Научная статья на тему 'Morphological and molecular characterisation of several known nematode species of the genera Criconema, Criconemoides and Mesocriconema (Tylenchida: Criconematidae) from the USA and South Africa'

Morphological and molecular characterisation of several known nematode species of the genera Criconema, Criconemoides and Mesocriconema (Tylenchida: Criconematidae) from the USA and South Africa Текст научной статьи по специальности «Биологические науки»

CC BY
64
16
i Надоели баннеры? Вы всегда можете отключить рекламу.
Журнал
Russian Journal of Nematology
WOS
Scopus
ВАК
Область наук
Ключевые слова
& Zheng / 2022 were transferred to the genus Criconemoides. Key words: Criconema annuliferum / Criconema mutabile / Criconemoides annulatus / Criconemoides informis / COI gene / D2-D3 of 28S rRNA gene / Mesocriconema nebraskense / Mesocriconema sphaerocephalum / Mesocriconema xenoplax / phylogeny / SEM

Аннотация научной статьи по биологическим наукам, автор научной работы — Esther Van Den Berg, Louwrens R. Tiedt, Sergei A. Subbotin

During nematological surveys in several locations of the USA and South Africa, and using integrative approach combining morphological and molecular analyses the following species were identified: Criconema annuliferum, Criconema mutabile, Criconemoides annulatus, C. informis, Criconemoides sp. A, Mesocriconema nebraskense, M. sphaerocephalum, M. xenoplax and Mesocriconema sp. A. These species were morphologically and morphometrically described and the SEM images were also given for some species. Molecular characterisations of the species using the D2-D3 expansions segments of 28S rRNA and COI mtDNA gene sequences were also provided. Based on the results of molecular dataset analysis, Discocriconemella sinensis Munawar, Cai, Subbotin &; Zheng, 2019 and D. parasinensis Li, Munawar, Castillo &; Zheng, 2022 were transferred to the genus Criconemoides.

i Надоели баннеры? Вы всегда можете отключить рекламу.
iНе можете найти то, что вам нужно? Попробуйте сервис подбора литературы.
i Надоели баннеры? Вы всегда можете отключить рекламу.

Морфологическая и молекулярная характеристика нескольких известных видов нематод родов Criconema, Criconemoides и Mesocriconema (Tylenchida: Criconematidae) из США и Южной Африки

В ходе нематологических обследований в нескольких регионах США и Южной Африки и с использованием интегративного подхода, сочетающего морфологический и молекулярный анализы, были обнаружены следующие виды: Criconema annuliferum, Criconema mutabile, Criconemoides annulatus, C. informis, Criconemoides sp. A, Mesocriconema nebraskense, M. sphaerocephalum, M. xenoplax и Mesocriconema sp. A. Эти виды были описаны морфологически и морфометрически, а также для некоторых видов даются СЭМ фотографии. Также предоставлены молекулярные характеристики видов с использованием последовательностей генов сегмента D2-D3 28S рРНК и COI мтДНК. По результатам молекулярного анализа Discocriconemella sinensis Munawar, Cai, Subbotin &; Zheng, 2019 и D. parasinensis Li, Munawar, Castillo &; Zheng, 2022 переведены в род Criconemoides.

Текст научной работы на тему «Morphological and molecular characterisation of several known nematode species of the genera Criconema, Criconemoides and Mesocriconema (Tylenchida: Criconematidae) from the USA and South Africa»

Russian Journal of Nematology, 2023, 31 (2), 139 - 159

Morphological and molecular characterisation of several known nematode species of the genera Criconema, Criconemoides and Mesocriconema (Tylenchida: Criconematidae) from the USA and

South Africa

Esther Van den Berg1, Louwrens R. Tiedt2 and Sergei A. Subbotin3' 4

'National Collection of Nematodes, Biosystematics Programme, ARC-Plant Protection Research Institute, Private Bag X134,

0121, Queenswood, South Africa ^Laboratory for Electron Microscopy, North-West University, Potchefstroom Campus, 2520, Potchefstroom, South Africa 3Plant Pest Diagnostic Centre, California Department of Food and Agriculture, 3294 Meadowview Road,

95832-1448, Sacramento, CA, USA 4Center of Parasitology of A.N. Severtsov Institute of Ecology and Evolution, Russian Academy of Sciences, Leninskii

Prospect 33, Moscow, 119071, Russia e-mail: sergei. a. subbotin@gmail. com

Accepted for publication 08 August 2023

Summary. During nematological surveys in several locations of the USA and South Africa, and using integrative approach combining morphological and molecular analyses the following species were identified: Criconema annuliferum, Criconema mutabile, Criconemoides annulatus, C. informis, Criconemoides sp. A, Mesocriconema nebraskense, M. sphaerocephalum, M. xenoplax and Mesocriconema sp. A. These species were morphologically and morphometrically described and the SEM images were also given for some species. Molecular characterisations of the species using the D2-D3 expansions segments of 28S rRNA and COI mtDNA gene sequences were also provided. Based on the results of molecular dataset analysis, Discocriconemella sinensis Munawar, Cai, Subbotin & Zheng, 2019 and D. parasinensis Li, Munawar, Castillo & Zheng, 2022 were transferred to the genus Criconemoides. Key words: Criconema annuliferum, Criconema mutabile, Criconemoides annulatus, Criconemoides informis, COI gene, D2-D3 of 28S rRNA gene, Mesocriconema nebraskense, Mesocriconema sphaerocephalum, Mesocriconema xenoplax, phylogeny, SEM.

During nematological surveys, several species of the genera Criconema Hofmänner & Menzel, 1914, Criconemoides Taylor, 1936 and Mesocriconema Andrassy, 1965 were found in South Africa and several states of the USA. Some species of these genera have been reported as parasitic and important pests of crops, causing damage of roots (Geraert, 2010). The genus Criconema currently contains more than 100 valid species (Geraert, 2010; Azimi & Pedram, 2020; Clavero-Camacho et al., 2022; Archidona-Yuste et al., 2023), the genus Criconemoides consists of nearly 50 species (Geraert, 2010; Munawar et al., 2020; Hosseinvand et al., 2023), whereas the genus Mesocriconema contains more than 90 species. Mesocriconema species are morphologocally similar to

Criconemoides species and differ from it by having an open vulva and submedian lobes arising from reduced pseudolips (Brzeski et al., 2002a, b; Geraert, 2010; Karani et al., 2020).

The objectives of this work were: i) to carry out a morphological and morphometric characterisation of several criconematids found from the USA and South Africa; ii) to provide molecular characterisation of these species using sequences of the D2-D3 expansion segments of the 28S nuclear ribosomal RNA and partial COI gene; and iii) to analyse phylogenetic relationships of several criconematids from the USA and South Africa within representatives of the genera Criconema, Criconemoides and Mesocriconema using these genes.

© Russian Society of Nematologists, 2023; doi: 10.24412/0869-6918-2023-2-139-159

MATERIALS AND METHODS

Nematode samples, light and scanning electron microscopic study. Soil samples were collected from the rhizosphere of different plans and locations as indicated in Table 1. Nematode specimens were extracted from samples using the rapid centrifugal-flotation method (Jenkins, 1964), fixed in 4% formalin or FPG (Netscher & Seinhorst, 1969), transferred to anhydrous glycerin (De Grisse, 1969) and mounted on Cobb's slides. Voucher nematode slides were deposited at the Nematology collection, ARC-Plant Protection Research Institute, Queenswood, South Africa. Measurements were made with a research microscope (Nikon Labophot-2) equipped with a drawing tube. Light micrographs were taken with an automatic Infinity 2 camera attached to a compound Olympus BX51 microscope equipped with Nomarski differential interference contrast. For scanning electron microscopy specimens were transferred to TAF (7 ml 40% formalin, 2 ml triethanolamine, and 91 ml distilled water) then dehydrated in increasing concentrations of alcohol in distilled water and finally into pure alcohol. Following conventional critical point drying and gold/palladium coating (15 nm), specimens were viewed with a FEI

ESEM Quanta 200 scanning electron microscope at 10 kV (Van den Berg et al., 2017).

DNA extraction, PCR, sequencing and phylogenetic analysis. DNA was extracted from several specimens of each species using the proteinase K protocol. Detailed protocols for DNA extraction, PCR, cloning and sequencing were as described by Subbotin (2021). The D2-D3 expansion segments of 28S rRNA gene and partial COI gene were amplified and sequenced. The following primers were used for amplification in the present study: D2-D3 of 28S rRNA gene - D2A (5'-ACA AGT ACC GTG AGG GAA AGT TG-3') and D3B (5'-TCG GA GGA ACC AGC TAC TA-3') (Subbotin et al., 2005) and partial COI gene -COIF5 (5'-AAT WTW GGT GTT GGA ACT TCT TGA AC-3') and COIR9 (5'-CTT AAA ACA TAA T GR AAA TGW GCW ACW ACA TAA TAA GTA TC-3') (Powers et al., 2014). The PCR products were purified using QIAquick (Qiagen) Gel or PCR extraction kits and submitted for direct sequencing or cloned using pGEM-T Vector System II kit (Promega). Sequencing was conducted at Quintara Biosciences. The obtained sequences were submitted to the GenBank database under the following accession numbers: OR157945-OR157956 (COI gene) and OR159851-OR159868 (28S rRNA gene) (Table 1).

Table 1. Criconematid species used in the present study.

