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Steinernema africanum n. sp. (Rhabditida, Steinernematidae), a New Entomopathogenic Nematode Species Isolated in the Republic of Rwanda

Ricardo A. R. Machado
Ricardo A. R. Machado
, 
Aashaq Hussain Bhat
Aashaq Hussain Bhat
, 
Joaquín Abolafia
Joaquín Abolafia
, 
Ebrahim Shokoohi
Ebrahim Shokoohi
, 
Patrick Fallet
Patrick Fallet
, 
Ted C. J. Turlings
Ted C. J. Turlings
, 
Eustachio Tarasco
Eustachio Tarasco
, 
Vladimír Půža
Vladimír Půža
, 
Joelle Kajuga
Joelle Kajuga
, 
Xun Yan
Xun Yan
 et 
Stefan Toepfer
Stefan Toepfer
   | 26 oct. 2022
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Article Category: Research Paper
Publié en ligne: 26 oct. 2022
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Reçu: 07 juin 2022
DOI: https://doi.org/10.2478/jofnem-2022-0049
Mots clés
© 2022 Machado et al. published by Sciendo
This work is licensed under the Creative Commons Attribution 4.0 International License.

Figure 1

Line drawings of Steinernema africanum n. sp. IJ. (A) Stoma and pharynx region. (B) Anterior end. (C) Entire IJ. (D) Posterior end. IJ, infective juvenile.
Line drawings of Steinernema africanum n. sp. IJ. (A) Stoma and pharynx region. (B) Anterior end. (C) Entire IJ. (D) Posterior end. IJ, infective juvenile.

Figure 2

Line drawings of first- and second-generation Steinernema africanum n. sp. females. (A–E) First-generation female: (A) Stoma and pharynx region; (B) Lip region and stoma; (C) Vagina region; (D) Posterior end; (E) Entire female. (F–I) Second-generation female: (F) Stoma and pharynx region; (G) Lip region and stoma; (H) Vagina region; (I) Posterior end.
Line drawings of first- and second-generation Steinernema africanum n. sp. females. (A–E) First-generation female: (A) Stoma and pharynx region; (B) Lip region and stoma; (C) Vagina region; (D) Posterior end; (E) Entire female. (F–I) Second-generation female: (F) Stoma and pharynx region; (G) Lip region and stoma; (H) Vagina region; (I) Posterior end.

Figure 3

Line drawings of first- and second-generation Steinernema africanum n. sp. males. (A–D) First-generation male: (A) Stoma and pharynx region; (B) Lip region and stoma; (C) Entire male; (D) Posterior end. (E–H) Second-generation male: (E) Stoma and pharynx region; (F) Lip region and stoma; (G) Entire male; (H) Posterior end.
Line drawings of first- and second-generation Steinernema africanum n. sp. males. (A–D) First-generation male: (A) Stoma and pharynx region; (B) Lip region and stoma; (C) Entire male; (D) Posterior end. (E–H) Second-generation male: (E) Stoma and pharynx region; (F) Lip region and stoma; (G) Entire male; (H) Posterior end.

Figure 4

Light microscope micrographs of first-generation adults and IJ of Steinernema africanum n. sp. (A, B) Stoma and pharynx region of female and male, respectively. (C–E) Lip region and stoma of female, male and IJ, respectively. (F) Vagina region. (G–I) Entire female, male and IJ, respectively. (J) Posterior end of a female. (K, L) Posterior end of males. Pictures are in right lateral view. IJ, infective juvenile.
Light microscope micrographs of first-generation adults and IJ of Steinernema africanum n. sp. (A, B) Stoma and pharynx region of female and male, respectively. (C–E) Lip region and stoma of female, male and IJ, respectively. (F) Vagina region. (G–I) Entire female, male and IJ, respectively. (J) Posterior end of a female. (K, L) Posterior end of males. Pictures are in right lateral view. IJ, infective juvenile.

Figure 5

Light microscope micrographs of second-generation adults and IJ of Steinernema africanum n. sp. (A–C) Stoma and pharynx region of female, male and IJ, respectively. Black arrow pointing the EP, white arrow pointing the bacteria sac. (D) Vagina region. (E, F) Entire female and male, respectively. (G, H) IJ bacterial sac and lateral field, respectively. (I–K) Posterior end of female, male, and IJ, respectively. Pictures are in right lateral view. EP, excretory pore; IJ, infective juvenile.
Light microscope micrographs of second-generation adults and IJ of Steinernema africanum n. sp. (A–C) Stoma and pharynx region of female, male and IJ, respectively. Black arrow pointing the EP, white arrow pointing the bacteria sac. (D) Vagina region. (E, F) Entire female and male, respectively. (G, H) IJ bacterial sac and lateral field, respectively. (I–K) Posterior end of female, male, and IJ, respectively. Pictures are in right lateral view. EP, excretory pore; IJ, infective juvenile.

