Table 1.
Morphometric characters of Mesocriconema abolafiai n. sp. (measurements are in μm and in the form of average ± s.d. (range)).
| Dehdasht population | Basht population | |||
|---|---|---|---|---|
| Characters | Holotype | 10 paratype females | 3 paratype males | 5 females |
| L | 540 | 540 ± 65 (402–612) | 471 ± 33.6 (435–502) | 519 ± 65 (425–605) |
| a | 14.9 | 14 ± 1.5 (10.3–15.3) | 24.6 ± 3.2 (20.9–27.1) | 12.6 ± 1.2 (10.9–14.4) |
| b | 4.8 | 4.8 ± 0.6 (3.2–5.7) | 24.6 | 4.3 ± 0.5 (3.5–5.0) |
| c | 14.5 | 15.8 ± 2.4 (12.1–19.5) | 10.5 ± 0.5 (9.9–11) | 19.1 ± 2.4 (16.3–21.9) |
| c´ | 1.4 | 1.3 ± 0 (1.3–1.4) | 3.3 ± 0.5 (2.9–3.9) | 1.3 ± 0 (1.1–1.4) |
| V | 90.5 | 90.3 ± 1.1 (87.8–91.9) | – | 90.1 ± 0.4 (89.8–90.8) |
| Stylet | 56.9 | 55.2 ± 1.3 (52.8–57.2) | – | 56.8 ± 2.3 (54–60) |
| Conus | 42.3 | 44 ± 6.3 (38.6–55.0) | – | 41.9 ± 1.6 (40.0–44.5) |
| m (conus/stylet %) | 74.4 | 80.1 ± 11.2 (73.1–100.0) | – | 73.7 ± 1.7 (71.9–76.3) |
| Pharynx | 110.7 | 113 ± 8 (103–124) | 59.3 | 119 ± 8 (105–126) |
| Post-vulval body length (VL) | 50.7 | 51.8 ± 4.9 (44.6–58.9) | – | 51.2 ± 6.7 (43–60) |
| Secretory-excretory pore | 109 | 108 ± 7 (99–118) | 106 ± 12 (92–113) | 108 ± 8 (97–118) |
| Lip region-vulva | 485.2 | 488 ± 62 (353–553) | 468 ± 58.6 (382–545) | |
| Lip region-anus | 498.9 | 506 ± 64.1 (372–575) | 426 ± 29.5 (396–455) | 492 ± 63.8 (399–577) |
| Vulva-anus | 24.0 | 22 ± 2.9 (17.1–26.0) | – | 24 ± 5.3 (17–32) |
| Tail length | 37.0 | 34.3 ± 3.6 (28.0–39.3) | 44.8 ± 4.5 (39.5–47.5) | 27.2 ± 2.9 (23–31) |
| Body width | 36.1 | 38.3 ± 2.4 (34.1–42.0) | 19.2 ± 1.6 (17.5–20.7) | 41.2 ± 4.3 (34–46) |
| Vulval body width (VB) | 30.4 | 30.1 ± 1.6 (27.2–32.3) | – | 30.7 ± 1.9 (27.5–32.5) |
| VL/VB | 1.7 | 1.7 ± 0.1 (1.6–1.8) | – | 1.6 ± 0.1 (1.5–1.8) |
| Annulus width | 5.2 | 5.4 ± 0.7 (4.1–6.5) | 2.8 ± 0.4 (2.3–3.2) | 5.7 ± 0.6 (4.9–6.8) |
| R | 104.0 | 104.2 ± 4 (97–113) | 132 | 97 ± 4.3 (90–101) |
| Rst | 16.0 | 15.5 ± 0.8 (14–17) | – | 14.5 ± 0.5 (14–15) |
| Rph | 26.0 | 25 ± 1.4 (23–28) | – | 24.8 ± 1.9 (23–28) |
| Rexp | 29 | 23.2 ± 4.2 (19–30) | 49.6 ± 2 (48–52) | 27.1 ± 4.2 (24–29) |
| RV | 12.0 | 12 ± 0.8 (11–14) | – | 12.8 ± 0.8 (12–14) |
| Ran | 8.0 | 9 ± 0.9 (8–10) | – | 7.4 ± 0.5 (7–8) |
| RVan | 3.0 | 3.6 ± 0.4 (3–4) | – | 3.6 ± 0.8 (3–5) |
| St/L × 100 | 10.6 | 10.3 ± 1.3 (9.2–13.6) | – | 11 ± 1.1 (9.4–12.7) |
| Spicules | – | – | 34.2 ± 0.7 (33.6–35.0) | – |
| Gubernaculum | – | – | 6.2 ± 0.9 (5.3–7.2) | – |

Figure 1:
Diagnostic drawings of Mesocriconema abolafiai n. sp. Female (A, C-E, H, I) and Male (B, F, G). A, F: entire body; B-D: anterior end and pharyngeal region; E: reproductive system; G-I: posterior end.