Species Locality Sample code GenBank accession numbers

D2-D3 of 28S rRNA gene COI gene

Criconema annuliferum USA, California, El Dorado County CD878, CD879 OR159851 OR157946

C. annuliferum USA, Washington CD920 OR159852 OR157945

C. mutabile South Africa, Gauteng Province, Pretoria, Roodeplaat Nature Reserve Tvl1982 - -

Criconemoides annulatus USA, California, El Dorado County CD879 OR159853 OR157947

C. informis USA, California, Sacramento County CD836 - OR157956

Criconemoides sp. A South Africa, KwaZulu Natal Province, Nottingham Road CD370, CD554; N775 OR159854-OR159857 OR157950, OR157951

Mesocriconema nebraskense USA, Kansas, Manhattan, Washington Marlatt park CD869 OR159862 OR157952

M. sphaerocephalum USA, Florida, Gainesville CD1183 OR159858, OR159859 OR157948, OR157949

M. xenoplax USA, California, Marin County CD859 OR159867, OR159868 -

M. xenoplax South Africa, Gauteng Province, Tarlton CD553; Tyl1976 OR159863, OR159864 OR157955

M. xenoplax USA, Kansas, Manhattan, Washington Marlatt park CD863, CD865 OR159865, OR159866 -

M. xenoplax South Africa, Mpumalanga Province, Groblersdal Tvl1928 - -

Mesocriconema sp. A USA, Florida, Gainesville CD1182 OR159860, OR159861 OR157953, OR157954

The newly obtained sequences for each gene were aligned using ClustalX 1.83 with default parameters with corresponding published gene sequences of selected species of the genera Criconemoides, Criconema and others (Subbotin et al., 2005; Powers et al., 2014, 2016, 2021; Van den Berg et al., 2017; Munawar et al., 2019; Clavero-Camacho et al., 2022; Li et al., 2022; Hosseinvand et al., 2020, 2023 and others). Outgroup taxa for each dataset were chosen according to the results of previously published data (Subbotin et al., 2005). Sequence datasets were analysed with Bayesian inference (BI) using MrBayes 3.1.2 (Ronquist & Huelsenbeck, 2003). BI analysis under the GTR + I + G model for each gene was initiated with a random starting tree and was run with four chains for 1.0 x 106 generations. The Markov chains were sampled at intervals of 100 generations. Two runs were performed for each analysis. After discarding burn-in samples and evaluating convergence, the remaining samples were retained for further analysis. The topologies were used to generate a 50% majority rule consensus tree. Posterior probabilities (PP) are given on appropriate clades. Pairwise divergences between taxa were computed as absolute distance values and as percentage mean distance values based on whole alignment, with adjustment for missing data using PAUP* (Swofford, 2003).

RESULTS AND DISCUSSION

Within studied samples using traditional morphological taxonomic characters integrated with molecular criteria, we distinguished seven valid criconematid species: Criconema annuliferum, Criconema mutabile, Criconemoides annulatus, Criconemoides informis, Mesocriconema nebraskense, M. sphaerocephalum, M. xenoplax, and two unidentified species: Mesocriconema sp. A and Criconemoides sp. A. Seven of these species were found in the USA and three species were reported from South Africa (Table 1). Short descriptions of some species are given below.

Criconema annuliferum (de Man, 1921) Micoletzky, 1925 (Figs 1 & 2)

Specimens of this species were found in several samples collected in Washington and California (Table 1). This species was originally obtained from soil covered with herbs and anemone in a forest near Breda, The Netherlands and from a municipal park

of Bergen op Zoom, The Netherlands and described as Hoplolaimus annulifer by de Man (1921). Micoletzky (1925) transferred it to the genus Criconema. Criconema annuliferum is widely distributed in several European countries, Africa, Asia, New Zealand and South America (Clavero-Camacho et al. , 2022). Several species had been synonymised with this species (Geraert, 2010). Using the integrative taxonomical analyses, Clavero-Camacho et al. (2022) and Archidona-Yuste et al. (2023) distinguished the C. annuliferum species complex, which includes four cryptic species: C. annuliferum, C. paraannuliferum Clavero-Camacho et al., 2022, C. plesioannuliferum Clavero-Camacho et al. , 2022 and C. pseudoannuliferum Archidona-Yuste et al., 2023.

Measurements. See Table 2.

Female. Female body almost smooth to slightly curved ventrad. Lip region with two annuli. First annulus with a smooth margin and projecting outward or slightly upward. Second lip annulus strong, pointed outward and with a slightly smaller diameter than first annulus. Labial plate rounded, protruding slightly above lip first lip annulus. Submedian lobes absent. All body annuli slightly retrorse with smooth margins. An occasional anastomosis posterior to vulva, otherwise no markings in the lateral field. Stylet long and slender with cupped basal knobs. Excretory pore situated from one to four annuli posterior to base of pharyngeal lobe. Hemizonid not seen except in one specimen, it is one annulus long and situated directly anterior to excretory pore. Hemizonion not seen. Spermatheca indistinct in all specimens. Anterior vulval lip slightly indented in middle, overlapping the posterior lip. Vagina sigmoid. Anus distinct four to five annuli from terminus. Tail tapering to a narrow rounded knob. Last annulus sometimes slightly irregular.

Males. Not found.

Juveniles. Not found.

Remarks. The present specimens are morphologically and morphometrically very similar to the specimens described by various authors from different locations (Raski & Golden, 1965; Gomez Barcina et al., 1989, 1991; Brzeski, 1998; Peneva et al., 2000; Etongwe et al., 2020) (Table 2).

Molecular characterisation. The D2-D3 of 28S rRNA (Fig. 3) and COI (Fig. 4) gene sequences of the Washington and California populations clustered with those of C. annuliferum populations from Belgium and Ireland. Intraspecific variation for the D2-D3 of 28S rRNA gene was 0.1-0.7% and for COI gene - 0-2.1%.

Fig. 1. Criconema annuliferum. CD920. Female. A: Anterior part of body; B: Ventral view of posterior region; C: Lateral view of posterior region; D: Annuli at midbody. CD879 Female. E: Anterior part of body; CD878. Female. F: Anterior part of body; G: Lateral view of posterior part of body; H: Ventral view of posterior part of body; I, J: Annuli at midbody. CD879 Female. K: Annuli at midbody; L: Lateral view of posterior part of body. Scale bar = 30 ^m.

Table 2. Measurements of Criconema annuliferum females compared with those from the literature. All measurements are in |im and in the form: mean ± SD (range).

Sample Character USA, Washington (CD920) USA, CaUfornia (CD878) USA, CaUfornia (CD879) Van den Berg & Heyns (1977) Gomez Barcina et al. (1989) References (a)

n 10 10 3 11 18 27

L 562 ±40 (507-616) 567 ±37.5 (510-626) 604-646 539 ± 17.1(503-558) 469 ±49 (348-541) 469-770

a 10.1 ± 1 (8.1-11.3) 9.4 ± 1.1 (8-11.5) 9.8-11.1 8 ±0.7 (7-10) 11 ±1.8 (9-14) 7.7-12

b 3.7 ±0.1 (3.5-3.9) 3.7 ±0.2 (3.5-4) 3.9-4.3 4 ±0.1 (3-4) 4 ±0.3 (3-4) 3-5

c 23 ±3(17.8-28.7) 31.4 ±2 (20.7-44.8) 27.6-41.1 27 ±4.7 (22-38) 16 ±4.4 (11-23) 17-34.3

o 7.6 ±0.5 (7-8.3) 8.8 ±0.7 (7.8-9.8) 8 8 ±0.9 (6-8) - -

DGO 8 ±0.4 (7.5-9) 9 ±0.4 (8-9.5) 8 8 ± 1 (6-9) - -

V 89 ±0.8 (88-90.5) 90 ± 1 (88-91) 90-92 89 ± 0.6 (89-90) 87 ± 11(85-89) 87-91

OV1 50 ± 10(41-67.5) 50.7 ±8.4 (40.5-65.4) 43-70 43 ± 8.2 (36-45) 42 ± 8.3 (32-60) -

OV length 281 ± 58.3 (227-380) 297 ±45.9 (250-307) 261-423 - - -

Stylet length 106.5 ±5.6 (98.5-116) 100 ±45.9 (93.5-109) 95-101 107± 2.6 (102-111) 100 ± 5.8 (88-107) 90-114

Metenchium length 88.5 ±4.8 (81.5-98.5) 82.5 ±3.9 (77-89) 76.5-84 87 ±2.1 (82-89) 86 ± 1.4(84-94) -

Telenchium length 17.5 ±2.1 (12.5-20) 18 ±0.9 (17-19) 17.5-20 20 ±0.5 (20-21) - -

in 83 ± 2.8 (76.3-87.3) 82 ±0.6 (81-83) 80.6-82.6 - - -

Stylet knob height 5 ± 0.6 (4-6) 5.5 ±0.7 (4.5-6.5) 6 5 ±0.5 (4-6) - -

Stylet knob width 12 ± 1 (10.5-13) 12 ± 0.5 (11-12.5) 12.5-13 12 ± 0.7 (11-13) 8-9 -

Exc. pore from anter. end 169.5 ± 17.6(144-195) 181 ± 12 (165-201.5) 179-187 165 ±34.4 (159-197) 158 ±9 (141-170) 150-185

Width at midbody 56 ± 7 (49-72) 61 ±6.2 (50-68) 58-62 67 ±4.8 (57-74) 41 ±4.2(36-50) -

Width at anus 27 ± 2.4 (23.5-32.5) 24 ±3.2 (19-29) 23.5 - - -

Width at vulva 42 ±4.2 (33-48.5) 45 ± 5.7 (34-50) 42.5-47 - - -

Annulus width 9.5 ±0.6 (9-11) 10 ± 1.1 (8-11) 9-10 9 ±0.8 (7-10) - 9-10

Tail length 24.5 ±2.1 (21.5-28.5) 19 ±3.8 (14-25.5) 15-22 21 ±3.2 (15-25) 32 ± 7.9 (22-47) 16-26

Pharynx length 151.5 ±7.6 (137.5-162.5) 154 ±9.2 (143-171) 147-157 150 ±7.8 (135-160) 132 ±9.5 (117-146) -

1st Up annulus diam. 22.5 ± 1.9(19-25) 22.5 ± 1.4(19-24) 22-24 24 ± 1.3 (22-26) 21 ±3 (15-25) -

