Plant-parasitic nematodes of potential phytosanitary importance, their main hosts and reported yield losses

Sunil Singh

The potential phytosanitary importance of all named plant-parasitic nematode species was determined by evaluating available information on species characteristics, association with economically-important crop hosts, and ability to act as vectors of viruses or form disease complexes with other pathogens. Most named species of plant-parasitic nematodes (PPN) are poorly known, recorded from a single location only, not associated with economically-important crops, and not known to be associated with other plant disease organisms. However, 250 species from 43 genera fulfilled one or more of the criteria to be considered to present a phytosanitary risk. The genera and number of species (in parentheses) considered as posing phytosanitary risk included: Achlysiella (1), Anguina (8), Aphasmatylenchus (1), Aphelenchoides (12), Aphelenchus (1), Belonolaimus (2), Bitylenchus (3), Bursaphelenchus (4), Cactodera (3), Ditylenchus (8), Dolichodorus (1), Globodera (3), Helicotylenchus (7), Hemicriconemoides (3), Hemicycliophora (3), Heterodera (25), Hirschmanniella (5), Hoplolaimus (5), Ibipora (3), Longidorus (10), Macroposthonia (2), Meloidogyne (38), Merlinius (3), Nacobbus (1), Neodolichodorus (2), Paralongidorus (2), Paratrichodorus (11), Paratylenchus (3), Pratylenchus (24), Punctodera (3), Quinisulcius (3), Radopholus (5), Rotylenchulus (3), Rotylenchus (1), Scutellonema (5), Sphaeronema (1), Subanguina (3), Trichodorus (5), Tylenchorhynchus (8), Tylenchulus (2), Vittatidera (1), Xiphinema (15) and Zygotylenchus (1). For each of the 250 species main hosts and yield loss estimates are provided with an extensive bibliography. Of the 250 species, only 126 species from 33 genera are currently listed as regulated pests in one or more countries worldwide. Almost all of these 250 species were also associated with economically important crops and some also acted as vectors for viruses.

Bulletin OEPP/EPPO Bulletin (2013) 43 (2), 334–374 ISSN 0250-8052. DOI: 10.1111/epp.12050 Plant-parasitic nematodes of potential phytosanitary importance, their main hosts and reported yield losses S. K. Singh1,2,3,4, M. Hodda1,4 and G. J. Ash2 1 CSIRO Ecosystem Sciences, Canberra, ACT, 2601, Australia; e-mail: sunil.singh@csiro.au Graham Centre for Agricultural Innovation (an alliance between Charles Sturt University and the NSW Department of Primary Industries), Wagga Wagga, NSW, 2678, Australia 3 Cooperative Research Centre for National Plant Biosecurity, Bruce, ACT, 2617, Australia 4 CSIRO Biosecurity Flagship, Canberra, ACT, 2601, Australia 2 The potential phytosanitary importance of all named plant-parasitic nematode species was determined by evaluating available information on species characteristics, association with economically-important crop hosts, and ability to act as vectors of viruses or form disease complexes with other pathogens. Most named species of plant-parasitic nematodes (PPN) are poorly known, recorded from a single location only, not associated with economicallyimportant crops, and not known to be associated with other plant disease organisms. However, 250 species from 43 genera fulfilled one or more of the criteria to be considered to present a phytosanitary risk. The genera and number of species (in parentheses) considered as posing phytosanitary risk included: Achlysiella (1), Anguina (8), Aphasmatylenchus (1), Aphelenchoides (12), Aphelenchus (1), Belonolaimus (2), Bitylenchus (3), Bursaphelenchus (4), Cactodera (3), Ditylenchus (8), Dolichodorus (1), Globodera (3), Helicotylenchus (7), Hemicriconemoides (3), Hemicycliophora (3), Heterodera (25), Hirschmanniella (5), Hoplolaimus (5), Ibipora (3), Longidorus (10), Macroposthonia (2), Meloidogyne (38), Merlinius (3), Nacobbus (1), Neodolichodorus (2), Paralongidorus (2), Paratrichodorus (11), Paratylenchus (3), Pratylenchus (24), Punctodera (3), Quinisulcius (3), Radopholus (5), Rotylenchulus (3), Rotylenchus (1), Scutellonema (5), Sphaeronema (1), Subanguina (3), Trichodorus (5), Tylenchorhynchus (8), Tylenchulus (2), Vittatidera (1), Xiphinema (15) and Zygotylenchus (1). For each of the 250 species main hosts and yield loss estimates are provided with an extensive bibliography. Of the 250 species, only 126 species from 33 genera are currently listed as regulated pests in one or more countries worldwide. Almost all of these 250 species were also associated with economically important crops and some also acted as vectors for viruses. Introduction Annual crop losses caused by plant-parasitic nematodes are estimated at 8.8–14.6% of total crop production and 100– 157 billion USD worldwide (Sasser & Freckman, 1987; Koenning et al., 1999; Abad et al., 2008; Nicol et al., 2011). Actual losses may be even higher because there is no data from many countries where nematological expertise is lacking. Yield loss data is also difficult to obtain because of the complex interactions of plants, nematodes, other soil organisms and soils (Seinhorst, 1982; Noling, 1986; Koenning et al., 1999; Coyne & Plowright, 2002). Often only the most noticeable localised damage is investigated and reported, while less obvious but more widespread damage is overlooked (Nicol et al., 2011). Substantial, crop losses from plant-parasitic nematodes (hereafter PPN), could be much greater if species currently causing localised damage became more widespread. Many PPN have low impact in their native range, but much 334 greater impact when introduced to new areas. Examples include: Bursaphelenchus xylophilus (Pine Wood Nematode) in Japan, China, Portugal and most recently in Spain (Mota et al., 1999; Braasch & Enzian, 2004; Cheng et al., 2007; Togashi & Jikumaru, 2007; Abelleira et al., 2011; Robertson et al., 2011), Heterodera glycines (Soybean Cyst Nematode) in USA (Winstead et al., 1955; Riggs, 1977); and the Potato Cyst Nematodes in Europe, the USA, Canada and Australia (Stone et al., 1977; Stanton, 1987; Turner & Evans, 1998; Hafez & Sundararaj, 2007; Sun et al., 2007). Cumulative losses from potato cyst nematodes introduction to Australia were estimated at 370 million AUD over 20 years (Hodda & Cook, 2009), and Pine Wood Nematode introduction to Europe estimated at 22 billion EUR over 22 years (Soliman et al., 2012). Quarantine measures against known damaging PPN are effective in preventing their spread, thus effectively and economically preventing crop losses (Lehman et al., 1996; Inserra et al., 2005; Sikora et al., 2005). The presence of ª 2013 The Authors. Journal compilation ª 2013 OEPP/EPPO, EPPO Bulletin 43, 334–374 Nematodes of phytosanitary importance PPN during quarantine inspections may also act as a bioindicator for consignments that do not meet the phytosanitary requirements of plants being grown in sterile environments and could be carrying other plant pathogens and microorganisms (Hockland & Anderson, 2012). Quarantine measures can also prevent spread of non-target species which are potentially invasive (Schrader & Unger, 2003). Quarantine and other phytosanitary measures are particularly important for PPN because other management methods such as chemical control or crop resistance can be far more costly and difficult to implement without other adverse effects (Hockland et al., 2006; Zasada et al., 2010a; Nicol et al., 2011). Furthermore, targeting PPN is important because of the wide range of survival adaptations and dispersal routes available (Singh, S.K., Hodda, M., Ash, G.J, Banks, N.C. unpublished data). An important first step in implementing quarantine measures is to determine which species should be regulated under international trade rules. The international plant pest convention defines a quarantine pest as ‘a pest of potential economic importance to the area endangered thereby and not yet present there, or present but not widely distributed and being officially controlled’ (FAO, 2011a). Organisms which meet this definition can be regulated. Countries determine their lists of regulated pests according to guidelines set by the International Plant Protection Convention (IPPC) and International Standards for Phytosanitary Measures No. 19 (FAO, 2011b). Countries may also establish lists of regulated nonquarantine pests. The IPPC defines these as follows: ‘A non-quarantine pest whose presence in plants for planting affects the intended use of those plants with an economically unacceptable impact and which is therefore regulated within the territory of the importing contracting party.’ All lists of regulated pests are dynamic, with species added or removed from lists as phytosanitary risks change or as new species emerge as pests (EPPO, 2007; FAO, 2011b). Species other than those on official lists of quarantine pests may also pose a biosecurity risk. Only a small proportion of nematode species have been described (Blaxter, 2011; Hodda, 2011). New species are being described regularly, and species new to science have been intercepted during phytosanitary inspections [e.g. Radopholus bridgei and Meloidogyne thailandica, both described from material intercepted in quarantine (Siddiqi & Hahn, 1995; Handoo et al., 2005)]. Records of non-compliance for PPN show species outside of official lists of regulated pests being frequently detected (Gao & Zhang, 1994; Mani, 1998; Queneherve et al., 1998; Plumas et al., 2002; Lal & Rajan, 2005; Lal & Lal, 2006; Roques & Auger-Rozenberg, 2006; McNeill et al., 2011). Information allowing assessment of the risks of the many PPN not on official lists of regulated pests is often not available (Singh, S.K., Hodda, M., Ash, G.J, Banks, N.C. unpublished data). To partially fill this gap, this paper presents a list of the 250 PPN likely to be 335 most important for quarantine at a global scale, selected from all named PPN. The authors present notes on their main hosts, information on yield losses where available and their quarantine status globally if known, together with an extensive bibliography. This paper also discusses how such a list could be used in the pest risk analysis (PRA) process. Methods Data on hosts, geographic distribution, yield loss and quarantine status were compiled from published records, the majority from online databases (Table 1). Species names of all known PPN were obtained from: Hunt (1993, 2008) for Aphelenchoidea and Longidoridae; Decraemer (1995) for Trichodoridae; Coomans et al. (2001) for Xiphinema; Ebsary (1991) and Siddiqi (2000) for Tylenchida; and Esser (1991) and Fortuner (1984) for general checklists of phytoparasitic nematodes. Synonyms were sourced from the database of nematode names created for the classification of phylum Nematoda (Hodda, 2011). CABI abstracts, Web of Knowledge, and Google Scholar were searched using these genus and species names. For species with over 200 hits, the search was refined using the advanced search option using the terms ‘distribution’, ‘hosts’, ‘yield’, ‘damage’, ‘disease’, ‘pathotype’ and ‘virus-vector’. Full texts were acquired where available, either online or through interlibrary loans. Records up to 31st May 2012 were included. To be considered of phytosanitary importance PPN species had to meet at least one of the following criteria: (1) Recorded in the peer-reviewed scientific literature as pathogenic (causing disease) or parasitic (infecting or associated as ectoparasites) of an economically-important crop host; (2) Recorded in the peer-reviewed scientific literature as acting as vectors of, or forming disease complexes with, other pathogens such as bacteria, fungi and viruses; (3) Currently on an official list of regulated pests for at least one country. Results Almost all of the 250 PPN species selected, satisfied criterion 1 [i.e. directly pathogenic or associated with economically important plants including crop, forest or pasture species (Table 2)]. The only exception, Ditylenchus weischeri which is a weed parasite; is included due to its recent taxonomic separation from the economically important D. dipsaci species complex. Half of these species (52%, N = 129) also satisfied criterion 2 (could form disease complexes with other organisms and or acted as vectors for viruses). A