Figure 6

Scanning electron microscope micrographs of first-generation adults of Steinernema africanum n. sp. (A, B) Lip region of a female and a male, respectively, in lateral view. Arrows pointing the amphids (C: cheilostom, G: gymnostom, S: stegostom, cp: cephalic papillae, and lp: labial papillae). (C) Male posterior end in sub-ventral view – GP, MP, ph. (D, E) Lip region of a female and a male, respectively, in frontal view. (F, G) EP of a female and a male, respectively. (H) Vulva. (I) Male posterior end in sub-ventral view – GP, MP, ph. (J) Male posterior end in sub-ventral view – GP, MP. (K) Female anus. Arrow pointing a post-anal pore. (L, M) Female tail in lateral and ventral views, respectively. Black arrow pointing a post-anal pore, white arrows pointing the phasmids. EP, excretory pore; GP, genital papillae; MP, mid-ventral papilla; ph, phasmid.
Scanning electron microscope micrographs of first-generation adults of Steinernema africanum n. sp. (A, B) Lip region of a female and a male, respectively, in lateral view. Arrows pointing the amphids (C: cheilostom, G: gymnostom, S: stegostom, cp: cephalic papillae, and lp: labial papillae). (C) Male posterior end in sub-ventral view – GP, MP, ph. (D, E) Lip region of a female and a male, respectively, in frontal view. (F, G) EP of a female and a male, respectively. (H) Vulva. (I) Male posterior end in sub-ventral view – GP, MP, ph. (J) Male posterior end in sub-ventral view – GP, MP. (K) Female anus. Arrow pointing a post-anal pore. (L, M) Female tail in lateral and ventral views, respectively. Black arrow pointing a post-anal pore, white arrows pointing the phasmids. EP, excretory pore; GP, genital papillae; MP, mid-ventral papilla; ph, phasmid.

Figure 7

Scanning electron microscope micrographs of second-generation adults and IJ of Steinernema africanum n. sp. (A–C) Lip region of female, male, and IJ in lateral view, respectively. Arrows pointing the amphids (C: cheilostom, G: gymnostom, S: stegostom, cp: cephalic papillae, lp: labial papillae). (D–F) Lip region of female, male, and IJ in frontal view, respectively. Black arrows point to the amphids, white arrow points to the dorsal gland opening. (G) EP of female. (H) Male posterior region in lateral view – GP, MP, ph. (I) EP of IJ. (J) Vulva. (K) IJ lateral field. (L, M) Female tail (arrows pointing the phasmids). (N) Male posterior region in lateral view. (O, P) IJ tail in lateral and ventral views, respectively. Arrows pointing the phasmids. EP, excretory pore; GP, genital papillae; MP, mid-ventral papilla; ph, phasmid.
Scanning electron microscope micrographs of second-generation adults and IJ of Steinernema africanum n. sp. (A–C) Lip region of female, male, and IJ in lateral view, respectively. Arrows pointing the amphids (C: cheilostom, G: gymnostom, S: stegostom, cp: cephalic papillae, lp: labial papillae). (D–F) Lip region of female, male, and IJ in frontal view, respectively. Black arrows point to the amphids, white arrow points to the dorsal gland opening. (G) EP of female. (H) Male posterior region in lateral view – GP, MP, ph. (I) EP of IJ. (J) Vulva. (K) IJ lateral field. (L, M) Female tail (arrows pointing the phasmids). (N) Male posterior region in lateral view. (O, P) IJ tail in lateral and ventral views, respectively. Arrows pointing the phasmids. EP, excretory pore; GP, genital papillae; MP, mid-ventral papilla; ph, phasmid.

Figure 8

Phylogenetic relationships between the newly described Steinernema africanum n. sp. and other Steinernema species. Maximum-likelihood phylogenetic tree reconstructed from: (A) the nucleotide sequences of the ITS rRNA. A total of 705 nucleotide positions, flanked by primers 18S and 26S, were analyzed; (B) the nucleotide sequences of the D2–D3 expansion segments of the 28S rRNA. A total of 786 nucleotide positions, flanked by primers D2F and 536, were analyzed; (C) the nucleotide sequences of the COI gene. A total of 567 nucleotide positions, flanked by primers LCO-1490 and HCO-2198, were analyzed; and (D) the nucleotide sequences of the mitochondrial 12S rRNA gene. A total of 467 nucleotide positions, flanked by primers 505F and 506R, were analyzed. Numbers at nodes represent bootstrap values based on 100 replications. Bars represent average nucleotide substitutions per sequence position. NCBI accession numbers of the nucleotide sequences used for the analyses are shown next to the species names. COI, cytochrome oxidase subunit I; ITS, internal transcribed spacer; NCBI, National Center for Biotechnology Information.
Phylogenetic relationships between the newly described Steinernema africanum n. sp. and other Steinernema species. Maximum-likelihood phylogenetic tree reconstructed from: (A) the nucleotide sequences of the ITS rRNA. A total of 705 nucleotide positions, flanked by primers 18S and 26S, were analyzed; (B) the nucleotide sequences of the D2–D3 expansion segments of the 28S rRNA. A total of 786 nucleotide positions, flanked by primers D2F and 536, were analyzed; (C) the nucleotide sequences of the COI gene. A total of 567 nucleotide positions, flanked by primers LCO-1490 and HCO-2198, were analyzed; and (D) the nucleotide sequences of the mitochondrial 12S rRNA gene. A total of 467 nucleotide positions, flanked by primers 505F and 506R, were analyzed. Numbers at nodes represent bootstrap values based on 100 replications. Bars represent average nucleotide substitutions per sequence position. NCBI accession numbers of the nucleotide sequences used for the analyses are shown next to the species names. COI, cytochrome oxidase subunit I; ITS, internal transcribed spacer; NCBI, National Center for Biotechnology Information.

Figure 9

Phylogenetic relationships between the Xenorhabdus symbiont isolated from Steinernema africanum n. sp. and other Xenorhabdus species. Phylogenetic trees were built based on core genome sequences. A total of 1,719,910 nucleotide positions were used in the analyses. Numbers at the nodes represent SH-like branch supports. Bar represents 0.05 nucleotide substitutions per sequence position. Accession numbers of the genome sequences used for the reconstruction are shown in Table S1 in Supplementary Material.
Phylogenetic relationships between the Xenorhabdus symbiont isolated from Steinernema africanum n. sp. and other Xenorhabdus species. Phylogenetic trees were built based on core genome sequences. A total of 1,719,910 nucleotide positions were used in the analyses. Numbers at the nodes represent SH-like branch supports. Bar represents 0.05 nucleotide substitutions per sequence position. Accession numbers of the genome sequences used for the reconstruction are shown in Table S1 in Supplementary Material.

