Steinernema sandneri n. sp. (Rhabditida: Steinernematidae), a new entomopathogenic nematode from Poland

Lis, Magdalena; Sajnaga, Ewa; Skowronek, Marcin; Wiater, Adrian; Rachwał, Kamila; Kazimierczak, Waldemar

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Steinernema sandneri n. sp. A: infective juvenile, anterior region; B: first-generation male, anterior region; C: first-generation female, tail region; D: first-generation female, vulval region; E: infective juvenile, tail region; F: first-generation male, tail region; G: spicule; H: gubernaculum. Scale bars as on images. Lateral views.

Steinernema sandneri n. sp. Differential interference contrast (A,C,E) and scanning electron (B,D,F) micrographs of infective juveniles. A – amphid openings, An – anus, BP – bacterial pouch, CP – cephalic papillae, EP – excretory pore, ES – esophagus, F – phasmid opening, H – hyaline part, LF – lateral fields, NR – nerve ring. Scale bars as on images.

Steinernema sandneri n. sp. Scanning electron micrographs of first-generation males. A,B: anterior region with excretory pore (EP), deirids (D), cephalic (CP) and labial papillae (LP). C,D: posterior region with genital papillae (arrows), single preanal papilla (S-PA) and mucron (Mu). Scale bars as on images.

Steinernema sandneri n. sp. Differential interference contrast micrographs of first- (A,B,E,F,G) and second-generation males (C,D). A,C: anterior region with esophagus (ES), excretory pore (EP) and nerve ring (NR). B,D: posterior region with spicules, gubernaculum, and mucron (Mu). E: spicules. F,G: gubernaculum (lateral and ventral view). Scale bars as on images.

Steinernema sandneri n. sp. Differential interference contrast (A,C,E) and scanning electron (B,D,F) micrographs of first-generation females. An – anus, EP – excretory pore, ES – esophagus, LP – labial papillae, NR – nerve ring, V – vulva. Scale bars as on images.

Phylogenetic tree of the phylogenetic relationships of S. sandneri n. sp. with other species of the genus Steinernema based on sequences of the ITS rDNA. Bootstrap values > 50% are indicated at the branching points. The scale bar indicates the number of nucleotide substitutions per site. The evolutionary history was inferred using the Maximum Likelihood method based on the HKY + G model. All positions containing gaps were eliminated. There were a total of 646 positions in the final dataset. Evolutionary analyses were conducted in MEGA6.

Phylogenetic tree of the phylogenetic relationships of S. sandneri n. sp. with other species of the genus Steinernema based on sequences of the D2D3 regions. Bootstrap values > 50% are indicated at the branching points. The scale bar indicates the number of nucleotide substitutions per site. The evolutionary history was inferred using the Maximum Likelihood method based on the GTR + G model. All positions containing gaps were eliminated. There were a total of 850 positions in the final dataset. Evolutionary analyses were conducted in MEGA6.

Phylogenetic tree of the phylogenetic relationships of S. sandneri n. sp. with other species of the genus Steinernema based on cox1 gene sequences. Bootstrap values > 50% are indicated at the branching points. The scale bar indicates the number of nucleotide substitutions per site. The evolutionary history was inferred using the Maximum Likelihood method based on the HKY + G + I model. All positions containing gaps were eliminated. There were a total of 567 positions in the final dataset. Evolutionary analyses were conducted in MEGA6.

Morphometrics (in μm) of different developmental stages of Steinernema sandneri n_ sp_ [mean ± SE (range)] [N = 25]_