total of 126 species from 33 PPN genera were currently recognised as regulated pest species in one or more countries, worldwide (Fig. 1). All PPN species reported as a regulated pest satisfied more than two criteria and were often associated with economically important ª 2013 The Authors. Journal compilation ª 2013 OEPP/EPPO, EPPO Bulletin 43, 334–374 336 S. K. Singh et al. Table 1 Online sources on organisms of phytosanitary importance Resource Notes on resource and their coverage of plant parasitic nematodes CABI Invasive Species Compendium The Invasive Species Compendium (ISC) includes known invasive species of all taxonomic groups affecting natural and managed ecosystems, except human pathogens, concentrating on those species that have the greatest impacts on livelihoods and environment. It also includes some species which currently have a low impact where they currently occur but may be a threat to other regions if introduced (L Charles, pers. comm.). There were 38 species of PPN with detailed datasheets listed on the ISC database. The database is open access. http:// www.cabi.org/ISC/ The Crop Protection Compendium (CPC) is an encyclopaedic resource that brings together a wide range of different types of science-based information on all aspects of crop protection. The crop protection compendium covers pests of agricultural and horticultural crops and, since 2004, forest trees. The species that have been selected for inclusion as full datasheets are those of major global or regional economic or phytosanitary importance. In addition to crops; weeds, invasive plants of rangelands and woody invasive plants are also included on the CPC. There were 97 species of PPN with detailed datasheets listed on the CPC database. Access to the detailed datasheets is restricted to users with a subscription. http://www.cabi.org/CPC/ The Distribution Maps of Plant Diseases (DMPD) ‘cover important diseases affecting agriculture, horticulture and forestry. Two sets of maps are produced each year, consisting mostly of new maps with a number of map revisions (where significant changes have merited a revision). There are 18 diseases per map set covering fungi, bacteria, viruses and, from 1999 onwards, nematodes (CABI, DMPD). There were 91 species distribution maps of PPN. Access to detailed maps is restricted to users with a subscription. http://www.cabi.org/DMPD/ The International Plant Protection Convention (IPPC) is an international agreement on plant health with 177 current signatories. It aims to protect cultivated and wild plants by preventing the introduction and spread of pests. Countries communicate their list of quarantine organisms, new pest records and phytosanitary alerts via the IPPC website http://www.ippc.int/ The EPPO PQR database provides data on all the pests of the EPPO A1 and A2 Lists, pests on the EPPO Alert List, and quarantine pests and invasive plants of interest to other regions of the world. For each pest, lists of host plants, commodities able to act as pathways in international trade, nomenclatural and taxonomic details, and geographical distribution are presented (EPPO PQR). PQR lists 77 species of PPN with quarantine categorization for 47 species. The database is open access and can be downloaded from the EPPO website http://www.eppo.int/DATABASES/pqr/pqr.htm The US National Agricultural Pest Information System (NAPIS) stores and manages pest survey data that is collected by Cooperative Agricultural Pest Surveys and other Plant Protection Quarantine survey programs. Information is provided on the distribution of plant pests of regulatory significance to the USA and the risks associated with entry, establishment and spread of animal, plant pests and noxious weeds to the USA. There are 63 species of PPN listed on the database with survey maps and host maps. The risk maps are specific to the USA and its states. http://pest.ceris.purdue.edu/index.php The University of Georgia (USA) Center for Invasive Species and Ecosystem Health website provides information on invasive species whose introduction causes or is likely to cause economic or environmental harm or harm to human health. There are 29 species of PPN listed as potentially invasive to North America http://www.invasive. org/species/nematodes.cfm The California Department of Agriculture website provides a rating system for pests based on the potential and actual harm to California’s agriculture and environment. Pest ratings for 109 species of PPN for the state of California, USA are given http://www.cdfa.ca.gov/plant/ppd/nematology/nema_by_rating.html The University of Nebraska website provides information on the top 15 species of PPN causing severe damage to crops which are the most widely regulated species worldwide. http://nematode.unl.edu/quaranem.htm Professional association website presenting results of a working group evaluation of phytosanitary risks to the USA from 68 species of PPN. Distribution and hosts are also provided. (Last updated September 2003) http://nematode. unl.edu/projectpest.htm Dutch Ministry of Economic Affairs sponsored database (Q-bank) providing descriptions of well-characterized regulated plants pests. It comprises ecological, morphological, physiological, and sequence data of materials that are available in physical collections of plant-pathogenic bacteria, fungi, insects, nematodes, phytoplasmas, viruses and viroids, and invasive plants. http://www.q-bank.eu/ The EU financed QBOL project aimed to develop rapid identification for prioritized quarantine organisms relevant for the EU, using DNA barcode sequences. There are 76 species of PPN on the nematode priority list. http://www.qbol.wur.nl/UK Website produced by South Pacific Commission. The Pacific Island Pest List database (PLD) provides records of pests that are currently known to affect agriculture, forestry and the environment in Pacific Island countries and territories (PLD). There are 31 species of PPN listed with information on their distribution and hosts. http://www. spc.int/pld/ CABI Crop Protection Compendium CABI Distribution Maps of Plant Diseases International Plant Protection Convention EPPO PQR database National Agricultural Pest Information System Invasive and Exotic Species of North America California Department of Food and Agriculture Nematodes of quarantinable concern Society of Nematologists Q-bank and QBOL Pacific Island Pest List database ª 2013 The Authors. Journal compilation ª 2013 OEPP/EPPO, EPPO Bulletin 43, 334–374 ª 2013 The Authors. Journal compilation ª 2013 OEPP/EPPO, EPPO Bulletin 43, 334–374 Table 2 List of plant-parasitic nematode species of phytosanitary importance Species Notes on main hosts and yield losses Achlysiella williamsi (Siddiqi, 1964) Hunt, Bridge & Machon, 1989 Anguina agropyri Kirjanova, 1955 Parasitises sugarcane (Hunt et al., 1989; Blair et al., 1999). Anguina agrostis (Steinbuch, 1799) Filipjev, 1936 Anguina australis Steiner, 1940 Anguina funesta Price, Fisher & Kerr, 1979 Anguina graminis (Hardy, 1850) Filipjev, 1936 Anguina microlaenae (Fawcett, 1938) Steiner, 1940 Anguina paludicola Bertozzi & Davies, 2009 Anguina tritici (Steinbuch, 1799) Filipjev, 1936 Aphasmatylenchus straturatus Germani, 1970 Aphelenchoides agarici Seth & Sharma, 1986 Aphelenchoides arachidis Bos, 1977 Aphelenchoides bicaudatus (Imamura, 1931) Filipjev & Schuurmans Stekhoven, 1941 Aphelenchoides blastophthorus Franklin, 1952 Aphelenchoides composticola Franklin, 1957 Aphelenchoides Aphelenchoides Buhrer, 1932 Aphelenchoides Aphelenchoides fragariae (Ritzema Bos, 1890) Christie, 1932 ritzemabosi (Schwartz, 1911) Steiner & sacchari Hooper, 1958 saprophilus Franklin, 1957 Yes Yes Yes Yes Yes Yes Yes Yes Yes 337 (continued) Nematodes of phytosanitary importance Aphelenchoides besseyi Christie, 1942 Acts as vector of bacterium Rathyaibacter (Evtushenko et al., 1994). The bacteria-nematode complex causes Annual Rye Grass Toxicity (ARGT) in livestock (Edgar et al., 1982). Acts as vector of bacterium Rathyaibacter (Stynes & Bird, 1980), part of bacteria-nematode ARGT disease complex. Acts as vector of bacterium Rathayibacter toxicus (syn. Clavibacter toxicus) on annual veldtgrass (Riley et al., 2001), part of bacteria-nematode ARGT disease complex. Acts as vector of bacterium Rathayibacter toxicus (Riley, 1995), part of bacteria-nematode ARGT disease complex. Acts as vector of bacterium Rathayibacter festucae on red fescue (Dorofeeva et al., 2002), part of bacteria-nematode ARGT disease complex. Potential vector of bacterium Rathayibacter toxicus (Riley & Barbetti, 2008), part of bacteria-nematode ARGT disease complex. Acts as vector of bacterium Rathayibacter toxicus (Bertozzi & Davies, 2009), part of bacteria-nematode ARGT disease complex. Important parasite of wheat in countries which do not have access to clean/certified nematode free seeds. Reduced wheat yields by 43–100% in India (Paruthi et al., 1987; Khan & Athar, 1996), 69–93% in Pakistan (Anwar & Inam-ul-Haq, 1992) and 32% in Turkey (Ozberk et al., 2011). Is of limited importance in countries using clean/certified nematode free seeds for planting. Pathogenic to peanuts and cowpeas (Baujard & Martiny, 1995b). This species is only known from the south western region of Burkina Faso and yield loss of peanuts can be as great as 70% in heavily infested fields (Dickson & De Waele, 2005). Parasitises edible mushrooms (Grewal & Siddiqi, 1993; Aman et al., 2002). Pathogenic to peanuts. Species is seed borne hence there is high risk of spread, recently reported from Egypt from peanuts (Montasser et al., 2008) and also intercepted in seeds imported to India (Lal & Lal, 2006). Pathogenic to rice and causes rice white tip disease. Species is seed borne and has been disseminated in most rice growing areas of Africa, North, Central and South America, Asia, Eastern Europe and Pacific Islands. Importance and severity can vary with country locality and rice environment. Rice yield loss varies from 10 to 50% (Muthukrishnan et al., 1974; Silva & Da Silva, 1992). Parasitises cultivated mushrooms (Grewal & Siddiqi, 1993), yield loss depends on nematode population density and can be up to 45% (McLeod, 1967). Pathogen of ornamental plant Scabiosa caucasica, causes destruction of inflorescence and distortion of laminae (Franklin, 1952). Little information is available on the species. Parasitises edible mushrooms Agaricus bisporus (Goodey, 1960; McLeod, 1967; Grewal & Siddiqi, 1993; Gitanjali & Nandal, 2005). Pathogenic to strawberry and reduced yields by up to 60% (Duggan, 1969; Bohmer, 1981). Pathogenic to strawberry and reduced yields by 65% (Bohmer, 1981) and on chrysanthemum reduced yields by 45–92% (Baranowski, 1976). Parasitises edible mushrooms (Grewal & Siddiqi, 1993). Caused yield losses of up to 100% (Aman et al., 2002). Parasitises garlic (Balasubramanian et al., 2002) and edible mushroom (McLeod, 1968; Grewal & Siddiqi, 1993). Regulated pest* 338 Table 2 (continued) Notes on main hosts and yield losses Aphelenchoides subtenuis (Cobb, 1926) Steiner & Buhrer, 1932 Aphelenchoides swarupi Seth & Sharma, 1986 Aphelenchus avenae Bastian, 1865 Parasitises bulbous crops such as Crocus, Allium and some species of Tulipa and Narcissus (van Leeuwen & Trompert, 2011). Parasitises edible mushroom Calocybe indica and reduced yield by 13–15% (Kumar et al., 2010). Parasitises edible mushroom (Grewal & Siddiqi, 1993) and reduced yield by 11% (Kumar et al., 2010). The species is also associated with a wide range of crop plants but is not pathogenic. Parasitises a wide range of cultivated crops (Christie et al., 1952; Esser, 1976), turf (Christie et al., 1954). Also forms disease complex with fungal pathogens (Holdeman & Graham, 1952). Pathogenic to potatoes (Crow et al., 2000b), turf (Crow, 2005), soybean (McSorley & Dickson, 1989). Can reduce the yield of cotton at low population densities or result in crop failure when population density is high (Crow et al., 2000a; Koenning et al., 2004). Species is listed as of quarantine concern (Karnkowski, 2007). Pathogenic to peanuts (Reddy et al., 1984; Dickson & De Waele, 2005) and is capable of parasitizing other crops (Thakar et al., 1986). Parasitises sugarbeet (Kalatur, 2008), wheat (Jones, 1979), turf (Blackburn et al., 1997) and flax (Skarbilovich, 1971). Parasitises small millet (Jain, 2009), wheat (Patel & Thakar, 1989) and maize (Jain, 1982). Yes Pathogenic to palm trees and causes red ring disease (Inserra et al., 2005). Pathogenic to pine trees; causes wilt disease however very little information is available on biology and ecology of this species (Smith et al., 2008). Pathogenic to Pinus thurnbergii and P. taiwanensis seedlings (Chen et al., 2010). Is able to carry bacteria responsible for the pine wilt disease complex (Zhao et al., 2009). The quarantine status of this species is not clear (See Society of Nematologists expert comments and discussion on this species) Pathogenic to pine trees; causes pine wilt disease (Kiyohara & Tokushige, 1971). Widely quarantined species (Dwinell & Nickle, 1989; Mota & Vieira, 2008; Zhao et al., 2008). Parasitises cactus (Zygocactus truncatus and Hylocereus trigonus) and commonly grown ornamental cacti (Paneque & Sampedro, 1986; Narbaev & Mirsalimova, 1989; Cho et al., 1995). Parasitises barley and is able to reproduce on weed hosts (Tovar-Soto et al., 2003; Tovar-Soto et al., 2007). Yes Belonolaimus gracilis Steiner, 1949 Belonolaimus longicaudatus† Rau, 1958 Bitylenchus brevilineatus (Williams, 1960) Jairajpuri, 1982 ª 2013 The Authors. Journal compilation ª 2013 OEPP/EPPO, EPPO Bulletin 43, 334–374 Bitylenchus dubius (B€utschli, 1873) Filipjev, 1934 Bitylenchus vulgaris (Upadhyay, Swarup & Sethi, 1972) Jairajpuri, 1982 Bursaphelenchus cocophilus (Cobb, 1919) Baujard, 1989 Bursaphelenchus hunanensis Yin Fang & Tarjan, 1988 Bursaphelenchus mucronatus Mamiya & Enda, 1979 Bursaphelenchus xylophilus (Steiner & Buhrer 1934) Nickle 1970 Cactodera cacti (Filipjev & Schuurmans-Stekhoven, 1941) Krall & Krall, 1978 Cactodera galinsogae Tovar-Soto, del-Prado-Vera, Nicol, Evans, Sandoval-Islas & Martinez-Garza, 2003 Cactodera rosae Cid del Prado & Miranda, 2008 Ditylenchus africanus Wendt, Swart, Vrain & Webster, 1995 Ditylenchus angustus (Butler, 1913) Filipjev, 1936 Ditylenchus destructor Thorne, 1945 Ditylenchus dipsaci† (K€uhn, 1857) Filipjev 1936 Ditylenchus gigas† Vovlas, Troccoli, Palomares-Rius, De Luca, Liebanas, Landa, Subbotin & Castillo 2011 Parasitises barley (Cid del Prado & Miranda, 2008). Parasitises groundnut seeds and pods. Downgrades groundnut kernel quality by 32–64% (Mc Donald et al., 2005). Important pest of rice. Caused yield losses of 10–100% (Hashioka, 1963; Cox & Rahman, 1980; Rahman et al., 1994; Latif et al., 2011). Important pest of a wide range of root and tuber crops including potato, sweet potato, garlic, onions, peanuts. Caused potato yield loss of up to 94% in Sweden (Andersson, 1971), 5–10% in Iran (Barooti, 1997). Controlling D. destructor in sweet potato resulted in yield increase (Jin et al., 2008). Important pest of a wide range of crops. Crop loss on onions and garlic can be as high as 60–80% (Sturhan & Brzeski, 1991), damages potato (Goodey, 1956), canola (Taylor & Szot, 2000). There are about 30 biological races which have different host ranges (Sturhan et al., 2008). Forms disease complexes with fungal pathogens (Hillnhutter et al., 2011). Pathogenic to beans. Recently-described species from the D. dipsaci species complex, previously referred to as the giant race of D. dipsaci (Vovlas et al., 2011). Yes Yes Yes Yes Yes Yes Yes (continued) S. K. Singh et al. Regulated pest* Species ª 2013 The Authors. Journal compilation ª 2013 OEPP/EPPO, EPPO Bulletin 43, 334–374 Table 2 (continued) Notes on main hosts and yield losses Ditylenchus medicaginis Wasilewska, 1965 Found on the leaves, stems and roots of Medicago sativa (commonly grown as pasture) (Wasilewska, 1965) and has been found associated with other crops such as wheat (Kheiri et al., 2002; Erum & Shahina, 2010). Parasitises edible mushroom (Goodey, 1960; Grewal & Siddiqi, 1993). Caused yield loss of up to 70% (Aman et al., 2002). Recently-described species parasitising Cirsium arvense (weed) previously reported as a race of D. dipsaci (Chizhov et al., 2010). Is of phytosanitary importance because of potential confusion with D. dipsaci. Pathogenic to celery, tomato and maize (Perry, 1953; Paracer et al., 1968). Important pest of potato with reported yield loss of up to 80% (Talavera et al., 1998). Yield loss varies depending on potato cultivar and species pathotypes (Evans & Franco, 1979). Important pest of potato with reported yield loss of up to 85% (Greco et al., 1984). Yield loss varies depending on potato cultivar, population density and species pathotypes (Evans & Franco, 1979). Important pest of tobacco with reported yield losses of 40–60% (LaMondia, 1995, 2002). Forms disease complex with Fusarium oxysporum (LaMondia & Taylor, 1987). Pathogenic to eggplant, tomato, wheat (Firoza & Maqbool, 1995), peanut, millet (Baujard & Martiny, 1995d), and fruit trees (Rossi & Camargo Barbosa Ferraz, 2005). Parasitizes a wide range of crop plants. Pathogenic to banana; causes root necrosis and stunted growth of banana (Bridge & Page, 1984). Also associated with fruit trees (Rossi & Camargo Barbosa Ferraz, 2005). Important pest of banana; caused yield loss of up to 29% (Speijer et al., 1999; Brentu et al., 2004). Parasitises olive trees (Inserra et al., 1979; Vovlas & Larizza, 1994). Yield loss estimates are not available. Parasitises corn (Vovlas & Inserra, 1985; Vovlas & Larizza, 1994), soybean (Elmiligy & Norton, 1973; Caveness et al., 1987), grapes (Pinochet et al., 1976) and pasture (Davis et al., 2004). Parasitises wheat (Jones, 1978), carnations (Khanna & Jyot, 2002) and marram grass (Reis et al., 2010). Parasitises olives (Abrantes et al., 1987) and sugarbeet (Spaull, 1982; Ebrahimi et al., 2004). Data on yield loss not available. Parasitises small millets (Jain, 2009), rice (Sharma et al., 1992a), sugarcane (Cadet & Albrecht, 1992), ornamental plants, medicinal plants (Rathour et al., 2003) and root crops (Ray et al., 1992). Parasitises litchi (Nath et al., 2008) and mango (Liu & Feng, 1995; Ni et al., 2004). Pathogenic to mango and litchi (Milne et al., 1971; Stokes, 1976; McSorley et al., 1981; McSorley, 1992; Saha et al., 2006; Chen et al., 2011). Parasitises grapes (Deimi & Mitkowski, 2010) and citrus (Crozzoli et al., 1998). Pathogenic to citrus (Inserra & Vovlas, 1977; Stokes, 1977; Lehman, 1981a), tomato and squash (McElroy & van Gundy, 1968). Associated with sugarcane (Moura & Almeida, 1982). Pathogenic to tomato (Chitambar, 1993, 1994) and celery (Emilse et al., 2011). Associated with maize (Jamali et al., 2004). Pathogenic to carrots in glasshouse experiment (McKewan, 1979) and cranberry (Bird, 1963). Important pest of wheat with reported yield loss of 20–50% (Meagher & Brown, 1974; Meagher, 1982). Pathogenic to pigeon pea, reduced yield by 20–67% and mungbean yield by 86% (Saxena & Reddy, 1987; Sharma et al., 1993a). Pathogenic to carrots, reduced yield by 50% (Greco, 1987; Bossis & Mugniery, 1989). Pathogenic to chickpeas and lentils, yields reduced by 20–100% depending on population density of nematode (Greco et al., 1988, 1993). Ditylenchus myceliophagus Goodey, 1958 Ditylenchus weischeri† Chizhov, Borisov & Subbotin, 2010 Dolichodorus heterocephalus Cobb, 1914 Globodera pallida† (Stone, 1973) Behrens, 1975 Globodera rostochiensis† (Wollenweber, 1923) Skarbilovich, 1959 Globodera tabacum (Lownsbery & Lownsbery, 1954) Skarbilovich, 1959 Helicotylenchus dihystera (Cobb, 1893) Sher, 1961 Helicotylenchus microcephalus Sher, 1966 Helicotylenchus multicinctus (Cobb, 1893) Golden, 1956 Helicotylenchus oleae Inserra, Vovlas & Golden, 1979 Helicotylenchus pseudorobustus (Steiner, 1914) Golden, 1956 Helicotylenchus varicaudatus Yuen, 1964 Helicotylenchus vulgaris Yuen, 1964 Hemicriconemoides cocophillus (Loos, 1949) Chitwood & Birchfield, 1957 Hemicriconemoides litchi Edward & Misra, 1964 Hemicriconemoides mangiferae Siddiqi, 1961 Hemicycliophora arenaria Raski, 1958 Hemicycliophora poranga Monteiro & Lordello, 1978 Hemicycliophora similis Thorne, 1955 Heterodera avenae† Wollenweber, 1924 Heterodera cajani Koshy, 1967 Heterodera carotae Jones, 1950 Heterodera ciceri Vovlas, Greco & Di Vito, 1985 Regulated pest* Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes 339 (continued) Nematodes of phytosanitary importance Species 340 Table 2 (continued) Notes on main hosts and yield losses ª 2013 The Authors. Journal compilation ª 2013 OEPP/EPPO, EPPO Bulletin 43, 334–374 Heterodera Heterodera Heterodera Heterodera Heterodera cruciferae Franklin, 1945 daverti Wouts & Sturhan, 1978 delvii Jairajpuri, Khan, Setty & Govindu, 1979 elachista Ohshima, 1974 fici Kirjanova, 1954 Heterodera Heterodera Heterodera Heterodera Heterodera Heterodera Heterodera Heterodera filipjevi (Madzhidov, 1981) Stelter, 1984 glycines† Ichinohe, 1952 goettingiana Liebscher, 1892 graminis Stynes, 1971 hordecalis Andersson, 1975 humuli Filipjev, 1934 latipons Franklin, 1969 medicaginis Kirjanova, 1971 Heterodera mediterranea Vovlas, Inserra & Stone, 1981 Heterodera oryzae Luc & Berdon Brizuela, 1961 Heterodera oryzicola Rao & Jayaprakash, 1978 Heterodera sacchari Luc & Merny, 1963 Heterodera schachtii Schmidt, 1871 Heterodera skohensis Kaushal, Sharma & Singh, 2000 Heterodera trifolii Goffart, 1932 Heterodera zeae Koshy, Swarup & Sethi, 1971 Hirschmanniella gracilis (de Man, 1880) Luc & Goodey, 1964 Hirschmanniella imamuri Sher, 1968 Hirschmanniella miticausa Bridge, Mortimer & Jackson, 1983 Hirschmanniella oryzae (van Breda de Hann, 1902) Luc & Goodey, 1964 Hirschmanniella spinicaudata (Schuurmans Stekhoven, 1944) Luc & Goodey, 1962 Pathogenic to cabbage, rape, radish, rutabaga (Krnjaic & Krnjaic, 1987; Evans & Webb, 1989; Chizhov et al., 2009). Pathogenic to carnations (Ambrogioni & D’Errico, 1994) and Egyptian clover (Massoud et al., 1988). Parasitises cereals (Jairajpuri et al., 1979; Krishna Prasad et al., 1980). Parasitises upland rice with reported yield loss of 16% (Ohshima, 1974; Shimizu, 1976). Pathogenic to figs (Braasch, 1973). Can also occur in glasshouses (Monteiro et al., 1977; Subbotin et al., 1989; Krnjaic′ et al., 1994). Pathogenic to wheat with reported yield loss of up to 48% (Hajihasani et al., 2010a). Important pest of soybean with reported yield loss of up to 70% (Ichinohe 1955). Also see (Wrather et al., 1997). Pathogenic to peas and broad beans; pea yield reduced by 68–85% (Greco et al., 1991; Greco & Di Vito, 1994). Parasitises turf grass (Cynodon dactylon) on bowling greens (Stynes, 1971). Data on economic importance is not available. Parasitises cereals, commonly associated with barley, rye, grasses (Andersson, 1975) and wheat (Lombardo et al., 2009). Pathogenic to hops (Hafez et al., 1999b; Hay & Pethybridge, 2003). Pathogenic to wheat, reduced yields by 55% (Hajihasani et al., 2010b). Pathogenic to lucerne (Medicago eusativa, M. falcata and M. lupulina); reduced yields by 46% at high population densities (Alpat’ev et al., 1980; Terent’eva, 1982; Artokhina, 1984). Pathogenic to olive and pistachio (Vovlas et al., 1981; Vovlas & Inserra, 1983; Castillo et al., 1999; Castillo & Vovlas, 2002). Pathogenic to upland rice (Lin, 1971; Ou, 1972; Tanaka, 1976). Pathogenic to rice (Jacob et al., 1988; Rao et al., 1988; Sharma, 2001). Parasitises banana (Charles & Venkitesan, 1993, 1994). Pathogenic to upland rice, reduced yields by 57% (Babatola, 1983; Lamberti et al., 1991; Coyne & Plowright, 2000). Damage to rice is severe under drought conditions (Audebert et al., 2000). Parasitises sugarcane (Salawu, 1992). Important pest of sugarbeet (Muller, 1999) and pathogenic on a number of crops. Reduced yield of spinach by 49%, table beet by 30%, rutabaga by 35% and cabbage by 24% (Olthof et al., 1974). Parasitises rice (Kaushal et al., 2000) Data on economic importance is not available. Important pest of clover (Mowat, 1974; Sikora, 1977; Yeates, 1978; Sarathchandra et al., 1995; Clements & Cook, 1998). Also damages sugarbeet (Heijbroek & Maas, 1978; Maas & Heijbroek, 1982), and carnations (Lamberti et al., 1987b). Pathogenic to maize, reduced yield by 13–73% (Hashmi et al., 1993; Ismail et al., 1996; Krusberg et al., 1997). Parasitises rice (Dash et al., 2008; Mei et al., 2009), sunflower (Khan et al., 2011), tall fescue (Prior et al., 2010b) and pondweed potamogeton (Prejs, 1986). Pathogenic to rice, reduced yields by 31–37% under experimental conditions (Kuwahara & Iyatomi, 1970; Babatola, 1979; Babatola & Bridge, 1979; Ichinohe, 1988). Pathogenic to taro (Colocasia esculenta) and causes corm rot or ‘mitimiti’ disease (Bridge & Page, 1984). Yield loss estimates not available. Important pest of rice, yield reduced by 25–39% (Babatola & Bridge, 1979; Jonathan & Velayutham, 1987; Cho et al., 1994; Ying et al., 1996; Youssef & El-Hamawi, 1996). Parasitises rice (Babatola & Bridge, 1979, 1980; Rinaudo & Germani, 1981). Regulated pest* Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes (continued) S. K. Singh et al. Species ª 2013 The Authors. Journal compilation ª 2013 OEPP/EPPO, EPPO Bulletin 43, 334–374 Table 2 (continued) Species Notes on main hosts and yield losses Hoplolaimus columbus Sher, 1963 Important pest of cotton (Noe & Imbriani, 1986; Koenning et al., 2004; Koenning & Bowman, 2005; Bond & Mueller, 2007), and soybean (Kinloch, 1980; Perez et al., 2003). Pathogenic to soybean (Rodriguez-Kabana & Thurlow, 1980; Weaver & Rodriguez-Kabana, 1987), pine (Stokes, 1982) and turf (Giblin-Davis et al., 1995; Nambiar et al., 2008). Also parasitises lucerne (Ng & Chen, 1985). Pathogenic to rice (Rao, 1970) and maize (Khan et al., 1992). Parasitises mango (Anita & Chaubey, 2003). Associated with sugarcane (Mehta et al., 1994), groundnut and chickpea cropping system (Yadav & Sehgal, 2010). Also interacts with fungal pathogens (Mehta et al., 1994). Parasitises coffee (Vovlas & Lamberti, 1985), banana (Mateille, 1992; Speijer et al., 2001) and cowpea (Baujard & Martiny, 1995c). Parasitises rice, cowpea, pepper, eggplant, tomato, black gram, soybean, pineapple, cocoa (Lamberti et al., 1993), pigeon pea (Sharma & Nene, 1988), mulberry (Toida & Keereewan, 1991) and sandalwood (Sivaprakash et al., 2009). Parasitises sugarcane (Monteiro & Lordello, 1977). Not much information available on the species. Parasitises sugarcane (Roman, 1968). Not much information available on the species. Pathogenic to turf and pasture grasses (Siviour, 1978; Siviour & McLeod, 1979). Parasitises celery and chicory (Bleve-Zacheo et al., 1977) and Chenopodium quinoa (Bleve-Zacheo et al., 1984). Also acts as vector of Artichoke Italian Latent Virus (Lamberti & Bleve-Zacheo, 1977). Acts as vector of Cherry Rosette Disease caused by nepovirus (Brown et al., 1994a; Kunz, 2003). Hoplolaimus galeatus (Cobb, 1913) Thorne, 1935 Hoplolaimus indicus Sher, 1963 Hoplolaimus pararobustus (Schuurmans Stekhoven & Teuniessen, 1938) Sher, 1963 Hoplolaimus seinhorsti Luc, 1958 Ibipora jara Monteiro & Lordello, 1977 Ibipora lineatus (Roman, 1964) Monteiro & Lordello, 1977 Ibipora lolii (Siviour, 1978) Siviour & Mcleod, 1979 Longidorus apulus Lamberti & Bleve-Zacheo, 1977 Longidorus arthensis Brown, Grunder, Hooper, Klingler & Kunz, 1994 Longidorus attenuatus Hooper, 1961 Longidorus diadecturus Eveleigh & Allen, 1982 Longidorus elongatus (de Man, 1876) Micoletzky, 1922 Longidorus macrosoma Hooper, 1961 Longidorus martini Merny, 1966 Longidorus pisi Edward, Misra & Singh, 1964 Macroposthonia onoensis (Luc, 1959) De Grisse & Loof, 1965 Macroposthonia xenoplax (Raski, 1952) De Grisse & Loof, 1965 Meloidogyne acronea Coetzee, 1956 Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes 341 (continued) Nematodes of phytosanitary importance Longidorus fasciatus Roca & Lamberti, 1981 Longidorus leptocephalus Hooper, 1961 Parasitises sugarbeet and associated with docking disorder (Whitehead & Hooper, 1970). Also acts as vector of tomato black ring virus (Gibbs & Harrison, 1964b; Harrison, 1964) and Artichoke Italian Latent Virus (Taylor et al., 1976). Vector of Peach Rosette Mosaic Virus (Allen et al., 1982; Eveleigh & Allen, 1982). Is subject to compulsory control in European Union (Karnkowski, 2004). Pathogenic to sugarbeet, caused yield loss of up to 60% (Sigareva & Filenko, 1983). Parasitises barley, potatoes, raspberry (Sharma, 1965; Brown & Sykes, 1971, 1975). Also acts as vector for Raspberry Ringspot Virus, Tomato Black Ring Virus and Spoon Leaf Virus (Taylor, 1962, 1970; Harrison, 1964; Taylor & Gordon, 1970). Vector of Artichoke Italian Latent Virus (Roca et al., 1982; Brown et al., 1997). Vector of Cherry Leaf Roll Virus (Jones et al., 1981) and feeds directly on the roots of Lolium perenne (Robertson et al., 1984). Vector of Raspberry Ringspot Virus (Bercks, 1968; Trudgill & Brown, 1978; Buser, 1999), Carnation Ringspot Virus (Fritzsche & Schmelzer, 1967) and Cherry Leaf Roll Virus (Jones et al., 1981). Also causes direct damage to rose (Winfield, 1974). Vector of Mulberry Ringspot Virus (Yagita & Komuro, 1972; Bellizzi, 2004). Associated with peas (Edward et al., 1964), soybean (Fourie et al., 2001), maize (De Waele & Jordaan, 1988a) and sugarcane (Spaull & Heyns, 1991). Parasitises pepper (Freire & Monteiro, 1978). Associated with sugarcane (Rossi et al., 1996), cocoa (Sharma & Loof, 1974) and fig (Campos, 1997). Associated with peach tree short life syndrome (Lownsbery et al., 1973; Nesmith et al., 1981; Nyczepir et al., 1983). Parasitises clover, carnation, tomato (Hussey et al., 1992) and other herbaceous plants (Zehr et al., 1986). Pathogenic to cotton and sorghum, reduced yields by 90% and 56% respectively (Page 1983 cited in CABI ISC). Regulated pest* 342 Table 2 (continued) Notes on main hosts and yield losses Meloidogyne africana Whitehead, 1960 Meloidogyne arabicida Lopez & Salazar, 1989 Parasitises coffee (Whitehead, 1959; Waikwa et al., 1978). Pathogenic to coffee (Lopez & Salazar, 1989) and forms disease complex with Fusarium oxysporum (Bertrand et al., 2000). Polyphagous and pathogenic to a wide range of crops. Reduced yield of groundnuts by 13–50% (Patel et al., 1996) and oriental melon by 45% (Kim & Ferris, 2002). Species is not included on list of regulated pests in many countries because of its widespread occurrence. Pathogenic to chickpeas (Greco et al., 1992, 1994). Forms disease complex by interacting with Fusarium oxysporum (Castillo et al., 2003). Pathogenic to tomato (Charchar & Eisenback, 2002) and is able to parasitise pea and tomato with the root knot resistance Mi gene (Charchar et al., 2010). Pathogenic to tea (Loos, 1953; Mehta & Somasekhar, 1998). Pathogenic to potato, reduces the market value of produce and due to quarantine status of the species there are trade restrictions on seed and ware potatoes (Finley, 1981; Braasch et al., 1996; Ingham et al., 2007a; Norshie et al., 2011). Parasitises citrus (Shaosheng et al., 1990; Zhang & Xu, 1994), limited information is available on this species. Pathogenic to coffee (Lordello & Zamith, 1960; Schmidt, 1969). Parasitises citrus (Zheng et al., 1994), limited information is available on this species. Polyphagous species; caused yield loss in tomato by 65% (Cetintas et al., 2007) and damage on guava (Iwahori et al., 2009; Tigano et al., 2010). Can reproduce on cultivars with the Mi resistance gene (Brito et al., 2007; Cetintas et al., 2007, 2008; Kiewnick et al., 2009). Also see (Castagnone-Sereno, 2012) on the quarantine significance of this species. Pathogenic to kiwi fruit (Carneiro et al., 2003), grape (Di Vito et al., 2009) and tomato (Strajnar et al., 2012). Potential quarantine species in Europe. Pathogenic to coffee; caused yield loss of 45% (Barbosa et al., 2004, 2010). Parasitises rubber trees (Schwob et al., 1999; Correa & Rodella, 2002). Pathogenic to potatoes and carrots; reduces produce market value (Karssen et al., 2009). Parasitises peach (Handoo et al., 2004) and ornamental plants (Brito et al., 2010). Is able to overcome Mi gene resistance in tomato (Stanley et al., 2009). Meloidogyne arenaria† (Neal, 1889) Chitwood, 1949 Meloidogyne artiellia Franklin, 1961 Meloidogyne brasilensis Charchar & Eisenback, 2002 ª 2013 The Authors. Journal compilation ª 2013 OEPP/EPPO, EPPO Bulletin 43, 334–374 Meloidogyne brevicauda Loos, 1953 Meloidogyne chitwoodi Golden, O’Bannon, Santo & Finley, 1980 Meloidogyne citri Zhang, Gao, & Weng, 1990 Meloidogyne coffeicola Lordello & Zamith, 1960 Meloidogyne donghaiensis Zheng, Lin, & Zheng, 1990 Meloidogyne enterolobii Yang & Eisenback, 1983 Meloidogyne ethiopica Whitehead, 1968 Meloidogyne exigua Goeldi, 1892 Meloidogyne fallax Karssen, 1996 Meloidogyne floridensis Handoo, Nyczepir, Esmenjaud, van der Beek, Castagnone-Sereno, Carta, Skantar & Higgins, 2004 Meloidogyne fujianensis Pan, 1985 Meloidogyne graminicola Golden & Birchfield, 1965 Meloidogyne hapla Chitwood, 1949 Meloidogyne incognita† (Kofold & White, 1919) Chitwood, 1949 Meloidogyne indica Whitehead, 1968 Meloidogyne javanica† (Treub, 1885) Chitwood, 1949 Parasitises citrus (Pan, 1985; Pan et al., 1999). Limited information is available on this species. Important pest of rice, caused yield losses of 11–73% in rice (Rao et al., 1988; Soriano et al., 2000; Soriano & Reversat, 2003; Jain et al., 2012). Polyphagous, pathogenic to many crops including tomato (yield reduced by up to 50%: (Barker et al., 1976) and lettuce (yield reduced by up to 64%: (Olthof & Potter, 1972; Viaene & Abawi, 1996). Polyphagous, pathogenic to many crops including vegetables (yield reduced by up to 90%: (Bhatti & Jain, 1979; Lamberti et al., 1988), tomato (yield reduced by up to 85%: (Barker et al., 1976) and cotton (yield reduced by up to 47% (Davis & May, 2005). Species is not included on list of regulated pests because of its widespread occurrence. Parasitises Amaranthus, watermelon, cowpea (Patel et al., 2003) and medicinal plants (Lin et al., 2004). Polyphagous, pathogenic to many crops including peanuts [yield reduced by up to 23% (Patel et al., 1996)] and banana [yield reduced by up to 30% (Brentu et al., 2004)]. Species and is not included on list of regulated pests because of its widespread occurrence. Regulated pest* Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes (continued) S. K. Singh et al. Species ª 2013 The Authors. Journal compilation ª 2013 OEPP/EPPO, EPPO Bulletin 43, 334–374 Table 2 (continued) Species Meloidogyne Meloidogyne Meloidogyne Meloidogyne Meloidogyne Notes on main hosts and yield losses jianyangensis Yang, Hu, Chen & Zhu, 1990 kongi Yang, Wang, & Feng, 1988 mali Itoh, Ohshima & Ichinoe, 1969 mingnanica Zhang, 1993 minor Karssen et al., 2004 Meloidogyne naasi Franklin, 1965 Meloidogyne oryzae Maas, Sanders & Dede, 1978 Merlinius brevidens (Allen, 1955) Siddiqi, 1970 Merlinius microdorus (Geraert, 1966) Siddiqi, 1970 Merlinius nanus (Allen, 1955) Siddiqi, 1970 Nacobbus aberrans† (Thorne, 1935) Thorne & Allen, 1944 Neodolichodorus australis Hodda & Nambiar, 2005 Neodolichodorus citri S’Jacob & Loof, 1996 Paralongidorus australis Stirling & McCulloch, 1984 Paralongidorus maximus (B€utschli, 1874) Siddiqi, 1964 Yes Yes Yes Yes Yes Parasitises peas, beans, tomato and tobacco (Charchar et al., 2008b). Parasitises tomato and carrots (Charchar et al., 2009). Pathogenic to rice (Lopez, 1984; Sancho et al., 1987). Parasitises black rice and sedge grass (Medina et al., 2009). Parasitises ginger (Handoo et al., 2005) species described from intercepted material however information on this species in its native range is not available. Parasitises tomato (Amin, 1993), cowpeas, lettuce (Ponte & Santos, 1981; Ponte, 1987), Bohemeria nivea (Mishra & Mandal, 1988), tea (Huan, 1983), and Paullinia cupana var. sorbilis (Ferraz & Campelo, 1988). Pathogenic to clover (Bernard & Eisenback, 1997; Zahid et al., 2001; Mercer, 2005; Bell et al., 2006). Parasitises soybean, broad bean, garden pea, Korean lespedeza, sweet clover, and common vetch (Bernard & Jennings, 1997). Pathogenic to wheat and associated with reduced yields (Jordaan et al., 1992; Smiley et al., 2006). Pathogenic to lettuce and strawberry (Szczygiel, 1981). Associated with Rumex sp., Acer sp. (Ivanova, 1978) and barley (Andersen, 1979). Parasitises watermelon (Tan & Okten, 2011). Associated with fruit trees (Liskova et al., 2007), cereals, pulses, vegetables (Kepenekci & Okten, 1996; Erdal et al., 2001) and bermuda grass (Ibrahim et al., 2000). Pathogenic to tomato, potato, beans, and sugarbeet. Reduced yields of tomato by up to 