Figure 2:
Light micrographs of Mesocriconema abolafiai n. sp. Female (A-J) and male (K-M). A: entire body; B, C and K: anterior end and pharyngeal region; D: cuticle at mid-body; E and F: cuticle at posterior end; G: vulval region and part of reproductive system; H-J and L: posterior end; M: spicule, gubernaculum and cloaca. (Scale bars: A = 50 µm; B-M = 10 µm).

Figure 3:
SEM micrograghs of Mesocriconema abolafiai n. sp. Female (A-L). A: anterior end (arrow pointing the secretory–excretory pore); B-E: lip region in sublateral, left subventral, frontal and right subventral views, respectively (arrows pointing the amphids); F: annuli; G: cuticle at mid-body; H: entire body; I-K: posterior end in lateral, subventral and terminal views, respectively (white arrow pointing the vulva, black arrow pointing the anus); L: excretory pore (arrow).
Table 2.
List of species, collection localities and GenBank accession numbers of individual specimens used in this study for phylogenetic analysis based on 28S rRNA gene.
| Species name | GeneBank accession no. | Locality | Species name | GeneBank accession no. | Locality |
|---|---|---|---|---|---|
| Aglenchus agricola | AY780979 | Belgium | Hemicycliophora typica | KF430515 | South Africa |
| Caloosia longicaudata | GU989627 | USA | H. wyei | KC329574 | USA |
| Criconema demani | MH828126 | Russia | H. wyei | KF430497 | USA |
| C. demani | MH828128 | Russia | Merlinius brevidens | KP313844 | Iran |
| C. mutabile | MK170079 | South Africa | Mesocriconema abolafiai n. sp. | MN334222 | Iran |
| Criconema sp. | FN433874 | USA | M. ornatum | AY780968 | Venezuela |
| Criconemoides brevistylus | JQ231183 | South Africa | M. solivagum | AY780969 | Russia |
| C. brevistylus | JQ231184 | South Africa | Mesocriconema sp. | AY780967 | Italy |
| C. brevistylus | KC937033 | China | M. sphaerocephalum | AB933464 | Japan |
| C. informis | KU722386 | Iran | M. sphaerocephalum | AB933465 | Japan |
| C. myungsugae | MH444641 | China | M. sphaerocephalum | AY780951 | Italy |
| C. obtusicaudatus | JQ231186 | South Africa | M. xenoplax | AB933468 | Japan |
| C. obtusicaudatus | JQ231187 | South Africa | M. xenoplax | AY780961 | Germany |
| Eutylenchus excretorius | AY780980 | Germany | M. xenoplax | AY780963 | USA |
| Hemicaloosia guangzhouensis | KT381016 | China | M. xenoplax | AY780965 | Italy |
| H. guangzhouensis | KT381017 | China | M. xenoplax | FN433855 | USA |
| H. vagisclera | JQ246422 | USA | M. xenoplax | FN433858 | USA |
| Hemicriconemoides gaddi | MK050500 | China | M. xenoplax | FN433859 | USA |
| H. rosae | MK371811 | India | M. xenoplax | KC538862 | USA |
| H. rosae | MK371813 | India | M. xenoplax | MG680454 | Portugal |
| H. silvaticus | KF856531 | Japan | Ogma civellae | AY780955 | Venezuela |
| H. strictathecatus | MH142613 | China | O. decalineatus | MF683230 | South Africa |
| H. wessoni | KF856521 | USA | Paratylenchus tenuicaudatus | KU291239 | Iran |
| Hemicycliophora conida | FN433875 | Belgium | Sphaeronema alni | AY780978 | Germany |
| H. epicharoides | KF430512 | Italy | Trophonema arenarium | AY780971 | Italy |
| H. gracilis | KF430482 | USA | Tylenchulus semipenetrans | KM598334 | Iran |
| H. halophila | KF430444 | New Zealand | T. semipenetrans | KM598335 | Iran |
| H. lutosa | GQ406240 | South Africa | T. semipenetrans | MH156801 | China |
| H. lutosa | GQ406241 | South Africa | T. semipenetrans | MH156802 | China |
| H. signata | MG019824 | Mozambique | Xenocriconemella macrodora | AY780960 | Italy |
| H. subbotini | MG701275 | China |

Figure 4:
Bayesian 50% majority rule consensus tree as inferred from the D2 to D3 expansion segments of 28S rRNA gene dataset of Criconematoidea under the general time reversible model of sequence evolution with correction for invariable sites and a gamma-shaped distribution (GTR + I + G). Posterior probabilities more than 50% are given for appropriate clades. The new obtained sequence in this study is indicated in bold. Scale bar=expected changes per site.