	First generation			Second generation
	Males		Females	Males	Females	Infective juveniles
Character	Holotype	Paratypes	Paratypes	Paratypes	Paratypes	Paratypes
Body length [L]	1,565.3	1,461 ± 22.1 (1,205.7–1,635.3)	4,628 ± 46.4 (4,244.0–5,014.0)	946 ± 13.8 (817.5–1,093.8)	2,120 ± 51.5 (1,640.6–2,753.2)	843.0 ± 13.9 (708.2–964.5)
Greatest body width [W]	143.0	155.1 ± 2.7 (123.8–177.7)	209.6 ± 3.4 (181.3–261.3)	70.1 ± 1.1 (54.9–79.5)	126.6 ± 3.3 (88.9–146.6)	27.4 ± 0.5 (23.0–31.9)
Anterior end to excretory pore [EP]	88.4	80.4 ± 1.5 (63.5–92.4)	84.4 ± 2.1 (61.4–101.6)	69.8 ± 1.5 (59.0–84.6)	72.1 ± 1.3 (57.3–88.4)	55.9 ± 0.8 (44.4–64.2)
Anterior end to nerve ring [NR]	121.9	126.0 ± 1.5 (112.0–138.1)	146.7 ± 1.2 (132.5–157.6)	97.7 ± 0.8 (86.4–105.8)	113.7 ± 1.1 (102.5–124.6)	102.6 ± 1.4 (82.6–117.9)
Anterior end to esophagus [ES]	155.9	157.2 ± 1.1 (147.6–169.6)	184.7 ± 1.0 (173.2–193.9)	120.5 ± 1.1 (109.0–128.7)	145.8 ± 1.4 (130.3–158.5)	138.4 ± 0.5 (122.5–150.5)
Testis reflection	461.6	452.1 ± 9.5 (359.3–537.7)	–	202.6 ± 13.2 (84.9–379.2)	–	–
Tail length [T]	45.1	41.2 ± 0.5 (35.4–45.5)	46.7 ± 1.6 (32.4–60.9)	42.3 ± 1.0 (31.7–52.1)	57.5 ± 1.4 (46.5–72.1)	75.2 ± 1.1 (64.4–86.4)
Anal body diameter [ABW]	50.1	54.1 ± 0.6 (49.9–59.2)	94.0 ± 2.9 (62.1–121.8)	36.8 ± 0.4 (30.7–40.8)	54.4 ± 1.5 (43.1–70.7)	17.3 ± 0.4 (14.6–23.8)
Spicule length [SL]	64.2	59.8±0.5 (52.6–65.3)	–	51.2 ± 0.9 (42.5–60.2)	–	–
Gubernaculum length [GL]	39.2	43.6 ± 0.5 (39.1–50.2)	–	30.4 ± 0.6 (24.2–39.5)	–	–
a [L/W]	10.9	9.5 ± 0.1 (8.5–11.0)	22.2 ± 0.4 (17.4–24.7)	13.6 ± 0.2 (12.0–16.4)	16.9 ± 0.4 (14.1–23.2)	30.9 ± 0.3 (27.2–33.8)
b [L/ES]	10.4	9.3 ± 0.1 (8.0–10.2)	25.1 ± 0.2 (23.5–27.2)	7.9 ± 0.1 (7.2–9.4)	14.5±0.3 (12.0–18.1)	6.1 ± 0.1 (5.5–6.9)
c [L/T]	34.7	35.6 ± 0.5 (31.2–41.9)	102.0 ± 3.8 (75.4–140.3)	22.7 ± 0.6 (17.2–28.5)	37.3 ± 1.1 (24.6–50.1)	11.2 ± 0.1 (10.5–13.2)
Hyaline% [(H/T) × 100]	–	–	–	–	–	33.6 ± 3.9 (22.7–39.9)
D% [(EP/ES) × 100]	56.7	51.2 ± 0.9 (42.1–59.3)	45.7 ± 1.1 (35.5–54.2)	58.0±1.2 (48.1–71.9)	49.5 ± 0.9 (36.2–58.1)	40.4 ± 0.4 (35.8–44.8)
E% [(EP/T) × 100]	196.0	195.8 ± 4.0 (160.2–240.9)	186.0 ± 7.9 (128.1–266.8)	167.4 ± 1.2 (128.2–222.8)	127.2 ± 3.9 (101.4–163.9)	74.4 ± 0.9 (62.6–85.8)
SW% [(SL/ABW) × 100]	128.1	110.9 ± 1.5 (97.0–126.9)	–	139.6 ± 3.2 (105.1–171.4)	–	–
GS% [(GL/SL) × 100]	61.1	79.1 ± 1.2 (60.8–82.8)	–	59.6 ± 1.1 (49.5–68.6)	–	–
V% [(Vulva – anterior end/L) × 100]	–	–	53.7 ± 0.3 (49.0–56.8)	–	54.3 ± 0.8 (39.3–59.1)	–

Percentage of similarity (upper triangle) and genetic distance measured by the number of nucleotide substitutions (lower triangle) in the sequences of cox1 gene of S_ sandneri n_ sp_ and other closely related Steinernema spp_

	Species	Acc. no.	1	2	3	4	5	6	7	8	9	10	11
1	S. sandneri n. sp.	MW078544	–	93.7	93.8	92.9	87.7	86.6	88.4	84.7	86.4	85.9	85.7
2	S. kraussei	JN683829	36	–	94.4	94.0	87.1	86.4	87.3	84.1	85.7	84.5	85.0
3	S. kraussei	AY943990	35	32	–	95.6	87.8	86.8	86.7	85.0	86.2	85.2	85.4
4	S. kraussei *	MW647850	40	34	25	–	86.2	86.4	86.9	83.6	85.7	84.5	85.4
5	S. silvaticum	MG547572	70	73	68	78	–	84.7	85.4	83.6	83.5	82.9	84.7
6	S. oregonense	AY943995	76	77	75	77	87	–	87.8	85.5	86.6	83.4	86.8
7	S. feltiae	JQ423217	66	72	75	74	83	69	–	85.9	88.0	84.0	96.9
8	S. jollieti	GU569068	87	90	85	93	93	82	80	–	85.0	84.7	84.1
9	S. weiseri	GU569075	77	81	78	81	99	76	68	85	–	82.7	84.8
10	S. kushidai	AY943991	80	88	84	88	97	94	91	87	98	–	84.5
11	S. monticolum	AY943994	81	85	83	83	87	75	74	90	86	88	–