Figure S1

Pairwise comparisons of the ITS nucleotide sequences. (A) Sequence similarities (%). (B) Number of nucleotide differences (bp). A total of 808 nucleotide positions, flanked by primers 18S and 26S, were analyzed. NCBI accession numbers of the nucleotide sequences used for the analyses are shown next to the species names. ITS, internal transcribed spacer; NCBI, National Center for Biotechnology Information.
Pairwise comparisons of the ITS nucleotide sequences. (A) Sequence similarities (%). (B) Number of nucleotide differences (bp). A total of 808 nucleotide positions, flanked by primers 18S and 26S, were analyzed. NCBI accession numbers of the nucleotide sequences used for the analyses are shown next to the species names. ITS, internal transcribed spacer; NCBI, National Center for Biotechnology Information.

Figure S2

Pairwise comparisons of the nucleotide sequences of the D2–D3 expansion segments of the 28S rRNA. (A) Sequence similarities (%). (B) Number of nucleotide differences (bp). A total of 786 nucleotide positions, flanked by primers D2F and 536, were analyzed. NCBI accession numbers of the nucleotide sequences used for the analyses are shown next to the species names. NCBI, National Center for Biotechnology Information.
Pairwise comparisons of the nucleotide sequences of the D2–D3 expansion segments of the 28S rRNA. (A) Sequence similarities (%). (B) Number of nucleotide differences (bp). A total of 786 nucleotide positions, flanked by primers D2F and 536, were analyzed. NCBI accession numbers of the nucleotide sequences used for the analyses are shown next to the species names. NCBI, National Center for Biotechnology Information.

Figure S3

Pairwise comparison of dDDH scores (%) of Xenorhabdus strains. A total of 1,719,910 nucleotide positions were used in the analyses. Accession numbers of gene sequences used are shown in Table S1 in Supplementary Material. dDDH, digital DNA–DNA Hybridization.
Pairwise comparison of dDDH scores (%) of Xenorhabdus strains. A total of 1,719,910 nucleotide positions were used in the analyses. Accession numbers of gene sequences used are shown in Table S1 in Supplementary Material. dDDH, digital DNA–DNA Hybridization.

Morphometrics of IJs and adult generations of Steinernema africanum n. sp.