Table 3.
List of species, collection localities and GenBank accession numbers of individual specimens used in this study for phylogenetic analysis based on 18S rRNA gene.
| Species name | GeneBank accession no. | Locality | Species name | GeneBank accession no. | Locality |
|---|---|---|---|---|---|
| Bakernema inaequale | MF094908 | USA | H. conida | AJ966471 | GenBank |
| Criconema mutabile | MF094914 | USA | H. conida | KJ934172 | USA |
| C. permistum | MF094899 | USA | H. conida | KJ934173 | USA |
| C. petasum | MF094927 | USA | H. subbotini | MG701280 | China |
| C. sphagni | MF094968 | USA | Lobocriconema sp. | MF094981 | USA |
| Criconemoides annulatus | MF095015 | USA | L. thornei | MF094928 | USA |
| C. annulatus | MF095024 | USA | L. thornei | MF094996 | USA |
| C. informis | MF094902 | USA | Merlinius joctus | FJ969128 | GenBank |
| C. informis | MF095025 | USA | Mesocriconema abolafiai n. sp. | MN334221 | Iran |
| C. parvus | MF795587 | China | M. curvatum | MF094891 | USA |
| Crossonema fimbriatum | MF095026 | USA | M. discus | MF094892 | USA |
| C. fimbriatum | MF094960 | USA | M. inaratum | MF094903 | USA |
| C. menzeli | MF094937 | USA | M. onoense | MF094909 | USA |
| Discocriconemella limitanea | MF795591 | China | M. ornatum | MF094893 | USA |
| D. limitanea | MF095031 | Costa Rica | M. rusticum | MF094965 | USA |
| Gracilacus paralatescens | MH200615 | China | Mesocriconema sp. | MF094967 | USA |
| G. wuae | MF095028 | Canada | Mesocriconema sp. | MF095012 | USA |
| Hemicaloosia graminis | JQ446376 | USA | M. sphaerocephalum | KJ934182 | USA |
| Hemicriconemoides. chitwoodi | KJ934162 | USA | M. xenoplax | KJ934180 | USA |
| Hemicriconemoides. fujianensis | MH444626 | China | M. xenoplax | KJ934177 | USA |
| H. kanayaensis | MG029558 | China | M. xenoplax | MF095021 | USA |
| H. kanayaensis | MG029559 | China | M. xenoplax | MF094992 | USA |
| H. parasinensis | MH444635 | China | Ogma decalineatus | MF094952 | USA |
| H. parataiwanensis | MG029556 | China | O. menzeli | EU669919 | GenBank |
| H. parataiwanensis | MG029557 | China | O. seymouri | MF094933 | USA |
| H. pseudobrachyurus | AY284622 | GenBank | Ogma sp. | KJ934175 | USA |
| Hemicriconemoides sp. | MF095013 | Thailand | Paratylenchus straeleni | AY284631 | GenBank |
| H. wessoni | KJ934163 | USA | Tylenchulus semipenetrans | MH136626 | China |
| H. wessoni | KJ934166 | USA | T. semipenetrans | AJ966511 | UK |
| Hemicycliophora aquatica | MF094911 | USA | Xenocriconemella macrodora | MF095001 | USA |
| H. conida | EU669914 | GenBank |

Figure 5:
Bayesian 50% majority rule consensus tree as inferred from the 18S rRNA gene dataset of Criconematoidea under the general time reversible model of sequence evolution with correction for invariable sites and a gamma-shaped distribution (GTR + I + G). Posterior probabilities more than 50% are given for appropriate clades. The new obtained sequence in this study is indicated in bold. Scale bar = expected changes per site.