Percentage of similarity (upper triangle) and genetic distance measured by the number of nucleotide substitutions (lower triangle) in the sequences of ITS rDNA of S_ sandneri n_ sp_ and other closely related Steinernema spp_

	Species	Acc. no.	1	2	3	4	5	6	7	8	9	10	11	12	13	14	15	16
1	S. sandneri n. sp.	MW078536	–	97.0	94.5	92.4	90.2	94.1	94.1	94.5	89.6	89.8	89.0	89.4	90.3	88.7	85.8	76.6
2	S. kraussei	AY230174	21	–	95.2	92.1	90.4	90.5	93.8	94.9	88.6	89.2	88.6	89.1	89.4	88.0	85.2	76.0
3	S. silvaticum	AY230162	37	31	–	92.0	90.4	90.2	92.6	94.6	88.8	89.4	89.2	88.9	89.6	88.0	84.8	75.8
4	S. cholashanense	EF431959	51	48	50	–	94.2	97.2	93.2	94.2	91.3	92.7	91.7	92.8	92.2	91.8	86.6	77.5
5	S. oregonense	AF122019	59	56	59	33	–	93.5	92.0	92.1	89.9	91.5	89.7	91.2	90.4	90.4	86.8	76.6
6	S. xueshanense	FJ660052	67	59	63	17	41	–	91.3	92.1	90.3	91.2	89.0	91.2	89.8	89.9	85.6	76.9
7	S. tielingense	GU994201	41	42	51	42	48	56	–	94.8	89.9	89.5	89.7	90.5	90.1	89.2	86.2	77.2
8	S. xinbinense	JN171593	35	34	36	35	47	50	34	–	90.8	90.8	90.2	91.5	91.0	90.6	86.6	77.0
9	S. feltiae	AF121050	73	73	74	57	60	61	68	60	–	95.8	91.4	94.0	93.4	94.4	88.9	76.1
10	S. ichnusae	EU421129	71	70	71	50	52	55	71	62	28	–	91.8	95.5	93.9	85.2	88.6	76.9
11	S. jollieti	AY171265	76	73	70	53	61	62	67	63	54	46	–	91.1	90.0	91.1	86.1	77.3
12	S. weiseri	AY171268	72	69	73	48	52	55	62	55	39	32	51	–	94.5	96.6	89.0	77.0
13	S. nguyeni	KP325084	67	69	70	53	60	61	67	61	47	43	56	39	–	92.9	87.9	76.4
14	S. litorale	AB243441	78	78	80	57	59	61	73	63	38	34	46	24	44	–	88.7	77.2
15	S. hebeiense	DQ105794	98	97	102	92	84	90	93	90	76	80	82	77	81	74	–	74.9
16	S. monticolum	AF122017	140	139	142	133	134	133	132	134	143	135	120	134	132	132	143	–

Percentage of similarity (upper triangle) and genetic distance measured by the number of nucleotide substitutions (lower triangle) in the sequences of D2D3 domain of 28 S rDNA of S_ sandneri n_ sp_ and other closely related Steinernema spp_

	Species	Acc. no.	1	2	3	4	5	6	7	8	9	10	11	12	13	14
1	S. sandneri n. sp.	MW078535	–	98.2	98.2	98.2	96.2	97.5	97.5	96.8	96.8	96.6	96.1	97.0	95.7	92.4
2	S. kraussei	AF331896	15	–	99.8	99.8	97.2	98.2	98.9	97.6	98.2	97.9	97.4	98.1	96.2	92.6
3	S. kraussei	GU569053	15	2	–	99.5	97.2	98.6	99.1	97.8	98.5	98.1	97.6	98.1	96.6	92.7
4	S. kraussei *	MW647849	15	2	4	–	97.2	98.2	98.9	97.7	98.2	97.8	97.3	97.0	95.7	92.9
5	S. silvaticum	MG547576	32	24	24	24	–	97.2	96.8	96.1	96.6	96.1	95.7	96.7	94.7	91.1
6	S. cholashanense	EF520284	21	14	12	14	24	–	98.4	97.5	98.1	97.7	97.2	97.8	96.4	92.7
7	S. oregonense	AF331891	21	9	7	11	27	14	–	97.9	98.7	98.2	98.0	98.6	96.9	93.4
8	S. xueshanense	FJ666053	27	19	19	19	33	20	18	–	98.0	97.9	97.3	98.1	96.6	92.7
9	S. feltiae	AF3311906	27	15	13	17	29	16	11	17	–	99.3	98.8	99.4	97.2	93.3
10	S. ichnusae	EU421130	28	18	16	18	32	19	14	17	5	–	98.6	98.9	97.2	93.3
11	S. jollieti	GU569051	32	22	20	24	36	23	16	22	9	12	–	98.9	96.7	93.1
12	S. weiseri	GU569059	26	16	16	18	28	19	12	16	5	8	9	–	97.1	93.7
13	S. texanum	EF152569	37	31	29	31	45	30	26	28	24	23	27	25	–	93.1
14	S. monticolum	EF439651	56	53	53	54	69	56	49	56	51	52	54	49	53	–