Characters Male first generation Male second generation Female first generation Female second generation Infective juvenile
Specimen type Holotype Paratype Paratype Paratype Paratype Paratype
n 1 15 15 15 15 15
Body length (L) 1,300 1,202.2 ± 141.4 (977‒1,400) 955.4 ±92.8 (831‒1,213) 3,319.1 ± 763.2(2,469‒5,033) 1,620.0 ±487.6 (924‒2,311) 750.9 ± 39.8 (690‒802)
a (L/BD) 9.9 10.3 ±0.9 (9.1‒11.6) 17.0 ±1.6 (14.3‒19.0) 18.0 ±3.9 (13.2‒27.0) 13.5 ±1.0 (12.1‒15.7) 28.0 ± 2.4 (23.2‒30.4)
b (L/NL) 8.8 8.4 ± 0.8 (7.2‒9.6) 7.3 ± 0.6 (6.6‒8.5) 17.4 ±3.7 (12.8‒23.9) 10.1 ±2.1 (7.2‒12.9) 4.9 ± 0.5 (4.3‒6.3)
c (L/TL) 28.3 29.0 ± 3.3 (24.5‒33.9) 29.7 ± 2.9 (25.8‒35.7) 74.1 ± 17.6 (50.6‒103.9) 33.5 ±11.7 (16.5‒50.9) 11.9 ± 1.3(10.3‒14.9)
c' (TL/ABW) 0.9 1.0 ±0.1 (0.9‒1.1) 1.0 ±0.1 (0.7‒1.1) 0.9 ±0.1 (0.7‒1.0) 1.5 ±0.3(1.2‒2.3) 3.7 ± 0.4 (2.9‒4.2)
T(MR/L × 100), 94 87.3 ± 11.8 (65‒99) 71.6 ± 13.4 (58‒96) 52.0 ± 2.7 (50‒57) 56.0 ± 1.9 (53‒58) ‒
V (VA/L × 100)
Lip region diameter 21 19.5 ±3.2(13‒25) 15.1 ±1.6 (11‒17) 24.9 ±3.3(19‒31) 22.3 ±6.5 (14‒34) 6.1 ± 0.7 (5‒7)
Stoma length 9 7.5 ±2.1 (5‒11) 6.7 ± 1.2 (5‒9) 12.8 ± 2.9 (8‒19) 11.1 ± 2.4 (8‒16) 17.6 ± 4.4 (10‒23)
Corpus length 84 83.5 ± 5.2 (72‒90) 69.9 ± 3.3 (65‒76) 91.9 ± 8.1 (77‒108) 83.0 ± 14.0 (62‒101) 85.5 ± 8.1 (72‒97)
Isthmus length 26 20.8 ± 3.3 (14‒26) 22.4 ± 2.7 (18‒28) 27.6 ± 5.6 (21‒40) 23.8 ± 7.1 (11‒35) 27.6 ± 5.4 (19‒37)
Bulb length 28 29.2 ± 3.7 (25‒38) 26.9 ± 2.6 (22‒32) 45.1 ± 6.6 (37‒58) 33.3 ± 4.7 (24‒42) 22.8 ± 2.0 (18‒26)
Cardia length 11 8.7 ±1.5 (6‒11) 5.8 ±1.8 (4‒11) 11.5 ± 3.1 (8‒19) 7.7 ± 2.7 (4‒12) ‒
NR to anterior 95 91.5 ± 8.1 (79‒104) 83.5 ± 7.6 (74‒101) 105.4 ± 14.9 (79‒130) 96.4 ± 19.8 (69‒124) 117.7 ± 11.5 (87‒132)
EP to anterior 105 93.4 ± 12.4 (69‒109) 91.2 ± 6.7 (83‒104) 89.4 ± 14.3 (67‒111) 83.6 ± 11.6 (61‒104) 59.3 ± 4.2 (54‒68)
PL 138 134.7 ± 5.4 (123‒142) 123.5 ± 5.6 (117‒135) 169.6 ± 8.8 (158‒188) 140.9 ± 19.7 (113‒167) 137.2 ± 9.2 (113‒153)
NL(Stoma+PL) 147 142.5 ± 4.4 (132‒147) 131.1 ±5.6 (124‒143) 185.3 ± 7.9 (170‒201) 152.6 ± 21.6 (122‒181) 154.4 ± 12.0 (123‒167)
BD at neck base 73 65.7 ± 10.8 (41‒76) 44.7 ± 11.4 (37‒73) 124.4 ± 9.3 (113‒153) 77.5 ± 17.4 (47‒105) 26.1 ±1.6(21‒29)
BD at midbody 131 114.8 ± 17.9 (65‒131) 54.1 ± 5.2 (39‒59) 181.0 ± 14.8 (154‒208) 114.7 ± 37.3 (59‒167) 26.9 ± 1.6 (25‒32)
Anal BD (ABW) 50 41.9 ± 5.5 (33‒50) 34.8 ± 5.7 (30‒48) 52.4 ± 8.2 (37‒70) 33.9 ± 5.8 (26‒43) 17.2 ± 1.1 (16‒20)
Vagina length ‒ ‒ ‒ 20.8 ± 6.3 (11‒33) 24.2 ± 11.9 (10‒49) ‒
VA end ‒ ‒ ‒ 1,767.1 ± 445.4 (1,284‒2,411) 905.9 ± 266.0 (516‒1,217) ‒
Rectum length (RL) ‒ ‒ ‒ 46.4 ± 8.8 (33‒63) 37.3 ± 14.3 (20‒64) 28.3 ± 3.9 (23‒38)
Tail length (TL) 46 41.1 ± 4.0 (34‒46) 33.9 ± 4.8 (28‒46) 44.6 ± 5.9 (35‒55) 49.7 ±5.7 (40‒60) 64.0 ± 7.1 (52‒72)
Tail hyaline length (H) ‒ ‒ ‒ 7.4 ± 2.6 (4‒11) 12.2 ± 3.8 (6‒19) 22.5 ± 3.7 (17‒27)
SL 72 71.1 ± 3.6 (65‒76) 59.5 ± 4.8 (53‒68) ‒ ‒ ‒
GL 42 42.5 ± 4.2 (32‒49) 32.6 ± 5.7 (25‒46) ‒ ‒ ‒
Stoma length/lip region width 0.4 0.4 ± 0.1 (0.2‒0.7) 0.4 ± 0.1 (0.3‒0.5) 0.5 ± 0.1 (0.3‒0.6) 0.5 ± 0.1 (0.3‒0.7) 2.9 ± 0.6 (1.5‒3.7)
NR% (NR/NL × 100) 65 63.9 ± 4.9 (56‒71) 63.7 ± 3.5 (60‒71) 56.9 ± 7.9 (43‒65) 63.2 ± 8.5 (43‒69) 76.3 ± 5.8 (69‒90)
D % (EP/NL × 100) 71 65.1 ± 7.2 (52‒74) 69.6 ± 2.7 (67‒75) 48.3 ± 8.4 (36‒62) 55.5 ± 9.7 (49‒79) 38.1 ± 2.7 (34‒46)
E% (EP/TL × 100) 228 224.6 ± 22.7 (197‒261) 284.3 ± 34.5 (243‒345) 204.8 ± 44.9 (132‒281) 170.9 ± 33.4 (122‒221) 94.7 ± 17.4 (79‒129)
Rectum % (RL/ABD × 100) ‒ ‒ ‒ 90.4 ± 19.7 (58‒112) 108.1 ±29.3 (74‒163) 164.3 ± 17.1 (128‒211)
H % (H/TL × 100) ‒ ‒ ‒ 15.7 ± 5.1 (8‒20) 21.6 ± 11.7 (12‒40) 35.2 ± 3.4 (28‒39)
SW% (SL/ABD × 100) 144.0 172.3 ± 21.3 (144‒197) 177.2 ± 19.8(142‒217) ‒ ‒ ‒
GS % (GL/SL × 100) 58 59.9 ± 5.9 (49‒68) 54.7 ± 7.1 (46‒68) ‒ ‒ ‒
Male reproductive system (MR) 1,218 1,044.3 ± 240.9 (639‒1,371) 692.9 ± 172.4 (553‒1,166)
Testis reflexion 281 280.2 ± 66.3 (116‒352) 159.7 ± 64.6 (89‒273) ‒ ‒ -
Mucron 5.1 4.8 ± 0.7 (4.0-5.3) 7.3 ± 2.5 (5-10) 4.4 ± 0.7 (3.7-5.0) Absent -

National Center for Biotechnology Information (NCBI) accession numbers of the sequences used in this study.

