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Bursaphelenchus Xylophilus ISPM 27 27 ANNEX 10 ENG DP 10: Bursaphelenchus xylophilus INTERNATIONAL STANDARD FOR PHYTOSANITARY MEASURES PHYTOSANITARY FOR STANDARD INTERNATIONAL DIAGNOSTIC PROTOCOLS Produced by the Secretariat of the International Plant Protection Convention (IPPC) This page is intentionally left blank This diagnostic protocol was adopted by the Standards Committee on behalf of the Commission on Phytosanitary Measures in January 2016. The annex is a prescriptive part of ISPM 27. ISPM 27 Diagnostic protocols for regulated pests DP 10: Bursaphelenchus xylophilus Adopted 2016; published 2016 CONTENTS 1. Pest Information ............................................................................................................................... 2 2. Taxonomic Information .................................................................................................................... 3 3. Detection ........................................................................................................................................... 3 3.1 Detection in trees ............................................................................................................... 4 3.2 Detection by the use of insect traps, trap logs and in samples from sawmills and timber yards ....................................................................................................................... 4 3.3 Direct detection in wood, wood products and solid wood packaging ............................... 5 3.4 Extraction of nematodes from wood samples ................................................................... 5 3.5 Extraction of nematodes from vector insects .................................................................... 5 4. Identification ..................................................................................................................................... 6 4.1 Morphological identification ............................................................................................. 7 4.1.1 Preparation of specimens .................................................................................................. 7 4.1.2 Key to species level ........................................................................................................... 8 4.1.3 Comparison of Bursaphelenchus xylophilus with similar species .................................. 12 4.2 Molecular identification .................................................................................................. 13 4.2.1 ITS-PCR RFLP ............................................................................................................... 13 4.2.2 Conventional PCR ........................................................................................................... 15 4.2.3 Real-time PCR ................................................................................................................ 16 4.2.4 RNA-based molecular tests for detection of living Bursaphelenchus xylophilus ........... 18 4.2.5 LAMP .............................................................................................................................. 19 4.2.6 Controls for molecular tests ............................................................................................ 20 4.2.7 Interpretation of results from PCR .................................................................................. 21 4.2.8 Sequencing ...................................................................................................................... 21 5. Records ........................................................................................................................................... 22 6. Contact Points for Further Information .......................................................................................... 22 7. Acknowledgements ........................................................................................................................ 23 8. References ...................................................................................................................................... 24 9. Figures ............................................................................................................................................ 29 International Plant Protection Convention DP 10-1 DP 10 Diagnostic protocols for regulated pests 1. Pest Information The pine wood nematode, Bursaphelenchus xylophilus (Steiner and Buhrer, 1934) Nickle 1970, is the causal agent of pine wilt disease. B. xylophilus is believed to be native to North America, where it is widely distributed in Canada and the United States (Ryss et al., 2005) and is apparently of limited distribution in Mexico (Dwinell, 1993). North American pine species are resistant or at least tolerant to B. xylophilus, but exotic species planted in North America, especially in the warmer southern areas of the United States, are killed when attacked by the nematode. B. xylophilus was carried to Japan at the beginning of the twentieth century, presumably on timber exported from North America, and it became one of the most damaging forest pests in the country, where it still causes remarkable losses of pine trees (Pinus densiflora, P. thunbergii and P. luchuensis) today. B. xylophilus was also introduced to China (including Taiwan) and Korea; it was found there in the mid to late 1980s. In 1999, B. xylophilus was found for the first time in Europe (Portugal) on P. pinaster, which is killed by the nematode within a few months after infestation (Mota et al., 1999; Fonseca et al., 2012). B. xylophilus has also been detected on P. nigra and P. radiata in Portugal and Spain, respectively (Inácio et al., 2014; Zamora et al., 2015). In 2008, B. xylophilus was found for the first time in Spain (Abelleira et al., 2011). B. xylophilus is transmitted from tree to tree by wood-inhabiting beetles of the genus Monochamus (Coleoptera: Cerambycidae) (Linit, 1990; Evans et al., 1996). The nematodes enter the bodies of the insects shortly after the latter emerge from pupation and just before they bore out of the host tree (Wingfield, 1987). The beetles fly to the crown of healthy trees and feed on the young shoots and leaves (maturation feeding). They then mate and the females search for a weakened tree or one that has died recently, or for trunks or bigger branches (including felling debris), depending on the Monochamus species, where they lay their eggs through the bark. The beetle larvae that hatch from the eggs feed in the cambial tissues just below the bark for several months. On reaching maturity, they bore deeper into the wood to pupate, and thus their life cycle is completed. B. xylophilus takes advantage of this life cycle to obtain transport to new host trees (Wingfield, 1987). Their introduction into the new tree may take place during oviposition by the beetle (this appears to be the only means of transmission for several species of Bursaphelenchus that colonize dead trees) (Edwards and Linit, 1992). B. xylophilus, however, seems to be unique among these species in that it can also be transmitted to a new tree during maturation feeding by beetles, and the development of pine wilt disease can occur as a consequence of transmission through the young shoots (Wingfield, 1987). When B. xylophilus is transmitted during oviposition, the nematodes remain relatively close to the site of introduction. But when transmission occurs through the young shoots and when the tree succumbs to pine wilt disease, the nematodes distribute throughout the whole tree, destroying wood tissues such as epithelial cells, parenchyma cells of axial and radial resin canals, cambium and phloem. B. xylophilus can also be found in roots, even when the above-ground part of the tree is already dead, dried out or felled. Whether the tree develops pine wilt disease depends on the tree species (in general only Pinus spp. of non-American origin are affected), its state of health and the climatic conditions (particularly temperature and water supply). These factors also influence the distribution of nematodes throughout the tree: their distribution can be localized or irregular and this needs to be taken into account in sampling strategies (Schröder et al., 2009). B. xylophilus can also be found in dead trees of Abies, Chamaecyparis, Cedrus, Larix, Picea and Pseudotsuga and other conifers (except Thuja spp.), but none of these genera is known to be affected by pine wilt disease, although pathogenicity tests on seedlings show remarkable reactions, including death (Evans et al., 1996). B. xylophilus is almost exclusively vectored by Monochamus species, with the vector species varying among the geographic regions; for example, M. alternatus in China and Japan, M. saltuarius in Japan, M. carolinensis in North America and M. galloprovincialis in Portugal. Occasionally, other beetles of the family Cerambycidae or other Coleoptera have been found to carry “dauer” juveniles of the DP 10-2 International Plant Protection Convention Diagnostic protocols for regulated pests DP 10 nematode on their bodies, but there is no evidence that they play a role as vectors in the dissemination of the nematode (Evans et al., 1996). Human activity is known to be the principal route for dispersal of B. xylophilus over greater distances and B. xylophilus and its vectors have been intercepted on a number of occasions in the international trade of wood, wood products and, most notably, solid wood packaging made from conifers. Therefore, the risk of further international
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