J. Crop Prot. 2019, 8 (3): 243-257______

Review Article The damage potential of pin , Micoletzky, 1922 sensu lato spp. (Nematoda: )

Reza Ghaderi

Department of Plant Protection, School of Agriculture, Shiraz University, Shiraz, Iran.

Abstract: The genus Paratylenchus sensu lato includes members belonging to the genera Paratylenchus sensustricto (species with 10 to 40µm long stylet), (species with 40-120µm long stylet), Gracilpaurus (species having cuticular punctuations) and Paratylenchoides (species having sclerotized cephalic framework). Long stylet species become swollen and feed as sedentary parasites of roots, some feed from cortex of perennial host roots, but most species feed as sedentary ectoparasites on roots. In other words, species with stylet shorter than 40µm commonly feed on epidermal cells, whilst the species with longer stylet nourish primarily in cortical tissue, without penetration into the plant tissue. In general, pin nematodes, Paratylenchus spp. are parasites of higher plants with a higher abundance in the rhizosphere of trees and perennials. In present review, an attempt is made to document published information on the pathogenicity and damage potential of the pin nematodes to plants.

Keywords: Gracilacus, damage, pathogenicity, perennials, pin nematodes, population, trees

Introduction12 Lisetskaya, 1963; 1965; Braun et al., 1966; Fisher, 1967; Ghaderi and Karegar, 2013), and The pin nematodes, Paratylenchus Micoletzky, in some nurseries of conifers, the density of

Downloaded from jcp.modares.ac.ir at 5:11 IRST on Sunday October 3rd 2021 1922 sensu lato, firstly have long been population was increased to more than 1000 considered as free-living nematodes, but further individuals per 100cm3 of soil (Ruehle, 1967; studies on their life cycle led researchers to find Rossner, 1969). Distribution and host evidence on their damage to plants (Solov’eva, preference of Paratylenchus species has already 1975). Paratylenchus species seem to be a been reviewed a few times (Loof, 1975; common component of the fauna of cultivated Brzeski, 1976; Bell and Watson, 2001; crops, plantations and natural vegetation Eroshenko and Volkova, 2005; Čermák and (Solov’eva, 1975), with a higher abundance in Renčo, 2010; Ghaderi et al., 2016). perennial plants, such as grass stands, hop These nematodes are described as a group gardens, orchards or forest trees and shelterbelts responsive to environmental fluctuations and (Čermák and Renčo, 2010). Large number of root development. Their shorter generation time these nematodes are common in the rhizosphere and smaller body size compared to other of fruit trees (Weischer, 1960; Nesterov and groups, allow faster buildup of their populations after environmental changes (Yeates and Lee, 1997). In addition, they are Handling Editor: Zahra Tanha Maaifie more resistant to dehydration and easily ______* dispersed by wind (Gaur, 1988). Usually Corresponding author, e-mail: [email protected] Received: 2 September 2018, Accepted: 26 February 2019 fourth-stage and sometimes third-stage Published online: 22 April 2019