83%, potato by 65%, beans by 36% and sugarbeet by 10–20% (Manzanilla-Lopez et al., 2002; Cristobal-Alejo et al., 2006). Pathogenic to carrots (Hodda & Nambiar, 2005). Associated with plum, peach, corn and Chenopodium (S’Jacob & Loof, 1996; Vovlas et al., 2003). Pathogenic to rice (Stirling & McCulloch, 1984; Lehman & Stirling, 1988; Stirling et al., 1989). Acts as vector of Cherry Leaf Roll Virus, Raspberry Ringspot Virus and Tomato Black Ring Virus (Jones et al., 1981, 1994; H€ ubschen et al., 2004). Pathogenic to Scots pine and European larch (Boag et al., 1977). Yes Yes Yes Yes 343 (continued) Nematodes of phytosanitary importance Meloidogyne paranaensis Carneiro, Carneiro, Abrantes, Santos & Almeida, 1996 Meloidogyne partityla Kleynhans, 1986 Meloidogyne phaseoli Charchar, Eisenback, Charchar & Boiteau, 2008 Meloidogyne pisi Charchar, Eisenback, Charchar & Boiteau, 2008 Meloidogyne polycephannulata Charchar, Eisenback, Vieira, Fonseca-Boiteau & Boiteau, 2008 Meloidogyne salasi Lopez, 1984 Meloidogyne thailandica Handoo, Skantar, Carta & Erbe, 2005 Meloidogyne thamesi Chitwood in Chitwood, Specht & Havis, 1952 (Goodey, 1963)‡ Meloidogyne trifoliophila Bernard & Eisenback, 1997 Pathogenic to mandarin orange (Yang et al., 1990; Zhu et al., 1991), limited information is available on this species. Parasitises citrus (Yang et al., 1986, 1991), limited information is available on this species. Pathogenic to apple and mulberry (Itoh et al., 1969; Inagaki, 1978; Toida, 1991). Parasitises citrus (Zhang, 1993; Zhang & Xu, 1994), limited information is available on this species. Pathogenic to turf grass, causes yellow patch disease on golf courses (Karssen et al., 2004). Parasitises potato (De Weerdt et al., 2011; Thoden et al., 2012). For a discussion on the phytosanitary importance of this species see Turner & Fleming (2005) and Morris et al., (2011). Polyphagous species, pathogenic to cereals, and reduced barley yields by up to 50% (York, 1980). Parasitises peas and beans (Gooris & D’Herde, 1969, 1972; Caubel et al., 1971; Ediz, 1972; Belair et al., 2006). Polyphagous species, pathogenic to rice and reduced yields by 10–15% in greenhouse experiments (Segeren & Sanchit, 1984). Additional hosts include grasses, wheat, tomato and potato (Maas et al., 1978). Pathogenic to coffee (Castro et al., 2003). Parasitises soybean, medicinal plants and weeds (Carneiro et al., 1996; Roese & Oliveira, 2004; Roese et al., 2004; Monarco et al., 2011). Pathogenic to pecan, parasitises walnut and hickory (Kleynhans, 1986; Starr et al., 1996; Nyczepir et al., 2002, 2006). Parasitises bean, tobacco, tomato and peas (Charchar et al., 2008a). Regulated pest* 344 Table 2 (continued) Notes on main hosts and yield losses Paratrichodorus allius (Jensen, 1963) Siddiqi, 1974 Acts as vector of Tobacco Rattle Virus which causes corky ringspot disease in potatoes (Mojtahedi & Santo, 1999; Gieck et al., 2007; Ingham et al., 2007b; Charlton et al., 2010). Associated with corn and wheat (Mojtahedi et al., 2002). Pathogenic to barley and wheat (Spaull, 1980; Spaull & Murphy, 1983; Spaull & Mewton, 1984). Acts as vector of Pea Early Browning and Tobacco Rattle Tobraviruses (Karanastasi et al., 2001; Karanastasi & Brown, 2004). Potential vector of Pea Early Browning and Tobacco Rattle Tobraviruses (Karanastasi & Brown, 2004). Associated with wheat (Roca & Arias, 1986). Pathogenic to sorghum, cowpea and eggplant (Baujard & Martiny, 1995a). Parasitises wheat (Jordaan et al., 1992), rice (Coyne et al., 2001), sugarcane (Blair & Stirling, 2007), pasture grasses Dactylis glomerata, Lolium multiflorum, L. perenne Festuca arundinacea (Bell & Watson, 2001) and turf grass (Crow, 2005). Associated with cotton (Bajaj & Bhatti, 1982), sugarcane (Maqbool & Hashmi, 1987) and litchi (Saha et al., 2006) Acts as vector of Tobacco Rattle Virus (Cooper & Thomas, 1970). Pathogenic to millet and sorghum (Baujard & Martiny, 1995a). Parasitises rice (Sharma et al., 1992a) and pasture grasses Dactylis glomerata, Lolium multiflorum, L. perenne (Bell & Watson, 2001). Acts as vector of Tomato Black Ring Virus, Tobacco Rattle Virus and Pea Early Browning Virus (Gibbs & Harrison, 1964b; Brown et al., 1989; Ploeg et al., 1992). Parasitises pine (Choleva & Samuleyan, 1996). Associated with potatoes (Leshcheva, 1982; De Pelsmaeker et al., 1985) and fruit trees (Liskova et al., 2007; Kumari, 2010). Associated with barley (Sheedy et al., 2010), sugarcane (Carneiro et al., 1982), grapevine (Wang et al., 1996; Aballay & Eriksson, 2006) and citrus (Park et al., 2008). Parasitises blueberry (Braasch, 1976; Forge et al., 2009; Zasada et al., 2010b). Associated with ornamentals, eucalyptus, pine tree nurseries (Ferraz et al., 1984; Braasch & Sturhan, 1991) and azaleas (Brinkman, 1977; Cotten & Hooper, 1991). Acts as vector of Tobacco Rattle Virus which causes corky ringspot disease of potato (De Pelsmaeker, 1987; Bos & Krikke, 1991; Riga & Neilson, 2005). Parasitises artichoke (Karanastasi et al., 2005). Acts as vector of Tobacco Rattle Virus (Roca & Rana, 1981). Associated with artichoke, tobacco and fruit trees (Roca & Lamberti, 1984). Pathogenic to parsley (Brzeski & Radzikowska, 1980) and celery (Brzeski, 1975). Parasitises peppermint (Monteiro, 1978). Pathogenic to dry land peas and lentils, associated with reduced yields (Riga et al., 2008), parsley (Sprau, 1969), mint (Lisetskaya, 1971) and roses (Macdonald, 1976). Associated with wheat (Jordaan et al., 1992), sugarcane (Decker et al., 1970), pineapples (Linford et al., 1949) anthuriums and other tropical ornamentals (Bala & Hosein, 1996). Parasitises soybean (Ferris & Bernard, 1962; Acosta, 1982), tomato (Dickerson, 1979) and raspberry (Doucet et al., 2005). Pathogenic to flowers, Alstroemeria cv. Jubilee (Amsing, 1996). Associated with tomato, potato, oats and carnations (Cotten et al., 1991). Pathogenic to soybean (McSorley & Dickson, 1989), maize (Inomoto, 2011), citrus (Inserra & Vovlas, 1977) and pineapple (Dias-Arieira et al., 2010). Associated with Miscanthus giganteus and Panicum virgatum used for biofuels (Mekete et al., 2011). Pathogenic to coffee (Inomoto et al., 2007; Trinh et al., 2011), citrus (Obannon & Tomerlin, 1973), banana (Brentu et al., 2004) and yams (Acosta & Ayala, 1975). Reduced yield of banana cv. Grand Naine by 34% (van den Bergh et al., 2006). Paratrichodorus anemones (Loof, 1965) Siddiqi, 1974 Paratrichodorus hispanus Roca & Arias, 1986 Paratrichodorus minor (Colbran, 1956) Siddiqi, 1974 Paratrichodorus mirzai (Siddiqi, 1960) Siddiqi, 1974 Paratrichodorus nanus (Allen, 1957) Siddiqi, 1974 ª 2013 The Authors. Journal compilation ª 2013 OEPP/EPPO, EPPO Bulletin 43, 334–374 Paratrichodorus pachydermus (Seinhorst, 1954) Siddiqi, 1974 Paratrichodorus porosus (Allen, 1957) Siddiqi, 1974 Paratrichodorus renifer Siddiqi, 1974 Paratrichodorus teres (Hooper, 1962) Siddiqi, 1974 Paratrichodorus tunisiensis (Siddiqi, 1963) Siddiqi, 1974 Paratylenchus bukowinensis Micoletzky, 1922 Paratylenchus hamatus Thorne & Allen, 1950 Paratylenchus minutus Linford in Linford, Oliveira & Ishii, 1949 Pratylenchus alleni Ferris, 1961 Pratylenchus bolivianus Corbett, 1983 Pratylenchus brachyurus (Godfrey, 1929) Filipjev & Schuumans Stekhoven, 1941 Pratylenchus coffeae (Zimmermann, 1898) Filipjev & Schuurmans-Stekhoven, 1941 Regulated pest* Yes Yes Yes Yes Yes Yes (continued) S. K. Singh et al. Species ª 2013 The Authors. Journal compilation ª 2013 OEPP/EPPO, EPPO Bulletin 43, 334–374 Table 2 (continued) Notes on main hosts and yield losses Pratylenchus convallariae Seinhorst, 1959 Pathogenic to Convallariae spp. (Cayrol & Ritter, 1962). Parasitises sweet potato (Huan & Xu, 1985), corn (Urek et al., 2003) and peach (Wu et al., 1993). Parasitises potato, corn, cereals, grasses, olives, roses, raspberry and other hosts (Castillo & Vovlas, 2007). Parasitises a wide range of crops including cotton, sugarcane, maize, oats, pearl millet, wheat, pigeon pea and peanuts (Sharma et al., 1992b; van Biljon & Meyer, 2000; Castillo & Vovlas, 2007). Parasitises a wide range of crops including cereals (Corbett, 1972), fruit trees and ornamental plants (Castillo & Vovlas, 2007). Parasitises a wide range of crops including corn, sweet potato, fruit trees and grasses (Castillo & Vovlas, 2007). Pathogenic to banana and forms disease complex with bacterial wilt pathogens (Peregrine & Bridge, 1992; Pattison et al., 2002; Talwana et al., 2003). Associated with grapevine, strawberry and other crops (Castillo & Vovlas, 2007). Parasitises corn, soybean, citrus, peach, apricot, and Pennisetum purpureum (Castillo & Vovlas, 2007). Pathogenic to tea (Gnanapragasam et al., 1987) pasture grasses (Inserra et al., 1996) and oranges (Ushiyama & Ogaki, 1970; Castillo & Vovlas, 2007). Parasitises chickpea, lentils, potatoes, carrots, wheat, and chrysanthemum (Orion & Glazer, 1987; Orion et al., 1988, 1995; Greco & Di Vito, 1994; Choi et al., 2006). Important pest of cereals (reducing wheat yields by up to 36%: (Smiley & Machado, 2009; Taylor et al., 1999), potato (Olthof, 1990; Hafez et al., 1999a), canola (Fatemy et al., 2006), oilseed rape (Kumari, 2012) and parasitises a wide range of other crops (Castillo & Vovlas, 2007). Pathogenic to a wide range of crops including cereals, corn potatoes, sweet potatoes, fruit trees, and ornamental plants (Castillo & Vovlas, 2007). Forms disease complex with fungal pathogen Verticillium dahlia to cause early dying complex in potato (Martin et al., 1982b). Parasitises wheat and other cereals (Corbett, 1969, 1970) and faba beans (Troccoli & Di Vito, 2002). Pathogenic to cereals, winter wheat and rye (Goffart, 1942; Stepanchuk, 1978). Parasitises a wide range of crops (Castillo & Vovlas, 2007). Forms disease complex with Fusarium moniliforme var. subglutinans (Revelo Moran et al., 1993). Parasitises a wide range of crops including corn, millet, soybean, tea, fruit trees, strawberry and date palms (Castillo & Vovlas, 2007). Associated with potato (Akgul et al., 2010). Pathogenic to potato (Martin et al., 1982a), and reduced maize yield by 26% under experimental conditions (Olowe, 2011). Parasitises a wide range of other crops (Castillo & Vovlas, 2007). Pathogenic to cotton and forms disease complex with Fusarium oxysporum f.sp. vasinfectum (Saadabi & Yassin, 2007). Parasitises pigeon pea and lubia bean (Yassin & Mohamed, 1980), potato, sweet potato, yams and sugarcane (Castillo & Vovlas, 2007). Parasitises cotton, millet and tobacco (Carta et al., 2002), soybean (Fourie et al., 2001), mango (Liu & Feng, 1995), pine (Savkina, 1989), sugarcane (van den Berg & Queneherve, 2000) and potato (Khan & Singh, 1974). Important pest of cereals and causes wheat yield loss of up to 70% (Taylor et al., 1999; Smiley et al., 2005; Thompson et al., 2008). Parasitises a wide range of other crops (Castillo & Vovlas, 2007). Pathogenic to plum (McKenry, 1989) and other Prunus species (Pinochet et al., 1996), strawberry (yields reduced by 80%: (Mohotti et al., 1997), chrysanthemum (Lee et al., 2008). Over 80 species of plants are reported as hosts (Castillo & Vovlas, 2007). Pratylenchus crenatus Loof, 1960 Pratylenchus delattrei (Luc, 1958) Handoo & Golden, 1989 Pratylenchus fallax Seinhorst, 1968 Pratylenchus flakkensis Seinhorst, 1968 Pratylenchus goodeyi Sher & Allen, 1953 Pratylenchus hexincisus Taylor & Jenkins, 1957 Pratylenchus loosi Loof, 1960 Pratylenchus mediterraneus Corbett, 1983 Pratylenchus neglectus (Rensch, 1924) Filipjev & Schuurmans Stekhoven, 1941 Pratylenchus penetrans (Cobb, 1917) Filipjev & Schuurmans Stekhoven, 1941 Pratylenchus pinguicaudatus Corbett, 1969 Pratylenchus pratensis (de Man, 1880) Filipjev, 1936 Pratylenchus pseudopratensis Seinhorst, 1968 Pratylenchus scribneri Steiner in Sherbakoff & Stanley, 1943 Pratylenchus sudanensis Loof & Yassin, 1971 Pratylenchus teres Khan & Singh, 1975 Pratylenchus thornei Sher & Allen, 1953 Pratylenchus vulnus Allen & Jensen, 1951 Regulated pest* Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes 345 (continued) Nematodes of phytosanitary importance Species 346 Table 2 (continued) Notes on main hosts and yield losses Pratylenchus zeae Graham, 1951 Pathogenic to maize (causing complete crop failure: (Patel & Patel, 2000), sugarcane (Tarte et al., 1977), and a wide range of other crops (Castillo & Vovlas, 2007). Pathogenic to maize (Suarez et al., 1985; Mundo et al., 1987; Tovar-Soto et al., 2006). Parasitises grasses (Mulvey & Stone, 1976). Associated with potato, recorded during a routine survey for PCN (Handoo et al., 2010). Can be confused with other regulated cyst nematodes. Parasitises grasses Poa annua (annual bluegrass), Poa pratensis (Merion Kentucky bluegrass), Lolium perenne (perennial ryegrass) and Festuca rubra rubra (Radice et al., 1984; Cook et al., 1992; Vandenbossche et al., 2011). Associated with potato and sugarbeet (Urek & Lapajne, 2001). Pathogenic on squash Cucurbita pepo (McSorley & Waddill, 1982), sorghum (Cuarezma-Teran & Trevathan, 1985). Associated with mango (McSorley et al., 1981), soybean (Niblack, 1988), cotton (Robbins et al., 1989) and horseradish (Walters et al., 2004). Parasitises coffee (Mekete et al., 2008), ornamental plants (Zhang et al., 1998), potato (Khan et al., 1990b), sunflower and maize (Doucet, 1986). Pathogenic to melon (Khan & Khanzada, 1990) and maize (Khan et al., 1988). Associated with ornamental plants (Hung et al., 2011). Pathogenic to coffee (Trinh et al., 2004, 2011, 2012). Punctodera chalcoensis Stone, Sosa Moss & Mulvey, 1976 Punctodera matadorensis Mulvey & Stone, 1976 Punctodera punctata (Thorne, 1928) Mulvey & Stone, 1976 Quinisulcius acutus (Allen, 1955) Siddiqi, 1971 Quinisulcius capitatus (Allen, 1955) Siddiqi, 1971 ª 2013 The Authors. Journal compilation ª 2013 OEPP/EPPO, EPPO Bulletin 43, 334–374 Quinisulcius curvus (Williams, 1960) Siddiqi, 1971 Radopholus arabocoffeae Trinh, Nguyen, Waeyenberge, Subbotin, Karssen & Moens, 2004 Radopholus citri Machon & Bridge, 1996 Radopholus duriophilus Nguyen, Subbotin, Madani, Trinh, & Moens, 2003 Radopholus nativus Sher, 1968 Radopholus similis† (Cobb 1893) Thorne, 1949 Rotylenchulus macrodoratus Dasgupta, Raski & Sher, 1968 Rotylenchulus parvus (Williams, 1960) Sher, 1961 Rotylenchulus reniformis† Linford & Oliveira, 1940 Rotylenchus robustus (de Man, 1876) Filipjev, 1936 Pathogenic to citrus (Machon & Bridge, 1996). Limited information available on this species. Pathogenic to durian and coffee (Nguyen et al., 2003; Trinh et al., 2004). Pathogenic to wheat (Riley & Kelly, 2001). Parasitises canola, triticale, oat, field pea, faba bean, durum wheat, narrowleafed lupin and chickpea (Vanstone, 2010). Important pest of banana (Gowen et al., 2005) and a wide range of crops, with over 250 plant hosts (Obannon, 1977). Has two pathotypes, the banana pathotype is more widespread and parasitises banana plus other crops, while the citrus pathotype parasitises only citrus and is restricted to Florida, USA (Huettel et al., 1984). Pathogenic to olives (Inserra & Vovlas, 1981; Lamberti, 1981). Parasitises Ceratonia siliqua (carob), fig, grape, Hedera helix (ivy), Laurus nobilis (laurel), Neriurn oleander (oleander), olive, Prunus amygdalus, (almond), P. armeniaca L. (apricot), P. domestica L. (plum), Quercus calliprinos and Q. farnetto (oak), soybean and carnations (Inserra & Vovlas, 1980; Vovlas, 1983; Vovlas & Vlachopoulos, 1991). Parasitises barley, bean, corn, cowpea, and bermuda grass (Dasgupta & Raski, 1968), soybean (Fourie et al., 2001), macadamia, pearl millet, potato, papaya, thyme, tobacco, tomato, sunn hemp, sugarcane and cotton (Robinson et al., 1998). Important pest of cotton (Koenning et al., 1999, 2004). Pathogenic to soybean (McGawley et al., 2011), castor (Neelu & Azam, 2012) and squash (McSorley & Waddill, 1982). Parasitises a wide range of crops with over 360 host species recorded (Robinson et al., 1998). Pathogenic to spruce (Hijink, 1969; Boag, 1982), pine (Rossner, 1969), olives (Abrantes et al., 1987), peas (Seinhorst, 1954), carrots (Boag, 1979) and lettuce (Lear et al., 1969). Regulated pest* Yes Yes Yes Yes Yes Yes Yes Yes Yes (continued) S. K. Singh et al. Species ª 2013 The Authors. Journal compilation ª 2013 OEPP/EPPO, EPPO Bulletin 43, 334–374 Table 2 (continued) Notes on main hosts and yield losses Scutellonema brachyurus (Steiner, 1938) Andrassy, 1958 Parasitises maize, sorghum, rice (De Waele & Jordaan, 1988a,b; van den Berg & De Waele, 1989), fruit trees (Rossi & Camargo Barbosa Ferraz, 2005) and lilyturf, Liriope muscari (Agudelo & Harshman, 2011). Associated with peach tree short life syndrome (Nesmith et al., 1981). Pathogenic to yams (Acosta & Ayala, 1975; Adesiyan et al., 1975), potato (Coyne et al., 2011), Sesamum indicum, Vigna unguiculata, pigeon pea, okra, tomato and melons (Adesiyan, 1976). Pathogenic to groundnut (Sharma et al., 1992c). Parasitises upland rice (Coyne et al., 2001), sunflower (Eldin & Siddiq, 1995), maize (Talwana et al., 2008), medicinal plant Aloe barbadensis (Kindelan et al., 1991) and grapes (Wang et al., 1991). Parasitises banana (Timm, 1965). Associated with trees, recovered from bushland soils (Reay, 1982) and fruit tree nursery soils (van den Berg, 1981). Parasitises yams (Park & Khan, 2007), potatoes, sweet potatoes (Njuguna & Bridge, 1998), pigeon pea (Sharma et al., 1993b), maize, pineapple, citrus (van den Berg & Heyns, 1973) and grape (Wang et al., 1991). Associated with Prunus amygdalus (Khan & Khan, 1985) and Ficus sp. (Melillo & Troccoli, 1993). Parasitises alder (Subbotin, 1989), downy birch (Prior et al., 2010a), and chestnut (Palomares-Rius et al., 2010). Parasitises leaves of tree Nothofagus obliqua (Vovlas et al., 2000; Baldini Urrutia & Aguayo Silva, 2007). Parasitises inflorescence of Hyparrheniae sp., H. collina, H. ruffa, H. nyassae, H. newtonii, H. variabilis, (Corbett, 1966). Limited information is available on this species. Pathogenic to turf grass Poa annua (Chizhov & Mar’enko, 1984; Mitkowski & Jackson, 2003; Mitkowski, 2007). Parasitises Ammophila breviligulata (Glover & Halisky, 1973), wheat and barley (Lu, 1983). Also associated with coryneform bacteria (Evtushenko et al., 1994, 2000). Parasitises bonsai trees (Hirata & Yuhara, 1986), coniferous trees (Kiyohara, 1970), pear (Zhao et al., 2005), China fir, peach, apricot, persimmon, and apple (Xu & Decraemer, 1995). Vector of Tobacco Rattle Virus which causes sparing disease of potatoes (Alphey et al., 1975; Brown et al., 1989). Pathogenic to barley (Spaull & Mewton, 1984). Parasitises potato, strawberry and hops (De Pelsmaeker & Coomans, 1985), fruit trees (Liskova et al., 2007), Populus sp. (Cooper & Sweet, 1976) and Cupressus sempervirens (G omezBarcina & Castillo, 1988). Vector of Pea Early Browning Virus (Harrison, 1966), Spinach Yellow Mottle Virus (Kurppa et al., 1981) and Tobacco Rattle Virus which causes sparing disease of potatoes (Brown & Sykes, 1973; Alphey et al., 1975; Ploeg et al., 1992). Vector of Tobacco Rattle Virus (van Hoof, 1967; Brown et al., 1996). Parasitises tobacco (Wyss, 1973, 1975), beans (Coosemans, 1993), potato, hops, strawberry (De Pelsmaeker & Coomans, 1985), and gladiolus (Cremer & Kooistra, 1964). Vector for Tobacco Rattle Virus (Brown et al., 1989), and Pea Early Browning Virus (Gibbs & Harrison, 1964a). Parasitises a wide range of plants including apple, barley, maize, pea, potato, rye, sugarbeet and wheat (Hooper, 1963; Gibbs & Harrison, 1964a; Pitcher & McNamara, 1971; Cooke, 1984). Parasitises rice (Xie et al., 2007), coffee (Mekete et al., 2008), red clover (Amosu & Taylor, 1974; Coates-Beckford, 1982) and citrus (Esser et al., 1993). Pathogenic to rice (Aly & Shaukat, 2000) and sugarcane (Hasselrot de Gomez et al., 1980; Bond et al., 2004). Parasitises corn (Chen et al., 2006), horseradish (Walters et al., 2004) and potatoes (Khan & Hussain, 2004). Scutellonema bradys (Steiner, 1937) Andrassy, 1958 Scutellonema clathricaudatum Whitehead, 1959 Scutellonema minutum Sher, 1964 Scutellonema unum Sher, 1964 Sphaeronema alni Turkina & Chizhov, 1986 Subanguina chilensis Vovlas Troccoli & Moreno, 2000 Subanguina hyparrheniae (Corbett, 1966) Fortuner & Maggenti, 1987 Subanguina radicicola (Greeff, 1872) Paramonov, 1967 Trichodorus cedarus Yokoo, 1964 Trichodorus cylindricus Hooper, 1962 Trichodorus primitivus (de Man, 1880) Micoletzky, 1922 Trichodorus similis Seinhorst, 1963 Trichodorus viruliferus Hooper, 1963 Tylenchorhynchus agri Ferris, 1963 Tylenchorhynchus annulatus (Cassidy, 1930) Golden, 1971 Regulated pest* Yes Yes Yes Yes Yes 347 (continued) Nematodes of phytosanitary importance Species 348 Table 2 (continued) Notes on main hosts and yield losses Tylenchorhynchus brassicae Siddiqi, 1961 Pathogenic to rice (Khan et al., 1990a), tomato (Ahmad & Khan, 1988), cauliflower (Khan et al., 1994), chickpeas (Tiyagi & Alam, 1989) and pigeonpeas (Tiyagi & Parveen, 1990). Pathogenic to cotton (Kheir et al., 1977; Embabi & Shohla, 1978). Parasitises chickpeas (Castillo et al., 1991), wheat (Koliopanos & Vovlas, 1977; Eissa & Moussa, 1982; Smiley et al., 2004), alfalfa, wild ryegrass, sweet corn, parsley and tomato (Edongali & Lownsbery, 1980). Parasitises sugarcane (Blair et al., 1999), tobacco (Aycock et al., 1976), potatoes (Olthof et al., 1982), pigeon pea (Ingram & Rodriguez-Kabana, 1977), azaleas (Cotten & Hooper, 1991), Japanese holly (Baicheva, 1974), pine (Ruehle, 1973), bentgrass and Bermuda grass (Lucas et al., 1974). Parasitises guava (Abivardi, 1973), strawberry (Nesterov & Koev, 1972), maize (Nesterov & Lizogubova, 1972), coconut (Valdez, 1980) and cotton (Tu et al., 1972). Parasitises maize (Mahapatra & Das, 1984), rice (Baqri & Ahmad, 2000), sugarcane (Ray et al., 1994), sweet potato, cassava, yam (Ray et al., 1992), fruit trees (Afshar et al., 2006), chickpeas (Ali, 1993), cabbage (Bilgrami, 1994) and ginger (Luqman Khan & Makhnotra, 1998). Parasitises wheat (Smolik, 1972), sorghum (Smolik, 1977), rice (Haidar et al., 1996), sugarcane (Hu & Chu, 1964; Haider et al., 1987), bluegrass, bentgrass (Smolik & Malek, 1973; Davis et al., 1994), chilli (Prasad et al., 1991) and banana (Choudhury & Phukan, 1992). Parasitises peach, Carolina ash and saltbush (Inserra et al., 1990; Eisenback et al., 2007), swamp plants; Aster elliottii, Liquidambar styraciflua, Borrichia arborescens and B. frutescens (Dow et al., 1990). Important pest of citrus, responsible for slow decline disease of citrus (Cohn, 1965; Timmer & Davis, 1982). Other hosts include Calodendrium capensis, Citrus volkameriana, Ruta bracteosa and R. graveolens (Cohn, 1966), Cydonia oblonga, Diospyros sp., Olea europaea, Philadelphus coronarius, Poncirus trifoliata and Vitis Vinifera (Kwaye et al., 2008). Hosts differ depending on the species pathotype (Murguia et al., 2005; Toktay et al., 2005; Kwaye et al., 2008). Parasitises corn and goosegrass (Bernard et al., 2010). Recently described genus and species closely-related to economically important cyst nematodes. Vector of nepoviruses; Cherry Rasp Leaf, Tobacco Ringspot, Tomato Ringspot (Brown et al., 1993), Soybean Severe Stunt Virus (Evans et al., 2007) and Peach Rosette Mosaic Virus (Allen et al., 1984). Parasitises a wide range of crop and weed hosts (Miller, 1980), including stone fruits, soybean, cotton, sugarcane, rice, tobacco and maize (CABI crop protection compendium). Species complex. Acts as vector of Cowpea Mosaic Virus (Caveness et al., 1975). Parasitises pomegranate (Aly & Shaukat, 2005), tomato, eggplant (Babu & Muthukrishnan, 1990), citrus (Edward & Rai, 1970), coconut (Ekanayake & Lamberti, 1987) and a wide range of other hosts (Lehman, 1981b). Parasitises cocoa (Afolami & Caveness, 1983), ornamental and flowering plants (Costa et al., 2003), citrus (Crozzoli et al., 1998), peach, grape (Maximiniano et al., 1998) and potato (Lordello, 1951). Vector of Raspberry Ringspot Virus (Fritzsche & Kegler, 1968; Romanenko, 1970). Parasitises pomegranate (Zhou et al., 2007), citrus (Lehman, 1981b; Hu, 1991), peach and grape (Maximiniano et al., 1998). Vector of Tomato Ringspot Virus (Brown et al., 1994b; Jones et al., 1995). Parasitises grape (Graham et al., 1988) and apple (Ebsary et al., 1989). Tylenchorhynchus clarus Allen, 1955 Tylenchorhynchus claytoni Steiner, 1937 Tylenchorhynchus cylindricus Cobb, 1913 Tylenchorhynchus mashhoodi Siddiqi & Basir, 1959 ª 2013 The Authors. Journal compilation ª 2013 OEPP/EPPO, EPPO Bulletin 43, 334–374 