2nd Up annulus diam. 21.5 ±2.1 (19-25.5) 20.5 ± 1.3(18.5-23) 20-21.5 22 ± 1.8(19-25) - -

1st body annulus diam. 30 ±2.3 (28-35.5) 31 ±2.6 (27-34) 31-32 34 ±2.6 (30-38) - -

2nd body annulus diam. 35.5 ±2.8 (31.5-41.5) 37 ±3.7 (31-41) 36.5-38 38 ± 11.5 (38-46) - -

R 65 ± 2.6 (62-70) 62 ± 2.4 (60-68) 63-66 66 ± 1.7(62-68) 65 ± 1.7(62-68) 61-74

RSt 15 ±0.9 (13-16) 13 ±0.7 (12-14) 11-13 14± 1.8(12-18) 15 ± 1.4(13-18) 12-15

ROes 19 ± 1.3 (18-21) 18 ± 1.5 (17-21) 17-18 19 ± 1.3 (17-21) 20 ± 1.7(16-24) 16-20

Rex 21 ± 1.2(19-23) 21 ± 1.2(20-24) 20-21 22 ± 1(20-23) 21 ± 1.5(19-24) 19-26

RV 8.5 ±0.5 (8-9) 8 ± 0.7 (7-9) 7-8 9 ±0.4 (8-9) 10 ± 0.8 (8-11) 6-11

RVan 3± 0.4 (3-4) 3± 0.8 (2-4) 3-4 3± 0.3 (3-4) 4 ± 1.2(2-6) 3-4

Ran 4 ±0.4 (4-5) 3 ± 0.6 (2-4) 3 5 ±0.5 (4-5) 7± 1.1 (5-8) 3-7

VL/VB 1.4 ± 0.08 (1.2-1.5) 1.3 ±0.1 (1.1-1.5) 1.2-1.4 1 1.8 ±0.1 (1.5-2) 1.1-1.7

St%L 19.1 ± 1.1 (16.3-20.3) 17.8 ± 1 (16.5-18.9) 15.7-17.8 20 ±0.7 (19-21) - 15.8-20.2

Note: (a) Raski and Golden (1965), Szczygiel (1974) and Peneva et al. (2000).

Fig. 2. Criconema annuliferum. CD878. Female. LM. A: Whole body; B: Anterior part of body; C: Posterior part of body. SEM. D, E: En face view of lip region; F: Annuli at midbody; G: Tail tip; H: Lateral view of posterior part of body. Scale bars: A-C = 60 ^m.

Criconema mutabile (Taylor, 1936) Raski & Luc, 1985 (Figs S1A-H & S2)

Taylor (1936) described this species from African marigold (Tagetes erecta L.) from glasshouses of the Department of Agriculture in

Washington D C., USA. Raski and Golden (1965) redescibed C. mutabile from the original materials used by Taylor (1936) and since then the species have been moved to a few other genera until 1985 when Raski and Luc (1985) in their reappraisal of the genus regarded mutabile as belonging to the genus Criconema. Several species have been consi-

r Mesocriconema sp.A (0R159860, CD1182a) ^ Mesocriconema sp.A (OR159861, CD1182b) _r Mesocriconema nebraskense (OR159862, CD869)

100 Mesocriconema nebraskense (MH013430) — Mesocriconema xenoplax (AY780963) r Mesocriconema xenoplax (OR159868, CD859, cl2) Mesocriconema xenoplax (MN783690) Mesocriconema xenoplax (OR159866, CD865, cl2) Mesocriconema xenoplax (OR159865, CD865, cl1) Mesocriconema xenoplax (OR159867, CD859, cl1) r Mesocriconema xenoplax (FN433860)

Mesocriconema xenoplax (OR159863, CD553, cl2)

- Mesocriconema xenoplax (OR159864, CD553, cl1) Mesocriconema xenoplax (OQ67427)

i— Mesocriconema sp. (MN462849)

— Mesocriconema ornatum (AY780968) - Mesocriconema antipolitanum (MN888461)

Criconemoides geraerti (MN738727) Criconemoides parasinensis comb. n. (MZ820651) Criconemoides parainformis (MN738723) Criconemoides sinensis comb. n. (MZ011403) Criconemoides informis (MN783679) Criconemoides neoinformis (ON3Q3845)

— Neobakernema variabiie (MF683232) Mesocriconema solivagum (AY780969) jj Mesocriconema sphaerocephalum (OR159859, CD1183a) — ■ Mesocriconema sphaerocephalum (OR159858, CD1183b)

Mesocriconema sphaerocephalum (AY780950) - Mesocriconema sphaerocephalum (AY780951)

- Criconemoides obtusicaudatus (JQ231187)

r Criconemoides sp.A (OR159854, CD370, cl2) Criconemoides sp.A (OR159855, CD370, cM)

iНе можете найти то, что вам нужно? Попробуйте сервис подбора литературы.

Criconemoides sp.A (OR159856, CD554, cl1) Criconemoides sp.A (OR159857, CD554, cl2)

Discocriconemella limitanea (MK253538) 1001— Criconemoides sp. (MW938506)

Criconemoides myungsugae (MZ041096) Criconemoides sp.* (OM938250) Criconemoides sp. (MW938508) - Criconemoides sp." (MK253542)

- Criconemoides annulatus (OR159853, CD879)

— Criconemoides persicus (ON3C13844) j— Criconemoides parvus (MN888467)

ioor— Cr r^-P- Cr, p-^pL Cric

'- Pr/onni

Criconemoides sp.** (MK253540) Criconemoides rotundicaudatus (MN738728)

- Criconemoides sp ** (MK253541)

- Criconemoides sp. (MW938510)

Criconema annuliferum (MN783702) Criconema annuliferum (OR159851, CD878) Criconema annuliferum (MN783699) Criconema annuliferum (OR159852, CD920) Criconema pseudoannuliferum (ON877364) Criconema pseudoannuliferum (ON877367 Criconema demani (MH828126) Criconema paraannuliferum (ON705079) ■ Criconema paraannuliferum (ON705071) - Xenocriconemella macrodora (AY780960)

71

100

100 -

100 ^

- Criconema silvum (MF683234)

Crossonema civellae (MN888468) - Criconema mutabiie (MK481268)

— Paratylenchus dianthus (KF242228)

- Paratylenchus straeleni (KF242236)

Paratylenchus hamatus (KF242209)

0.1

Fig. 3. Phylogenetic relationships within some representatives of the family Criconematidae. Bayesian 50% majority rule consensus tree from two runs as inferred from analysis of the D2-D3 of 28S rRNA gene sequence alignment under the GTR + I + G model. Posterior probabilities equal to, or more than, 70% are given for appropriate clades. New sequences are indicated in bold letters. * - identified as Criconemoides annulatus in the GenBank by Zhao (unpublished); ** - identified as Mesocriconema sp. by Munawar et al. (2019).

-Criconema sp. (MN710764)

-Criconema sphagni (MN710795)

21-Criconema longulum (MN710700)

-Criconema sp. (MN710756)

-Criconema acriculum (MN710669)

-Criconema petasum (MN710751 )

100 -Criconema mutabiie (MN710704)

_!!_ -Criconema sp. (MN710777)

-Criconema permistum (OK561943)

L 100i— Criconema sp. (MN710768) n l— Criconema plesioannuliferum (ON648875)

-Criconema paraannuliferum (ON648825)

Criconema annuliferum (OR157945, CD920, Washington, USA) — Criconema annuliferum (MN782388, Belgium) Criconema annuliferum (MN782389, Belgium)

-—— Criconema annuliferum (MN782387, Belgium)

Criconema annuliferum (OR157946, CD878, California, USA) Criconema annuliferum* (MN710781, Ireland) Circonema annuliferum (MN782395, Belgium)

-Criconema crotaloides (MN710681)

_Ji I-Criconema sp. (MN710770)

-Criconema sp. (MN710775)

_9oj-Criconema loofi (MN710703)

-Criconema pseudoannuliferum (ON899944)

-Paratylenchus baldaccii (MZ262220)

Paratylenchus hamatus (MW797016)

Fig. 4. Phylogenetic relationships within some representatives of the genus Criconema. Bayesian 50% majority rule consensus tree from two runs as inferred from analysis of the COI gene sequence alignment under the GTR + I + G model. Posterior probabilities equal to, or more than, 70% are given for appropriate clades. New sequences are indicated in bold letters. * - identified as Criconemoides sp. in the GenBank and Powers et al. (2021).

dered as synonyms of C. mutabile (Geraert, 2010). Criconema mutabile was found in many different countries and it was reported very often from agricultural soil and natural veldt in South Africa. Recently, this species was morphologically and molecularly characterised by Shokoohi et al. (2020) from South Africa and Iran. The present specimens were collected in 2009 in the Gauteng Province from Roodeplaat Nature Reserve, 2 m from the side of the Roodeplaat dam amongst grass and weeds (Tvl1982) and used for morphological study only.

Measurements. See Table S1.

Female. Body form straight to slightly curved ventrad. Lip region with two annuli, first one pointing outward and second slightly retrorse. First with a smaller diameter than second. Lip annuli not set off from body annuli. SEM photographs show labial area with pseudolips projecting well above first lip annulus. All body annuli retrorse with smooth margins. Lateral field not demarcated except for rarely with a few irregularities at midbody or one or two anastomosis posterior to vulva and one with a few indents anterior to vulva. Stylet well developed with cupped basal knobs. Excretory pore

situated from four annuli anterior to three annuli posterior to posterior margin of pharyngeal lobe. Hemizonid one annulus long and situated from opposite to four annuli anterior to excretory pore. Hemizonion not seen. Spermatheca small, round to oblong and empty. Vulval lips not protruding and closed. Anterior lip not indented or overhanging the posterior lip. Vagina straight. Anus six to eight annuli from tail tip. Tail narrowing gradually to a rounded tip with a few small lobes.

