Morphological and molecular characterisation of some Hemicriconemoides species (Nematoda: Criconematidae) together with a phylogeny of the genus

in Nematology
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Sheathoid nematodes of the genus Hemicriconemoides are migratory root-ectoparasites of many plants including various agricultural crops and fruit trees. They are generally found inhabiting warm areas of the world and presently consist of 52 valid species. In this study we provide morphological and molecular characterisation of 12 species of this genus viz.: H. alexis, H. brachyurus, H. californianus, H. chitwoodi, H. macrodorus, H. minutus, H. ortonwilliamsi, H. promissus, H. silvaticus, H. strictathecatus, H. wessoni and Hemicriconemoides sp. originating from China, Greece, Japan, Myanmar, Spain, South Africa and the USA. Morphological descriptions, measurements, light and scanning electron microscopic observations and drawings are given for several species. Phylogenetic relationships within Hemicriconemoides, as inferred from the analyses of the D2-D3 of 28S rRNA and ITS-rRNA gene sequences, resulted in trees with three major clades that corresponded with species groupings based on morphology of the lip pattern and vulval flap. PCR with species-specific primers were developed for H. californianus, H. chitwoodi and H. strictathecatus.

Morphological and molecular characterisation of some Hemicriconemoides species (Nematoda: Criconematidae) together with a phylogeny of the genus

in Nematology

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References

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Figures

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    Anterior regions of Hemicriconemoides species. A: H. alexis, MN, USA (CD1163); B: H. alexis, topotype Greece; C: H. brachyurus, Japan; D: H. californianus, CA, USA (CD1021); E, F: H. chitwoodi, topotype FL, USA (CD1185); G: H. chitwoodi, Japan (intercepted in Italy); H: H. chitwoodi, Japan; I, J: H. macrodorus, topotype, Spain; K, L: H. minutus, topotype FL, USA; M: H. ortonwilliamsi, Spain; N, O: H. ortonwilliamsi, Italy. (Scale bar = 50 μm.)

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    Anterior regions of Hemicriconemoides species. A, B: H. promissus, Spain; C: H. silvaticus, Japan; D: H. strictathecatus, China (intercepted in Italy); E-H: H. strictathecatus, South Africa; I-J: H. wessoni, FL, USA. (Scale bar = 50 μm.)

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    Posterior regions of Hemicriconemoides species. A: H. alexis, topotype Greece; B: H. alexis, MN, USA (CD1163); C: H. brachyurus, Japan; D, E: H. californianus, CA, USA (CD1021); F: H. chitwoodi, Japan (intercepted in Italy); G: H. chitwoodi, Japan; H: H. chitwoodi, topotype FL, USA (CD1185); I, J: H. macrodorus, topotype, Spain; K, L: H. minutus, topotype FL, USA; M: H. ortonwilliamsi, Italy; N, O: H. ortonwilliamsi, Spain. (Scale bar = 50 μm.)

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    Posterior regions of Hemicriconemoides species. A, B: H. promissus, Spain; C, D; H. silvaticus, Japan; E: H. strictathecatus, China (intercepted in Italy); F-H: H. strictathecatus, South Africa; I, J: H. wessoni, FL, USA. (Scale bar = 50 μm.)

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    Hemicriconemoides californianus female CA, USA (CD847). A: Anterior region; B: Posterior region; C: Annuli at mid-body. CD1021. D, E: Anterior region; F: Posterior region; G: Annuli at mid-body; H, I: Posterior regions. (Scale bar = 30 μm.)

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    Hemicriconemoides californianus female CA, USA (CD847). A: Anterior region; B: Annuli at mid-body; C: Posterior region, ventral view.

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    Hemicriconemoides chitwoodi topotype female FL, USA (CD1185). A: Anterior region; B, C: Posterior regions; D: Annuli at mid-body; E: Lip region of another female. Hemicriconemoides minutus topotype female FL, USA (CD1181). F: Anterior region; G, H: Posterior regions; I: Annuli at mid-body; J: Tail of another female. J4 Juvenile (CD1181). K: Tail region; L: Annuli in posterior part of body. (Scale bar = 30 μm.)