243 The damage potential of pin nematodes ______J. Crop Prot.

juveniles serve as resistant (or resting) stage of reviewed. Associated plants and hosts are Paratylenchus species which they may have a mentioned for each species, and symptoms of well-developed stylet, reduced stylet or no the nematode infection are discussed as stylet (Brzeski and Háněl, 1999). Additional described and reported in literature. However, it effects of different climatic factors on should be noted that information on population of these nematodes have been pathogenicity is available only for few species, discussed earlier (Reuver, 1959; Fisher, 1965; which are discussed in alphabetical order as 1967; Wu and Hown, 1975; Brzeski, 1991; Bell follows. and Watson, 2001). Paratylenchus bukowinensis Micoletzky, 1922 Feeding behavior of pin nematodes Brzeski (1971) observed P. Bukowinensis did The feeding behavior somewhat differs among not cause visible injuries to cabbage roots; Paratylenchus species. It seems that long stylet however, the fresh weight of roots was reduced bearing species become swollen as sedentary markedly, and that of aerial parts was decreased feeders, some of them feed from deeper layers in slightly. Although P. bukowinensis may cause cortex of perennial host roots; most others are yield decrease of cabbage, this species seems ectoparasites on roots (Ghaderi et al., 2016). In most injurious to root crops of Apiaceae. In pot other words, species with stylet shorter than experiments (Brzeski, 1976), hosts of P. 40µm commonly feed on epidermal cells and bukowinensis were found in the families root hairs (Lindford et al., 1949; Rhoades and Brassicaceae and Apiaceae including carrot, Linford, 1961; Brzeski et al., 1975; Wang et al., , parsley, cabbage and rutabaga. No hosts 2016), but species with longer style tend to feed were found among examined species of primarily on cortical tissue (Inserra and Vovlas, Solanaceae, Papillionaceae, Asteraceae, 1977; Cid del prado Vera and Maggenti, 1988; Chenopodiaceae or Poaceae. Troccoli et al., 2002; Inserra et al., 2003). The Parsley and carrot roots were misshapen, long and robust stylet enables these species to shorter or forked (Weischer, 1961) and penetrate several cells deep and become finally, the whole plant may be completely permanently attached to the root surface in a destroyed (Brzeski, 1976). Celery roots sedentary manner without penetration of their developed many lateral roots and some body into the root tissues (Inserra and Vovlas, necrosis appeared which might eventually

Downloaded from jcp.modares.ac.ir at 5:11 IRST on Sunday October 3rd 2021 1977).The long stylet also enable spin nematodes destroy the whole root system. The tolerance to parasitize deep root tissues of trees and shrubs, limit of celery was about 70 nematodes in but they can change their host preference from 100cm3 of soil and the minimum yield was woody species to several grasses and herbs with about 60% of that of non-infested treatments changes in the soil ecosystems and nutrient (Brzeski, 1975). cycling (Čermák and Renčo, 2010). Feeding Brzeski (1976) noted that females, second behavior of Paratylenchus spp. has been studied and third stage juveniles (J2s and J3s) of P. in more detail by some researchers (Linford, bukowinensis, the only stages that feed, are 1942; Linford et al., 1949; Rhoades and Linford, found feeding mainly on the epidermal cells, 1961; Brzeski et al., 1975; Inserra and Vovlas, and sometimes on cells two layers deeper in the 1977; cid del prado Vera and Maggenti, 1988; root parenchyma of parsley and cabbage. The Troccoli et al., 2002; Inserra et al., 2003; Wang fourth stage juveniles (J4s), have thin, short et al., 2016). stylet and a reduced pharynx and do not feed. The root diffusates stimulate the molting of J4s Damage potential, host range and symptoms to adults; although some molting occurs in the of pin nematodes infection spring in the absence of root exudates. In the present paper, the damage potential and Population increase of 700% in one season on pathogenicity of pin nematodes to plants is parsley has been recorded.

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Paratylenchus dianthus Jenkins & Taylor, walnut roots appear to be immune to the closely 1956 related genus, Cacopaurus pestis. Thorne (1943) P. dianthus has been reported to retard growth was unable to find any evidence that C. pestis of carnation (Dianthus caryophyllus L.) in USA could attack the roots of black walnut. However, (Jenkins and Taylor, 1956). Root and soil it remains to be determined if P. epacris could samples were collected in three commercial attack English walnut roots (Allen and Jensen, greenhouses from carnations exhibiting poor 1950). growth. Each of four pots containing three plants were inoculated with approximately one Thorne & Allen, 1950 thousand individuals of this species. After 50 P. hamatus has been reported as a contributing days, examination of the soil and roots from factor to, if not the primary cause of, the fig tree each pot revealed a population increase of 700 decline (leaf drop) in some fig Ficus carica L. percent. The nematodes inoculated into fallow orchards in California, USA. In the affected pots failed to survive, indicating the host trees, the first observed symptom is the lighter preference of the species. It seems probable that color of the leaves, which gradually becomes this species was one of the contributing factors more pronounced until the leaves die and fall. to poor growth in the three greenhouses Fruits on these trees are undersized and sampled. generally fall along with the leaves. A slow In another study, P. dianthus was decline of the entire tree takes place, determined to be an important pest of carnation culminating the dieback of twigs and small in the Naples province, Italy. Fumigation prior limbs. However, conditions in the observed to planting prevented damage until the second orchards may have been aggravated by growing year when nematode populations built inadequate irrigation during the hot summer up to damaging levels (Pennacchio et al., 1985). months. Populations of P. hamatus in the Under greenhouse conditions and during a105- infected orchards ranged from 6 to 3000 per days period, P. dianthus increased from 100 to 400 gram of soil. Many specimens of P. 37000 with carnation, to 1509000 with celery, hamatus were observed attached by their stylets and to 820000 with jasmine tobacco as host to the rootlets (Thorne and Allen, 1950). Some plants (Rhodes and Linford, 1961). years later, the deterioration of the condition of pear orchards in California was correlated with

