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Description of Pratylenchus Hispaniensis N. Sp. from Spain And

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Description of Pratylenchus Hispaniensis N. Sp. from Spain And Nematology, 2010, Vol. 12(3), 429-451 Description of Pratylenchus hispaniensis n. sp. from Spain and considerations on the phylogenetic relationship among selected genera in the family Pratylenchidae ∗ Juan E. PALOMARES-RIUS 1,PabloCASTILLO 1, , Gracia LIÉBANAS 2,NicolaVOVLAS 3, Blanca B. LANDA 1,JuanA.NAVA S -CORTÉS 1 and Sergei A. SUBBOTIN 4,5 1 Institute of Sustainable Agriculture (IAS), Spanish National Research Council (CSIC), Alameda del Obispo s/n, Apdo. 4084, 14080 Córdoba, Spain 2 Department of Animal Biology, Vegetal Biology and Ecology, University of Jaén, Campus ‘Las Lagunillas’ s/n, Edificio B3, 23071 Jaén, Spain 3 Istituto per la Protezione delle Piante (IPP), Sezione di Bari, Consiglio Nazionale delle Ricerche (C.N.R.), Via G. Amendola 122/D, 70126 Bari, Italy 4 Plant Pest Diagnostic Center, California Department of Food and Agriculture, 3294 Meadowview Road, Sacramento, CA 95832-1448, USA 5 Centre of Parasitology of A.N. Severtsov Institute of Ecology and Evolution of the Russian Academy of Sciences, Leninskii Prospect 33, Moscow, 117071, Russia Received: 31 July 2009; revised: 12 October 2009 Accepted for publication: 13 October 2009 Summary – A new amphimictic species, Pratylenchus hispaniensis n. sp., parasitising the roots of gum cistus in Andújar (Jaén), southern Spain, is described. The new species is characterised by the presence of numerous males and by the female having a lip region with three annuli, a divided face, a robust stylet (14.5-17.0 μm) with rounded knobs, lateral fields with four lines, V = 80-84, a round spermatheca full of sperm, well developed post-vulval uterine sac and an obliquely truncate tail with irregularly annulated terminus. Morphologically this species is related to P. bhatti, P. kralli, P. mediterraneus, P. pseudofallax and P. thornei. A phenetic study of the 25 most useful diagnostic morphological and allometric characters for Pratylenchus species was done using multivariate factor and linear discriminant analyses. In the factor analysis the first seven factors accounted for 71.1% of the total variance of the characters selected. These factors were related to female tail, pharyngeal overlap, reproductive behaviour, stylet length, L/post-vulval uterine sac ratio, body length and number of lip annuli. Discriminant analysis differentiated Pratylenchus spp. from the three valid species of Zygotylenchus. The results of the phylogenetic analysis based on sequences of the D2-D3 expansion regions of 28S, partial 18S and ITS rRNA genes confirmed the close relationship of P. hispaniensis n. sp. with P. mediterraneus and inferred molecular affinity with P. brzeskii, P. neglectus and P. thornei, in spite of variation in the position of P. hispaniensis n. sp. in the clades. Additional phylogenetic analyses based on the same sets of sequences for P. hispaniensis n. sp., Zygotylenchus guevarai and other Pratylenchidae indicated that Pratylenchus includes several paraphyletic lineages; however, likelihood tests did not reject monophyly of the genus. The inclusion of Pratylenchus, Zygotylenchus, Hirschmanniella, Nacobbus and Apratylenchus in Pratylenchidae was supported. Keywords – Cistus ladanifer, D2-D3 region, gum cistus, molecular, morphology, morphometrics, root-lesion nematodes, SEM, Shimodaira-Hasegawa test, taxonomy. The genus Pratylenchus Filipjev, 1936 contains more these characters and the many incomplete descriptions than 70 species of root-lesion nematodes. The morpholo- published in the literature (Castillo & Vovlas, 2007). As gical identification and delimitation of these species re- a result, the actual number of valid species has varied ac- mains problematic due to their high morphological plas- cording to author (see Siddiqi, 1986, 2000; Ryss, 1988, ticity, the small number of diagnostic features available 2002a, b; Frederick & Tarjan, 1989; Handoo & Gol- at species level, the intraspecific variability of some of den, 1989; Loof, 1991; Duncan et al., 1999; Castillo & ∗ Corresponding author, e-mail: [email protected] © Koninklijke Brill NV, Leiden, 2010 DOI:10.1163/138855409X12559479585043 Also available online - www.brill.nl/nemy 429 J.E. Palomares-Rius et al. Vovlas, 2007). Species differentiation is also difficult in limitation within and between Pratylenchus and Zygoty- the genus Zygotylenchus Siddiqi, 1963, a well defined lenchus taxa based on a multivariate analyses statistical genus commonly occurring in the Mediterranean region approach using the most useful diagnostic morphological and other geographical areas (Siddiqi, 2000; Urek et al., and allometric characters for Pratylenchus species. 