Strain Genome
Xenorhabdus sp. XENO-1 JAMGSK01
X. beddingii Q58T MUBK01
X. bovienii T228T JANAIF01
X. budapestensis DSM 16342T NIBS01
X. cabanillasii USTX62T QTUB01
X. doucetiae FRM16T FO704550
X. eapokensis DL20T MKGQ01
X. ehlersii DSM 16337T NIBT01
X. griffiniae BMMCB LDNM01
X. hominickii KE01T NJAI01
X. indica DSM 17382T NKHP01
X. innexi DSM 16336T NIBU01
X. ishibashii GDh7T NJAK01
X. japonica DSM 16522T FOVO01
X. khoisanae MCB LFCV01
X. koppenhoeferi USNJ01T FPBJ01
X. kozodoii SaVT NJCX01
X. lircayensis VLST JACOII01
X. mauleonii VC01T NITY01
X. miraniensis Q1T NITZ01
X. nematophila ATCC 19061T FN667742
X. poinarii G6T FO704551
X. stockiae TH01T NJAJ01
X. szentirmaii DSM 16338T NIBV01
X. thuongxuanensis 30TX1T MKGR01
X. vietnamensis VN01T MUBJ01

Comparative morphometrics of Steinernema africanum n. sp. IJs.

Species Reference L BD EP NR NL TL a b C c' D% E% H%
S. africanum Present study 751 27 59 117 154 64 28 4.9 12 3.7 38 94 35
(690‒802) (25‒32) (54‒68) (87‒132) (123‒167) (52‒72) (23‒30) (4.3‒6.3) (10‒15) (2.9‒4.2) (34‒46) (79‒129) (28‒39)
S. akhursti Qiu et al. (2005) 812 33 59 90 (83‒95) 119 73 24 6.8 11 3.5 47 77 52
(770‒835) (33‒35) (55‒60) (115‒123) (68‒75) (23‒26) (6.6‒7.2) (10‒12) (3.3‒3.7) (45‒50) (73‒86) (49‒56)
S. Ma et al. (2012) 843 30 62 87 (72‒97) 125 73 28 6.8 12 4.3 49 81 39
cholashanense (727‒909) (26‒35) (59‒65) (110‒138) (60‒80) (24‒34) (6.1‒7.2) (10‒14) (3.5‒5.0) (46‒53) (76‒91) (33‒47)
S. citrae Stokwe et al. (2011) 754 (623‒849) 26 (23‒28) 56 (49‒64) 98 (83‒108) 125 (118‒137) 71 (63‒81) 30 (25‒34) 6.0 (5.1‒7.1) 15 (13‒17) NA 44 (39‒58) 110 (85‒132) 43 (37‒50)
S. feltiae Nguyen et al. 849 29 63 113 136 86 30 6.4 10 4.8 46 74 44
(2007) (766‒928) (22‒32) (58‒67) (108‒117) (130‒143) (81‒89) (27‒34) (5.8‒6.8) (9.4‒11) (4.5‒5.1) (44‒50) (67‒81) (37‒51)
S. hebeiense Chen et al. 658 26 48 78 (73‒83) 107 66 26 6.2 10 NA 45 72 43
(2006) (610‒710) (23‒28) (43‒51) (100‒111) (63‒71) (24‒28) (5.7‒6.7) (9.4‒11) (40‒50) (65‒80) (32‒50)
S. ichnusae Tarasco et al. 866 31 63 102 138 81 28 6.3 11 (9‒12) 4.6 46 77 48
(2008) (767‒969) (27‒35) (59‒68) (94‒108) (119‒148) (76‒89) (24‒32) (5.6‒6.9) (4.2‒5.1) (42‒49) (68‒83) (44‒50)
S. jollieti Spiridonov et al. (2004) 711 23 60 NA 123 68 31 5.7 10.5 4.5 (NA) 48 88 (NA) 55
(625‒820) (20‒28) (53‒65) (115‒135) (60‒73) (25‒34) (4.9‒6.4) (9‒12) (46‒50) (46‒60)
S. kraussei Nguyen et al. 951 33 63 105 134 79 29 (NA) 7.1 (NA) 12.1 (NA) 3.9 (NA) 47 (NA) 80 (NA) 38
(2007) (797‒1,102) (30‒36) (50‒66) (99‒111) (119‒145) (63‒86) (35‒40)
