Deladenus nitobei n. sp. (Tylenchomorpha: Allantonematidae) isolated from Sirex nitobei (Hymenoptera: Siricidae) from Aomori, Japan, a new member of the siricidicola superspecies

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Deladenus nitobei n. sp., a parasite of a woodwasp species, Sirex nitobei, is described based on its typological characters and molecular profiles of part of the small subunit D2-D3 expansion segments of the large subunit and internal transcribed spacer region of the ribosomal RNA gene, as well as part of the mitochondrial cytochrome oxidase subunit I (mtCOI) gene. Adult host woodwasps emerging from dead Japanese red pine logs, Pinus densiflora, collected at Aomori, Japan, were dissected and examined for nematode association. The new species was isolated from the body cavity and reproductive system of S. nitobei as large parasitic females and small parasitic juveniles. The nematodes were cultured successfully on 1.0% malt extract agar medium, inoculated with Sirex-associated fungus, Amylostereum areolatum. The mycophagous adult nematodes were characterised by the relative position of the excretory pore, located at 25 (19-28) and 25 (18-30) μm anterior to the hemizonid in the male and female, respectively, and a broad female tail with a rounded distal end. Typologically, the new species forms a cryptic species complex with D. siricidicola and D. canii. In addition, the new species and D. siricidicola share the same host wasp, tree and fungal species in Japan. However, the cryptic species can be separated from each other based on the described morphological and molecular sequence differences in the mtCOI gene.

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References
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Figures
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    Mycophagous male of Deladenus nitobei n. sp. A: Entire body, right lateral view; B: Anterior region, right lateral view; C: Stylet; D: Body surface showing deirid, lateral field and annulations; E: Lateral field at mid-body; F: Anterior end of testis; G: Posterior part of testis and vas deferens producing large amoeboid sperm in right lateral view; H: Middle-posterior part of testis producing small sperm; I: Vas deferens containing small sperm, right lateral view; J, K: Male tail, right lateral view (J) and ventral view (K); L: Spicule and gubernaculum, right lateral view.

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    Mycophagous female of Deladenus nitobei n. sp. A: Entire body, right lateral view; B: Lip region, right lateral view; C: Anterior part, right lateral view; D: Posterior part of ovary containing well developed oocytes; E: Posterior part of gonad (spermatheca to vagina/vulval region), right lateral view: F, G: Vulval and anal region, right lateral view (F) and ventral view (G).

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    Infective and parasitic females of Deladenus nitobei n. sp. A: Entire infective female, left lateral view; B: Lip-stylet region of infective female, left lateral view; C: Anterior part of infective female, right lateral view; D: Body surface showing deirid, lateral field and annulations; E: Lateral field at mid-body; F: Anterior part of gonad (ovary to anterior end of uterus); G, H: Vulval and anal region, left lateral view (G) and ventral view (H); I: Entire body of parasitic female (the specimen was twisted at mid-body, showing right lateral side of anterior part and left lateral side of posterior part); J-M: Embryo in uterus from anterior to posterior part, where the uterus contains single- and two-cell stages in anterior region (J), developing embryo in multiple-celled stage (K) or forming juveniles (L) in middle section, and hatched juveniles (M) in posterior region; N: Anterior end of parasitic female, right lateral view; O: Vulval and anal region of parasitic female, left lateral view.

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    Mycophagous male of Deladenus nitobei n. sp. A-D: Anterior region, right lateral view in different focal planes; E-G: Close-up of nerve ring-pharyngeal gland part, left lateral view in different focal planes; H: Anterior part of testis; I: Middle-posterior part of testis producing small sperm; J: Vas deferens containing small sperm, right lateral view; K-M: Male tail, right lateral view showing tail tip (K), spicule and gubernaculum (L) and bursa (M). Abbreviations: d = deirid; ep = excretory pore; hem = hemizonid; nr = nerve ring; pgn = pharyngeal gland cell nucleus. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/15685411.

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    Mycophagous female of Deladenus nitobei n. sp. A, B: Anterior region, left lateral view in different focal planes showing anterior end of ovary (A), lateral field and relative position of deirid (B); C: Lip region, right lateral view; D, E: Anterior (D) and middle (E) part of ovary containing developing oocytes; F: Posterior part of ovary, oviduct and anterior end of spermatheca; G: Spermatheca containing large amoeboid sperm; H: Posterior part of body, left lateral view; I: Tail, left lateral view. Abbreviations: a = anus; c = stylet conus; cr = crustaformeria; d = deirid; ep = excretory pore; hem = hemizonid; nr = nerve ring; od = oviduct; s = stylet shaft; sk = stylet knob; sp = spermatheca; ut = uterus; vo = vulval opening. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/15685411.

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    Infective female of Deladenus nitobei n. sp. A-D: Anterior region, left lateral view in different focal planes; E: Lip region and stylet, left lateral view. Abbreviations: c = stylet conus; d = deirid; dpg = dorsal pharyngeal gland orifice; ep = excretory pore; hem = hemizonid; lfs = start of lateral field; nr = nerve ring; s = stylet shaft; svpg = subventral pharyngeal glands orifice. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/15685411.

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    Infective female of Deladenus nitobei n. sp. A, B: Anterior region of ovary in different focal planes showing relative position of ovary and pharyngeal gland cell nucleus; C: Mid-region of gonad (posterior end of ovary to anterior part of uterus); D, E: Vulva to anus region, left lateral view in different focal planes. Abbreviations: a = anus; pgn = subventral pharyngeal gland nucleus; vo = vulval opening. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/15685411.

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    Parasitic female of Deladenus nitobei n. sp. A: Anterior region, right lateral view; B-D: Head to pharynx region in different focal planes; E: Body surface showing weak and coarse annulation; F-H: Embryos deposited in anterior to posterior part of uterus; I: Vulval and anal region, left lateral view. Abbreviations: a = anus; ag = anterior glands; d = dorsal pharyngeal gland orifice; ec = excretory cell; ep = excretory pore; od = oviduct; ov = ovary; sb = stylet base; sp = spermatheca; st = stylet tip; sv = subventral pharyngeal glands orifice; ut = uterus; vo = vulval opening. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/15685411.

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    Molecular phylogenetic relationship among several tylenchid insect-parasitic species. The Maximum Likelihood tree was inferred from near-full-length SSU. The substitution model selected was GTR + I + G (lnL=7760.16, AIC = 15726.31) and parameters are: Pinvar = 0.451; Shape = 0.639. Bootstrap support exceeding 50% is shown for appropriate clades. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/15685411.

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    Molecular phylogenetic relationship among several Deladenus species. The Maximum Likelihood tree was inferred from mitochondrial cytochrome oxidase subunit I sequences. The substitution model selected was GTR + G (lnL=1853.06, AIC = 3998.13) and shape parameter is 0.175. Bootstrap support exceeding 50% is shown for appropriate clades. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/15685411.

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    Molecular phylogenetic relationship among several Deladenus species. The Maximum Likelihood tree was inferred from internal transcribed spacer region of ribosomal RNA. The substitution model was selected by JMODELTEST as HKY + G (lnL=2898.38, AIC = 6090.22) and shape parameter is 0.175. Bootstrap support exceeding 70% is shown for appropriate clades. This figure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/15685411.

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