Table 4.
List of species, collection localities and GenBank accession numbers of individual specimens used in this study for phylogenetic analysis based on ITS rRNA gene.
| Species name | GeneBank accession no. | Locality | Species name | GeneBank accession no. | Locality |
|---|---|---|---|---|---|
| Coslenchus rhombus | MK874505 | South Africa | Mesocriconema curvatum | MF094891 | USA |
| Criconema silvum | MF683236 | South Africa | M. inaratum | HM116070 | USA |
| C. silvum | MF683237 | South Africa | M. inaratum | HM116069 | USA |
| Criconemoides brevistylus | KC937032 | China | M. inaratum | HM116058 | USA |
| C. brevistylus | JQ231188 | South Africa | M. inaratum | HM116055 | USA |
| C. myungsugae | MH444640 | China | M. inaratum | HM116052 | USA |
| C. myungsugae | MH444639 | China | M. inaratum | HM116051 | USA |
| C. obtusicaudatus | JQ231189 | South Africa | M. nebraskense | MH013431 | USA |
| C. obtusicaudatus | JQ231190 | South Africa | M. nebraskense | KY574844 | USA |
| Crossonema sp. | MK292124 | USA | M. nebraskense | KY574860 | USA |
| Hemicriconemoides californianus | KF856558 | USA | M. nebraskense | KY574861 | USA |
| H. californianus | KF856560 | USA | M. nebraskense | KY574862 | USA |
| H. chitwoodi | KF856543 | USA | M. nebraskense | KY574863 | USA |
| H. fujianensis | MH444616 | China | M. nebraskense | KY574864 | USA |
| H. kanayaensis | EF126179 | Taiwan | M. nebraskense | KY574865 | USA |
| H. kanayaensis | MG029566 | China | M. onoense | JQ708120 | USA |
| H. kanayaensis | MG029568 | China | M. ornatum | JQ708124 | USA |
| H. ortonwilliamsi | KF856552 | Spain | M. ozarkiense | JQ708122 | USA |
| H. paracamelliae | MG029560 | China | Mesocriconema sp. | KY574858 | USA |
| H. promissus | KF856555 | Spain | Mesocriconema sp. | KY574857 | USA |
| H. rosae | MK371815 | India | Mesocriconema sp. | KY574856 | USA |
| Hemicriconemoides sp. | KM516185 | USA | M. xenoplax | JQ708112 | USA |
| H. strictathecatus | KF856565 | South Africa | M. xenoplax | HM116073 | USA |
| H. strictathecatus | MH142617 | China | M. xenoplax | HM116057 | USA |
| H. strictathecatus | KM516186 | USA | M. xenoplax | MF095021 | USA |
| H. strictathecatus | KM516190 | USA | M. xenoplax | MF094992 | USA |
| H. strictathecatus | KM516191 | USA | M. xenoplax | MF094915 | USA |
| Hemicycliophora californica | KF430576 | USA | M. xenoplax | MF094916 | USA |
| H. gracilis | MG019827 | USA | Neobakernema variabile | MF683239 | USA |
| H. raskii | KF430577 | USA | N. variabile | MF683238 | USA |
| H. thienemanni | KF430569 | Russia | Ogma decalineatus | MF683235 | USA |
| Mesocriconema abolafiai n. sp. | MN334228 | Iran | Paratylenchus hamatus | KF242257 | USA |
| M. curvatum | HM116062 | USA | Tylenchulus semipenetrans | JN112274 | USA |
| M. curvatum | HM116064 | USA | T. semipenetrans | FJ588909 | China |
| M. curvatum | HM116066 | USA | T. semipenetrans | MH124562 | China |
| M. curvatum | HM116067 | USA | T. semipenetrans | MH124561 | China |
| M. curvatum | HM116068 | USA |

Figure 6:
Bayesian 50% majority rule consensus tree as inferred from the ITS rRNA gene dataset of Criconematoidea under the symmetrical model of sequence evolution with a gamma-shaped distribution (SYM + G). Posterior probabilities more than 50% are given for appropriate clades. The new obtained sequence in this study is indicated in bold. Scale bar = expected changes per site.
Table 5.