Downloaded from jcp.modares.ac.ir at 5:11 IRST on Sunday October 3rd 2021 Paratylenchus epacris (Allen & Jensen, 1950) the concomitant infection of P. hamatus and Goodey, 1963 pythiaceous fungi (French et al., 1964). This P. epacris was found attacking black walnut nematode was found in 116 out of the 121 roots in California, USA. Although trees were orchards investigated, and in 85 orchards large also infected with a root-lesion nematode, and an populations were found. The density reached up accurate estimation of the damage was not to 2500 individuals in 250cm3 of soil. However, possible, but some evidences indicated that P. the number of Paratylenchus in the roots was epacris might contribute to the disease very low and only in one case, 143 individuals symptoms observed in the infected trees. were found in one gram of roots. Numerous colonies of adult females and P. hamatus was also found to be associated juveniles were observed with their stylet with 60 declined citrus trees in Shiraz, Iran imbedded in the root tissues, and eggs were (Abivardi, 1970). Populations from various observed attached to the debris usually present samples ranged from 225 to over 300 nematodes around the colonies, and males were obtained by per 100cc soil, but no direct evidence of feeding washing infested soil through a series of graded on the roots of sour orange and lime seedlings screens and from scrapings made from the bark was observed under the microscope. However, of infected roots (Allen and Jensen, 1950). This second stage juveniles (J2s) of the citrus is especially interesting, considering that black nematode were also present in higher

245 The damage potential of pin nematodes ______J. Crop Prot.

populations, which ranged from 420 to 650 Experiments indicated that the nematode nematodes per 100cc soil, thus the observed population increases rapidly up to the beginning decline cannot be attributed only to P. hamatus. of flowering, it then decreases during It has also been shown that high populations of flowering, and increases again after flowering, P. hamatus can reduce flower production and reaching up to 100000 individuals on a single quality of roses (MacDonald, 1976). This species plant. At the end of the experiment, with the seems to be also damaging to grapevine orchards decrease in the fresh\dry weight of plant, (Raski and Lider, 1959; Philis, 2003). In an population of the nematode was considerably established vineyard, the nematicides cadusafos reduced. It is presumed that P. hamatus reduces and carbofuran controlled Xiphinema index the yield of peppermint and adversely affects Thorne & Allen, 1950, Mesocriconemax enoplax the quality of the essential volatile oil (Raski, 1952) Loof & de Grisse, 1989 and P. (Faulkner, 1964). In the USSR, P. hamatus was hamatus, and subsequently increased yields also found constantly in large numbers (up to (Philis, 2003); however, the exact role of P. 1000 individuals in 15 gram of soil) in the hamatus cannot be determined as the observed subsoil of peppermint fields in Moldavia yield loss may have been related to the other two (Lisetskaya, 1968; 1969). nematodes which are considered as important parasites of grapevine in literature. Andrassy, 1959 There are also cases in which P. hamatus Andrassy (1985) observed a heavy infestation has considerably reduced the yield of vegetable of P. microdorus in Hungary which could delay crops. According to Lownsbery et al. (1952), growth of red clover and lettuce. The leaves of this nematode severely infected celery in affected plants were smaller and their lateral Connecticut, USA, and under greenhouse roots were fewer in number than healthy plants. conditions it showed marked pathogenicity. Some other reports further proved the damage Methyl bromide treatment of soils infested with of this species to monocotyledons, especially this nematode increased the weight of pot- grasses). Brzeski (1998) reported P. microdorus grown celery by four times when compared as a common species in meadows, sometimes with the control. Rich (1955) reported P. also found in the rhizosphere of corn plants. He hamatus attacking the roots of celery in New suggested that Poacae are probably the main Hampshire, USA and causing severe stunting host. Ciobanu et al. (2003) also noted that this

