Multi-locus phylogenetic analyses uncover species boundaries and reveal the occurrence of two new entomopathogenic nematode species,  n. sp. and  n. sp.

Ricardo A.R. Machado; Aashaq Hussain Bhat; Joaquín Abolafia; Arthur Muller; Pamela Bruno; Patrick Fallet; Carla C.M. Arce; Ted C.J. Turlings; Julio S. Bernal; Joelle Kajuga; Bancy Waweru; Stefan Toepfer

Accès libre

Multi-locus phylogenetic analyses uncover species boundaries and reveal the occurrence of two new entomopathogenic nematode species, Heterorhabditis ruandica n. sp. and Heterorhabditis zacatecana n. sp.

Journal of Nematology

Édition 53 (2021): Edition 1 (January 2021)

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Publié en ligne: 11 nov. 2021

Pages: 1 - 42

DOI: https://doi.org/10.21307/jofnem-2021-089

Mots clés
Biocontrol agents, Dichotomous key, Entomopathogenic nematodes, Nematode morphology, Photorhabdus, Phylogenetics, Species description,

© 2021 Ricardo A.R. Machado et al., published by Sciendo.

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivatives 4.0 International License.

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.

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).

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).

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).

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.

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).

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.

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).

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.

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).

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.

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.

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.

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

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

Morphometrics of infective juveniles and adult generations of Heterorhabditis ruandica n. sp.

	Male
Characters	Holotype	Paratypes	Hermaphrodite (1^st Gen) paratypes	Female (2^nd 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)

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

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

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)

eISSN:: 2640-396X
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Multi-locus phylogenetic analyses uncover species boundaries and reveal the occurrence of two new entomopathogenic nematode species, Heterorhabditis ruandica n. sp. and Heterorhabditis zacatecana n. sp.

Publié en ligne: 11 nov. 2021

Pages: 1 - 42

DOI: https://doi.org/10.21307/jofnem-2021-089

Mots clés
Biocontrol agents, Dichotomous key, Entomopathogenic nematodes, Nematode morphology, Photorhabdus, Phylogenetics, Species description,

© 2021 Ricardo A.R. Machado et al., published by Sciendo.

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivatives 4.0 International License.

Figure 1:

Figure 2:

Figure 3:

Figure 4:

Figure 5:

Figure 6:

Figure 7:

Figure 8:

Figure 9:

Figure 10:

Figure 11:

Figure 12:

Figure 13:

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)

Comparative morphometrics of infective juveniles of Heterorhabditis ruandica n. sp., H. zacatecana n. sp., and of different closely related Heterorhabditis species.

Morphometrics of infective juveniles and adult generations of Heterorhabditis ruandica n. sp.

Comparative morphometrics of adult females of Heterorhabditis ruandica n. sp., H. zacatecana n. sp., and of different closely related Heterorhabditis species.

Comparative morphometrics of hermaphrodite females of Heterorhabditis ruandica n. sp., H. zacatecana n. sp., and of different closely related Heterorhabditis species.

Morphometrics of infective juveniles and adult generations of Heterorhabditis zacatecana n. sp.

Multi-locus phylogenetic analyses uncover species boundaries and reveal the occurrence of two new entomopathogenic nematode species, Heterorhabditis ruandica n. sp. and Heterorhabditis zacatecana n. sp.

Publié en ligne: 11 nov. 2021

Pages: 1 - 42

DOI: https://doi.org/10.21307/jofnem-2021-089

Mots clésBiocontrol agents, Dichotomous key, Entomopathogenic nematodes, Nematode morphology, Photorhabdus, Phylogenetics, Species description,

© 2021 Ricardo A.R. Machado et al., published by Sciendo.

This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivatives 4.0 International License.

Figure 1:

Figure 2:

Figure 3:

Figure 4:

Figure 5:

Figure 6:

Figure 7:

Figure 8:

Figure 9:

Figure 10:

Figure 11:

Figure 12:

Figure 13:

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)

Comparative morphometrics of infective juveniles of Heterorhabditis ruandica n. sp., H. zacatecana n. sp., and of different closely related Heterorhabditis species.

Morphometrics of infective juveniles and adult generations of Heterorhabditis ruandica n. sp.

Comparative morphometrics of adult females of Heterorhabditis ruandica n. sp., H. zacatecana n. sp., and of different closely related Heterorhabditis species.

Comparative morphometrics of hermaphrodite females of Heterorhabditis ruandica n. sp., H. zacatecana n. sp., and of different closely related Heterorhabditis species.

Morphometrics of infective juveniles and adult generations of Heterorhabditis zacatecana n. sp.

Mots clés
Biocontrol agents, Dichotomous key, Entomopathogenic nematodes, Nematode morphology, Photorhabdus, Phylogenetics, Species description,