Male. Not found.

Juvenile. Not found.

Remarks. The specimens are similar to those from all the previous descriptions of the species (Table S1). Molecular analysis of this sample is not given.

Criconemoides annulatus Cobb in Taylor, 1936 (Figs S3 & 5)

Nine specimens of this species were collected in California. Criconemoides annulatus was originally collected from soil around Scrub Oak (Quercus ilicifolia Wangenh.) near Red Butte, Montana, USA, described by Taylor (1936) and re-described using the type specimens and collections from California and Idaho by Raski and Golden (1965). Criconemoides annulatus was recorded from various areas in North America, also Canada, Asia and Europe. Several species had been synonymised with C. annulatus (Choi et al., 2000; Brzeski et al., 2002a; Geraert, 2010; Powers et al., 2021).

Measurements. See Table S2.

Female. Body slightly curved ventrad. Lip region with three annuli, first quite smaller than following two, all three slightly retrorse and not set off from body. Labial area low and not projecting above the first annulus. Lip region covered with bacteria and not much can be seen in the en face view. All body annuli slightly retrorse with smooth to slightly wavy margins. In a few cases the annuli appeared to have very fine longitudinal lines. Anastomosis rare, sometimes a few are visible on the first few annuli or a few may be present posterior to the vulva. Stylet long, sturdy with cupped basal knobs. Excretory pore situated from one annulus anterior to four annuli posterior to basal margin of basal pharyngeal lobe. Hemizonid rarely seen but where seen, it is one annulus long and situated opposite the excretory pore. Hemizonion not seen. Spermatheca indistinct and not containing sperm cells. Vulva an oval open slit. Vagina straight. Anus distinct two to five annuli from tail tip. Tail rounded with small irregular amalgamated lobes.

Male. Not found.

Juvenile. Very similar to female. Lateral field with a few irregularities and a few anastomosis. Posterior margins of annuli slightly crenate.

Remarks. These specimens compare well with those from the literature. Several authors reported a large variation in some of the morphological characters in this species such as stylet length, number of body annuli, Rex and RV values etc. (Raski & Golden, 1965; Popovici, 1988; Choi et al., 2000). Table S2 gives measurements of the populations of this species provided by various authors as compared with original ones.

Molecular characterisation. The D2-D3 of 28S rRNA gene sequence of the California population of C. annulatus clustered with that of the Washington population (Fig. 3) and they were different in 2.7% (15 bp). The COI gene sequence of the California population of C. annulatus clustered with those of the Utah, Wyoming, Colorado, South Dakota and Quebec populations (Fig. 6). Intraspecific COI gene sequence variation reached 11.2% (81 bp).

Criconemoides informis (Micoletzky, 1922) Taylor, 1936 (Figs S4 & 7A-D)

Specimens of this species were collected in California, Sacramento County. Unfortunately, most specimens were covered with bacteria and not well preserved and mounted, and only two specimens could be measured. This species was described by Micoletzky (1922) from soil about the roots of aspen (Populus tremuloides Michx.) near Idaho Springs, Clear Creek, Colorado. Then this species was frequently reported from Europe, North America and Asia by various authors (Loof, 1965; Gomez Barcina et al., 1989; Choi et al., 2000; Eskandari et al., 2010; Geraert, 2010). Hosseinvand et al. (2023) distinguished the Criconemoides informis group with several species: C. informis, C. amorphous De Grisse, 1967; C. parainformis Munawar et al. , 2020; C. neoinformis Hosseinvand et al., 2023 and C. geraerti Munawar et al., 2020.

Measurements. See Table S2.

Female. Body slightly curved ventrad. Lip region with two annuli, first pointing slightly upward and second pointing outward. First lip annulus with a smaller diameter than second. SEM shows labial disc protruding above first lip annulus, sub median lobes more broad and rounded, labial plates not very well developed. Diameter of first lip annulus markedly smaller than that of the second lip annulus. Lip annuli not set off from body annuli. All

Fig. 5. Criconemoides annulatus. CD879. LM. Female. A. Whole body; B-D: Anterior part of body; E-G: Posterior part of body. SEM. H, I: Lateral view of anterior part of body; J, M: En face view of lip region; K: Annuli at midbody; L: Ventral view of tail region. Scale bars: A = 30 ^m, B-G = 50 ^m.

S3

01 t1

100 j Mesocriconema xenoplax (KJ787885) 98 P Mesocriconema xenoplax (OR157955, CD553)

lr Mesocriconema xenoplax (KY574637) L Mesocriconema xenoplax (KY574634) Lr Mesocriconema xenoplax (OP431940) Mesocriconema xenoplax (OP418003) !2j- Mesocriconema xenoplax (OK561968) Mesocriconema xenoplax (KY574627) — Mesocriconema xenoplax (MN711215) Mesocriconema xenoplax (KY574643) -)— Mesocriconema xenoplax (MN304982) L Mesocriconema xenoplax (KY574647) r Mesocriconema xenoplax (KJ787902) jr Mesocriconema xenoplax (KY574642) L Mesocriconema xenoplax (KJ787903) Mesocriconema xenoplax (MN711243) Mesocriconema xenoplax (MN304979)

Mesocriconema nebraskense (KJ787959) — Mesocriconema nebraskense (KY574695) Mesocriconema nebraskense (KJ787997) Mesocriconema nebraskense (KY574696) Mesocriconema nebraskense (OR157953, CD869) Mesocriconema nebraskense (MN734380) Mesocriconema nebraskense (MN711088) Mesocriconema sp. (KY574808) Mesocriconema sp. (KJ787999) Mesocriconema sp. (KJ787922, Alabama, USA) Mesocriconema sp.A (OR157954, CD1182b) Mesocriconema sp.A (OR157953, CD1182a) Mesocriconema sp. (KJ787917, South Carolina, USA) Mesocriconema sp. (KJ787918, South Carolina, USA)

- Mesocriconema sp. (KY574828)

Mesocriconema sp. (KJ787928) r Mesocriconema p. (KJ788060) f1 Mesocriconema sp. (KY574756) '— Mesocriconema sp. (KJ788019)

'oo r Mesocriconema inaratum (KJ787931) L Mesocriconema inaratum (KJ787957) Mesocriconema ericaceum (KX290525) Mesocriconema discus (KJ787866) Mesocriconema rusticum (KJ787855)

Neobakernema variabilis (MF683242) Criconemoides informis (MN305136) Criconemoides informis (MN305133) Criconemoides informis (MN305128) Criconemoides informis (MN305134) r Criconemoides informis (MN305121) 1 Criconemoides informis (MN305107) Criconemoides informis (MN305132) Criconemoides informis (MN305138) Criconemoides informis (MN305084) Criconemoides informis (MN305137)

Criconemoides informis (MN710862) " j Criconemoides parainformis (MN791124) 1 Criconemoides parainformis (MN791126) Criconemoides sinensis comb. n. (MK249990)

Criconemoides informis (MT328867)

■- Criconemoides informis (OR157956, CD836)

Criconemoides neoinformis (ON322563) Criconemoides informis (MN710860)

ioo r~ Criconemoides annulatus (MN710852) '— Criconemoides annulatus (MN710854) Criconemoides annulatus (OR157947, CD879) Criconemoides annulatus (MF770956) Criconemoides annulatus (MN710858) — Criconemoides annulatus (MN710845) 100 I Criconemoides persicus (ON322562)

1 C.rinnnpmniriw neirGiniis

Criconemoides persicus (ON322561) j Criconemoides annulatus (MN710856) 1 Criconemoides annulatus (MN710857)

2j Criconemoides sp.A (OR157950, CD370) ' Criconemoides spA (OR157951, CD554) — Criconemoides myungsugae (MH496164) Criconemoides rotundicaudatus (MN791128) Criconemoides parvus (MF770968)

Mesocriconema sphaerocephalum (MN711165, Florida, USA) Mesocriconema sphaerocephalum (KY574841, Florida, USA) Mesocriconema sphaerocephalum (OR157948, CD1183b) Mesocriconema sphaerocephalum (OR157949, CD1183a) Mesocriconema sphaerocephalum (KU236638, Puerto Rico) Mesocriconema sphaerocephalum (MF770901, Puerto Rico) Mesocriconema sphaerocephalum (MN711166, Florida, USA) Mesocriconema sphaerocephalum (KY574840, Alabama, USA) Mesocriconema sphaerocephalum (MF770898. Nebraska, USA) Mesocriconema sphaerocephalum (KY574837, Nebraska, USA) ioo | I Mesocriconema sphaerocephalum (KY574838, Nebraska, USA)

Mesocriconema sphaerocephalum (KY574839, Nebraska, USA) Mesocriconema sphaerocephalum (MN711173, Missouri, USA) Mesocriconema sphaerocephalum (MN711169, Botswana)

Paratylenchus sp. (MN711374)

Paratylenchus projectus (MW238473)

Mesocriconema sphaerocephalum (MN711170, Botswana) Tylenchulus semipenetrans (OP726467)

0.1

Fig. 6. Phylogenetic relationships within some representatives of the genus Criconemoides and Mesocriconema. Bayesian 50% majority rule consensus tree from two runs as inferred from analysis of the COI gene sequence alignment under the GTR + I + G model. Posterior probabilities equal to, or more than, 70% are given for appropriate clades. New sequences are indicated in bold letters.

Fig. T. Criconemoides informis. CD836. Female. SEM. A: En face view of lip region; B: Lateral view of lip region; C: Annuli at midbody; D: Ventral view of posterior part of body; Mesocriconema sphaerocephalum. CD1183. Female. SEM. E: En face view of lip region; F: Lateral view of anterior part of body; G: Annuli at midbody; H: Dorsal view of posterior part of body.

body annuli slightly retrorse and rounded with very slight irregular margins from about middle of body, becoming more irregular towards vulva and posterior to the vulva they can be very irregular. Anastomosis very rare. Stylet robust with cupped basal knobs. Excretory pore situated from two annuli anterior to opposite base of pharyngeal lobe. Hemizonid and hemizonion not seen. Spermatheca not seen in one specimen but in the other it was large, oblong and filled with sperm cells. Vulval lips closed. Anterior lip not indented and not overhanging the posterior lip. Vagina straight. Anus distinct, four annuli from tail tip. Tail tapering to a narrow tip with two or three irregular, indistinct lobes.

