 74-82 | 100-170 | Egypt | Abd-Elgawad and Ameen (2005) |
| H. floridensis | 554-609 | 19-23 | 101-122 | 68-107 | 123-142 | 91-113 | 25-32 | 3.9-4.9 | 5.3-6.6 | 7.2* | 71–90 | 95–134 | USA | Nguyen et al. (2006) |
| H. georgiana | 547-651 | 17-26 | 97-113 | 74-94 | 110-139 | 86-108 | 23–34 | 4.1-5.3 | 5.5-6.9 | 6.8* | 70-93 | 106 | USA | Nguyen et al. (2008) |
| H. hambletoni + | – | – | – | – | – | – | – | – | – | – | – | – | Brazil | Pereira (1937) |
| H. hoptha $ | – | – | – | – | – | – | – | – | – | – | – | – | USA | Turco (1970) |
| H. indica | 479-573 | 19-22 | 88-107 | 72-85 | 109-123 | 93-109 | 25-27 | 4.3-4.8 | 4.5-5.6 | – | 79-90 | 83-103 | India | Poinar et al. (1992) |
| 511-546 | 21-24 | 92-108 | 63–73 | 86-103 | 24–34 | 22-25 | 5-6 | 4.6-5.4 | 2.8-5.2 | 77-96 | 100-118 | India | Kajol et al. (2020) | |
| 516-598 | 21-25 | 98-123 | 82-101 | 102-129 | 80-112 | 24-27 | 4.5-5.4 | 4.9-5.7 | 5.6-8.1 | 83-97 | 93–136 | India | Bhat et al. (2021b) | |
| as H. brevicaudis # | 528-632 | 20-24 | 104-116 | 96-104 | 120-136 | 68-80 | – | – | 6.6-8.6 | 6.3* | 81* | 150-180 | China | Liu (1994) |
| as H. gerrardi # | 551-682 | 18-29 | 92-111 | 81-105 | 110-130 | 76-141 | 23-32 | 16-23 | 11-21 | 6.8* | 73-92 | 73-138 | Australia | Plichta et al.(2009) |
| as H. hawaiiensis # | 506-631 | 21-28 | 116-175 | 79-103 | 115-181 | 82-108 | – | – | – | 6.0* | 77* | 88* | USA | Gardner et al. (1994) |
| as H. pakistanense # | 558-624 | 19-23 | 95-106 | 73-90 | 113-125 | 95-110 | 25-29 | 4.7-5.3 | 5.4-6.2 | 5.4* | 78-97 | 95-107 | Pakistan | Shahina et al. (2017) |
| H. marelatus | 588-700 | 24-32 | 81-113 | 83-113 | 121-139 | 99-117 | 21-29 | 4.7-5.4 | 5.5-6.6 | 3.0* | 60-86 | 89-110 | USA | Liu and Berry (1996) |
| 567-780 | 16-24 | 88-94 | 71-88 | 110-119 | 50-71 | 31-35 | 5.1-6.5 | 10.8-11.5 | – | 73-88 | 130-142 | USA | Stock (1997) | |
| as H. hepialius # | 540-600 | 34-39 | 84-112 | 80-101 | 106-130 | 49-60 | 5-7 | 4-5 | 9–12 | – | 79-98 | 100-200 | USA | Stock et al. (1996) |
| H. megidis | 736-800 | 27-32 | 123-142 | 104-115 | 147-160 | 112–128 | 23-38 | 4.6-5.9 | 6.1-6.9 | 6.3* | 81–91 | 103-120 | USA | (Poinar et al., 1987 |
| H. mexicana | 530-620 | 20-24 | 83-109 | 74-88 | 104-142 | 91-106 | 24-28 | 4.2-5.1 | 5.5-6.3 | 8.3* | 72-86 | 87-111 | Mexico | Nguyen et al. (2004) |
| H. noenieputensis | 484-578 | 21-25 | 88-105 | 69-96 | 79-115 | 78-95 | 21-27 | 4.3-5.2 | 5.5-6.8 | 3.4-4.3 | 81-95 | 99-125 | S. Africa | Malan et al. (2014) |
| H. poinari $ | 350–410 | 18-22 | – | – | – | 15-22 | – | – | – | – | – | – | USA | Kakulia and Mikaia (1997) |
| H. ruandica Rw14_NC4a | 496-591 | 18-27 | 70-89 | 52-64 | 103-131 | 49-64 | 20-29 | 4.1-5.4 | 7.6-8.6 | 3.4-5.8 | 66-98 | 112-168 | Rwanda | This study |
| H. safricana | 550-676 | 19-23 | 103-122 | 86-101 | 125-141 | 86-108 | 25-32 | 3.9-4.9 | 5.4-7.5 | 8.7* | 80-90 | 99–133 | S. Africa | Malan et al. (2008) |
| H. taysearae | 332-499 | 17-23 | 74-113 | 58-87 | 96–130 | 44-70 | 18-27 | 3.4-4.2 | 6.5-8.7 | 3.7* | 71-96 | 110-230 | Egypt | Shamseldean et al. (1996) |
| as H. sonorensis # | 495-570 | 19-32 | 97-116 | 87-98 | 110-131 | 91-125 | 19-26 | 4.4-5.4 | 4.0-6.5 | 6.7* | 78-110 | 81-111 | Mexico | Stock et al. (2009) |
| H. zealandica | 570-740 | 22-30 | 94-123 | 90-107 | 135-147 | 87-119 | 25 | 4.9 | 6.7 | – | 73-92 | 103-109 | N. Zealand | Poinar (1990) |
| as H. heliothidis # | 570-740 | 22-30 | 94-123 | 90-107 | 135-147 | 87-119 | 25 | 4.9 | 6.7 | – | 73-92 | 103-109 | N. Zealand | Wouts (1979) |
| H. zacatecana MEX-39 | 493-578 | 23-27 | 72-99 | 69-72 | 96-124 | 52-63 | 19-24 | 4.4-5.9 | 8.2-10.5 | 4.3–6.7 | 68-120 | 128-184 | Mexico | This study |
Figure 5:
Line drawings of Heterorhabditis zacatecana n. sp. (A) A hermaphroditic female. (B) Pharyngeal region of a hermaphroditic female. (C) Anterior part of the reproductive system of a hermaphroditic female. (D) Posterior end of a hermaphroditic female. (E) An amphimictic female. (F) Pharyngeal region of an amphimictic female. (G) Posterior end of an amphimictic female. (H) A male adult. (I) Pharyngeal region of a male adult. (J) Posterior end of a male adult. (K) Pharyngeal region of an infective juvenile. (L) Posterior end of an infective juvenile. (M) An infective juvenile.