S. kushidai Mamiya (1998) 589 26 46 76 (70‒84) 111 50 23 5.3 11.7 NA 41 92 (NA) NA
(424‒662) (22‒31) (42‒50) (106‒120) (44‒59) (19‒25) (4.9‒5.9) (10‒13) (38‒44)
S. litorale Yoshida, (2004) 909 31 61 96 125 83 30 7.3 11 4.5 49 73 33 (NA)
(834‒988) (28‒33) (54‒69) (89‒104) (114‒133) (72‒91) (27‒31) (6.7‒7.9) (10‒11.9) (3.8‒5.4) (44‒56) (68‒84)
S. nguyeni Malan et al. 737 25 52 80 (74‒86) 110 67 29 6.7 11 4.3 48 79 27
(2016) (673‒796) (22‒28) (47‒58) (101‒121) (61‒73) (27‒33) (6.2‒7.4) (10‒12) (2.8‒4.8) (43‒57) (70‒86) (20‒31)
S. oregonese Liu and Berry, (1996) 980 34 66 NA 132 70 30 7.6 (6‒8) 14 4.7 (NA) 50 100 31
(820‒1,110) (28‒38) (60‒72) (116‒148) (64‒78) (24‒37) (12‒16) (40‒60) (90‒110) (30‒33)
S. populi Tian et al. (2022) 1,095 36 77 106 149 64 30 7.4 17 2.8 52 121 35
(973‒1,172) (33‒41) (70‒86) (98‒113) (134‒159) (55‒72) (24‒33) (6.8‒8.5) (15‒20) (2.4‒3.3) (47‒61) (105‒140) (26‒44)
S. puntauvense Uribe-Lorfo 670 33 25 54 (46‒69) 94 54 20 6.1 12 NA 42 44 54
et al. (2007) (631‒728) (31‒38) (20‒30) (81‒103) (51‒59) (17‒23) (7.1‒7.9) (11‒13) (25‒50) (35‒56) (52‒55)
S. sandneri Lis et al. (2021) 843 27 56 103 138 75 31 6.1 11.2 NA 40 74 34
(708‒965) (23‒32) (44‒64) (83‒118) (123‒151) (64‒86) (27‒34) (5.5‒6.9) (11‒13) (36‒45) (63‒86) (23‒40)
S. sangi Phan et al. (2001) 753 35 52 91 (78‒97) 127 81 22 5.9 9.3 4.5 (NA) 40 62 49
(704‒784) (30‒40) (46‒54) (120‒138) (76‒89) (19‒25) (5.6‒6.3) (9‒10) (36‒44) (56‒70) (44‒52)
S. silvaticum Sturhan et al. (2005) 860 30 62 96 121 75 29 7.3 11.4 4.0 50 ‒ 46
(670‒975) (26‒35) (51‒73) (75‒109) (100‒141) (63‒86) (23‒33) (6.3‒7.7) (10‒13) (3.1‒4.9) (46‒56) (37‒53)
S. tielingense Ma et al. (2012) 915 35 69 98 128 81 295 73 11 4 49 23 58
(824‒979) (32‒38) (64‒73) (90‒105) (120‒135) (74‒85) (27‒31) (67‒79) (10‒12) (3.5‒4.6) (44‒56) (68‒84) (53‒64)
S. texanum Nguyen et al. (2007) 756 30 59 92 115 73 25 6.5 10.4 3.3 51 81 59
(732‒796) (29‒34) (52‒62) (84‒102) (111‒120) (60‒79) (22‒27) (6.2‒7.0) (10‒13) (3.3‒4.6) (46‒53) (76‒88) (53‒69)
S. xinbinense Ma et al. (2012) 694 30 51 86 (75‒90) 116 73 24 6.1 (5‒7) 9.7 4.2 (3‒5) 44 71 35
(635‒744) (28‒31) (46‒53) (109‒125) (65‒78) (21‒25) (8‒11) (40‒47) (65‒78) (30‒42)
S. xueshanense Mrácek et al. 860 30 67 91 (81‒96) 135 87 28 6.4 9.9 4.6 50 78 51
(2009) (768‒929) (29‒33) (60‒72) (130‒143) (80‒92) (26‒32) (5.8‒7.0) (9‒11) (3.8‒5.1) (46‒52) (70‒90) (46‒55)
S. weiseri Mrácek et al. 740 25 57 84 (72‒92) 113 60 29 6.6 12 3.7 51 95 (NA) 22
(2003) (586‒828) (24‒29) (43‒65) (95‒119) (49‒68) (25‒33) (5.7‒7.2) (10‒14) (3.2‒4.1) (44‒55) (18‒24)