Tylenchorhynchus nudus Allen, 1955 Tylenchulus palustris Inserra, Vovlas, O’Bannon & Esser, 1988 Tylenchulus semipenetrans† Cobb, 1913 Vittatidera zeaphila Bernard, Handoo, Powers, Donald & Heinz, 2010 Xiphinema americanum† Cobb, 1913 Xiphinema basiri Siddiqi, 1959 Xiphinema brasiliense Lordello, 1951 Xiphinema brevicolle Lordello & Da Costa, 1961 Xiphinema bricolensis Ebsary, Vrain & Graham, 1989 Regulated pest* Yes Yes Yes Yes Yes (continued) S. K. Singh et al. Species ª 2013 The Authors. Journal compilation ª 2013 OEPP/EPPO, EPPO Bulletin 43, 334–374 Table 2 (continued) Notes on main hosts and yield losses Xiphinema californicum Lamberti & Bleve-Zacheo, 1979 Vector of Tomato Ringspot, Tobacco Ringspot and Cherry Leaf Rasp Viruses (Hoy et al., 1984; Brown et al., 1993). Parasitises forest trees (Lownsbery & Lownsbery, 1985) and associated with wheat (Erum & Shahina, 2010). Potential threat to stone and pome fruits (Tacconi & Talame, 1995). Acts as vector of Arabic Mosaic Virus (Harrison & Winslow, 1961), Raspberry Ringspot Virus (Fritzsche & Kegler, 1968) and Strawberry Latent Ringspot Virus (Trudgill et al., 1981). Parasitises strawberry, raspberry, ryegrass, hops, barley and wheat (Flegg et al., 1970; Cotten, 1975; Griffiths et al., 1982). Parasitises tomato (Bleve-Zacheo et al., 1987), rice (Lamberti et al., 1987a, 1991), soybean (Lamberti et al., 1993), cocoa, citrus (Lamberti et al., 1992a), cowpea, eggplant, tomato, okra (Lamberti et al., 1992b), black pepper (Lamberti et al., 1983) and banana (Adiko, 1988). Vector of Grapevine Fanleaf Virus (Hewitt et al., 1958; Taylor & Raski, 1964). Parasitises a wide range of plants including grape, fig, ornamental plants, vegetables (Lehman, 1981b; Coiro et al., 1990; Arias & Fresno, 1994; van Zyl et al., 2012). Species complex. Pathogenic to grape (Lal et al., 1982) and coconut (Ekanayake & Lamberti, 1987). Parasitises lily bulbs (Saigusa & Yamamoto, 1971), fruit trees (Chen et al., 2004), Pennisetum purpureum (Fang, 1994), bonsai trees (Hirata & Yuhara, 1986), citrus (Zhou et al., 2007) and pomegranate (Hu, 1991). Vector of Grapevine Fanleaf Virus (Cohn et al., 1970; Dalmasso et al., 1972). Parasitises grape (Arias et al., 1994; Avgelis & Tzortzakakis, 2001), walnut (Ciancio et al., 1996), citrus (Edongali & El-Majberi, 1988) and fruit trees (KatalanGateva, 1980). Parasitises grape (Kumari, 2004; Gangl et al., 2009), apricot (Ivanova & Choleva, 1999), walnut (Liskova & Brown, 1998), strawberry (Samaliev & Mohamedova, 2011), cherry, cypress and fig (Koliopanos & Vovlas, 1977). Vector of Tobacco Ringspot, Tomato Ringspot, Cherry Leaf Rasp, and Peach Rosette Mosaic Virus (Forer et al., 1981; Brown et al., 1994b; Stobbs & Schagen, 1996; Sirca et al., 2007). Parasitises fruit trees (Georgi, 1988; van Driel et al., 1990; Islam et al., 1996; Gutierrez-Gutierrez et al., 2011). Parasitises citrus (1994). Associated with oak (Lamberti & Bleve-Zacheo, 1979). Belongs to the economically important Xiphinema americanum group (Gozel et al., 2006). Limited information is available on the biology and ecology of this species. Parasitises grape (Samota et al., 1994; Coiro et al., 2000), apricot (Ivanova & Choleva, 1999), walnut (Liskova & Brown, 1998), ornamental plants (Brzeski et al., 1978) and fruit trees (Choleva et al., 1984). Pathogenic to celery, carrot, pea (Vovlas et al., 1976) and Viola odorata (Ambrogioni & Rapetti, 1992). Parasitises a wide range of plants; faba bean, chickpea, lentils (Vovlas & Inserra, 1977; Di Vito et al., 1994; Troccoli & Di Vito, 2002), sugarbeet (Ebrahimi et al., 2004), lucerne (van den Berg, 1989), ornamental plants (Deimi et al., 2008), and forest trees (Talavera et al., 1999). Xiphinema diversicaudatum (Mikoletzky, 1927) Thorne, 1939 Xiphinema ifacolum Luc, 1961 Xiphinema index Thorne & Allen, 1950 Xiphinema insigne Loos, 1949 Xiphinema italiae Meyl, 1953 Xiphinema pachtaicum (Tulaganov, 1938) Kirjanova, 1951 Xiphinema rivesi Dalmasso, 1969 Xiphinema tarjanense Lamberti & Bleve-Zacheo, 1979 Xiphinema vuittenezi Luc, Lima, Weisher & Flegg, 1964 Zygotylenchus guevarai (Tobar Jimenez, 1963) Braun & Loof, 1966 Regulated pest* Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes *On an official list of regulated pests for at least one country globally. †See Discussion on phytosanitary importance of PPN with pathotypes/races and species complexes. ‡Hunt & Handoo (2009) consider that M. thamesi is a synonym of M. arenaria. However, measurements reported in Whitehead (1968) and Amin (1993) indicate differences from those of M. arenaria. M. thamesi also has different host reactions compared to M. arenaria. In the absence of more detailed published information supporting synonymising M. thamesi with M. arenaria, we retain M. thamesi. Nematodes of phytosanitary importance Species 349 350 S. K. Singh et al. Number of species 25 20 15 10 5 Meloidogyne Heterodera Pratylenchus Xiphinema Aphelenchoides Hirschmanniella Longidorus Ditylenchus Paratrichodorus Anguina Globodera Hoplolaimus Rotylenchulus Bursaphelenchus Helicotylenchus Punctodera Radopholus Scutellonema Trichodorus Aphasmatylenchus Belonolaimus Cactodera Dolichodorus Hemicriconemoides Hemicycliphora Macroposthonia Merlinius Nacobbus Quinisulcius Subanguina Tylenchorhynchus Tylenchulus Zygotylenchus 0 PPN genus Fig. 1 Number of regulated species per PPN genus. plants. Over a third (35%, N = 88) species satisfied all three criteria. The information from the sources in Table 1 varied in quality and consistency. The reasons for selecting particular species were not always explicitly stated. The CABI databases, EPPO PQR database, Society of Nematologists, University of Nebraska nematodes of quarantinable concern (See Table 1), were especially comprehensive information sources. Except for a few PPN species and countries worldwide (UK, EPPO countries, USA and Australia), detailed risk assessments leading to the quarantine listing of a species were not publicly available. In most other instances, the broad definition of a pest species under IPPC and ISPM were used to justify the quarantine status of a species. The quarantine listing of PPN species varied between countries. PPN species well known for their damaging impacts (including for example: Anguina tritici, Aphelenchoides besseyi, A. fragariae, Bursaphelenchus xylophilus, B. cocophilus, Ditylenchus destructor, D. dipsaci, Globodera pallida, G. rostochiensis, Heterodera glycines, H. schachtii, Nacobbus abberans, Radopholus similis, Xiphinema americanum and X. index [Source: Nematodes of quarantinable concern, Table 1)] were the most widely regulated. Other species, although damaging, are not regulated in many countries often because they are usually widespread. However, some countries regulate specific races e.g. of Meloidogyne incognita, M. javanica, M. arenaria which are not present in their territory. Distinguishing races is a major challenge for inclusion of races on list of regulated pests and further research is needed to develop rapid and specific methods for their distinction (see Discussion). International standards (ISPM) on pest risk analysis have been developed in the IPPC framework to determine the phytosanitary risks and biosecurity importance of species. The scarcity of publicly available documentation on pest risk analysis from countries worldwide and lack of transparency in the process leads to differences in how the phytosanitary importance of species are rated. For instance, some countries consider Bursaphelenchus mucronatus as of phytosanitary importance, but others do not. This is partially due to uncertainty on the whether the species can cause disease. Recent studies have demonstrated that the species can carry pathogenic bacteria (Zhao et al., 2009), hence it was included on our list under criterion 2. Discussion Importance of economic impact in assessing the phytosanitary risk of a nematode species A small percentage of the approximately 3400 known species of PPN (Hodda, 2011) are widespread and cause significant losses to crop production (Sasser, 1988; Koenning et al., 1999; Nicol et al., 2011). Of the remainder, their importance as plant pathogens is unknown; some species have limited distributions and cause localised damage to plants, and some species are recorded only once from their type host and locality. Hence, based on distribution, phytosanitary measures for PPN seem generally justifiable, depending on whether potential impacts outweigh costs. Assessment of potential impacts on economically-important crops is an important component in determining the phytosanitary importance of a pest species. However, yield loss estimates are available mainly for species whose pathogenicity or disease-causing abilities are already well known (Table 2). Data on the economic impacts of the majority of PPN remains sparse (Nicol et al., 2011). The available data can also be difficult to interpret and compare. Yield loss caused by PPN is often used to determine economic importance, but there are limitations in its calculation. Yield loss calculations from different studies and countries do not necessarily use the same methods, with some reporting damage as percentage yield loss and others reporting as tonnes per hectare or as percentage yield gained after application of nematicides or as correlations of yield gains with declining PPN abundance. Indirect losses ª 2013 The Authors. Journal compilation ª 2013 OEPP/EPPO, EPPO Bulletin 43, 334–374 Nematodes of phytosanitary importance (see below) also generally remain unaccounted for in most yield loss figures. Yield loss estimates based on work done long ago and under different nematode management regimes are also likely to require updating. Many frequently-used nematicides, such as Methyl Bromide, have been removed recently from the market (Noling & Becker, 1994) and alternate nematode management practices have been developed (Trudgill, 1991; Chitwood, 2002; Zasada et al., 2010a). PPN yield loss estimates from old publications may therefore need re-evaluation. In addition, PPN management practices differ between countries, depending on availability of nematicides, resistant varieties, and expertise in PPN management (Sasser & Krishnappa, 1980; Sharma, 1997; De Waele & Elsen, 2007). For example, in developing countries, nematicide use is limited and PPN yield loss estimates could be much higher compared to developed countries, which use nematicides widely to suppress crop losses (Sasser & Freckman, 1987; Nicol et al., 2011). The list of 250 PPN species in this paper, although representative of the major nematode genera, may not cover all species of phytosanitary importance. Some genera such as Meloidogyne, Heterodera and Pratylenchus are well represented in the list with over 20 species each (Table 2). All these genera are regarded as of economic importance (Evans et al., 1993; Luc et al., 2005; Castillo & Vovlas, 2007), and this is reflected in large numbers of publications on species from these genera. Lack of published information on the pathogenicity and economic importance of many less well-known species makes it difficult to predict the potential impacts of these species. Although over 100 valid species are known from genera such as Xiphinema, Tylenchorhynchus and Hemicycliophora (Siddiqi, 2000; Coomans et al., 2001) the economic importance of only a few species from these genera has been investigated. 