Downloaded from jcp.modares.ac.ir at 5:11 IRST on Sunday October 3rd 2021 and chlorosis. In the Federal Republic of species prefers grasses. Talavera and Navas Germany, this species caused heavy damage to (2002) found some Paratylenchus species (P. carrots (Weischcr, 1957). According to the microdorus, P. similis, P. nanus and P. results, the critical number of P. hamatus in ciccaronei) were the most abundant (98% 500cm3 of soil, causing damage to carrot, was frequency) and prevalent (average 146 3000 to 4000 nematodes. The extent of damage nematodes per 100cm3 soil) plant-parasitic depended also on the phase of development of nematodes associated with pastures and the plants; and younger plants were damaged grassland s in southern Spain. They also noted more readily than older ones. At the end of the that P. microdorus populations were maintained vegetative period, even a few thousand s of or increased by all grasses or legumes tested in Paratylenchus cannot cause perceivable a pot experiment. They concluded that damage to carrot (Weischer, 1964). In the Paratylenchus spp. well exploit the summer Wisconsin State, USA, P. hamatus was found drought in semi-arid grassland s for their to be a parasite of peppermint (Faulkner, 1964). survival in better competition with other In highly infested fields, the plant growth was nematodes and thus, they are predominant in retarded, flowering delayed, the root system the region. Dominance of Paratylenchus in dry was weaker than that of the healthy plants and l and pastures has also been reported by Yeates the plants showed symptoms of withering. (1984) and Nombela et al. (1999).

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Paratylenchus minutus Linford in Linford, bluegrass Poa annua L. was considered as a Oliveira & Ishii, 1949 poor host and the two C4 grasses, paspalum It seems that pineapple plantation environment Paspalum dilatatum Poir. and summer grass is especially favorable to P. minutus (Linford, Digitarias anguinalis (L.) Scop. were non host 1942; Linford et al., 1949). Linford (1942) plants. There was no significant difference in found large numbers of this nematode attached the proportion of life stages between good and to pineapple rootlets. Furthermore, Linford et poor hosts of the nematode. Furthermore, they al. (1949) observed that the old plantations of considered apple, cherry, grapevine, potato, pineapple in Hawaiian region frequently carrot, celery, corn and also 35 woody and contained 100 to over 900 nematodes per gram bushy plants as other hosts of this species. of soil, and up to 23800 nematodes per gram of Brzeski (1998) also reported P. nanusas a root. They also stated that P. minutus might be common species in meadow soils, and the able to feed and reproduce on roots of 24 other rhizosphere of cereals. However, Ciobanu et al. plants including weeds, crop plants, and (2003) found this species in forests on brown ornamentals, when they were grown in acid soils located at high altitudes and miniature root-observation boxes of infested therefore, they suggested that this species is not soil. Although the lack of visible pathological restricted to lowland habitats and grass symptoms in cells fed on, and the occurrence of vegetation. large populations on roots of apparently normal The effects of the host on reproduction or pineapple plants, tend to indicate that P. morphometric characters of P. nanus has been minutusis non-pathogenic, the authors finally studied in few works. In an infection of the noted that such a conclusion would be garden balsam Impatiens balsamina L. by 4000 premature and needs further confirmations. individuals, the number of nematodes increased by 23 times during two months (Odihirin and Paratylenchus nanus Cobb, 1923 Jenkins, 1965). The symptoms of infection on Populations of P. nanus were for the first time plants were: growth arrest, yellowing, late recovered from the rhizosphere of apparently flowering and reduction in the weight of shoots healthy roots of the Californian laurel tree (by 19 to 30%) and roots (by 2.3 times). In the (Cobb, 1923) and the necrotic parts of the roots another study (Fisher, 1965), approximately of elegant zinnia (Steiner, 1924). Raski (1975) 1000 adults and fourth-stage juveniles of this

