European Journal of Plant Pathology

, Volume 133, Issue 1, pp 89–99 | Cite as

Pine Wilt Disease: a threat to European forestry

  • Cláudia Vicente
  • Margarida Espada
  • Paulo Vieira
  • Manuel MotaEmail author


Bursaphelenchus xylophilus, the pinewood nematode (PWN) and causal agent of Pine Wilt Disease (PWD), was detected for the first time, in 1999, in Portugal, and in Europe. Despite the efforts of the Portuguese National Forestry and Quarantine Authorities, the disease has spread to new forest areas in the centre of mainland Portugal, in 2008, and to the island of Madeira, in 2009. More recently, two foci of PWD were reported from Spain. The free circulation of non-treated wood and wood products, as part of global trade, may explain this phenomenon, which constitutes a threat to other European forestlands. This review gathers the most recent up-to-date information about PWD in Europe, as well as in other countries, presenting integrative management procedures for detection and control, and the scientific knowledge generated over the last decade, to understand the complex biological system behind the disease.


Bursaphelenchus xylophilus Monochamus galloprovincialis Pinewood nematode Pinus Portugal 


  1. Abelleira, A., Picoaga, A., Mansilla, J. P., & Aguin, O. (2011). Detection of Bursaphelenchus xylophilus, causal agent of pine wilt disease on Pinus pinaster in Northwestern Spain. Plant Disease, 95(6), 776.CrossRefGoogle Scholar
  2. Akbulut, S., & Stamps, W. T. (2011). Insect vectors of the pinewood nematode: a review of the biology and ecology of Monochamus species. Forest Pathology. doi: 10.1111/j.1439-0329.2011.00733.x.
  3. Anon. (2011). Forests in Europe. FOREST EUROPE (The Ministerial Conference on the Protection of Forests in Europe) (
  4. Barbosa, P., Lima, A. S., Vieira, P., Dias, L. S., Tinoco, M. T., Barroso, J. G., et al. (2010). Nematicidal activity of essential oils and volatiles derived from Portuguese aromatic flora against the pinewood nematode, Bursaphelenchus xylophilus. Journal of Nematology, 42, 8–16.Google Scholar
  5. Braasch, H. (2009). Re-establishment of Devibursaphelenchus Kakuliya, 1967 (Nematoda, Aphelenchoididae) and proposal for a new combinationof several Bursaphelenchus species. Journal of Nematode Morphology and Systematics, 12(1), 1–5.Google Scholar
  6. Braasch, H., Swart, A., Tribe, G., & Burgermeister, W. (1998). First record of Bursaphelenchus leoni in South Africa and comparison with some other Bursaphelenchus spp. Bulletin OEPP, EPPO Bulletin, 28(1–2), 211–216.CrossRefGoogle Scholar
  7. Burg, R. W., Miller, B. M., Baker, E. E., Birnbaum, J., Currie, J. A., Harman, R., et al. (1979). Avermectins, new family of potent anthelmintic agents: producing organism and fermentation. Antimicrobial Agents and Chemotherapy, 15, 361–367.PubMedGoogle Scholar
  8. Burgermeister, W., Metge, K., Braasch, H., & Buchbach, E. (2005). ITS-RFLP patterns for differentiation of 26 Bursaphelenchus species (Nematoda: Parasitaphelenchidae) and observations on their distribution. Russian Journal of Nematology, 13(1), 29–42.Google Scholar
  9. Burgermeister, W., Braasch, H., Metge, K., Gu, J., Schroeder, T., & Woldt, E. (2009). ITS-RFLP analysis, an efficient tool for differentiation of Bursaphelenchus species. Nematology, 11(5), 649–668.CrossRefGoogle Scholar
  10. Cao, A. X., Liu, X. Z., Zhu, S. F., & Lu, B. S. (2005). Detection of the pinewood nematode, Bursaphelenchus xylophilus, using a real-time polymerase chain reaction assay. Phytopathology, 95(5), 566–571.PubMedCrossRefGoogle Scholar
  11. Castagnone, C., Abad, P., & Castagnone-Sereno, P. (2005). Satellite DNA-based species-specific identification of single individuals of the pinewood nematode Bursaphelenchus xylophilus (Nematoda: Aphelenchoididae). European Journal of Plant Pathology, 112, 191–193.CrossRefGoogle Scholar
  12. CEC (Comission of the European Communities) (2009). Commission staff working document concerning the control of the pinewood nematode in the forestry sector, in the European Union ( Brussels, Belgium.