Comparative morphometrics of first‒generation Steinernema africanum n. sp. males. All measurements are in micrometer (except ratios and percentages).

Species L BD EP NR NL TL SL GL a b c c' D% SW% GS%
S. africanum 1,202 115 93 92 143 41 71 43 10 8 29 1.0 65 172 60
(977‒1,400) (65‒131) (69‒109) (79‒104) (132‒147) (34‒46) (65‒76) (32‒49) (9‒12) (7‒12) (25‒34) (0.9‒1.1) (52‒74) (144‒197) (49‒68)
S. akhursti 1,589 131 102 136 182 35 90 64 NA NA NA NA 56 180 71
(1,350‒1,925) (115‒150) (93‒113) (120‒163) (168‒205) (30‒40) (85‒100) (58‒68) (52‒61) (140‒200) (65‒77)
S. 1,428 137 99 106 152 35 66 39 11 9.3 41 0.7 64 115 71
cholashanense (1,070‒1,778) (73‒204) (75‒135) (91‒126) (135‒173) (29‒43) (60‒71) (32‒45) (9‒24) (8‒11) (36‒51) (0.6‒0.9) (50‒85) (92‒144) (61‒85)
S. citrae 1,154 103 81 106 139 25 65 44 NA NA NA NA 58 198 68
(1,028‒1,402) (87‒113) (64‒92) (92‒119) (123‒155) (17‒31) (57‒80) (32‒59) (47‒67) (156‒233) (48‒89)
S. feltiae 1,612 (1,414‒1,817) 75 (60‒90) 115 (110‒126) 69 (55‒87) a 170 (164‒180) 89 (37‒43) 70 (65‒77) 41 (34‒47) 11.5 (NA) 9.5 (NA) 41.3 (NA) 0.8 (NA) 60 (51‒64) 113 (99‒130) 59 (52‒61)
S. hebeiense 1,177 86 64 84 (78‒93) 126 30 57 46 14 9 39 0.8 51 140 80
(1,036‒1,450) (74‒98) (58‒73) (118‒132) (24‒35) (51‒63) (38‒50) (12‒17) (8‒11) (30‒49) (0.6‒0.9) (48‒59) (120‒170) (60‒90)
S. ichnusae 1,341 137 101 NA 165 40 66 44 22 8.2 34 0.8 62 139 67
(1,151‒1,494) (73‒204) (94‒108) (135‒173) (33‒48) (64‒67) (43‒46) (20‒29) (7‒9) (29‒39) (0.8‒0.9) (59‒65) (120‒162) (64‒69)
S. jollieti 1,662 115 98 NA 156 33 64 54 15 11 51 0.8 (NA) 64 145 (NA) 84 (NA)
(1,296‒1,952) (98‒135) (83‒110) (110‒168) (24‒38) (55‒70) (45‒60) (12‒19) (8‒14) (53‒86) (53‒83)
S. kraussei 1,400 128 81 105 153 39 49 33 11 9 (NA) 37 (NA) 0.9 (NA) 53 (NA) 110 (NA) 67 (NA)
(1,200‒1,600) (110‒144) (73‒99) (95‒122) (137‒178 (36‒44) (42‒53) (29‒37)
S. kushidai 1,400 97 84 129 167 33 63 44 NA NA NA NA 51 150 (NA) 70 (NA)
(1,200‒1,900) (75‒156) (71‒105) (120‒137) (156‒189) (30‒40) (48‒72) (39‒60) (42‒59)
S. litorale 1,360 96 96 114 147 34 75 53 14 9.3 41 0.8 40 174 71
(1,230‒1,514) (82‒111) (77‒107) (94‒128) (133‒163) (26‒41) (67‒89) (44‒64) (12‒16) (8‒10) (33‒56) (0.6‒0.9) (34‒56) (154‒200) (62‒81)
S. nguyeni 997 82 59 91 124 21 66 43 12 8 46 0.7 48 215 66
(818‒1,171) (58‒106) (47‒71) (70‒103) (112‒144) (18‒25) (58‒75) (30‒55) (11‒15) (7‒10) (38‒53) (0.6‒0.8) (38‒57) (185‒279) (46‒81)
S. oregonense 1,680 138 112 111 154 29 71 56 NA NA NA 0.6 (NA) 73 151 (NA) 79 (NA)
(1,560‒1,820) (105‒161) (95‒139) (101‒133) (139‒182) (24‒32) (65‒73) (52‒59) (64‒75)
S. 1,591 119 94 115 140 33 77 34 NA NA NA NA 67 170 65
puntauvense (1,010‒1,931) (101‒139) (68‒114) (104‒128) (130‒159) (28‒40) (71‒81) (30‒40) (45‒85) (140‒200) (55‒75)
S. sandneri 1,461 155 80 126 157 41 60 44 10 9.3 37 NA 51 111 79
(1,206‒1,635) (124‒178) (64‒92) (112‒138) (148‒170) (35‒46) (53‒65) (39‒50) (9‒11) (8‒10) (31‒42) (42‒59) (97‒127) (61‒83)
S. sangi 1,774 159 82 126 166 32 63 40 NA NA NA NA 49 150 60
(1,440‒2,325) (120‒225) (67‒99) (109‒166) (150‒221) (27‒42) (58‒80) (34‒46) (42‒63) (120‒160) (50‒70)
S. silvaticum 1,090 65 79 119 142 34 51 37 17 7.7 34 1.0 60 155 (NA) 73 (NA)
(975‒1,270) (52‒78) (71‒92) (90‒126) (116‒168) (20‒47) (42‒64) (30‒43) (14‒20) (8‒9) (24‒55) (0.8‒1.4) (45‒63)
S. tielingense 1,778 129 114 112 160 26 88 62 11 11 70 0.5 71 191 73
(1,430‒2,064) (111‒159) (94‒133) (96‒132) (145‒173) (22‒33) (79‒98) (49‒70) (11‒18) (9‒13) (57‒85) (0.3‒0.6) (64‒78) (176‒212) (59‒82)
S. texanum 1,296 99 90 104 135 23 60 45 NA NA NA NA 67 157 75
(1,197‒1,406) (81‒116) (79‒100) (94‒114) (123‒147) (19‒30) (55‒66) (39‒53) (58‒73) (127‒203) (62‒84)
S. xinbinense 1,265 103 68 106 149 37 56 35 12 8.5 34 0.9 45 137 63
(1,133‒1,440) (90‒126) (57‒75) (91‒120) (138‒159) (30‒41) (49‒62) (30‒41) (11‒13) (7‒9) (31‒39) (0.7‒1.0) (41‒50) (114‒156) (54‒72)
S. 1,589 144 128 NA 160 38 76 49 NA NA NA NA 80 152 64
xueshanense (1,313‒2,040) (97‒159) (113‒137) (151‒175) (29‒48) (66‒91) (41‒60) (73‒87) (93‒172) (58‒95)
S. weiseri 1,180 112 70 99 141 25 68 53 11 8 48 0.7 49 180 80
(990‒1,395) (84‒138) (57‒84) (94‒115) (134‒154) (19‒32) (62‒72) (46‒57) (9‒12) (7‒10) (36‒64) (0.6‒0.9) (39‒60) (150‒240) (70‒85)

Comparative morphometrics of first‒generation Steinernema africanum n. sp. females. All measurements are in micrometer (except ratios and percentages).