Steinernema sandneri n_ sp_ – permanent slides description and designation numbers in the collection of Museum and Institute of Zoology, Polish Academy of Sciences, Warsaw, Poland_

Slide description	Slide ID
Slide no. 1 – Steinernema sandneri n. sp. (Rhabditida: Steinernematidae) (male), holotype, natural host unknown, isolated from soil samples: 51°46’55”N 22°42’35”E in 2017	MIZ PAN WARSZAWA 2-2021/1
Slide no. 2 – Steinernema sandneri n. sp. (Rhabditida: Steinernematidae), 55 infective juveniles, paratype, natural host unknown, isolated from soil samples: 51°46’55”N 22°42’35”E in 2017	MIZ PAN WARSZAWA 2-2021/2
Slide no. 3 – Steinernema sandneri n. sp. (Rhabditida: Steinernematidae), 36 infective juveniles, paratype, natural host unknown, isolated from soil samples: 51°46’55”N 22°42’35”E in 2017	MIZ PAN WARSZAWA 2-2021/3
Slide no. 4 – Steinernema sandneri n. sp. (Rhabditida: Steinernematidae), 10 (males), first generation, paratype, natural host unknown, isolated from soil samples: 51°46’55”N 22°42’35”E, in 2017	MIZ PAN WARSZAWA 2-2021/4
Slide no. 5 – Steinernema sandneri n. sp. (Rhabditida: Steinernematidae), 10 (males), first generation, paratype, natural host unknown, isolated from soil samples: 51°46’55”N 22°42’35”E, in 2017	MIZ PAN WARSZAWA 2-2021/5
Slide no. 6 – Steinernema sandneri n. sp. (Rhabditida: Steinernematidae), 10 (males), first generation, paratype, natural host unknown, isolated from soil samples: 51°46’55”N 22°42’35”E, in 2017	MIZ PAN WARSZAWA 2-2021/6
Slide no. 7 – Steinernema sandneri n. sp. (Rhabditida: Steinernematidae), 10 (males), first generation, paratype, natural host unknown, isolated from soil samples: 51°46’55”N 22°42’35”E, in 2017	MIZ PAN WARSZAWA 2-2021/7
Slide no. 8 – Steinernema sandneri n. sp. (Rhabditida: Steinernematidae), 10 (males), first generation, paratype, natural host unknown, isolated from soil samples: 51°46’55”N 22°42’35”E in 2017	MIZ PAN WARSZAWA 2-2021/8
Slide no. 9 – Steinernema sandneri n. sp. (Rhabditida: Steinernematidae), 5 (females), first generation, paratype, natural host unknown, isolated from soil samples: 51°46’55”N 22°42’35”E in 2017	MIZ PAN WARSZAWA 2-2021/9
Slide no. 10 – Steinernema sandneri n. sp. (Rhabditida: Steinernematidae), 5 (females), first generation, paratype, natural host unknown, isolated from soil samples: 51°46’55”N 22°42’35”E in 2017	MIZ PAN WARSZAWA 2-2021/10
Slide no. 11 – Steinernema sandneri n. sp. (Rhabditida: Steinernematidae), 5 (females), first generation, paratype, natural host unknown, isolated from soil samples: 51°46’55”N 22°42’35”E in 2017	MIZ PAN WARSZAWA 2-2021/11
Slide no. 12 – Steinernema sandneri n. sp. (Rhabditida: Steinernematidae), 5 (females), first generation, paratype, natural host unknown, isolated from soil samples: 51°46’55”N 22°42’35”E, in 2017	MIZ PAN WARSZAWA 2-2021/12
Slide no. 13 – Steinernema sandneri n. sp. (Rhabditida: Steinernematidae), 5 (females), first generation, paratype, natural host unknown, isolated from soil samples: 51°46’55”N 22°42’35”E in 2017	MIZ PAN WARSZAWA 2-2021/13
Slide no. 14 – Steinernema sandneri n. sp. (Rhabditida: Steinernematidae), 13 (females), second generation, paratype, natural host unknown, isolated from soil samples: 51°46’55”N 22°42’35”E in 2017	MIZ PAN WARSZAWA 2-2021/14
Slide no. 15 – Steinernema sandneri n. sp. (Rhabditida: Steinernematidae), 15 (females), second generation, paratype, natural host unknown, isolated from soil samples: 51°46’55”N 22°42’35”E in 2017	MIZ PAN WARSZAWA 2-2021/15
Slide no. 16 – Steinernema sandneri n. sp. (Rhabditida: Steinernematidae), 9 (females), second generation, paratype, natural host unknown, isolated from soil samples: 51°46’55”N 22°42’35”E in 2017	MIZ PAN WARSZAWA 2-2021/16
Slide no. 17 – Steinernema sandneri n. sp. (Rhabditida: Steinernematidae), 13 (males), second generation, paratype, natural host unknown, isolated from soil samples: 51°46’55”N 22°42’35”E in 2017	MIZ PAN WARSZAWA 2-2021/17
Slide no. 18 – Steinernema sandneri n. sp. (Rhabditida: Steinernematidae), 15 (males), second generation, paratype, natural host unknown, isolated from soil samples: 51°46’55”N 22°42’35”E in 2017	MIZ PAN WARSZAWA 2-2021/18
Slide no. 19 – Steinernema sandneri n. sp. (Rhabditida: Steinernematidae), 15 (males), second generation, paratype, natural host unknown, isolated from soil samples: 51°46’55”N 22°42’35”E in 2017	MIZ PAN WARSZAWA 2-2021/19