  13. Cheng, H. R., Lin, M., Li, W., & Fang, Z. (1983). The occurrence of a pine wilting disease caused by a nematode found in Nanjing. Forest Pest and Disease, 4, 1–5.Google Scholar
  14. Costanza, R., d’Arge, R., de Groot, R., et al. (1997). The value of the world´s ecosystem services and natural capital. Nature, 387, 253–260.CrossRefGoogle Scholar
  15. Elbadri, G. A. A., Lee, D. W., Park, J. C., Yu, H. B., Choo, H. Y., Lee, S. M., et al. (2008). Nematocidal screening of essential oils and herbal extracts against Bursaphelenchus xylophilus. Plant Pathology Journal, 24, 178–182.CrossRefGoogle Scholar
  16. Evans, H., McNamara, D., Braasch, H., Chadouef, J., & Magnusson, C. (1996). Pest risk analysis (PRA) for the territories of the European Union (as PRA area) on Bursaphelenchus xylophilus and its vectors in the genus Monochamus. Bulletin OEPP/EPPO Bulletin, 26, 199–249.Google Scholar
  17. Fielding, N. J., & Evans, H. F. (1996). The pine wood nematode Bursaphelenchus xylophilus (Steiner and Buhrer) Nickle (= B. lignicolus Mamiya and Kiyohara): an assessment of the current position. Forestry, 69(1), 35–46.CrossRefGoogle Scholar
  18. Fonseca, L., Vieira dos Santos, M. C., Santos, M. S. N. A., Curtis, R. H. C., & Abrantes, I. (2008). Morpho-biometrical characterisation of Portuguese Bursaphelenchusxylophilus isolates with mucronate, digitate or round tailed females. Phytopatologia Mediterranea, 47, 223–233.Google Scholar
  19. Fonseca, L., Lopes, A., Cardoso, J., Pestana, M., Abreu, F., Nunes, N., et al. (2010). The pinewood nematode, Bursaphelenchus xylophilus, in Madeira Island. Paper presented at the 30th International Symposium of European Society of Nematologists September 19–23, Vienna, Austria.Google Scholar
  20. François, C., Castagnone, C., Boonham, N., Tomlinson, J., Lawson, R., Hockland, S., et al. (2007). Satellite DNA as a target for TaqMan real-time PCR detection ofthe pinewood nematode, Bursaphelenchus xylophilus. Molecular Plant Pathology, 8(6), 803–809.PubMedCrossRefGoogle Scholar
  21. Gu, J., Braasch, H., Burgermeister, W., & Zhang, J. (2006). Records of Bursaphelenchus spp. intercepted in imported packaging wood at Ningbo, China. Forest Pathology, 36, 323–333.CrossRefGoogle Scholar
  22. Gu, Y. Q., Mo, M. H., Zhou, J. P., Zou, C. S., & Zhang, K. Q. (2007). Evaluation and identification of potential organic nematicidal volatiles from soil bacteria. Soil Biology and Biochemistry, 39, 2567–2575.CrossRefGoogle Scholar
  23. Guo, D. S., Cong, P. J., Li, L., & Zhao, B. G. (2002). Determination of bacterial number carried by a pine wood nematodes and culture of sterilized nematodes on calli of Pinus Thunbergii. Journal of Qingdao University (in Chinese), 15, 29–31.Google Scholar
  24. Han, Z. M., Hong, Y. D., & Zhao, B. G. (2003). A study on pathogenicity of bacteria carried by pine wood nematodes. Journal of Phytopathology, 151, 683–689.CrossRefGoogle Scholar
  25. Hu, Y. Q., Kong, X. C., Wang, X. R., Zhong, T. K., Zhu, X. W., Mota, M. M., et al. (2011). Direct PCR-based method for detecting Bursaphelenchus xylophilus, the pine wood nematode in wood tissue of Pinus massoniana. Forest Pathology, 41, 165–168.CrossRefGoogle Scholar
  26. Huang, X. W., Tian, B. Y., Niu, Q. H., Yang, J. K., Zhang, L. M., & Zhang, K. Q. (2005). An extracellular protease from Brevibacillus laterosporus G4 without parasporal crystal can serve as a pathogenic factor in infection of nematodes. Research in Microbiology, 156, 719–727.PubMedCrossRefGoogle Scholar