Male. Not found.

Juvenile. Two juveniles found but in very bad condition and not good for measuring. One specimen had 66 body annuli, rounded, retrorse with slight irregular margins from about midbody to tail terminus where they were slightly more irregular in the last four annuli.

Remarks. These two specimens compared well to the various descriptions of the species (Table S2).

Molecular characterisation. Only COI gene sequence was obtained for this sample. The sequence of Californian C. informis clustered with that of the Russian population of this species (Fig. 6) and they were different in 9.7% (62 bp).

Criconemoides sp. A from South Africa

This species was found in South Africa, KwaZulu Natal Province. Specimens of this sample were used for molecular analysis only. Phylogenetic position of this sample within the genus Criconemoides using D2-D3 of 28S rRNA and COI gene sequences is not well resolved and given in Figure 3 and 6.

Mesocriconema nebraskense Olson et al., 2017

(Figs S5A-D & 9E-H)

Mesocriconema nebraskense was described in Spring Creek Prairie, Nebraska located in the Central Tall Grasslands ecoregion of North America. This species was found in several states of the USA (Powers et al., 2014, 2021; Olson et al., 2017) and recently from grasses in Korea (Mwamula & Lee, 2021).

Measurements. See Table 3.

iНе можете найти то, что вам нужно? Попробуйте сервис подбора литературы.

Female. Body slightly curved ventrad. Lip region with two annuli. Stylet strong and well developed with cupped basal knobs. Vulva an open slit with anterior lip bearing two rounded

projections. Vagina always distinctly sigmoid. Anus distinct four to eight annuli from tail tip. Tail form varying form round to conoid.

Male. Not found.

Juvenile. Not found.

Remarks. Description of this population is similar to that provided by Olson et al. (2017). This species differs from M. xenoplax by shorter body and stylet lengths for females (Table 3).

Molecular characterisation. The D2-D3 of 28S rRNA gene sequence of M. nebraskense from Kansas differ only in 0.1% (1 bp) from that of North Dakota population (Yan et al., 2018), whereas COI gene sequence was identical to that of population from Aurora Prairie, South Dakota published by Olson et al. (2017). Phylogenetic position of this species is given in Figures 3 and 6.

Mesocriconema sphaerocephalum Taylor, 1936

(Figs S1I-L & 7E-H)

Mesocriconema sphaerocephalum was described by Taylor (1936) from soil around roots of grass in Trinidad in the West Indies, redescribed by Raski and Golden (1965). This species was reported from North and South America, Europe, tropical Africa and Asia (Orton Williams, 1972; Geraert, 2010). The present specimens were collected from Florida, USA.

Measurements. See Table S3.

Female. Body slightly curved ventrad. SEM photos show a slightly elevated labial disc with a prominent mouth opening, distinct amphid openings, four small distinct, rounded submedian lobes. Labial plates not very prominent. Lip region with two annuli, first smaller than second, not set off from body annuli. All body annuli slightly retrorse with smooth margins. Lateral field area marked by numerous anastomosis and broken lines creating a zig zag effect. Stylet robust with cupped basal knobs. Excretory pore situated from opposite to two annuli posterior to base of pharyngeal lobe. Hemizonid and hemizonion not seen. Spermatheca indistinct and empty. Vulval lips closed. Vagina straight. Anus distinct, one to three annuli from tail tip. Tail tapering very slightly to a broadly rounded tip with one or two lobes on the terminus.

Male. Not found.

Juvenile. Not found.

Remarks. These specimens compare very well with the numerous descriptions of the species, although some of the characters vary quite a bit in different regions of the world, such as having longer stylets in Spain (Gomez Barcina et al., 1991) compared to the shorter stylets of Venezuelan

identical or 0.3% (2 bp) to those from Puerto Rico (KU236638) and Florida (KY574841) published by Powers et al. (2016) and Olson et al. (2017) and differed in 14.0-17.5% (100-125 bp) from those from Missouri and Nebraska (Fig. 6).

Mesocriconema xenoplax Raski, 1952 (Figs S6, 8 & 9A-D)

Mesocriconema xenoplax was originally described by Raski (1952) from the roots of grape (Vitis vinifera L. var. sultanina) grown on a Vitis longii rootstock, east of Fresno, Fresno County, California, USA and reported from many localities in the USA and worldwide (Geraert, 2010; Powers et al., 2014). Mesocriconema xenoplax was found in two samples from South Africa and two samples from the USA (Table 1). The specimens from Tvl1928 are some of the most recent specimens of this species found in South Africa and they are included in this morphological study only.

Measurements. See Table 3.

Fig. 8. Mesocriconema xenoplax. LM. CD865. Female. A. Whole body; B-D: Anterior part of body; E-G: Posterior part of body. Scale bars = 50 цт.

specimens (Crozzoli & Lamberti, 2001) which were 44-53 ra 67-77 pm. Orton Williams (1972) give a stylet range of 30-69 pm. After analysing intraspecific morphological and morphometric variation, De Grisse and Loof (1970) found that populations of this species from the temperate zones had a slightly longer stylet than those from tropical countries. The number of annuli also varied quite a lot from 52 to 82, tail length from 4.5 to 19.5 pm with the c value ranging from 19 to 122. The South African specimens normally had much more anastomosis and breaks in the lateral field than the present specimens.

Molecular characterisation. After analysing the D2-D3 of the 28S rRNA gene sequences of M. sphaerocephalum from Italy and Venezuala, Subbotin et al. (2005) concluded that they were different and, perhaps, belonged to two sibling species. The D2-D3 of the 28S rRNA gene sequences of Florida population differed in 0-0.2% (0-1 bp) from that of Venezuala population and in 4.5-4.7% (25-26 bp) from that of Italian populations (Fig. 3). The COI gene sequences of Florida population were

■ji

Species Mesocriconema xenoplax M. itebraskense Mesocriconema sp.A.

Sample Character USA, CaUfornia (CD859) USA, Kansas (CD865) South Africa (Tvll928) USA, Kansas (CD869) USA, Florida (CD1182)

n 7 6 9 4 12

L 561 ± 85.5 (495-746) 544 ± 6 (489-669) 578 ± 51.2 (515-685) 482 ±26 (458-519) 500 ± 55.1(435-626)

a 9.6 ±0.7 (8.8-10.8) 12.2 ±0.8 (11.3-13.3) 9.4± 0.9 (8.2-11.2) 9.7 ±0.7 (9.1-10.7) 11.2 ± 1.2(9.5-12.9)

b 3.6 ±0.7 (3.5-3.8) 4.3 ±0.4 (3.7-4.5) 3.9 ±0.3 (3.4-4.4) 4.3 ±0.1 (4.2-4.5) 3.9 ±0.3 (3.5-4.5)

c 21.9 ±2.6 (17.7-25.3) 23.8 ±4.6 (17.7-29.4) 21.2 ±2.4 (18.7-24) 19.5 ± 1.6(14.3-21.5) 18.8 ±2.3 (14.5-21.8)

o 9.6 ± 1.5 (7.8-11.3) 12.2 ±2.6 (10.1-16.2) 12.3 ± 1.2(11-14.4) 15.1 ±0.7(14.3-16) 14.6 ± 1.6(11.3-17.4)

DGO 7.5 ± 1.6(6-9) 7 ±0.8 (6-8) 10.3 ±0.8 (9.6-11.8) 8 ±0.6 (7.5-8.5) 7.5 ±0.7 (7-9)

V 92 ±0.7 (91-94) 94 ± 1(93-96) 94 ± 0.6 (93-95.5) 92 ±0.2 (92-92.5) 93 ± 0.5 (92-94)

OV1 - 47.3 ± 8.7 (40.2-59.4) 65 ± 13.2(43.8-85.4) 48 ± 1.1 (47.5-49) 44.2 ±4.1 (38.1-48.4)

OV length - 261 ±36.2 (213-301) 367 ± 87.4(300-574.5) 239 ±20.8 (225-254) 228 ±26.1 (180-252)

Stylet length 75.5 ±2.9 (72-79.5) 60 ±9.9 (50-73) 84 ±3 (79.5-88) 50.5 ± 2 (48.5-53) 52.5 ± 2.2 (48-56)

Metenchium length 57 ± 1.8 (54.5-58) 43 ±7.8 (36-56) 64.5 ±2.6 (61-69) 36.5 ± 1.5 (34.5-38) 38 ± 1.3 (34.5-40)

Telenchium length 18.5 ± 1.3(17-21) 17 ± 2.3 (13-19) 19.5 ±0.9 (18.5-20.5) 13.2 ± 1 (12.5-15) 15 ± 1(13-17)

in 75.5 ±0.9 (74.1-76.5) 72.8 ± 1.6(72-73.8) 77 ±0.9 (75.6-78.3) 72 ± 1.7(70-74) 71.9 ± 1 (69.8-74)

Stylet knob height 5.5 ±0.4 (5-6) 5 ± 0.8 (4-7) 5.5 ±0.5 (4.5-6) 5 ± 1.2(4-6.5) 5 ±0.7 (4-6.5)

Stylet knob width 11.5 ±0.6 (10.5-12) 11 ±2 (9-14) 12 ± 0.5 (11-12.5) 9 ±0.5 (8.5-9.5) 9.5 ±0.8 (7.5-10.5)

Exc. pore from anter. end 157 ± 22.9 (137.5-195.5) 143 ± 18.5 (123.5-173.5) 163 ± 15.4(148-193.5) 128 ±6.8 (121.5-136) 135 ± 17.4 (112.5-175)