Downloaded from jcp.modares.ac.ir at 5:11 IRST on Sunday October 3rd 2021 listed alfalfa and several other plants as possible species were added to each pot with apple and hosts from the USA and Canada. Corbett (1978) apricot seedlings as host plants and allowed to stated that it is possible to find an enormous develop for four months. The two hosts had no population up to 250 thousand of P. nanus per a effect on the morphometric characters of litre of soil in the rhizosphere of perennial females but apricot seedlings allowed the plants, particularly orchards. It appears that development of longer males. grasses are good hosts of P. nanus, as confirmed by Viketoft et al. (2005) and Paratylenchus neoamblycephalus Geraert, Viketoft (2008) who found orchard grass 1965 Dactylis glomerata L. and timothy-grass There is a well-documented and helpful piece Phleum pretense L. as the best hosts for this of information on the pathogenicity of P. species. In glasshouse tests of 15 pasture plants, neoamblycephalus by Braun and Lownsberry common in New Zeal and, Bell and Watson (1975). They indicated that elimination of the (2001) indicated that all good hosts nematode from soil by fumigation with 1,2- (reproduction factor > 1) of P. nanus were dibromoethane, stimulated the growth of grasses, namely orchard grass D. glomerata, Myrobalan plum seedlings. Addition of a Italian ryegrass Lolium multiflorum Lam. and suspension of P. neoamblycephalus to perineal ryegrass L. perenne L.. Annual Myrobalan seedlings inhibited their growth

247 The damage potential of pin nematodes ______J. Crop Prot.

compared to non-inoculated control treatments. root systems were unusually clean and white, Roots of Myrobalan seedlings inoculated with with less developed lateral roots, and maximum the nematode were smaller, darker, and had population in pots reached about 40000 fewer feeder roots compared to those of control nematodes per root system. The symptoms on plants. Nematodes were observed feeding tall fescue were slight stunting, increase in root ectoparasitically, but their head was embedded weight and the number of tillers. In agreement in root cortex. They were associated with small with the suitability of tobacco as a host for this lesions and dead lateral roots. Clusters of species, Loof (1975) noted that in the field nematodes were common at ruptures in the where the type population of P. projectus was epidermis, and where lateral roots emerged. found, tobacco culture had been abandoned Inhibition of Myrobalan growth by P. because of low yields. In another study (Olthof, neoamblycephalus was greater at 20 and 27 ºC 1979), common bean was recommended for than at 30 ºC, and was not affected by pH over rearing large numbers of P. projectus under the range 4.5 to 6.5. Rose, apricot, , and greenhouse condition. all varieties and hybrids of cherry (Prunus Smolik et al. (1983) indicated that the most cerasifera Ehrh.) tested, were hosts for this dominant species of nematodes in sunflower species. None of the herbaceous plants fields of South Dakota, USA, was P. projectus; examined were hosts for the nematode, and late-season populations frequently ranged from some trees (e. g., walnut and fig) were either 1500 to 4000 per 100cm3 of soil. Smolik and non or very poor hosts. However, the authors Walgenbach (1984) noticed that sunflower did not obtain any success in culturing the appears to be an excellent host for P. projectus, nematode on various herbaceous plants or and it is probable that P. projectus control was Myrobalan callus tissue. Reuver (1959), Geraert in part responsible for the yield increases (1965) and Fisher (1967) reported apple as a observed in the nematicide application host for P. neoamblycephalus in Europe and treatment. However, Smolik (1987) concluded Australia, but Braun and Lownsberry (1975) that P. projectus is was only a mild parasite of noted that apple does not seem to be a host for sunflower in his greenhouse study, and large Californian population of P. neoamblycephalus. populations would be necessary for substantial They argued that this may indicate the existence plant growth reduction. P. projectus of races for this species. significantly (P < 0.05) reduced sunflower seed