  27. Hunt, D. J. (2008). A checklist of the Aphelenchoidea (Nematoda: Tylenchina). Journal of Nematode Morphology and Systematics, 10, 99–135.Google Scholar
  28. Jones, J., Moens, M., Mota, M., Li, H., & Kikuchi, T. (2008). Challenges for molecular plant pathology over the next ten years Bursaphelenchus xylophilus: opportunities in comparative genomicsand molecular host–parasite interactions. Molecular Plant Pathology, 9(3), 357–368.PubMedCrossRefGoogle Scholar
  29. Kang, J. S., Choi, K. S., Shin, S. C., Moon, I. L., Lee, S. G., & Lee, S. H. (2004). Development of an efficient PCR-based diagnosis protocolfor the identification of the pinewood nematode, Bursaphelenchus xylophilus (Nematoda:Aphelenchoididae). Nematology, 6(2), 279–285.CrossRefGoogle Scholar
  30. Kang, J. S., Lee, H., Moon, I. S., Lee, Y., Koh, Y. H., Je, Y. H., et al. (2009). Construction and characterization of subtractive stage-specific expressed sequence tag (EST) libraries of the pinewood nematode Bursaphelenchus xylophilus. Genomics, 94, 70–77.PubMedCrossRefGoogle Scholar
  31. Kanzaki, N. (2008). Taxonomy and systematics of the nematode genus Bursaphelenchus (Nematoda: Parasitaphelenchidae). In B. Zhao, K. Futai, J. R. Sutherland, & Y. Takeuchi (Eds.), Pine Wilt Disease (pp. 44–66). Dordrecht: Springer.CrossRefGoogle Scholar
  32. Kawazu, K. (1998). Pathogenic toxins of pine wilt disease. Kagaku to Seibutsu, 36, 120–124.CrossRefGoogle Scholar
  33. Kikuchi, T., Jones, J. T., Aikawa, T., Kosaka, H., & Ogura, N. (2004). A family of glycosyl hydrolase family 45 cellulases from the pinewood nematode Bursaphelenchus xylophilus. FEBS Letters, 572, 201–205.PubMedCrossRefGoogle Scholar
  34. Kikuchi, T., Shibuya, H., & Jones, J. T. (2005). Molecular and biochemical characterization of an endo-β-1,3-glucanase from the pinewood nematode Bursaphelenchus xylophilus acquired by horizontal gene transfer from bacteria. Biochemichal Journal, 389, 117–125.Google Scholar
  35. Kikuchi, T., Shibuya, H., Aikawa, T., & Jones, J. T. (2006). Cloning and characterization of pectate lyases expressed in the esophageal gland of the pine wood nematode Bursaphelenchus xylophilus. Molecular Plant-Microbe Interactions, 19(3), 280–287.PubMedCrossRefGoogle Scholar
  36. Kikuchi, T., Aikawa, T., Kosaka, H., Pritchard, L., Ogura, N., & Jones, J. T. (2007). Expressed sequence tag (EST) analysis of the pine wood nematode Bursaphelenchus xylophilus and B. mucronatus. Molecular & Biochemical Parasitology, 155, 9–17.CrossRefGoogle Scholar
  37. Kikuchi, T., Li, H., Karim, N., Kennedy, M. W., Moens, M., & Jones, J. T. (2009a). Identification of putative expansin-like genes from the pine wood nematode, Bursaphelenchus xylophilus, and evolution of the expansin gene family within the Nematoda. Nematology, 11, 355–364.CrossRefGoogle Scholar
  38. Kikuchi, T., Aikawa, T., Oeda, Y., Karim, N., & Kanzaki, N. (2009b). A rapid and precise diagnostic method for detecting the pinewood nematode Bursaphelenchus xylophilus by Loop-Mediated Isothermal Amplification. Phytopathology, 99(12), 1365–1369.PubMedCrossRefGoogle Scholar
  39. Kikuchi, T., Cotton, J. A., Dalzell, J. J., Hasegawa, K., Kanzaki, N., et al. (2011). Genomic Iinsights into the origin of parasitism in the emerging plant pathogen Bursaphelenchus xylophilus. PLoS Pathogens, 7(9), e1002219. doi: 10.1371/journal.ppat.1002219.PubMedCrossRefGoogle Scholar