Width at midbody 59 ± 6.8 (50.5-69) 46 ±6.3 (38-55) 64 ±5.2 (56-72) 50 ± 4 (44-53) 45 ±3.7 (39-51.5)

Width at anus 38 ±5 (31.5-45) 31 ±3.3 (28-36) 39 ± 2.2 (36-42) 34.5 ±2.6 (31.5-37.5) 33 ± 2.6 (28-37.5)

Width at vulva 45 ± 5.9 (37-54) 35 ±4.8 (31-43) 44.5 ±2.9 (40.5-48) 40 ±2.1 (37.5-42.5) 36.5 ±3.1 (32-41)

Annulus width 6± 1 (5-8) 6 ±0.7 (5.5-7) 6.5 ±0.5 (6-7.5) 5 ±0.3 (4.5-5) 5.9 ±0.7 (5-7.5)

Tail length 26 ±5.4 (21-36) 23 ±3.5 (20-29) 28 ± 4.3 (22-32) 25 ±3.2 (22-29.5) 27 ± 5 (21-39)

Pharynx length 144.5 ±9.6 (135-160) 129 ± 17.6(115-150) 152 ± 5.5 (144-161) 112 ±8.2 (104.5-122) 127 ± 13.2(110-152)

1st Up annulus diam. 14.5 ± 1.7(12.5-17) 15 ±0.5 (14-16) 19 ±2 (16-21.5) 14 ± 1.2 (12.5-15.5) 17 ± 1.4 (14.5-19.5)

2nd Up annulus diam. 20 ±2 (18-23) 19 ± 1.3 (18-21) 24 ± 1.7(22-26.5) 17.5 ± 1 (17-19) 22 ± 1.6(18.5-25)

1st body annulus diam. 23.5 ± 1.9(21.5-26) 23 ±2.5 (21-26.5) 28 ± 1.8(26-32) 21 ± 1.2 (20-23) 25 ± 1.8(23-29)

2nd body annulus diam. 26 ± 1.8 (23.5-29.5) 26 ±3.4 (23-32) 32.5 ±2.4 (29-36) 24.5 ± 1 (23.5-26) 28 ± 1.9(26-32)

R 103 ±2.6 (100-107) 99 ±3.4 (94-104) 100 ±3.1 (95-105) 104 ± 1.5 (103-106) 90 ±2.1 (87-94)

RSt 17 ± 1.4(16-20) 14± 1.3 (13-16) 17 ± 1.1 (15-18) 13 ± 1(12-14) 11 ±0.7(10-12)

Roes 31 ± 1.3 (29-32) 25 ± 1.7(24-28) 28 ± 1.1 (26-29) 26 (n= 1) 23 ± 2 (20-27)

Rex 31 ±0.8 (30-32) 28 ± 1.2(27-30) 30 ± 1.2(28-32) 29 ±0.5 (29-30) 25 ± 1.1 (24-27)

RV 8 ± 1.7(7-11) 8± 1.3 (6-10) 7 ±0.4 (6-7) 8.5 ±0.6 (8-9) 7 ±0.8 (6-9)

RVan 1.5 ±0.8 (1-3) 2 ± 1(1-3) 0-1 1 0-1

Ran 6 (5-7) 6± 1.5 (4-7) 6 ±0.9 (5-7) 6.5 ± 0.6 (6-7) 6± 1.1 (4-8)

VL/VB 1± 0.2 (0.7-1.1) 0.9 ±0.2 (0.7-1.1) 0.8 ±0.1 (0.6-1) 0.9 ±0.05 (0.9-1) 1± 0.08 (0.8-1.1)

St%L 13.6 ± 1.5 (10.4-14.6) 10 ±0.6 (10-11) 14.4 ± 1.2(12.4-16.1) 10.5 ±0.3 (10.2-10.8) 10.6 ± 1 (8.5-12)

o

o

0

s

1

c

in >

(S>

0

1

3:

o

02

Fig. 9. Mesocriconema xenoplax. SEM. CD865. Female. A: Lateral view of anterior part of body; B: Annuli at midbody; C: En face view of lip region; D: Lateral view of posterior part of body. Mesocriconema nebraskense. SEM. CD869 Female. E, G: En face view of two lip regions; F: Ventral view of posterior part of body; H: Lateral view of anterior part of body. Mesocriconema sp. A. SEM. CD1182. Female. I: En face view of lip region. J, K: Lateral view of anterior part of body of two females; L: Ventral view of posterior part of body.

Female. Body slightly curved ventrad. Lip region with two annuli, first one pointing more outward and second one retrorse, not set off from body. First annulus usually divided into two parts by a deep lateral indentation but in some populations the indentation is not so distinct. Submedian lobes prominent, rounded and 2.5 to 3 pm in diameter. Labial plates normally large and prominent and in some cases appearing as an additional annulus. All body annuli retrorse with smooth margins, sometimes slightly irregular. Anastomosis rare. Stylet strong and well developed with cupped basal knobs. Excretory pore situated from one annulus anterior to six annuli posterior to base of basal pharyngeal lobe. Hemizonion not seen. Spermatheca mostly indistinct or small, round and empty. Vulva an open slit with anterior lip bearing two rounded projections. Vagina always distinctly sigmoid. Anus distinct four to eight annuli from tail tip. Tail form varying form round to conoid with one flat lobe or a few rounded lobes on the tip.

Male. Not found.

Juvenile. Not found.

Remarks. These specimens compared very well with those from the literature although some variation was observed. Orton Williams (1972) discussed the great variation in the arrangement of the submedian lobes and labial plates. Popovici and Ciobanu (2000) mentioned that all their females had sperm filled spermathecae. Some populations had fewer body annuli than others (Peneva et al., 2000). Crozoli and Lamberti (2001) stated that anastomosis was common along the whole length of the body.

Molecular characterisation. The D2-D3 of the 28S rRNA gene sequences were obtained from three samples and they were different from that from Belgium in 0.2-1.3%. Phylogenetic relationships of this species with other criconematids are given in Figure 3. COI gene sequence analysis revealed that the South African sample belonged to the haplotype 10 as it has been defined by Powers et al. (2014) (Fig. 6).

Mesocriconema sp. A (Figs S5E-H & 9I-L)

This sequence of the sample from Gainesville, Florida, USA is similar to those of Mesocriconema ornata (Raski, 1958) Loof & De Grisse, 1989 haplotype group 15 as identified by Powers et al. (2014). Powers et al. (2014) distinguished three haplotype groups 1, 15 and 16 within this species and stated that groups 1 and 15 both conformed to the morphospecies description and were only isolated from agricultural soils, yet the mean

pairwise p-distance of the COI haplotypes of the two groups was 21.6%. Because, the sequences of group 1 were significantly different from others, we could consider it as belonging to a true Mesocriconema ornata, because these samples were collected near the type locality, whereas others are considered to be representatives of an undescribed Mesocriconema sp. A.

Measurements. See Table 3.

Female. Body slightly curved ventrad. Lip region with two annuli. Body annuli retrorse with smooth margins, sometimes slightly irregular. Stylet strong and well developed with cupped basal knobs. Excretory pore situated from one annulus anterior to six annuli posterior to base of basal pharyngeal lobe. Spermatheca mostly indistinct or small, round and empty. Anus distinct four to eight annuli from tail tip. Tail form varying from round to conoid with one flat lobe or a few rounded lobes on the tip.

Male. Not found.

Juvenile. Not found.

Remarks. Measurements of this sample are similar to those of haplotype 15 provided by Powers et al. (2014).

Molecular characterisation. This species is related to M. nebraskense (Figs 3 & 6). The provided COI sequences are identical to that of a sample identified as Mesocriconema ornata from Alabama by Powers et al. (2014).

Phylogenetic relationships reconstructed in this study using the D2-D3 of 28S rRNA gene sequences are congruent to those presented by other authors (Munawar et al., 2020; Hosseinvand et al., 2020, 2023; Li et al., 2022; Nguyen et al., 2022) and also revealed that genera Criconemoides, Criconema, Discocriconemella and Mesocriconema are not monophyletic. Nguyen et al. (2022) concluded that key morphological characters used in the classification of Criconematidae are the consequence of convergent evolution. Thus, the results of molecular dataset analysis show that the classification of Criconematidae needs to be revised.

The genus Discocriconemella was proposed by De Grisse and Loof (1965) when revising Criconemoides and up to now it included 31 species, which were characterised by the presence of a disc-shaped head (Geraert, 2010; Munawar et al., 2019; Li et al., 2022). Based on the results of molecular dataset analysis, Powers et al. (2014) transferred Discocriconemella inarata to the genus Mesocriconema and named as M. inaratum. The present analysis showed that the D2-D3 of 28S rRNA gene sequences of D. sinensis Munawar, Cai, Subbotin & Zheng, 2019 and D. parasinensis Li, Munawar, Castillo & Zheng, 2022 clustered with

those of Criconemoides species belonging to the C. informis group and based on this grouping, these species are considered here as representative of this genus: Criconemoides sinensis (Munawar et al., 2019) comb. n. and Criconemoides parasinensis (Li et al., 2022) comb. n., both described from China. The positions of these species within the genus Criconemoides were already noticed and discussed by Munawar et al. (2019) and Li et al. (2022).

The diagnosis for the genus Criconemoides provided by Siddiqi (2000) and Geraert (2010) is shorten and modified and given below.

Genus Criconemoides

Diagnosis. Criconematidae. Female: Small to moderately large (about 0.3-1 mm), sausage- or ring-like when relaxed. Annuli crenate, rough or smooth, with round to pointed edges. Cephalic annuli two to three, smaller than and not differentiated or separated by a collar from body annuli. Some species with cephalic annulus appearing as a large, anteriorly flattened disc. Submedian lobes absent or present. Stylet moderately long usually rigid. Vulva lips closed or open. Vagina straight. Tail short, conoid, convex-conoid or hemispheroidal. Male: Cephalic region rounded or conoid, lateral field with three to four incisures. Bursa distinct, subterminal.