351 effects of the nematode alone. Furthermore, the amount of damage may differ according to the nematode pathotype, population levels, crop species or cultivar, nematode/farm management practices, edaphic factors, and climatic conditions (Seinhorst, 1970; Lehman et al., 1971; Barker & Olthof, 1976; Green & Dennis, 1981; Noe & Barker, 1985; Oteifa, 1997; Schouten & Beniers, 1997). Damage symptoms and impacts are not always obviously associated with PPN, being frequently misidentified as due to drought, nutrient deficiency or other causes (Barker et al., 1994). There is also little data on the impact of PPN species on biodiversity. Even though there are provisions under ISPM and IPPC to assess the effects of a species on biodiversity, it is seldom included in assessment of phytosanitary importance of PPN. Therefore a wide range of potential damage and interactions of PPN with other disease-causing organisms were taken into account while compiling the list of species. Another reason for taking a broad approach is that apparently benign PPN can emerge as pests with changes in cropping patterns, nematode management, climate or arrival in new regions (Nicol et al., 2011). For instance, Meloidogyne enterolobii (syn. M. mayaguensis), is able to overcome the resistance of tomato and pepper genotypes carrying the Mi-1, N and Tabasco resistance genes widely used for nematode management (Brito et al., 2007; Kiewnick et al., 2009; Castagnone-Sereno, 2012). Hence, it has recently been added to the EPPO A2 list of pests recommended for regulation due to its pathogenicity and potential for spread in the EPPO region (EPPO, 2011). Globally few PPN species are currently treated as regulated non-quarantine pests, although regulated non-quarantine status could be an effective way of preventing further spread of recently established exotic PPN species. Nematode disease complexes and indirect impacts Assessing risks from nematode pathotypes and species complexes Impacts can also be difficult to estimate or severely underestimated because PPN may injure plants in many different ways. One way is by direct feeding action (Endo, 1975; Bridge & Starr, 2007), but this may take many different forms including direct damage, root galls, root stunting or withdrawal of resources from other parts of the plant (Norton & Niblack, 1991). Another way PPN injure plants is through disease complexes formed with other organisms, including other PPN (Powell, 1971; Sidhu & Webster, 1974; Davis & Webster, 2005). This type of damage may also take many forms. Species from the genera Longidorus, Paralongidorus, Paratrichodorus, Trichodorus and Xiphinema can act as vectors for several important plant viruses (Hewitt et al., 1958; Decraemer, 1995; Taylor & Brown, 1997). Root damage caused by migratory endoparasites from the genera Pratylenchus and Radopholus allows ingress of damaging rots. The effects on the plant of both organisms together in all these cases are greater than the Another aspect to consider when assessing phytosanitary status is intra-specific variation. This is often not considered because most phytosanitary regulation is on the taxonomic level of the species. However, species from the economically-important genera Belonolaimus, Ditylenchus, Globodera, Heterodera, Meloidogyne, Nacobbus, Radopholus, Rotylenchulus and Tylenchulus all have pathotypes or races with distinctive host responses and differences in host range (Abugharbieh & Perry, 1970; Michell et al., 1973; Gottlieb et al., 1986; Mojtahedi et al., 1988; Toktay et al., 2005; Anthoine & Mugniery, 2006; Sturhan et al., 2008; Robertson et al., 2009). Under the IPCC, absence of pathotypes in a country is justification for implementing quarantine measures against exotic pathotypes [FAO, 2011c; EFSA (Panel on Plant Health), 2012]. For example, only the sugarbeet pathotype of Nacobbus aberrans is present in the USA, while the potato pathotype is absent, and hence the potato pathotype is a regulated pest in the USA (Inserra et al., ª 2013 The Authors. Journal compilation ª 2013 OEPP/EPPO, EPPO Bulletin 43, 334–374 352 S. K. Singh et al. 2004). The known highly damaging species such as Globodera pallida, G. rostochiensis, Heterodera avenae and Ditylenchus dipsaci which have numerous pathotypes therefore may require further phytosanitary risk assessment specific to pathotypes to prevent the spread of exotic pathotypes and justify regulatory measures. It is not known how widespread pathotype variability is in the many less well investigated PPN, but if it is as common in the best-known species, then this is another reason to take the broadest approach for assessing phytosanitary risk from PPN. The recent description of the new species Ditylenchus gigas and D. weischeri, previously considered as part of D. dipsaci species complex (Chizhov et al., 2010; Vovlas et al., 2011), highlight the importance of research into taxonomy and specific identification methods, especially for species complexes and races. With more specific identification methods, the distributions and phytosanitary importance of closely related or cryptic species and races could be assessed more precisely. Synonymising species names can also have important implications on the phytosanitary status of a species. For example, until recently M. enterolobii and M. mayaguensis were considered separate species, but their recent synonymization meant that when distribution, host range and other information published under both names were consolidated, the apparent risk increased substantially (see CastagnoneSereno, 2012; Karssen et al., 2012). Therefore information published under species synonyms also needs to be considered when assessing phytosanitary importance. Biosecurity implications Despite these limitations, making the best systematic predictions of impact and risk possible, based on as much data as can be obtained is preferable to the alternative of empirical measurements of actual damage following real introductions of potentially damaging species. Where real introductions of exotic PPN have occurred and been measured, the impacts have mostly been substantial. Of course, by this stage, eradication is seldom an option (Hodda et al., 2008). More recently PPN species have been used as bioindicators during quarantine inspections. In the UK, Helicotylenchus dihystera is often intercepted with planting materials supposedly grown in sterile conditions, indicating that phytosanitary standards were not met (Hockland & Anderson, 2012). Other microscopic plant pathogens such as fungi, bacteria and viruses may also use similar pathways to PPN (Grousset et al., 2012), so targeting the generally larger PPN during quarantine inspections has assisted in reducing the risks from other microscopic quarantine organisms generally. Phytosanitary risks are specific to countries or regions. Regulated organisms differ between countries depending on species distributions and regulatory or biosecurity policy in different countries. Formal pest risk analysis processes include multiple stages (initiation, pest categorization, risk assessment and risk management) and so require considerable time and resources to scientifically assess and determine the regulatory measures commensurate with the risks posed by each species (Petter et al., 2010). Yet, it is important that this process is followed because the regulatory status of a species has important trade implications: countries can impose trade restrictions if certain species are present in the exporting country (Gebrehiwet et al., 2007; Cook et al., 2011). Conclusions The authors hope that the list provided in this paper will assist assessment of the risks from species (including pathotypes and races) in at least the first, initiation stage of the pest risk analysis process globally. It may assist in deciding on which species to build diagnostic and detection capacity. The systematic list should be useful in providing preliminary information and guidance to nematologists in many countries assessing the risks from PPN. The authors hope that the criteria presented for assessment also facilitate pest risk analysis and prompt appropriate gathering of additional data, thus enhancing nematode biosecurity globally. Acknowledgements The authors would like to thank D Paini and J Roberts (CSIRO Ecosystem Sciences) for reading and commenting on the manuscript. The authors and the editorial team of the EPPO Bulletin would like to thank S Hockland, Emeritus Fellow of the Food and Environment Research Agency and Independent Consultant in Plant Nematology, for her review of the paper. The authors also thank the editorial team of the EPPO Bulletin for their helpful comments. The authors acknowledge the support of the Australian Government’s Cooperative Research Centres Program. matodes parasites des plantes Les ne ^ tes d’importance phytosantaire, leurs ho es principaux, et les pertes de recolte releve L’importance phytosanitaire potentielle de l’ensemble des especes de nematodes parasites des vegetaux a ete determinee en evaluant les informations disponibles sur les caracteristiques de chacune des especes, leur association avec des cultures economiquement importantes, et leur capacite a ^etre vecteurs de virus ou de former des complexes de maladies avec d’autres agents pathogenes. La plupart des especes de nematodes parasites des plantes decrites sont mal connues, ont ete observees en un seul endroit, ne sont pas associes avec des cultures economiquement importantes, et ne sont pas connues pour ^etre associees avec d’autres organismes phytopathogenes. Cependant, 250 especes appartenant a 43 genres ont satisfait au moins un des criteres a prendre en compte pour ª 2013 The Authors. Journal compilation ª 2013 OEPP/EPPO, EPPO Bulletin 43, 334–374 Nematodes of phytosanitary importance ^etre considere comme presentant un risque phytosanitaire. Les genres et les nombres d’especes (entre parentheses) consideres comme presentant un risque phytosanitaire comprenaient: Achlysiella (1), Anguina (8), Aphasmatylenchus (1), Aphelenchoides (12), Aphelenchus (1), Belonolaimus (2), Bitylenchus (3), Bursaphelenchus (4), Cactodera (3), Ditylenchus (8), Dolichodorus (1), Globodera (3), Helicotylenchus (7), Hemicriconemoides (3), Hemicycliophora (3), Heterodera (25), Hirschmanniella (5), Hoplolaimus (5), Ibipora (3), Longidorus (10), Macroposthonia (2), Meloidogyne (38), Merlinius (3), Nacobbus (1), Neodolichodorus (2), Paralongidorus (2), Paratrichodorus (11), Paratylenchus (3), Pratylenchus (24), Punctodera (3), Quinisulcius (3), Radopholus (5), Rotylenchulus (3), Rotylenchus (1), Scutellonema (5), Sphaeronema (1), Subanguina (3), Trichodorus (5), Tylenchorhynchus (8), Tylenchulus (2), Vittatidera (1), Xiphinema (15), and Zygotylenchus (1). Pour chacune des 250 especes, les h^otes principaux et des estimations de pertes de rendement sont fournies avec une bibliographie detaillee. Parmi les 250 especes, seulement 126 especes parmi 33 genres sont actuellement listees comme des organismes reglementes dans un ou plusieurs pays dans le monde. Presque toutes ces 250 especes etaient associees a des cultures economiquement importantes, et certaines agissaient egalement comme vecteur pour des virus. Gapapbnbpy⁄obe ya pacneybzx yevanols, bve⁄obe goneywbakmyoe pyaxeybe lkz abnocaybnapbb, bx ocyodyse pacneybz-xopzeda b coo,oaevse gonepb ypo;aqyocnb Goneywbakmyoe pyaxeybe lkz abnocaybnapbb ra;louo bp yrapayysx dblod yevanol ogpelekzkocm gynev oweyrb cyoecndy⁄obx layysx go xaparnepbcnbrav dbla, codvecnbvocnb c 'royovbxecrb pyaxbvsvb gonepzvb ypo;aqyocnb b bx cgoco,yocnb leqcndodanm d raxecnde gepeyocxbrod dbpycod bkb ,okepyendopysx rovgkercod d coxenaybb c lpyubvb ganoueyavb. Xaoe dceuo wbnbpyevse dbls gapapbnbpy⁄obx ya pacneybzx yevanol (PPN) gkoxo bpyxeys, papeubcnpbpodays oyb kbim d olyoq noxre, ye cdzpays c 'royovbxecrb pyaxbvsvb rykmnypavb b ye bpdecnyo, coxena⁄ncz kb oyb c lpyubvb ,okepyendopysvb dpelysvb opuaybpvavb pacneybq. Olyaro 250 dblod bp 43 polod ondexakb olyovy bkb yecrokmrbv rpbnepbzv, ronopse xpedans abnocaybnapysv pbcrov. Pols b xbcko dblod (d rpyuksx cro,rax), ronopse cxbna⁄ncz rar gpelcnadkz⁄obe abnocaybnapysq pbcr gpbdolzncz: Achlysiella (1), Anguina (8), Aphasmatylenchus (1), Aphelenchoides (12), Aphelenchus (1), Belonolaimus (2), Bitylenchus (3), Bursaphelenchus (4), Cactodera (3), Ditylenchus (8), Dolichodorus (1), Globodera (3), Helicotylenchus (7), Hemicriconemoides (3), Hemicycliophora (3), Heterodera (25), Hirschmanniella (5), Hoplolaimus (5), Ibipora (3), Longidorus (10), Macroposthonia (2), Meloidogyne (38), Merlinius (3), 353 Nacobbus (1), Neodolichodorus (2), Paralongidorus (2), Paratrichodorus (11), Paratylenchus (3), Pratylenchus (24), Punctodera (3), Quinisulcius (3), Radopholus (5), Rotylenchulus (3), Rotylenchus (1), Scutellonema (5), Sphaeronema (1), Subanguina (3), Trichodorus (5), Tylenchorhynchus (8), Tylenchulus (2), Vittatidera (1), Xiphinema (15), a nar;e Zygotylenchus (1). 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