  40. Kiyohara, T., & Bolla, R. I. (1990). Pathogenic variability among populations of the pinewood nematode, Bursaphelenchus xylophilus. Forest Science, 36, 1061–1076.Google Scholar
  41. Kiyohara, T., & Tokushige, Y. (1971). Inoculation experiments of a nematode, Bursaphelenchus sp., onto pine trees. Journal of the Japanese Forestry Society, 53, 210–218.Google Scholar
  42. Kong, J. O., Lee, S. M., Moon, Y. S., Lee, S. G., & Ahn, Y. J. (2006). Nematicidal activity of plant essential oils against Bursaphelenchus xylophilus. Journal of Asia-Pacific Entomology, 9, 173–178.CrossRefGoogle Scholar
  43. Kurinobu, S. (2006). Current status of resistance breeding of Japanese pine species to Pine Wilt Disease. Proceedings of the International Symposium on Pine Wilt Disease held at Korea Forest Research Institute, Seoul, Korea in December 10, 2006: 83–94.Google Scholar
  44. Kuroda, K., & Ito, S. (1992). Migration speed of pine wood nematodes and activities of other microbes during the development of pine-wilt disease in Pinus thunbergii. Journal of the Japanese Forestry Society, 74, 383–389.Google Scholar
  45. Kusunoki, M. (1987). Symptom development of pine wilt disease histopathological observations with electron microscope. Annals of the Phytopathological Society of Japan, 53, 622–629.CrossRefGoogle Scholar
  46. Kwon, H. R., Choi, G. J., Choi, Y. H., Jang, K. S., Sung, N.-D., Kang, M. S., et al. (2010). Suppression of pine wilt disease by an antibacterial agent, oxolinic acid. Pest Management Science, 66, 634–639.PubMedGoogle Scholar
  47. Leal, I., Green, M., Allen, E., Humble, L., & Rott, M. (2005). An effective PCR-based diagnostic method for the detection of Bursaphelenchus xylophilus (Nematoda: Aphelenchoididae) in wood samples from lodgepole pine. Nematology, 7(6), 833–842.CrossRefGoogle Scholar
  48. Leal, I., Green, M., Allen, E., Humble, L., & Rott, M. (2007). Application of a real-time PCR method for the detection of pinewood nematode, Bursaphelenchus xylophilus, in wood samples from lodgepole pine. Nematology, 9(3), 351–362.CrossRefGoogle Scholar
  49. Li, X., Zhuo, K., Luo, M., Sun, L., & Liao, J. (2011). Molecular cloning and characterization of a calreticulin cDNA from the pinewood nematode Bursaphelenchus xylophilus. Experimental Parasitology, 128, 121–126.PubMedCrossRefGoogle Scholar
  50. Lin, S., Jian, H., Zhao, H., Yang, D., & Liu, Q. (2011). Cloning and characterization of a venom allergen-like protein gene cluster from the pinewood nematode Bursaphelenchus xylophilus. Experimental Parasitology, 127, 440–447.PubMedCrossRefGoogle Scholar
  51. Linit, M. (1988). Nematode-vector relationships in the pine wilt disease system. Journal of Nematology, 20, 227–235.PubMedGoogle Scholar
  52. Linit, M. J. (1990). Transmission of pinewood nematode through feeding wounds of Monochamus carolinensis (Coleoptera: Cerambycidae). Journal of Nematology, 22, 231–236.PubMedGoogle Scholar
  53. Maehara, N. (2008). Reduction of Bursaphelenchus xylophilus (Nematoda: Parasitaphelenchidae) population by inoculating Trichoderma spp. into pine wilt-killed trees. Biological Control, 44, 61–66.CrossRefGoogle Scholar
  54. Mamiya, Y. (1984). The pine wood nematode. In W. R. Nickle (Ed.), Plant and insect nematodes (pp. 589–627). New York and Basel: Marcel Dekker, Inc.Google Scholar