Figure 6:
Light microscope micrographs of Heterorhabditis zacatecana n. sp. (A) An amphimictic female (black arrow pointing the vulva, white arrow pointing the anus). (B) Pharyngeal region of an amphimictic female. (C) Posterior end of an amphimictic female. (D) A male adult. (E) Pharyngeal region of a male adult. (F) Posterior end of a male adult (arrows pointing at the genital papillae).
Figure 7:
Light microscope micrographs of Heterorhabditis zacatecana n. sp. (A) A hermaphroditic female. (B) Pharyngeal region of a hermaphroditic female. (C) Posterior end of a hermaphroditic female. (D) A sheathed third stage juvenile (J2). (E) Pharyngeal region of a sheathed third stage juvenile (J2). (F) Posterior end of a sheathed third stage juvenile (J3). (G) A non-sheathed third stage juvenile (J3). (H) Pharyngeal region of a non-sheathed third stage juvenile (J3). (I) Posterior end of a non-sheathed third stage juvenile.
Figure 8:
Scanning electron microscope (SEM) micrographs of Heterorhabditis zacatecana n. sp. (A, B) Lip region in lateral and frontal views, respectively, of a hermaphroditic female. (C) Broken cuticle of a hermaphroditic female with a juvenile emerging. (D) Vulva of a hermaphroditic female (pointed by a white arrow). (E) Tail of a hermaphroditic female in lateral view. (F, G) Lip region of a female adult in lateral and frontal views, respectively. (H) Excretory pore of a female adult (pointed by a white arrow). (I) Vulva of a female adult. (J) Tail of a female adult in ventral view. (K, L) Lip region of a male adult in lateral and frontal views, respectively. (M, N) Posterior end of a male adult in lateral and ventral views, respectively (arrows pointing at the genital papillae). (O) Lip region of a second-stage juvenile (J2) in lateral view. (P) Cuticle of a second-stage juvenile (J2) (arrow pointing the excretory pore). (Q) Tail of a second-stage juvenile (J2) in lateral and ventral views, respectively. (R) Lip region of a third-stage juvenile (J3) in dorsal view (arrow pointing the frontal tooth). (S) Cuticle of a third-stage juvenile (J3) (arrow pointing the excretory pore).
Figure 9:
Maximum-likelihood phylogenetic tree reconstructed from: (A) the sequences of the cytochrome c oxidase I (COI) of different Heterorhabditis species. A total of 343 nucleotide positions, flanked by primers HCF and HCR, were analyzed; and (B) the concatenated sequences of the following genes/genetic regions of different Heterorhabditis species: the D2–D3 expansion segments of the 28S rRNA (D2–D3), the internal transcribed spacer (ITS) of the rRNA (ITS), and the cytochrome c oxidase I (COI). A total of 1673 concatenated nucleotide positions were included in the reconstruction. Accession numbers of the nucleotide sequences used for the analyses are shown in Table S3. *For H. marelatus, H. indica, and H. mexicana, the sequences that were concatenated are derived from different nematode isolates. Heterorhabditis safricana, and H. tayserae were not included as their COI or their D2–D3 sequences, respectively, are not publicly available. Numbers at nodes represent bootstrap values based on 100 replications. Bars represent average nucleotide substitutions per sequence position.
Figure 10:
Polymorphism in the sequences of the ITS region (A, B), the D2–D3 region (C), and the COI gene (D-F) showing taxonomically relevant nucleotide positions for Heterorhabditis nematodes of the “Bacteriophora-group”. Nucleotide position numbers of rRNA genes are according to the sequences of C. elegans N2 (NCBI accession number: MN519140) and of mitochondrial genes are according to the sequences of C. elegans N2 (NCBI accession number: AY171203).
Figure 11:
Pairwise nucleotide similarities (%) in the sequences of the cytochrome c oxidase I (COI) gene of different Heterorhabditis species. A total of 344 nucleotide positions, flanked by primers HCF and HCR, were analyzed. Accession numbers of gene sequences used are shown in Table S3.
Figure 12:
Phylogenetic reconstruction based on core genome sequences of Photorhabdus bacterial strains. Numbers at the nodes represent SH-like branch supports. Bar represents average nucleotide substitutions per sequence position. Accession numbers of the genome sequences used for the reconstruction are shown in Table S4.
Figure 13:
Dichotomous key to identify the species of the genus Heterorhabditis based on morphological and morphometrical characters of L3 juveniles, of male and female adults, and of hermaphroditic females.