Percentage of similarity (upper triangle) and genetic distance measured by the number of nucleotide substitutions (lower triangle) in the sequences of ITS rDNA regions of S_ sandneri n_ sp_, S_ kraussei and S_ silvaticum isolates, the closest relatives_

	Species	Acc. no.	1	2	3	4	5	6	7	8	9	10	11	12	13	14	15	16
1	S. sandneri	MW078536	–	97.0	97.7	97.7	97.5	97.5	97.2	97.7	97.5	96.6	96.5	96.3	96.0	95.0	94.9	94.5
2	S. kraussei	AY230174	21	–	98.1	98.1	97.9	97.9	98.1	97.2	95.7	98.1	98.2	97.5	97.5	95.4	95.3	94.9
3	S. kraussei	AY171270	17	13	–	99.2	98.5	98.5	99.6	98.6	98.2	100	99.3	99.4	98.3	96.1	96.0	95.6
4	S. kraussei	AY171248	16	12	5	–	99.0	99.0	99,0	99.0	97.3	99.2	98.9	98.6	97.6	95.7	95.6	95.2
5	S. kraussei*	KY819012	18	14	11	6	–	99.7	98.2	99.6	96.6	98.5	98.9	97.9	97.2	95.3	95.2	94.8
6	S. kraussei	AY171264	18	14	11	6	2	–	98.2	99.9	86.6	98.5	99.2	97.9	96.9	95.3	95.2	94.8
7	S. kraussei	AB243442	20	13	3	6	13	13	–	98.2	97.8	99.6	98.9	99.0	98.3	96.3	96.1	95.7
8	S. kraussei	AY230175	17	14	10	6	3	1	13	–	96.8	98.6	99.3	98.1	96.9	95.5	95.3	94.9
9	S. kraussei	JN683825	17	14	1	6	12	12	4	11	–	99.2	98.5	99.5	98.3	96.1	96.0	95.6
10	S. kraussei	AY171258	17	13	0	5	11	11	3	10	1	–	99.3	98.6	98.3	96.1	96.0	95.6
11	S. kraussei	AY230176	18	12	5	7	8	6	8	5	6	5	–	98.0	97.6	95.6	95.5	95.0
12	S. kraussei*	MW647848	20	16	3	8	14	14	6	13	4	3	8	–	97.8	95.6	95.5	95.0
13	S. kraussei	AY230161	28	15	11	15	19	21	11	21	10	11	16	14	–	95.3	95.1	94.7
14	S. silvaticum	MG543845	34	30	26	28	32	32	25	31	24	26	30	29	31	–	99.9	99.4
15	S. silvaticum	AY171255	35	31	27	29	33	33	26	32	25	27	31	30	32	2	–	99.6
16	S. silvaticum	AY230162	37	33	29	31	35	35	28	34	26	29	32	32	34	3	2	–