  55. Mamiya, Y. (2004). Pine wilt disease in Japan. In M. Mota & P. Vieira (Eds.), The pinewood nematode, Bursaphelenchus xylophilus. Nematology Monographs and Perspectives 1 (pp. 9–20). Leiden: Brill Academic Publishers.Google Scholar
  56. Matsunaga, K., & Togashi, K. (2004). A simple method for discriminating Bursaphelenchus xylophilusand B. mucronatus by species-specific polymerasechain reaction primer pairs. Nematology, 6(2), 273–277.CrossRefGoogle Scholar
  57. Mota, M., & Vieira, P. (2008). Pine wilt disease: a worldwide threat to forest ecosystems. Dordrecht: Springer. 428 pp. ISBN 978-1-4020-8454-6.Google Scholar
  58. Mota, M., Braasch, H., Bravo, M. A., Penas, A. C., Burgermeister, W., Metge, K., et al. (1999). First report of Bursaphelenchus xylophilus in Portugal and in Europe. Nematology, 1, 727–734.CrossRefGoogle Scholar
  59. Mota, M., Futai, K., & Vieira, P. (2009). Pine wilt disease and the pinewood nematode. In A. Ciancio & K. G. Mukerji (Eds.), Integrated Management of Fruit Crops and Forest Nematodes, vol. IV (pp. 253–274). Dordrecht: Springer.CrossRefGoogle Scholar
  60. Naves, P., Kenis, M., & Sousa, E. (2005). Parasitoids associated with Monochamus galloprovincialis (Oliv.) (Coleoptera; Cerambycidae) within the pine wilt nematode-affected zone in Portugal. Journal of Pesticide Science, 78, 57–62.Google Scholar
  61. Naves, P. M., Camacho, S., Sousa, E. M., & Quartau, J. A. (2006). Entrance and distribution of the pine wood nematode Bursaphelenchus xylophilus on the body of its vector of Monochamus galloprovincialis (Coleoptera: Cerambycidae). Entomologia Generalis, 29(1), 071–080.Google Scholar
  62. Naves, P., Sousa, E., & Quartau, J. A. (2006a). Feeding and oviposition preferences of Monochamus galloprovincialis for certain conifers under laboratory conditions. Entomologia Experimentalis et Applicata, 120(2), 99–104.CrossRefGoogle Scholar
  63. Naves, P., Sousa, E., & Quartau, J. A. (2006b). Reproductive traits of Monochamus galloprovincialis (Coleoptera: Cerambycidae) under laboratory conditions. Bulletin of Entomological Research, 96, 289–294.PubMedCrossRefGoogle Scholar
  64. Naves, P. M., Camacho, S., Sousa, E. M., & Quartau, J. A. (2007). Transmission of the pine wood nematode Bursaphelenchusxylophilus through feeding activity of Monochamusgalloprovincialis (Col., Cerambycidae). Journal of Applied Entomology, 131(1), 21–25.CrossRefGoogle Scholar
  65. Oka, Y., Nacar, S., Putievsky, E., Ravid, U., Yaniv, Z., & Spiegel, Y. (2000). Nematicidal activity of essential oils and their components against the root-knot nematode. Phytopathology, 90, 710–715.PubMedCrossRefGoogle Scholar
  66. Oku, H., Shiraishi, S., Kurozumi, S., & Ohta, H. (1980). Pine wilt toxin, the metabolite of a bacterium associated with a nematode. Naturwissenschaften, 67, 198–199.CrossRefGoogle Scholar
  67. Park, I. K., Park, J. Y., Kim, K. H., Choi, K. S., Choi, I. H., Kim, C. S., et al. (2005). Nematicidal activity of plant essential oils and components from garlic (Allium sativum) and cinnamon (Cinnamomum verum) oils against the pinewood nematode (Bursaphelenchus xylophilus). Nematology, 7, 767–774.CrossRefGoogle Scholar
  68. Penas, A. C., Dias, L. S., & Mota, M. M. (2002). Precision and selection of extraction methods of aphelenchid nematodes from maritime pine wood, Pinus pinaster L. Journal of Nematology, 34(1), 62–65.PubMedGoogle Scholar