Criconemoides sinensis (Munawar et al., 2019) comb. n.

= Discocriconemella sinensis Munawar, Cai, Subbotin & Zheng, 2019

Criconemoides parasinensis (Li et al., 2022) comb. n.

= Discocriconemella sinensis Li, Munawar, Castillo & Zheng, 2022

Among the species of the genus Discocriconemella, Orton Williams (1981) and Vovlas (1992) distinguished four groups based on configuration of the cephalic disc. Group 1 included species with round cephalic disc with an uninterrupted margin and Criconemoides sinensis comb. n., C. parasinensis comb. n. and Mesocriconema inaratum belonged to this group. Discocriconemella limitanea (Luc, 1959) De Grisse & Loof, 1965, the type species of this genus, belonged to group 2 having disc with ventral and dorsal deep indentations and this species formed a separate lineage within criconematids in phylogenetic trees. Munawar et al. (2019) showed that D. hengsungica Choi & Geraert, 1975 belonging to group 4 with round disc with paired

dorsal and ventral projections also formed a separate lineage, which was sister to Xenocriconemella macrodora (Taylor, 1936) De Grisse & Loof, 1965. Species belonging to group 3 with disc indented medially and laterally giving a four-lobed appearance are not still molecularly characterised. Thus, disc-shaped head appeared several times in criconematid evolution and can not be used as reliable taxonomic character for genus differentiation. We agree with the proposal made by Munawar et al. (2019) that it needs detailed molecular characterisation of other Discocrico-nemella species in order to facilitate their phylogenetic grouping and then use these results to revise this genus.

iНе можете найти то, что вам нужно? Попробуйте сервис подбора литературы.

SUPPLEMENTAL MATERIALS

Figure S1. Criconema mutabile and Mesocriconema sphaerocephalum. Figure S2. Criconema mutabile. Figure S3. Criconemoides annulatus. Figure S4. Criconemoides informis. Figure S5. Mesocriconema nebraskense and Mesocriconema sp. A.

Figure S6. Mesocriconema xenoplax. Table S1. Measurements of Criconema mutabile. Table S2. Measurements of Criconemoides annulatus and C. informis.

Table S3. Measurements of Mesocriconema sphaerocephalum.

http ://www.russjnematology.com/Articles/rjn312 /Paper9VandenBerg-SUPPL.pdf

REFERENCES

Archidona-Yuste, A., Palomares-Rius, J.E., Clavero-Camacho, I., Cantalapiedra-Navarrete, C., Liebanas, G. & Castillo, P. 2023. A blind-identification test on Criconema annuliferum (de Man, 1921) Micoletzky, 1925 species complex corroborate the hyper-cryptic species diversity using integrative taxonomy. Plants 12: 1044. DOI: 10.3390/plants12051044 Azimi, S. & Pedram, M. 2020. Description of Criconema iranicum n. sp. (Nematoda: Criconematidae) from Iran. Journal of Crop Protection 9: 497-505. Brzeski, M.W. 1998. Morphological observations on three species of Mesocriconema Andrassy, 1962 and nomenclatorial note on M. goodeyi (Jairajpuri, 1963) Loof & De Grisse, 1989 (Nematoda: Criconematidae). Journal of Nematode Morphology and Systematics 1: 47-56. Brzeski, M.W., Choi, Y.E. & Loof, P.A. 2002a. Compendium of the genus Criconemoides Taylor,

1936 (Nematoda: Criconematidae). Nematology 4: 325-339. DOI: 10.1163/156854102760199178 Brzeski, M.W., Loof, P.A. & Choi, Y.E. 2002b. Compendium of the genus Mesocriconema Andrassy, 1965 (Nematoda: Criconematidae). Nematology 4: 341-360. DOI: 10.1163/156854102760199187 Chaves, E. 1983. Criconematidae (Nematoda) from Argentina. Nematologica 29: 404-424. DOI: 10.1163/187529283X00285 Choi, Y.E., Brzeski, M.W. & Kim, J.-L. 2000. Observations of some species of Criconemoides Taylor, 1936 (Nematoda: Criconematidae) with proposals of new synonyms. Nematology 2: 273-284. DOI: 10.1163/156854100509141 Clavero-Camacho, I., Palomares-Rius, J.E., Cantalapiedra-Navarrete, C., Castillo, P. & Archidona-Yuste, A.A. 2022. Proposed new species complex within the cosmopolitan ring nematode Criconema annuliferum (de Man, 1921) Micoletzky, 1925. Plants 11: 1977. DOI: 10.3390/plants11151977 Crozzoli, R. & Lamberti, F. 2001. Known and new species of Mesocriconema Andrassy, 1965 (Nematoda: Criconematidae) from Venezuela. Russian Journal of Nematology 9: 85-105. Crozzoli, R. & Lamberti, F. 2002. Species of Criconema Hofmänner & Menzel, 1914 and Ogma Southern, 1914 occurring in Venezuela with description of Ogma araguaensis sp.n. (Nematoda: Criconematidae). Russian Journal of Nematology 10: 89-98. De Grisse, A.T. 1969. Redescription ou modification de quelques techniques utilisées dans l'étude des nematodes phytoparasitaires. Mededelingen van de Rijksfaculteit der Landbouwwetenschappen Gent 34: 351-369. De Grisse, A.T. & Loof, P.A.A. 1965. Revision of the genus Criconemoides (Nematoda). Overdruk uit de Mededelingen van Landbouwhoeschool en de Opzoekingsstations van de Staat Gent 30: 577-603. De Grisse, A. & Loof, P.A.A. 1970. Intraspecific variation in some Criconematidae (Nematoda). Mededeelingen Faculteit Landbouwetenschappen Rijksuniversiteit Gent 35: 41-63. De Man, J.G. 1921. Nouvelles recherches sur les nematodes libres terricoles de la Hollande. Capita Zoologica 1: 1-62. Eskandari, A., Karegar, A., Pourjam, E., van den Berg, E. & Tiedt, L.R. 2010. Additional data on some poorly known species of Criconemoides Taylor, 1936 (Nematoda: Criconematidae). Nematology 12: 505-518. DOI: 10.1163/138855409X125068559 79596 Etongwe, C.M., Singh, P.R., Bert, W. & Subbotin, S.A. 2020. Molecular characterisation of some plant-parasitic nematodes (Nematoda: Tylenchida) from Belgium. Russian Journal of Nematology 28: 1-28. DOI: 10.24411/0869-6918-2020-10001

Geraert, E. 2010. Criconematidae of the world -Identification of the Family Criconematidae (Nematoda). Belgium, Academia Press, 615 pp.

Gomez Barcina, A., Vovlas, N., Castillo, P. & Gonzales-Pais, M.A. 1991. Morphometries and SEM observations of four criconematid species from Spain. Nematologica Mediterranea 19: 121-127.

Gomez Barcina, A., Castillo, P. & Gonzales-Pais, M.A. 1989. Nematodos fitoparasito de la subfamilia Criconematinae Taylor, 1936 en la Sierra de Cazorla. Revista Ibérica de Parasitologia 49: 241-255.

Heyns, J. 1970a. South African Criconematinae. Part 2. Genera Criconema, Hemicriconemoides and some Macroposthonia (Nematoda). Phytophylactica 2: 129-136.

Heyns, J. 1970b. South African Criconematidae. Part 3. More species of Hemicriconemoides and Macroposthonia (Nematoda). Phytophylactica 2: 243-250.

Hosseinvand, M., Eskandari, A. & Ghaderi, R. 2020. Morphological and molecular characterisation of three known species of Criconematoidea from Iran. Nematology 22: 745-758. DOI: 10.1163/1568541100003337

Hosseinvand, M., Eskandari, A., Palomares-Rius, J.E., Castillo, P., Abolafia, J. & Ghaderi, R. 2023. Morphological and molecular characterisation of a new cryptic species of Criconemoides informis group, C. neoinformis n. sp., and C. persicus n. sp., with notes on C. avicenniae. Nematology 25: 13-32. DOI: 10.1163/15685411-bja10204

Jenkins, W.R. 1964. A rapid centrifugal-flotation method for separating nematodes from soil. Plant Disease Reporter 48: 692.

Karani, H.M., Eskandari, A., Ghaderi, R. & Karegar, A. 2020. Description and molecular phylogeny of Mesocriconema abolafiai n. sp. (Nematoda: Criconematidae) from iran. Journal of Nematology 52: 1-17. DOI: 10.21307/jofnem-2020-048

Li, J., Munawar, M., Castillo, P. & Zheng, J. 2022. Morpho-molecular and ultrastructural characterization of Discocriconemella parasinensis n. sp. from Zhejiang province, China. Journal of Nematology 54: 20220011. DOI: 10.2478/jofnem-2022-0011

Loof, P.A.A. 1965. Zur Taxonomie von Criconemoides rusticus (Micoletzky) und informis (Micoletzky). Mitteilungen aus dem Zoologischen Museum in Berlin 41: 183-192.

Micoletzky, H. 1922. Die freilebenden Erd-Nematoden mit besonderer Berücksichtigung der Steiermark und der Bukowina, zugleich mit einer Revision sämtliger, nicht mariner, freilebender Nematoden in Form von Genus-Beschreibungun und Bestimmingsschlüsseln. Archiv für Naturgeschichte, Abteilung A 87 (1921): 1-650.