Details on taxa used in the molecular analyses_

		GeneBank accession no.
Species	Isolate name/geographic origin	ITS rDNA	28S rDNA	cox1
Steinernema sandneri n. sp.	S17-050, Poland	MW078536	MW078535	MW078544
Steinernema affine	B1, England		AF331899
Steinernema affine	The Netherlands	AY171298
Steinernema akhursti	China	DQ375757
Steinernema bicornutum	Serbia		AF331904
Steinernema bicornutum	Yugoslavia	AF121048
Steinernema cameroonense	OB, Cameroon	JX985267
Steinernema carpocapsae	Russia	AY171282
Steinernema cholashanense	Tibet, China	EF431959	EF520284
Steinernema citrae	141-C, South Africa	EU740970	GU004534
Steinernema costaricense	Costa Rica		EF187017
Steinernema feltiae	Bodega Bay, USA		AF331906
Steinernema feltiae	SN, USA	AF121050
Steinernema feltiae	3, Portugal			JQ423217
Steinernema glaseri	NC, USA		AF331908
Steinernema glaseri	NJ, USA	AF122015
Steinernema hebeiense	G6, China	DQ105794
Steinernema hermaphroditum	VK-2013, India	KC252604
Steinernema ichnusae	Sardinia, Italy	EU421129	EU421130
Steinernema jollieti	Monsanto, USA		GU569051	GU569068
Steinernema jollieti	73, USA	AY171265
Steinernema kraussei	Westphalia, Germany	AY230175	AF331896	AY943990
Steinernema kraussei	Altai 35, Russia	AY171270
Steinernema kraussei	Nash, UK	AY230176
Steinernema kraussei	Italy	AY230174
Steinernema kraussei	Iceland	AY171248
Steinernema kraussei	20F, Portugal	JN683825
Steinernema kraussei	D, Switzerland	AY171258
Steinernema kraussei	Russia	AY171264
Steinernema kraussei	HkHm22, Japan	AB243442
Steinernema kraussei	Skr-LUB, Lublin, Poland	KY819012
Steinernema kraussei	B2, UK	AY230161
Steinernema kraussei	20F, Portugal			JN683829
Steinernema kraussei	Quebec, Canada		GU569053
Steinernema kraussei	SKR S11-50, Poland	MW647848	MW647849	MW647850
Steinernema kushidai	Hamakita, Japan	AB243440
Steinernema kushidai	N22, Japan			AY943991
Steinernema kushidai	Japan		AF331897
Steinernema litorale	AiAt199, Japan	AB243441
Species	Isolate name/geographic origin	GeneBank accession no.
		ITS rDNA	28S rDNA	cox1
Steinernema monticolum	Korea, South Korea	AF122017	EF439651
Steinernema monticulum	Mt. Chiri, South Korea			AY943994
Steinernema nguyeni	F2, South Africa	KP325084
Steinernema oregonense	Oregon, USA	AF122019
Steinernema oregonense	OS-10, USA		AF331891	AY943995
Steinernema sangi	Vietnam	AY355441	GU569057
Steinernema scarabaei	New Jersey, USA		AY172023
Steinernema scarabaei	Chile	FJ263673
Steinernema silvaticum	S16/019, Poland	MG543845	MG547576	MG547572
Steinernema silvaticum	B, Germany	AY171255
Steinernema silvaticum	B3, UK (type)	AY230162
Steinernema texanum	Texas, USA	EF152568	EF152569
Steinernema tielingense	LFS65, China	GU994201	GU994202
Steinernema weiseri	F, Germany	AY171268
Steinernema weiseri	Turkey		GU569059	GU569075
Steinernema xinbinense	LFS8, China	JN171593
Steinernema xinbinense	LFS40, China		GU994202
Steinernema xueshanense	Yunnan, China	FJ666052	FJ666053
Caenorhabditis elegans	N2 Bristol, USA			NC001328
Caenorhabditis elegans		X03680	X03680

Comparative morphometrics of third-stage infective juveniles of S_ sandneri n_ sp_ and related Steinernema spp_