  69. Penas, A. C., Correia, P., Bravo, M., Mota, M., & Tenreiro, R. (2004). Species of Bursaphelenchus Fuchs, 1937 (Nematoda: Parasitaphelenchidae) associated with maritime pine in Portugal. Nematology, 6, 437–453.Google Scholar
  70. Penas, A. C., Bravo, M. A., Naves, P., Bonifácio, L., Sousa, E., & Mota, M. (2006). Species of Bursaphelenchus Fuchs, 1937 (Nematoda: Parasitaphelenchidae) and other nematode genera associated with insects from Pinus pinaster in Portugal. Annals of Applied Biology, 148(2), 121–131.CrossRefGoogle Scholar
  71. Penas, A. C., Bravo, M. A., Valadas, V., & Mota, M. (2007). Detailed morphobiometric studies of Bursaphelenchus xylophilus andcharacterisation of other Bursaphelenchus species (Nematoda:Parasitaphelenchidae) associated with Pinus pinaster in Portugal. Journal of Nematode Morphology and Systematics, 10(2), 137–163.Google Scholar
  72. Proença, D. N., Francisco, R., Santos, C. V., Lopes, A., Fonseca, L., Abrantes, I. M., et al. (2010). Diversity of bacteria associated with Bursaphelenchusxylophilus and other nematodes isolated from Pinus pinaster trees with pine wilt disease. PLoS One, 5, e15191.PubMedCrossRefGoogle Scholar
  73. Robertson, L., Cobacho Arcos, S., Escuer, M., Santiago Merinos, R., Esparrago, G., Abelleira, A., et al. (2011). Incidence of the pinewood nematode Bursaphelenchus xylophilus Steiner & Buhrer, 1934 (Nickle, 1970) in Spain. Nematology, 13(6), 755–757.CrossRefGoogle Scholar
  74. Robinet, C. H., Roques, A., Pan, H., Fang, G., Ye, J., Zhang, Y., et al. (2009). Role of human-mediated dispersal in the spread of the pinewood nematode in China. PLoS One, 4(2), e4646. doi: 10.1371/journal.pone.0004646.PubMedCrossRefGoogle Scholar
  75. Rodrigues, J. (2008). National erradication programme for the pinewood nematode in Portugal. In M. Mota & P. Vieira (Eds.), Pine wilt disease: a worldwide threat to forest ecosystems (pp. 5–14). Dordrecht: Springer.CrossRefGoogle Scholar
  76. Ryss, A., Vieira, P., Mota, M., & Kulinich, O. (2005). A synopsis of the genus Bursaphelenchus Fuchs, 1937 (Aphelenchida: Parasitaphelenchidae) with keys to species. Nematology, 7, 393–458.CrossRefGoogle Scholar
  77. Schröder, T., McNamara, D. G., & Gaar, V. (2009). Guidance on sampling to detect pine wood nematode Bursaphelenchus xylophilus in trees, wood and insects. Bulletin OEPP/EPPO Bulletin, 39, 179–188.Google Scholar
  78. Shimazu, M. (2006). Current status on research and management of pine wilt disease in Japan. In. Current status on research and management of pine wilt disease, International Symposium, October 20, 2006 (1–18) Korea Forest Research Institute, Seoul, Korea.Google Scholar
  79. Shin, S., & Han, H. (2006). Current status on research and management of pine wilt disease in Korea. In Current status on research and management of pine wilt disease, International Symposium, October 20, 2006 (pp. 31–44).Korea Forest Research Institute, Seoul, Korea, October.Google Scholar
  80. Shinya, R., Morisaka, H., Takeuchi, Y., Ueda, M., & Futai, K. (2010). Comparison of the surface coat proteins of the pine wood nematode appeared during host pine infection and in vitroculture by a proteomic approach. Nematology, 100, 1289–1297.Google Scholar
  81. Sniezko, R. A. (2006). Resistance breeding against nonnative pathogens in forest trees, current successes in North America. Canadian Journal Plant Pathology, 28, s270–s279.CrossRefGoogle Scholar
