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Parapristionchus Giblindavisi N. Gen., N. Sp. (Rhabditida: Diplogastridae) Isolated from Stag Beetles (Coleoptera: Lucanidae) in Japan

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Parapristionchus Giblindavisi N. Gen., N. Sp. (Rhabditida: Diplogastridae) Isolated from Stag Beetles (Coleoptera: Lucanidae) in Japan Nematology, 2012, Vol. 14(8), 933-947 Parapristionchus giblindavisi n. gen., n. sp. (Rhabditida: Diplogastridae) isolated from stag beetles (Coleoptera: Lucanidae) in Japan Natsumi KANZAKI 1,ErikJ.RAGSDALE 2, Matthias HERRMANN 2,WernerE.MAYER 2,†, ∗ Ryusei TANAKA 1 and Ralf J. SOMMER 2, 1 Forest Pathology Laboratory, Forestry and Forest Products Research Institute, 1 Matsunosato, Tsukuba, Ibaraki 305-8687, Japan 2 Max Planck Institute for Developmental Biology, Department of Evolutionary Biology, Spemannstraße 37, Tübingen, Germany Received: 3 January 2012; revised: 1 March 2012 Accepted for publication: 1 March 2012; available online: 15 May 2012 Summary – A new species of diplogastrid nematode, isolated in a previous survey of nematodes associated with stag beetles in Japan, is described as Parapristionchus giblindavisi n. gen., n. sp. Parapristionchus n. gen. differs from other diplogastrid genera chiefly by its stomatal morphology. Distinguishing the genus are the presence of a claw-like dorsal tooth in both the eurystomatous and stenostomatous forms and the division of the cheilostom into 12 plates lacking apical flaps. According to phylogenetic analysis of nine ribosomal protein gene sequences, Parapristionchus n. gen. shows deep divergence from other known genera. Molecular evidence strongly supports P. giblindavisi n. gen., n. sp. + Pristionchus spp. as monophyletic with respect to all other diplogastrids examined. Congruent with a clade of P. giblindavisi n. gen., n. sp. + Pristionchus spp. is the shared presence of a bifurcate P7 genital papilla. Discovery and description of a close sister group to Pristionchus, a model biological system, enables character polarisation in macroevolutionary studies of Pristionchus nematodes. Keywords – Dorcus rubrofemoratus, molecular, morphology, morphometrics, new genus, new species, phylogeny, taxonomy. The nematode family Diplogastridae Micoletzky, 1922 genera according to phylogenetic analysis of D2/D3 large currently includes 30 genera (Sudhaus & Fürst von subunit (LSU) rRNA and near full length small subunit Lieven, 2003; Kanzaki et al., 2009a; Mayer et al., 2009; (SSU) rRNA. ‘Pristionchus sp.’ was clearly differentiated Fürst von Lieven et al., 2011; Susoy & Herrmann, 2012), from the Pristionchus clade by both morphological and species of which are generally regarded to be insect- molecular criteria, although it was closer to Pristionchus associated nematodes. Kanzaki et al. (2011) surveyed ne- than any other genus included in that study. matodes associated with stag beetles (Lucanidae) in Japan The growing use of Pristionchus Kreis, 1932 as a model and reported seven diplogastrid species from eight species biological system (Sommer, 2009) highlights the need of stag beetles. Isolated nematodes included Koerneria for resolved systematics of the genus. Studies of macro- luziae (Körner, 1954) Meyl, 1960, K. lucani (Körner, evolution, which place P.pacificus Sommer, Carta, Kim & 1954) Meyl, 1960, Pseudodiplogasteroides compositus Sternberg, 1996 as a focal point, require proper outgroups Körner, 1954, Pseudodiplogasteroides sp., Rhabditidoi- for polarising character transitions across Pristionchus des sp., Pristionchus cf. pacificus, and ‘Pristionchus sp.’ species. Although ingroup relationships of the genus are Among these seven species, the taxonomic affiliation of increasingly resolved (Mayer et al., 2007, 2009), stud- two species, K. luziae and ‘Pristionchus sp.’, were uncer- ies have not agreed on placing an outgroup to the genus tain, as both were separated from their respective assigned (Mayer et al., 2007, 2009; van Megen et al., 2009; Kion- † Deceased 23 January 2012. * Corresponding author, e-mail: [email protected] © Koninklijke Brill NV, Leiden, 2012 DOI:10.1163/156854112X635878 933 N. Kanzaki et al. tke & Fitch, 2010). Alleviating this disparity, the iso- and sequence amplified for the latter isolate is given in late reported as ‘Pristionchus sp.’wasdiscoveredasa Mayer et al. (2007). taxon that breaks up the phylogenetic distance between For phylogenetic analysis, nine ribosomal protein genes Pristionchus and other diplogastrid genera (Kanzaki et were amplified and sequenced, generating a total align- al., 2011). This species is described herein as Parapris- ment of 5996 coding nucleotides. Genes included in anal- tionchus giblindavisi n. gen., n. sp. based on its apo- ysis were rpl-2, rpl-10, rpl-14, rpl-23, rpl-29, rpl-35, morphic morphological characters and phylogenetic posi- rps-14, rps-27 and rps-28. Sequences for ribosomal pro- tion as inferred from sequences of nine ribosomal protein tein