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    Hemicriconemoides chitwoodi topotype female FL, USA (CD1185). A: Anterior region with lateral view of lip region; B: En face view of lip region; C: Annuli at mid-body; D: Anterior region with sheath pushed anteriorly over lip region; E: Posterior region, ventral view.

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    Hemicriconemoides minutus topotype female FL, USA (CD1181). A, B: Anterior region; C: En face view of lip region; D: Annuli at mid-body; E: Posterior region with vulva and two small vulval flaps laterally with rounded tail.

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    Hemicriconemoides strictathecatus female South Africa (Tvl1948). A: Anterior region; B, C: Posterior regions; D: Lip region of another female; E: Annuli at mid-body. Female (N826). F: Anterior region; G: Posterior region; H: Anastomosing annuli at mid-body; I, J: Tail regions; K: Annuli at mid-body. Male (N826). L: Anterior region; M: Tail region; N: Lateral field at mid-body. (Scale bar = 30 μm.)

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    Hemicriconemoides strictathecatus female South Africa (N826). A: Anterior region with lateral view of lip region; B: Another view of lip region; C: Lip region with sheath pushed forward; D: En face view of lip region; E: Annuli at mid-body; F-H: Various tail regions.

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    Hemicriconemoides wessoni female FL, USA (CD1107). A: Anterior region; B: Annuli at mid-body; C-F: Various tail regions. Female (CD1054). G: Anterior region; H: Annuli at mid-body; I-L: Various tail regions. (Scale bar = 30 μm.)

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    Hemicriconemoides wessoni female FL, USA (CD1107). A: Lateral view of lip region; B: En face view of lip region with sheath pushed anteriorly; D: Annuli at mid-body; E: Tail region with distinct vulval flaps. Female FL, USA (CD1154). C: En face view of lip region; F: tail region, ventral view with vulval opening and vulval flaps.

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    Phylogenetic relationships within populations and species of Hemicriconemoides as inferred from Bayesian analysis using the D2-D3 of 28S rRNA gene sequence dataset with the GTR + I + G model. Posterior probability more than 70% is given for appropriate clades. Newly obtained sequences are indicated in bold.

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    Phylogenetic relationships within populations and species of Hemicriconemoides as inferred from Bayesian analysis using the ITS-rRNA gene sequence dataset with the GTR + I + G model. Posterior probability more than 70% is given for appropriate clades. Newly obtained sequences are indicated in bold.  – several nucleotides from 3′ end were excluded from the original sequence due to possible mistakes in sequence reading; 1 – identified as H. californianus in GenBank and the article by Chen et al. (2007); 2 – identified as H. strictathecatus in GenBank and as H. litchi in the article by Chen et al. (2011); 3 – identified as H. strictathecatus in GenBank and as H. mangiferae in the article by Chen et al. (2011).

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    Phylogenetic relationships within populations and species of Hemicriconemoides as inferred from Bayesian analysis using the combined dataset (the D2-D3 of 28S rRNA and the ITS-rRNA gene sequences) with mapping of lip region structure, vulval region and tail terminus shapes. Posterior probabilities for BI (GTR + I + G model) and bootstrap values for MP are given for appropriate clades. Lip pattern: A1: first type; A2: second type. Vulval flaps: B1: absent; B2: present. Tail terminus shape: C1: pointed; C2: rounded.  Lip region structure of H. silvaticus is unknown. Paratylenchus species were used as outgroups.

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    Gel with specific amplicons obtained in the results of PCR with species-specific primers. A: PCR with the Hemicriconemoides californianus-specific primer (TW81 + H_califor); B: PCR with the H. strictathecatus-specific primer (TW81 + H_strict); C: PCR with the H. chitwoodi-specific primer (TW81 + H_chitw). Lanes: M = 100 bp DNA ladder (Promega, USA); 1 = H. californianus CA, USA (CD847); 2 = H. californianus CA, USA (CD907); 3 = H. strictathecatus Venezuela (839); 4 = H. strictathecatus South Africa (CD896); 5 = H. chitwoodi topotype FL, USA (CD1185); 6 = H. minutus topotype FL, USA (CD1181); 7 = H. wessoni FL, USA (CD1107); 8 = control without DNA.

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