  82. Sousa, E., Bravo, M. A., Pires, J., Naves, P., Penas, A. C., Bonifácio, L., et al. (2001). Bursaphelenchus xylophilus (Nematoda: Aphelenchoididae) associated with Monochamus galloprovincialis (Coleoptera: Cerambycidae) in Portugal. Nematology, 3, 89–91.CrossRefGoogle Scholar
  83. Sousa, E., Naves, P., Bonifácio, L., Bravo, M. A., Penas, A. C., Pires, J., et al. (2002). Preliminary survey for insects associated with Bursaphelenchus xylophilus in Portugal. Bulletin OEPP/EPPO Bulletin, 32, 499–502.Google Scholar
  84. Steiner, G., & Buhrer, E. M. (1934). Aphelenchoides xylophilus, n. sp. a nematode associated with blue-stain and other fungi in timber. Journal of Agricultural Research, 48(10), 949–951.Google Scholar
  85. Suzuki, K. (2002). Pine wilt disease - a threat to pine forest in Europe. Dendrobiology, 48, 71–74.Google Scholar
  86. Takai, K., Suzuki, T., & Kawazu, K. (2003). Development and preventive effect against pine wilt disease of a novel liquid formulation of emamectin benzoate. Pest Management Science, 59, 365–370.PubMedCrossRefGoogle Scholar
  87. Takeuchi, Y., & Futai, K. (2009). Diagnosis and quantification of pine wood nematode, Bursaphelenchus xylophilus (Steiner and Buhner), in wood of Pinus thunbergii with real-time PCR. Nematological Research, 39(1), 9–16.CrossRefGoogle Scholar
  88. Takeuchi, Y., Kanzaki, N., & Futai, K. (2005). A nested PCR-based method for detecting the pine wood nematode, Bursaphelenchus xylophilus, from pine wood. Nematology, 7(5), 775–782.CrossRefGoogle Scholar
  89. Tamura, H. (1983). Pathogenicity of aseptic Bursaphelenchus xylophilus and associated bacteria to pine seedlings. Japanese Journal of Nematology, 13, 1–5.Google Scholar
  90. Tian, B., Yang, J., & Zhang, K. Q. (2007). Bacteria used in the biological control of plant-parasitic nematodes: populations, mechanisms of action, and future prospects. FEMS Microbiology Ecology, 61, 197–213.PubMedCrossRefGoogle Scholar
  91. Toda, T., & Kurinobu, S. (2001). Genetic improvement in pine wilt disease resistance in Pinus thunbergii: the effectiveness of pre-screening with an artificial inoculation at the nursery. Journal of Forest Research, 6, 197–201.CrossRefGoogle Scholar
  92. Tzean, S. & Jan, S. (1985a). The occurrence of pinewood nematode, Bursaphelenchus xylophilus, in Taiwan. In Proceedings 6th ROC Symposium of Electron Microscopy (pp. 38–39).Google Scholar
  93. Tzean, S., & Jan, S. (1985b). Pine wilt disease caused by pinewood nematode (Bursaphelenchus xylophilus) and its occurrence in Taiwan. Phytopathologist and Entomologist, NTU, 12, 1–19.Google Scholar
  94. Valadas, V., Barbosa, P., Espada, M., Oliveira, S., & Mota., M. M. (2011). The pinewood nematode, Bursaphelenchus xylophilus, in Portugal: possible introductions and spread routes of a serious biological invasion revealed by molecular methods. Paper presented at the XLIII ONTA Annual Meeting, Coimbra, Portugal, September 2011.Google Scholar
  95. Vicente, C., Nascimento, F., Espada, M., Mota, M., & Oliveira, S. (2011). Bacteria associated with the pinewood nematode Bursaphelenchus xylophilus collected in Portugal. Antonie van Leeuwenhoek Journal of Microbiology, 100(3), 477–481. doi: 10.1007/s10482-011-9602-1.CrossRefGoogle Scholar
  96. Vieira, P., Burgermeister, W., Mota, M., Metge, K., & Silva, G. (2007). Lack of genetic variation of Bursaphelenchus xylophilus in Portugal revealed by RAPD-PCR analyses. Journal of Nematology, 39(2), 118–126.PubMedGoogle Scholar