genes have been deposited in GenBank under ac- genes. cession numbers JQ677914-JQ677920. All information regarding genes, primers, and PCR conditions is given Materials and methods in Mayer et al. (2009). Species selected for analysis in- clude three undescribed species isolated from Japan (Pris- NEMATODE ISOLATION AND CULTIVATION tionchus sp. 10, Pristionchus sp. 14, and Pristionchus sp. 15), which are the most basal known representatives of the Parapristionchus giblindavisi n. gen., n. sp. was iso- genus Pristionchus and thus likely to be the Pristionchus lated from an adult of Dorcus rubrofemoratus Vollen- species closest to P. giblindavisi n. gen., n. sp. Molecu- hoven (Coleoptera: Lucanidae) collected in Iwaizumi, lar sequence data for Pristionchus species were collected Iwate Prefecture, Japan, in September 2010. The strain and reported by Mayer et al. (2007). Other species in- has been kept in laboratory culture on NGM agar plates cluded represent other diplogastrid and outgroup genera seeded with Escherichia coli strain OP50, under the cul- for which homologous sequence data are already pub- ture code and freezing voucher number RS5555. Detailed lished (Mayer et al., 2009). Isolation details for included isolation conditions for the strain are described in Kanzaki Pristionchus species other than P. giblindavisi n. gen., n. et al. (2011). sp. are given in Mayer et al. (2007) and Kanzaki et al. (2012). Isolation details for species of other genera are MORPHOLOGICAL OBSERVATION AND PREPARATION given in Mayer et al. (2009). OF TYPE MATERIAL The concatenated dataset of ribosomal protein genes One- to 2-week-old cultures of P. giblindavisi n. gen., was aligned in ClustalX and then analysed in MEGA5.05 n. sp. were prepared for morphological observation. Light (Tamura et al., 2011) after complete deletion of gapped microscopic observations were conducted using live ne- or ambiguous positions. Substitution models were evalu- matodes, which were handpicked from culture plates. To ated in MEGA5. The model with the lowest Bayesian In- + + prepare type material, nematodes were isolated from cul- formation Criterion (BIC) score Tamura-Nei93 I G, tures, rinsed in distilled water to remove bacteria, heat- was chosen for analysis. Phylogenetic analysis was per- killed at 65°C, fixed in 5% formalin, and processed formed using the maximum likelihood (ML) criterion as through a glycerin and ethanol series using Seinhorst’s implemented in MEGA5. The taxon Rhabditoides inermis method (see Hooper, 1986). (SB328) was specified as outgroup, as informed by more broadly representative phylogenetic analyses (van Megen MOLECULAR CHARACTERISATION AND PHYLOGENY et al., 2009). Node support was evaluated by 500 boot- strap pseudoreplicates. In a previous study, all isolates other than that from Okinawa (see below) were confirmed to be the same Results species by their D2/D3 LSU and near full length SSU * rRNA sequences (Kanzaki et al., 2011), each of which Parapristionchus n. gen. was identical across isolates. Details about primers and sequences amplified are given therein. To confirm that DESCRIPTION the isolate collected from Okinawa belongs to the same Diplogastridae. Body cylindrical, stout. Cuticle thick, species as the others, an 830 bp fragment of the SSU with fine annulation and clear longitudinal striations. Lips rRNA gene was extracted, amplified, and sequenced for the isolate. The SSU rRNA sequence was identical to * Derived from the Greek παρα (‘near’) + Pristionchus,a that of the above isolates and is not discussed further closely related genus according to morphological and molecular herein. Information regarding primers, PCR conditions, evidence. 934 Nematology Parapristionchus giblindavisi n. gen., n. sp. from Japan not clearly separated from each other or from rest of body. stomatous form. According to phylogenetic analysis of Six short and papilliform labial sensilla present in male SSU rRNA sequences, Parapristionchus n. gen. and Pris- and female, and four small, papilliform cephalic papil- tionchus form a well-supported monophyletic clade but lae present in male, as typical for diplogastrid nema- are clearly distinguished from each other by deep branch todes. Stomatal dimorphism with stenostomatous (narrow lengths (Kanzaki et al., 2011). mouth) and eurystomatous (wide mouth) forms occurring Parapristionchus n. gen. is morphologically close to in both male and female. In both forms, cheilostom con- Micoletzkya Weingärtner, 1955 but is distinguished from sisting of 12 well-cuticularised and conspicuous plates, the latter genus by its cheilostomatal structure in both the and a thick, well-cuticularised gymnostom forming a steno- and eurystomatous form, having 12 vs six plates short tube or ring. Stegostom bearing a conspicuous and and by the stegostom in both mouth forms, the postdental movable dorsal, claw-like tooth in both forms. Stegos- part of which in Parapristionchus n.
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