Downloaded from jcp.modares.ac.ir at 5:11 IRST on Sunday October 3rd 2021 yields in this study. Yield reductions occurred Jenkins, 1956 in both fertilized and unfertilized treatments The pathogenicity of this species has been and ranged from 12 to 33%. Application of investigated more than any other species of the fertilizer did not affect P. projectus damage to pin nematodes. A large number of studies were sunflower. Populations of the nematode conducted on grasses and legumes, although increased 20 to 126 fold over 14 weeks. information is available on the pathogenicity of Population increase of the nematode on P. projectus to some other plants including sunflower was highest at 20 and 25 ºC, and tobacco, bean and sunflower. Some of the populations did not increase above initial conducted researches are reviewed in Loof inoculum levels at 10, 15, or 35 ºC. Both seed (1975). yield and final populations of P. projectus were Coursen et al. (1958) provided a list of 10 significantly (P < 0.01) greater in the fertilized non-hosts and 42 host species. Coursen and treatments. He also noted that early planting Jenkins (1958) carried out pot experiments with (mid-April to early May) of sunflower may tobacco and tall fescue Festuca elatior L. They reduce P. projectus damage to this crop found that inoculated tobacco plants were (Smolik, 1987). shorter than the control plants, and had stunted Excluding the above-mentioned studies, top growth and reduced internode length. The other investigations are mainly focused on the

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effects of P. projectus on legumes or grasses. grass Lolium multiflorum Lam. were among Rhoades and Linford (1961) inoculated 100 some of the best hosts of P. projectus, but Paratylenchus projectus of mixed stages to 4- Millet Panicum capillare L. and rice cutgrass inch pots in which various plants were growing Leersia oryzoides (L.) Sw. were among the in sterilized potting mixture. After 105 days, the poorest hosts. Among the legumes, birdsfoot number of nematodes were 38000 with red trefoil Lotus corniculatus L. was the best host, clover Trifolium pretense L., 147000 with and alfalfa Medicago sativa L., and red clover timothy-grass Phleum pretense L., 181000 with Trifolium pratense L. were the poorest hosts. celery Apium graveolens var. dulce pers., and Although P. projectus appears to multiply on 2637000 with jasmine tobacco Nicotiana many plants, the best grass hosts supported alatavar.gr and iflora, Link and Otto. In North higher nematode populations than the best Carolina, McGlohon et al. (1961) found that P. legumes (Townshend and Davidson, 1989). projectus causes significant decrease in top In a pot experiment, populations of P. weight of a legume, Lespedeza stipulacea projectus became established on 10 grasses and Maxim., and damages the root system severely, two legumes (Wood, 1973).Reproduction but yield reduction was not observed in this occurred on all of the hosts examined. Non- poor host. Shesteperov (1971) observed that in feeding, preadult fourth stage juveniles the Moscow region of the USSR a complex of comprised about 50% of most populations. In plant-parasitic nematodes, among them P. another greenhouse study (Sohlenius et al., projectus, delays growth and development of 2011), P. projectus increased with time in all red clover. In addition, the number of plants and treatments except for timothy-grass Phleum of the leaves per plant were reduced. pretense L., Alsike clover Trifolium hybridum Furthermore, the leaves were reduced in size L. and the control treatments. The nematode and dry weight and winter hardiness were increased in several treatments, including white decreased. The susceptibility to pathogenic clover T. repens L., where it reached extremely organisms was also increased. high numbers. Contrary to this, it was almost In his greenhouse experiments, Townshend totally absent under T. hybridum treatment. (1972) found an increase from 37 to 3200 However, it has already been shown that the nematodes per 25 gram of soil under timothy- nematode population increased markedly in the grass Phleum pretense var. Climax, and from grass plots in a Swedish study of arable l and