Summary of the diagnostic characteristics of 12 species of Mesocriconema recently described worldwide. For other species, see (Brzeski et al., 2002). (St measurements in µm, L measurements in mm).
| Species | St | R | Rex | RV | RVan | Ran | V | VL/VB | Anas. | A. M. | Vagina | A. V. lip | Tail shape | L | S. M. L. | Original des. |
|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
| M. abolafiai n. sp. | 52–60 | 90–113 | 19–30 | 11–14 | 3–5 | 7–10 | 87–90 | 1.5–1.8 | 1 | smo.–irr. | straight | simple | con–acute | 0.4–0.61 | flat | Present study |
| M. apurense | 48–52 | 140–147 | 36–38 | 9–12 | 1–2 | 8–9 | 93–95 | – | 0 | smo. | straight | lobulated, without projections | con.–rounded | 0.38–0.45 | rounded | Crozzoli and Lamberti (2001) |
| M. campbelli | 58–62 | 102–108 | 25–28 | 11–15 | – | 5–7 | 89–93 | 1.2–1.8 | rare | smo.–ser. | straight | serrated | con.–pointed | 0.37–0.45 | – | Wouts (2006) |
| M. juliae | 79–86 | 89–95 | 26 | 8–9 | 1–2 | 5–6 | 91–92 | – | few | smo. | straight | with two projections | conical | 0.46–0.51 | flat | Crozzoli and Lamberti (2001) |
| M. lamothei | 65–70 | 89–98 | 17–31 | 6–8 | 0–2 | 4–8 | 91–95 | rare | smo. | slightly curved | with two projections | con.–truncate | 0.40–0.49 | rounded | Cid del Prado Vera (2009) | |
| M. lobellum | 51–60 | 85–92 | 25 | 6–7 | 2–3 | 3–4 | 92–93 | 0.9–1.2 | rare | – | straight | simple | rounded | 0.41–0.53 | rounded | Pramodini et al. (2007) |
| M. malagutii | 45–49 | 108–114 | 33–35 | 8–9 | 2 | 4–6 | 93–94 | – | many | smo. | straight | lobulated, without projections | con.–truncate | 0.34–0.40 | rounded | Crozzoli and Lamberti (2001) |
| M. nebraskense | 45–59 | 84–113 | 24–31 | 6–11 | 2–3 | 3–7 | 90–96 | 0.7–1.6 | 1–4 | smo. | straight | with two projections | rounded | 0.39–0.60 | rounded | Olson et al. (2017) |
| M. ovospermatum | 82 | 109 | 38 | 9 | 3 | 6 | 93 | – | 0 | finely crenate | straight | ? | conical | 0.22 | ? | Mohilal and Dhanachand (1998) |
| M. ozarkiense | 49–61 | 107–119 | 27–34 | 10–14 | 2–4 | 6–10 | 89–93 | 1.0–1.4 | 0–1 | som.–irr. | straight | simple | conical | 0.38–0.51 | flat | Cordero et al. (2012) |
| M. theobromae | 47–51 | 73–74 | 24–26 | 7 | 1 | 5 | 93–94 | – | many | som. | straight | lobulated, without projections | conical | 0.27–0.32 | rounded | Crozzoli and Lamberti (2001) |
| M. waitha | 66–78 | 117-140 | 26–31 | 9–10 | 4–5 | 4–6 | 92–94 | 0.9–1.0 | many | som.-irr. | sigmoid | simple | rounded | 0.43-0.53 | flat? | Pramodini et al. (2006) |
1 Notes: St, stylet; Anas., Anastomoses; A. M., Annuli margin; A. V. lip, Anterior vulval lip; S. M. L, Submedian lobes; smo., smooth; ser., serrate; irr., irregular; con, conical; Original des., Original description.
Note1: Mesocriconema bakeri (Wu, 1965; Loof and De Grisse, 1989); M. calvatum (Eroshenko, 1982; Loof and De Grisse, 1989); M. hymenophorum (Wouts and Sturhan, 1999); M. longistyletum (De Grisse and Maas, 1970; Loof and De Grisse, 1989); M. paramonovi (Razzhivin, 1974; Loof and De Grisse, 1989); M. variabile (Raski and Golden, 1966; Brzeski et al., 2002) and M. yukonense (Ebsary, 1982, 1991) that have been listed in Brzeski et al. (2002), transferred to Neobakernema (Ebsary, 1981b) by Geraert (2010).
Note2: Mesocriconema incrassatum (Raski and Golden, 1966; Loof and De Grisse, 1989) that has been listed in Brzeski et al. (2002), transferred to Lobocriconema (De Grisse and Loof, 1965) by Geraert (2010).