  97. Wang, X. R., Zhu, X. W., Kong, X. C., & Mota, M. M. (2010a). A rapid detection of the pinewood nematode, Bursaphelenchusxylophilus in stored Monochamus alternatus by rDNAamplification. Journal of Applied Entomology, 135, 156–159.CrossRefGoogle Scholar
  98. Wang, Y. Q., Zhang, L. Y., Lai, D., & Xu, H. H. (2010b). The nematicidal and proteomic effects of Huanong AVM (analog of avermectin) on the pine-wilt nematode, Bursaphelenchus xylophilus. Pesticide Biochemistry and Physiology, 98, 224–230.CrossRefGoogle Scholar
  99. Wang, C. Y., Fang, Z. M., Wang, Z., Zhang, D. L., Gu, L. J., Lee, M. R., et al. (2011). Biological control of the pinewood nematode Bursaphelenchusxylophilus by application of the endoparasitic fungus Esteya vermicola. BioControl, 56, 91–100.CrossRefGoogle Scholar
  100. Xie, L. Q., & Zhao, B. G. (2008). Post-inoculation population dynamics of Bursaphelenchusxylophilus and associated bacteria in pine wilt disease on Pinus thunbergii. Journal of Phytopathology, 156, 385–389.CrossRefGoogle Scholar
  101. Yang, B. (2004). The history, dispersal and potential threat of pine wood nematode in China. In M. Mota & P. Vieira (Eds.), The pinewood nematode, Bursaphelenchus xylophilus. Nematology Monographs and Perspectives 1 (pp. 21–24). The Netherlands: Brill Academica Publishers.Google Scholar
  102. Yano, M. (1913). Investigation on the cause of pine mortality in Nagasaki Prefecture. Sanrinkoho, 4, 1–14 (in Japanese).Google Scholar
  103. Yi, C., Byun, B., Park, J., Yang, S., & Chang, K. (1989). First finding of the pine wood nematode, Bursaphelenchus xylophilus (Steiner et Buhrer) Nickle and its insect vector in Korea. Research Reports of the Forestry Research Institute Seoul, 38, 141–149.Google Scholar
  104. Zhang, K., Lui, H., Sun, J., Lui, J., Fei, K., Zhang, C., et al. (2008). Molecular phylogeny of geographical isolates of Bursaphelenchus xylophilus: implications on the origin and spread of this species in China and worldwide. Journal of Nematology, 40(2), 127–137.PubMedGoogle Scholar
  105. Zhao, B., & Lin, F. (2005). Mutualistic symbiosis between Bursaphelenchus xylophilus and bacteria of the genus Pseudomonas. Forest Pathology, 35, 339–345.CrossRefGoogle Scholar
  106. Zhao, B. G., Guo, D. S., & Gao, R. (2000). Observation of the site of pine wood nematodes where bacteria are carried with SEM and TEM. Journal of Nanjing Forest University (in Chinese), 24, 69–71.Google Scholar
  107. Zhao, B. G., Wang, H. L., Han, S. F., & Han, Z. M. (2003). Distribution and pathogenicity of bacteria species carried by Bursaphelenchus xylophilus in China. Nematology, 5, 899–906.CrossRefGoogle Scholar
  108. Zhao, B., Liu, Y., & Lin, F. (2006). Mutual influences in growth and reproduction between pine wood nematode Bursaphelenchusxylophilus and bacteria it carries. Frontiers of Forestry in China, 3, 324–327.CrossRefGoogle Scholar
  109. Zhao, B., Liu, Y., & Lin, F. (2007). Effects of bacteria associated with pine wood nematode (Bursaphelenchusxylophilus) on development and egg production of the nematode. Journal of Phytopathology, 155, 26–30.CrossRefGoogle Scholar

Copyright information

© KNPV 2011

Authors and Affiliations

  • Cláudia Vicente
    • 1
  • Margarida Espada
    • 1
  • Paulo Vieira
    • 1
  • Manuel Mota
    • 1
    Email author
  1. 1.NemaLab/ICAAMUniversidade de ÉvoraÉvoraPortugal

Personalised recommendations