	Morphometric character ^a
Species	L	W	EP	NR	ES	T	a	b	c	D%	E%	Reference
S. kushidai	589 (424–662)	26 (22–31)	46 (42–50)	76 (70–84)	111 (106–120)	50 (44–59)	22.5 (19–25)	5.3 (4.9–5.9)	11.7 (10–13)	41 (38–44)	92 (NA)	Mamiya, (1988)
S. hebeiense	658 (610–710)	26 (23–28)	48 (43–51)	78 (73–83)	107 (100–111)	66 (63–71)	26 (24–28)	6.2 (5.7–6.7)	10 (9.4–11)	45 (40–50)	72 (65–80)	Chen et al. (2006)
S. puntauvense	670 (631–728)	33 (31–38)	25 (20–30)	54 (46–69)	94 (81–103)	54 (51–59)	20 (17–23)	6.1 (7.1–7.9)	12 (11–13)	42 (25–50)	44 (35–56)	Uribe–Lorío et al. (2007)
S. xinbinense	694 (635–744)	30 (28–31)	51 (46–53)	86 (75–90)	116 (109–125)	73 (65–78)	24 (21–25)	6.1 (5–7)	9.7 (8–11)	44 (40–47)	71 (65–78)	Ma et al. (2012)
S. jollieti	711 (625–820)	23 (20–28)	60 (53–65)	NA	123 (115–135)	68 (60–73)	31 (25–34)	5.7 (4.9–6.4)	10.5 (9.0–11.7)	48 (46–50)	88 (NA)	Spiridonov et al. (2004b)
S. nguyeni	737 (673–796)	25 (22–28)	52 (47–58)	80 (74–86)	110 (101–121)	67 (61–73)	29 (27–33)	6.7 (6.2–7.4)	11 (10–12)	48 (43–57)	79 (70–86)	Malan et al. (2016)
S. weiseri	740 (586–828)	25 (24–29)	57 (43–65)	84 (72–92)	113 (95–119)	60 (49–68)	29 (25–33)	6.6 (5.7–7.2)	12 (10–14)	51 (44–55)	95 (NA)	Mráček et al. (2003)
S. sangi	753 (704–784)	35 (30–40)	52 (46–54)	91 (78–97)	127 (120–138)	81 (76–89)	22 (19–25)	5.9 (5.6–6.3)	9.3 (8.7–10.2)	40 (36–44)	62 (56–70)	Phan et al. (2001)
S. citrae	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 (13–14)	44 (39–58)	110 (85–132)	Stokwe et al. (2011)
S. texanum	756 (732–796)	30 (29–34)	59 (52–62)	92 (84–102)	115 (111–120)	73 (60–79)	25 (22–27)	6.5 (6.2–7.0)	10 (9.6–12.5)	51 (46–53)	81 (76–88)	Nguyen et al. (2007)
S. akhursti	812 (770–835)	33 (33–35)	59 (55–60)	90 (83–95)	119 (115–123)	73 (68–75)	24 (23–26)	6.8 (6.6–7.2)	11 (10–12)	47 (45–50)	77 (73–86)	Qiu et al. (2005)
S. sandneri n. sp.	843 (708–965)	27 (23–32)	56 (44–64)	103 (83–118)	138 (123–151)	75 (64–86)	31 (27–34)	6.1 (5.5–6.9)	11.2 (11–13.2)	40 (36–45)	74 (63–86)	–
S. cholashanense	843 (727–909)	30 (26–35)	62 (59–65)	87 (72–97)	125 (110–138)	73 (60–80)	28 (24–34)	6.8 (6.1–7.2)	12 (10–14)	49 (46–53)	81 (76–91)	Nguyen et al., (2008)
S. feltiae	849 (766–928)	29 (22–32)	63 (58–67)	113 (108–117)	136 (130–143)	86 (81–89)	30 (27–34)	6.4 (5.8–6.8)	10 (9.4–11)	46 (44–50)	74 (67–81)	Nguyen et al., (2007)
S. silvaticum	860 (670–975)	30 (26–35)	62 (51–73)	96 (75–109)	121 (100–141)	75 (63–86)	29 (23–33)	7.3 (6.3–7.7)	11.4 (9.9–13.1)	50 (46–56)	–	Sturhan et al. (2005)
S. xueshanense	860 (768–929)	30 (29–33)	67 (60–72)	91 (81–96)	135 (130–143)	87 (80–92)	28 (26–32)	6.4 (5.8–7.0)	9.9 (9.0–11)	50 (46–52)	78 (70–90)	Mrácˇek et al. (2009)
S. ichnusae	866 (767–969)	31 (27–35)	63 (59–68)	102 (94–108)	138 (119–148)	81 (76–89)	28 (24–32)	6.3 (5.6–6.9)	11 (8.8–12)	46 (42–49)	77 (68–83)	Tarasco et al. (2008)
S. litorale	909 (834–988)	31 (28–33)	61 (54–69)	96 (89–104)	125 (114–133)	83 (72–91)	29.5 (27–31)	7.3 (6.7–7.9)	11 (9.7–11.9)	49 (44–56)	73 (68–84)	Yoshida (2004)
S. tielingense	915 (824–979)	35 (32–38)	69 (64–73)	98 (90–105)	128 (120–135)	81 (74–85)	26 (23–28)	7 (6–8)	11 (9–13)	55 (47–61)	88 (85–94)	Ma et al. (2012)
S. kraussei	951 (797–1,102)	33 (30–36)	63 (50–66)	105 (99–111)	134 (119–145)	79 (63–86)	29 (NA)	7.1 (NA)	12.1 (NA)	47 (NA)	80 (NA)	Nguyen et al. (2007)
S. oregonense	980 (820–1,110)	34 (28–38)	66 (60–72)	NA	132 (116–148)	70 (64–78)	30 (24–37)	7.6 (6–8)	14 (12–16)	50 (40–60)	100 (90–110)	Liu and Berry (1996)

Comparative morphometrics of first-generation males of S_ sandneri n_ sp_ and related Steinernema spp_