Micoletzky, H. 1925. Die freilebenden Süsswasser- und Moornematoden Dänemarks. Det Kongelige Danske Videnskabelige Selskap Skrifter, Naturvidenskabelige ogMathematiske Afdeling 8: 57-310. Munawar, M., Cai, R., Subbotin, S.A. & Zheng, J.W. 2019. Description of Discocriconemella sinensis n. sp. (Nematoda: Criconematidae) from the rhizosphere of Camellia sinensis in China. Nematology 21: 779792. DOI: 10.1163/15685411-00003252 Munawar, M., Miao, W., Cai, R., Tian, Z., Castillo, P. & Zheng, J. 2020. Species diversity of ring nematodes of the genus Criconemoides (Nematoda: Criconematidae) based on three new species from China, using integrative taxonomy. European Journal of Plant Pathology 157: 119-139. DOI: 10.1007/ s10658-020-01990-2 Mwamula, A.O. & Lee, D.W. 2021. Occurrence of plant-parasitic nematodes of turfgrass in Korea. The Plant Pathology Journal 5: 446-454. DOI: 10. 5423/PPJ.0A.04.2021.0059 Netscher, C. & Seinhorst, J.W. 1969. Propionic acid better than acetic acid for killing nematodes. Nematologica 15: 286. Nguyen, H.T., Nguyen, T.D., Le, T.M.L., Trinh, Q.P. & Bert, W. 2022. Remarks on phylogeny and molecular variations of criconematid species (Nematoda: Criconematidae) with case studies from Vietnam. Scientific Reports 12: 14832. DOI: 10.1038/s41598-022-18004-2

Olson, M., Harris, T., Higgins, R., Mullin, P., Powers, K., Olson, S. & Powers, T.O. 2017. Species delimitation and description of Mesocriconema nebraskense n. sp. (Nematoda: Criconematidae), a morphologically cryptic, parthenogenetic species from North American grasslands. Journal of Nematology 49: 42-66. Orton Williams, K.J. 1972. Macroposthonia xenoplax. C.I.H. Descriptions of Plant-parasitic Nematodes. Set 1, No. 12.

Orton Williams, K.J. 1973. Macroposthonia sphaerocephala. C.I.H. Descriptions of Plant-parasitic Nematodes. Set 2, No. 28. Orton Williams, K.J. 1981. Revision of the genus Discocriconemella De Grisse & Loof, 1965 and the erection of the new genus Acrozostron (Nematoda: Criconematoidea). Systematic Parasitology 2: 133138. DOI: 10.1007/bf00009901 Peneva, V., Neilson, R., Boag, B. & Brown, D.J. 2000. Criconematidae (Nematoda) from oak forests in two nature reserves in Russia. Systematic Parasitology 46: 191-201. DOI: 10.1023/A:1006338019502 Popovici, I. 1988. Intraspecific variation in Criconemoides annulatus Taylor, 1936 (Nematoda: Criconematidae) from Romania. Revue Roumaine de Biologie. Série de Biologie Animale 33: 11-14.

Popovici, I. & Ciobanu, M. 2000. New records of some Criconematidae (Nematoda) from Rumania. Journal of Nematode Morphology and Systematics 10: 135-140. Powers, T.O., Bernard, E.C., Harris, T., Higgins, R., Olson, m., Lodema, M., Mullin, P., Sutton, L. & Powers, K.S. 2014. COI haplotype groups in Mesocriconema (Nematoda: Criconematidae) and their morphospecies associations. Zootaxa 3827: 101146. DOI: 10.11646/zootaxa.3827.2.1 Powers, T.O., Bernard, E.C., Harris, T., Higgins, R., Olson, M., Olson, S., Lodema, m., Matczyszyn, J., Mullin, P., Suttoni, L. & Powers, K.S. 2016. Species discovery and diversity in Lobocriconema (Criconematidae: Nematoda) and related plant-parasitic nematodes from North American ecoregions. Zootaxa 4085: 301-344. DOI: 10.11646/zootaxa.4085.3.1 Powers, T.O., Harris, T.S., Higgins, R.S., Mullin, P.G. & Powers, K.S. 2021. Nematode biodiversity assessments need vouchered databases: A BOLD reference library for plant-parasitic nematodes in the superfamily Criconematoidea. Genome 64: 232-241. DOI: 10.1139/gen-2019-0196 Raski, D.J. 1952. On the morphology of Criconemoides Taylor, 1936, with descriptions of six new species (Nematoda: Criconematidae). Proceedings of the Helminthological Society of Washington 19: 85-99. Raski, D.J. & Golden, M.A. 1965. Studies on the genus Criconemoides Taylor, 1936 with descriptions of eleven new species and Bakernema variabile n. sp. (Criconematidae: Nematoda). Nematologica 11: 501565. DOI: 10.1163/187529265X00690 Raski, D.J. & Luc, M. 1985. A reappraisal of the genus Criconema Hofmänner & Menzel, 1914 (Nematoda: Criconematidae). Revue de Nématologie 7 (1984): 323-334.

Ronquist, F. & Huelsenbeck, J.P. 2003. MRBAYES 3: Bayesian phylogenetic inference under mixed models. Bioinformatics 19, 1572-1574. DOI: 10.1093/ bioinformatics/btg180 Sakwe, P.N. & Geraert, E. 1993. Criconematidae Taylor, 1936 (Nematoda) from Cameroon. Afro-Asian Journal of Nematology 3: 22-38. Shokoohi, E., Mashela, P.W. & Panahi, H. 2020. Criconema mutabile (Nematoda: Criconematidae) from Iran and South Africa. Biologia 75: 1143-1153. DOI: 10.2478/s11756-019-00364-2 Siddiqi, M.R. 2000. Tylenchida: Parasites of Plants and

Insects. UK, CAB International. 833 pp. Subbotin, S.A. 2021. Molecular identification of nematodes using polymerase chain reaction (PCR). In: Techniques for work with plant and soil nematodes (R.N. Perry, D.J. Hunt & S.A. Subbotin Eds). pp. 218-239. Wallingford, UK, CAB International. DOI: 10.1079/9781786391759.0012A Subbotin, S.A., Vovlas, N., Crozzoli, R., Sturhan, D.,

Lamberti, F., Moens, M. & Baldwin, J.G. 2005. Phylogeny of Criconematina Siddiqi, 1980 (Nematoda: Tylenchida) based on morphology and D2-D3 expansion segments of the 28S-rRNA gene sequences with application of a secondary structure model. Nematology 7: 927-944. DOI: 10.1163/ 156854105776186307 Swofford, D.L. 2003. PAUP*: phylogenetic analysis using parsimony (*and other methods), version 4.0b 10. USA, Sinauer Associates. Szczygiel, A. 1974. Plant parasitic nematodes associated with strawberry plantations in Poland. Zeszyty Problemowe Postepöw NaukRolniczych 154: 1-132. Tabolin, S.B., Migunova, V.D., Volkov, Y.A. &. Volkova, M.V. 2020. Morphological and molecular characterisation of Criconemoides informis (Micoletzky, 1922) from Southern Russia. Russian Journal of Nematology 28: 79-80. DOI: 10.24411/0869-6918-2020-10007 Taylor, A.L. 1936. The genera and species of the Criconematinae, a sub-family of the Anguillulinidae (Nematoda). Transactions of the American Microscopical Society 55: 391-421. Van den Berg, E. 1980. Studies on some Criconematoidea (Nematoda) from South Africa with a description of Ogma rhombosquamatum (Mehta & Raski, 1971) Andrassy, 1979. Phytophylactica 12: 15-23.

Van den Berg, E. 1984. New and known species of some genera of the Criconematidae (Nematoda) from South Africa. Phytophylactica 16: 33-44.

Van den Berg, E. & Heyns, J. 1977. Descriptions of new and little known Criconematidae from South Africa (Nematoda). Phytophylactica 9: 95-101.

Van den Berg, E., Mekete, T. & Tiedt, R.L. 2004. New records of Criconematidae from Ethiopia. Journal of Nematode Morphology and Systematics 6 (2003): 161-174.

Van den Berg, E., Tiedt, L.R. & Subbotin, S.A. 2017. Morphological and molecular characterisation of some Criconematidae (Nematoda, Tylenchida): Ogma decalineatus (Chitwood, 1957) Andrassy, 1979, Criconema silvum (van den Berg, 1984) Raski & Luc, 1985 and Neobakernema variabile (Raski & Golden, 1966) Ebsary, 1981 from South Africa and the USA. Russian Journal of Nematology 25: 85-92.

Vovlas, N. 1992. Taxonomy of Discocriconemella (Nematoda: Criconematoidea) with a redescription of D. mauritiensis. Journal of Nematology 24: 391-398.

Yan, G., Plaisance, A., Huang, D., Baidoo, R., Ransom, J.K. & Handoo, Z.A. 2018. First report of the ring nematode Mesocriconema nebraskense from a corn field in North Dakota. Journal of Nematology 50: 531-532. DOI: 10.21307/jofnem-2018-043

E. Van den Berg, L.R. Tiedt and S.A. Subbotin. Морфологическая и молекулярная характеристика нескольких известных видов нематод родов Criconema, Criconemoides и Mesocriconema (Tylenchida: Criconematidae) из США и Южной Африки.

Резюме. В ходе нематологических обследований в нескольких регионах США и Южной Африки и с использованием интегративного подхода, сочетающего морфологический и молекулярный анализы, были обнаружены следующие виды: Criconema annuliferum, Criconema mutabile, Criconemoides annulatus, C. informis, Criconemoides sp. A, Mesocriconema nebraskense, M. sphaerocephalum, M. xenoplax и Mesocriconema sp. A. Эти виды были описаны морфологически и морфометрически, а также для некоторых видов даются СЭМ фотографии. Также предоставлены молекулярные характеристики видов с использованием последовательностей генов сегмента D2-D3 28S рРНК и COI мтДНК. По результатам молекулярного анализа Discocriconemella sinensis Munawar, Cai, Subbotin & Zheng, 2019 и D. parasinensis Li, Munawar, Castillo & Zheng, 2022 переведены в род Criconemoides.

i Надоели баннеры? Вы всегда можете отключить рекламу.