2003; Lisková et al., 2007). It contains only three valid Sequence analyses of nuclear ribosomal RNA genes species, Z. guevarai (Tobar Jiménez, 1963) Braun & Loof, have been used for molecular characterisation and recon- 1966, Z. natalensis Van den Berg, 2003, and Z. taomasi- struction of phylogenetic relationships of Pratylenchus nae (de Guiran, 1963) Braun & Loof, 1966, plus another, spp. (Al-Banna et al., 1997; Duncan et al., 1999; Carta poorly defined, species, Z. biterminalis Razjivin & Milan, et al., 2001; De Luca et al., 2004; Subbotin et al., 2008). 1978, which has been considered incertae sedis (Siddiqi, These studies have clarified the taxonomical status of a 2000). Morphologically, Zygotylenchus is an easily recog- large number of root-lesion nematodes, although many nised taxon, although its validity has not previously been species have yet to be so characterised. corroborated by molecular data. There has been consensus among taxonomists in con- Configuration of the lip pattern has been used suc- sidering the two genera mentioned above along with cessfully in the separation of Pratylenchus species. Lip Achlysiella Hunt, Bridge & Machon, 1989, Apratylen- patterns of en face with SEM were used to reveal phy- choides Sher, 1973, Apratylenchus Trinh, Waeyenberge, logenetic relationships in several plant-parasitic nema- Nguyen, Baldwin, Karssen & Moens, 2009, Hirschman- tode genera (Baldwin, 1992), as well as in the Praty- niella Luc & Goodey, 1964, Hoplotylus s’Jacob, 1960, lenchus (Corbett & Clark, 1983). They were coincident Nacobbus Thorne & Allen, 1944, Pratylenchoides with major clades of selected species, including Praty- Winslow, 1958, Radopholus Thorne, 1949 and Zygradus lenchus brachyurus (Godfrey, 1929) Filipjev & Schuur- Siddiqi, 1991 as members of the family Pratylenchidae mans Stekhoven, 1941, P. coffeae (Zimmermann, 1898) Thorne, 1949 (Luc, 1987; Siddiqi, 2000; Trinh et al., Filipjev & Schuurmans Stekhoven, 1941, P. goodeyi Sher 2009). Recent molecular and phylogenetic studies con- & Allen, 1953, P. loosi Loof, 1960, P. panamaensis Sid- cerning the relationships within these genera using se- diqi, Dabur & Bajaj, 1991 (= P. gutierrezi Golden, López quences of the D2-D3 expansion regions of 28S rRNA & Vilchez, 1992), P. pratensis (de Man, 1880) Filipjev, (Subbotin et al., 2006; Trinh et al., 2009) and 18S rRNA 1936, and P. pseudocoffeae Mizukubo, 1992, obtained (Holterman, 2007; Bert et al., 2008) have indicated that from Bayesian analysis of D2-D3 expansion segments of Radopholus and Pratylenchoides stood apart from other 28S rRNA and partial 18S rRNA (Duncan et al., 1999; genera and did not belong to this family. These findings Handoo et al., 2008; Subbotin et al., 2008). may cast doubt on the inclusion of Achlysiella, Hoploty- Only recently has there been an interest in applying lus and Zygradus in the Pratylenchidae since they are mor- morphology for cladistic and phenetic analysis of the phologically related to Radopholus and Pratylenchoides. genus Pratylenchus. Duncan et al. (1999) showed that During nematode surveys conducted in cultivated and principal component analysis (PCA) of one morpholo- natural environments in southern Spain, an amphimictic gical (smooth face vs divided face) and three allomet- root-lesion nematode was detected in roots of gum cis- ric characters (length of stylet, ratios V and a) revealed tus, a flowering ornamental shrub used for the extraction congruence with phylogenetic relationships inferred from of essential oils. Preliminary morphological examinations analysis of 28S rDNA sequences based on 32 Praty- indicated that this amphimictic species did not fit any de- lenchus isolates with two lip annuli and amphimictic scription of known Pratylenchus species and appeared to reproduction (i.e., presence of males). Ryss (2002a, b) be morphologically related to P. mediterraneus Corbett, provided a comprehensive morphological analysis of 49 1983, P. neglectus (Rensch, 1924) Filipjev & Schuurmans species and 26 characters of Pratylenchus and, simultane- Stekhoven, 1941 and P. thornei Sher & Allen, 1953. In or- ously, Carta et al. (2002) published a morphologically- der to verify the taxonomic status of this species, we con- based phylogenetic tree for 11 Pratylenchus species. ducted a morpho-biological and molecular study of this However, these statistical approaches have not been ap- unknown Pratylenchus, which is described herein as P. plied to compare and differentiate species groups in Praty- hispaniensis n. sp. These morpho-biological and molec- lenchus or to validate differences with Zygotylenchus. ular analyses were expanded to other valid members of Therefore, one of the main objectives of the present study the family Pratylenchidae and included two Californian was to provide insights into the genus and species de- populations of P. neglectus and P. thornei, and an Italian 430 Nematology Morphology and phylogeny of Pratylenchus and
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