	Morphometric character ^a
Species	SL	GL	W	D%	SW%	GS%	MUC ^b	n
S. sandneri n. sp.	60 (53–65)	44 (39–50)	155 (124–178)	51 (42–59)	111 (97–127)	79 (61-–83)	P	25
S. akhursti	90 (85–100)	64 (58–68)	131 (115–150)	56 (52–61)	180 (140–200)	71 (65–77)	P	20
S. cholashanense	66 (60–71)	39 (32–45)	137 (73–204)	64 (50–85)	115 (92–144)	71 (61–85)	P	20
S. citrae	65 (57–80)	44 (32–59)	103 (87–113)	58 (47–67)	198 (156–233)	68 (48–89)	P	20
S. costaricense	92 (81–101)	46 (41–51)	128 (89–157)	53 (51–66)	160 (150–170)	49 (45–55)	A	19
S. feltiae	70 (65–77)	41 (34–47)	75 (60–90)	60 (51–64)	113 (99–130)	59 (52–61)	P	25
S. hebeiense	57 (51–63)	46 (38–50)	86 (74–98)	51 (48–59)	140 (120–170)	80 (60–90)	A	20
S. ichnusae	66 (64–67)	44 (43–46)	137 (73–204)	62 (59–65)	139 (120–162)	67 (64–69)	A	20
S. jollieti	64 (55–70)	54 (45–60)	115 (98–135)	64 (53–83)	145 (NA)	84 (NA)	A	12
S. kraussei	49 (42–53)	33 (29–37)	128 (110–144)	53 (NA)	110 (NA)	67 (NA)	P	NA
S. kushidai	63 (48–72)	44 (39–60)	97 (75–156)	51 (42–59)	150 (NA)	70 (NA)	A	20
S. litorale	75 (67–89)	53 (44–64)	96 (82–111)	40 (34–56)	174 (154–200)	71 (62–81)	P	25
S. nguyeni	66 (58–75)	43 (30–55)	82 (58–106)	48 (38–57)	215 (185–279)	66 (46–81)	P	20
S. oregonense	71 (65–73)	56 (52–59)	138 (105–161)	73 (64–75)	151 (NA)	79 (NA)	A	20
S. puntauvense	77 (71–81)	34 (30–40)	119 (101–139)	67 (45–85)	170 (140–200)	65 (55–75)	P	19
S. sangi	63 (58–80)	40 (34–46)	159 (120–225)	49 (42–63)	150 (120–160)	60 (50–70)	P	20
S. silvaticum	51 (42–64)	37 (30–43)	65 (52–78)	60 (45–63)	NA	NA	P	26
S. texanum	60 (55–66)	45 (39–53)	99 (81–116)	67 (58–73)	157 (127–203)	75 (62–84)	A	20
S. tielingense	88 (79–98)	62 (49–70)	129 (111–159)	71 (64–78)	191 (176–212)	73 (59–82)	A	20
S. weiseri	68 (62–72)	53 (46–57)	112 (84–138)	49 (39–60)	180 (150–240)	80 (70–85)	A	20
S. xinbinense	56 (49–62)	35 (30–41)	103 (90–126)	45 (41–50)	137 (114–156)	63 (54–72)	P	20
S. xueshanense	76 (66–91)	49 (41–60)	144 (97–159)	80 (73–87)	152 (93–172)	64 (58–95)	A	20

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Steinernema sandneri n. sp. (Rhabditida: Steinernematidae), a new entomopathogenic nematode from Poland

Figures & Tables

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Figure 3.

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Figure 7:

Figure 8:

Morphometrics (in μm) of different developmental stages of Steinernema sandneri n_ sp_ [mean ± SE (range)] [N = 25]_

Percentage of similarity (upper triangle) and genetic distance measured by the number of nucleotide substitutions (lower triangle) in the sequences of cox1 gene of S_ sandneri n_ sp_ and other closely related Steinernema spp_

Percentage of similarity (upper triangle) and genetic distance measured by the number of nucleotide substitutions (lower triangle) in the sequences of ITS rDNA of S_ sandneri n_ sp_ and other closely related Steinernema spp_

Percentage of similarity (upper triangle) and genetic distance measured by the number of nucleotide substitutions (lower triangle) in the sequences of D2D3 domain of 28 S rDNA of S_ sandneri n_ sp_ and other closely related Steinernema spp_

Steinernema sandneri n_ sp_ – permanent slides description and designation numbers in the collection of Museum and Institute of Zoology, Polish Academy of Sciences, Warsaw, Poland_

Percentage of similarity (upper triangle) and genetic distance measured by the number of nucleotide substitutions (lower triangle) in the sequences of ITS rDNA regions of S_ sandneri n_ sp_, S_ kraussei and S_ silvaticum isolates, the closest relatives_

Details on taxa used in the molecular analyses_

Comparative morphometrics of third-stage infective juveniles of S_ sandneri n_ sp_ and related Steinernema spp_

Comparative morphometrics of first-generation males of S_ sandneri n_ sp_ and related Steinernema spp_

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