Species L BD EP NR NL TL a b c c' V ABD D% Mucron
S. africanum 3,205 180 89 105 185 45 18 17 74 0.9 52 52 48 Present
(2,469‒4,215) (154‒194) (67‒111) (79‒130) (170‒201) (35‒55) (13‒27) (13‒24) (51‒104) (0.7‒1.0) (50‒57) (37‒70) (36‒62)
S. akhursti 7,283 239 126 164 239 49 29b (NA) 31b (NA) 141b 0.6b(NA) 51 86 53 (NA) Present
(5,625‒9,000) (200‒270) (113‒138) (150‒175) (213‒258) (38‒63) (NA) (48‒53) (68‒100)
S. 4,692 255 129 190 196 57 13 25 83 0.8 53 77 50 Present
cholashanense (3,232‒6,363) (156‒332) (111‒148) (176‒223) (181‒231) (46‒70) (13‒23) (18‒32) (62‒119) (0.6‒1.0) (50‒57) (54‒105 (29‒65)
S. citrae 3,087 175 75 151 215 44 NA NA NA NA 54 62 37 Present
(2,038‒4,019) (137‒212) (54‒90) (130‒179) (189‒220) (33‒60) (50‒59) (43‒79) (27‒46)
S. feltiae 3,380 204 82 84 (70—97)a 237 52 17 14 65 1.0 56 52 46 Present
(3,095‒3,774) (170‒254) (68‒97)a (197‒304) (39‒70) (14‒20) (12‒17) (49‒88) (0.7‒1.2)a (44‒57) (47‒62) (40—54)a
S. hebeiense 3,465 167 65 104 147 35 21 24 103 0.7 54 53 45 Absent
(3,972‒4,254) (142‒245) (48‒95) (88‒123) (133‒158) (25‒50) (17‒25) (21‒29) (67‒129) (0.5‒0.9) (50‒57) (45‒65) (36‒66)
S. ichnusae 5,514 269 126 NA 239 60 21 23 93 0.8 53 80 53 Present
(4,547‒6,186) (242‒323) (106‒156) (215‒262) (51‒79) (17‒24) (21‒26) (68‒113) (0.6‒1.0) (51‒57) (70‒94) (47‒63)
S. jollieti 5,148 259 111 NA 214 43 20 26 128 NA 51 NA 52 Present
(3,746‒6,030) (219‒298) (96‒136) (184‒310) (31‒55) (15‒24) (19‒31) (72‒185) (44‒56)
S. kraussei 4,200 240 87 137 192 48 17 22 88 NA 54 45 45 Present
(2,500‒5,400) (153‒288) (66‒99) (127‒146) (178‒205) (33‒59) (39‒50)
S. kushidai 3,500 175 91 124 227 38 NA NA NA NA 56 64 40 Absent
(2,100‒4,700) (54‒59) (78‒105) (111‒144) (204‒255) (30‒45) (54‒59) (54‒84) (37‒46)
S. litorale 4,462 191 88 146 196 39 23 23 117 0.6 56 62 45 Present
(3,930‒5,048) (175‒215) (65‒105) (130‒165) (185‒213) (25‒60) (21‒26) (20‒26) (78‒157) (0.5‒0.9) (0.5‒0.9) (55‒75) (33‒57)
S. nguyeni 4,775 178 72 113 169 32 21 22 119 0.7 56 178 43 Absent
(2,290‒5,361) (130‒216) (49‒98) (84‒139) (137‒194) (20‒67) (15‒30) (15‒30) (53‒165) (0.6‒1.1) (52‒63) (130‒216) (30‒56)
S. oregonense 5,200 242 103 147 210 37 NA NA NA 0.7 (NA) 52 56 49 Absent
(4,400‒6,200) (217‒268) (217‒268) (129‒162) (186‒220) (28‒46) (46‒56) (42‒79) (43‒57)
S. 6,198 198 70 135 192 49 NA NA NA NA 53 76 37 Present
puntauvense (3,687‒8,335) (181‒221) (51‒85) (123‒146) (141‒206) (41‒66) (51‒55) (57‒102) (25‒45)
S. sandneri 4,628 210 84 147 185 147 22 25 102 NA 54 94 46 P‒esent
(4,244‒5,014) (181‒261) (61‒102) (132‒158) (173‒194) (32‒61) (17‒25) (24‒27) (75‒140) (49‒57) (62‒122) (36‒54)
S. sangi 6,030 336 101 158 229 49 NA NA NA NA 51 111 44 P‒esent
(4,830‒7,200) (270‒360) (80‒121) (140‒170) (216‒240) (936‒62) (43‒530) (84‒140) (35‒51)
S. silvaticum 2,150 116 69 113 146 45 19 15 49 1.3 52 35 47 Absent
(1,520‒3,290) (50‒175) (50‒175) (50‒175) (121‒188) (33‒79) (15‒41) (10‒18) (34‒80) (1.0‒1.8) (44‒57) (26‒53) (33‒79)
S. texanum 3,058 163 88 122 172 40 NA NA NA NA 52 60 NA Absent
(2,720‒3,623) (130‒202) (78‒107) (111‒135) (160‒189) (30‒52) (50‒55) (50‒71)
S. tielingense 6,190 251 84 136 202 45 23 29 117 0.6 51 69 41 Absent
(4,028‒8,538) (200‒307) (82‒103) (111‒144) (186‒263) (40‒69) (17‒32) (21‒45) (72‒158) (0.5‒0.9) (49‒54) (56‒92) (32‒49)
S. xinbinense 4,037 176 80 126 186 40 22 22 106 0.6 52 61 40 Present
(3,025‒5,121) (159‒200) (70‒87) (106‒141) (167‒192) (30‒53) (19‒25) (17‒26) (79‒123) (0.5‒0.8) (46‒57) (50‒67) (38‒45)
S. 5,092 257 148 NA 230 55 NA NA NA NA 54 84 NA Present
xueshanense (4,181‒8,181) (182‒343) (117‒148) (196‒274) (43‒66) (52‒62) (38‒72)
S. weiseri 4,610 223 80 125 184 42 21 25 111 0.7 53 63 NA Present
(3,780‒5,940) (202‒263) (75‒86) (108‒154) (162‒226) (38‒59) (17‒29) (22‒31) (87‒156) (0.5‒0.8) (50‒58) (51‒80)
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