Downloaded from jcp.modares.ac.ir at 5:11 IRST on Sunday October 3rd 2021 67 to 866 under trefoil Lotus corniculatus var. (Sohlenius et al., 1987). Ina series of grassland Viking after seven months. Townshend and s of different ages, Wasilewska (1997) found a Potter (1976) found that legumes are good hosts very high abundance of P. projectus in of P. projectus with the exception of alfalfa that permanent grassland with an age of more than was a poor host. Among grasses, timothy-grass 20 years. Korthals et al. (2001) also found an Phleum pretense L. was a good host, orchard increased number of this species in plots grass D. glomerata L. a fair host, and brome changed from monoculture field crops into grass Bromus inermis Leyss. a poor host. Oat more permanent or highly diverse plant and rye were good hosts among the cereals, communities. barley and wheat were fair hosts, and corn was It appears that P. projectus can be a poor host. In other experiment, Townshend et considered as an important plant-parasitic al. (1973) indicated that P. projectus increased nematode of forage fields in Canada. Webster to much greater numbers under forage crops in et al. (1972) noticed that the occurrence of P. the greenhouse than are normally found in the projectus appears to be associated with a field. Among the grasses, squirrel-tail grass widespread disease of alfalfa in in Alberta, Hordeum jubatum L., Echinochloa pungens var. Canada, called "alfalfa sickness". Affected Wieg and ii, barnyard grass E. crusgalli L., plants were stunted, spindly, yellowish-green in quaking grass Briza maxima L. and Italian rye- color, and poorly nodulated. Amending soil

249 The damage potential of pin nematodes ______J. Crop Prot.

with macronutrients (NPK) and micronutrients the cell walls and middle lamellae were did not significantly improve the growth of partially dissolved because of nematode affected plants. Beneficial effects with treating migration and feeding. "sick" soils with steam and with Metham sodium have been reported. Examined alfalfa Other species plants and soil adhering to their root systems Some information is available on the damage consistently revealed higher counts of potential of other Paratylenchus species, Paratylenchus in soils from areas of poor although usually there is no direct evidence on growth compared to areas of good growth. the pathogenicity or damage level of these Counts of Paratylenchus varied from zero to nematodes to the associated plants. High more than 7000 per kg soil. Twenty-three populations of P. besokianus Bally & Reydon, percent of the soils contained more than 4000 1931 were recovered in the necrotic parts of the Paratylenchus per kg of soil (Webster et al., roots of coffee tree (Bally and Reydon, 1931). 1972). In another study in Alberta, Canada Boag (1974) observed high population densities (Webster and Hown, 1973), sampled locations of an unknown species of Paratylenchus in the in the Peace River had low numbers of P. Quercus spp. rhizosphere in Scotland. Wu and projectus with only 25% of the samples Hown (1975) noticed that rhubarb Rheum showing greater than 1000 and only 4% with rhabarbarum L. plants grown in a small pot, more than 10000 nematodes per kg of soil. In containing soil infested with P. neoprojectus contrast, an area in central Alberta had 56% of Wu & Hown, 1975, were not vigorous and the counts greater than 1000 and 20% greater appeared to become unhealthy as they grew than 10000 per kg of soil. In two other surveys older. Furthermore, the nematode population conducted in Canada, P. projectus was found in was extremely heavy and countless thous and s 85-90% of the forage fields in the province of of nematodes at different stages were present. Ontario, and in 61-63% in the provinces of The authors concluded that this crowded Quebec and New Brunswick (Townshend et al., environment and declining condition of the 1973; Willis et al., 1976). plants might affect the development of the nematode ovary, which was usually shorter. Paratylenchus shenzhenensis Wang, Xie, Li, Microscopic observations were made on Xu, Yu & Wang, 2013 feeder roots collected from an olive orchard in

Downloaded from jcp.modares.ac.ir at 5:11 IRST on Sunday October 3rd 2021 High population densities of P. shenzhenensis Italy naturally infected with P. peraticus (7600 nematodes per 100cm3 of soil) were (Raski, 1962) Siddiqi & Goodey, 1964. During considered to be the cause of severe damage to the observations, active vermiform juveniles, anthurium Anthurium and raeanum in immature females, and males were detected Shenzhen, Guangdong Province, China (Wang only in the soil, while mature females were et al., 2016). Commercial fields of this plant found attached to the roots. This species showed a patchy distribution of chlorotic induced feeding tubes in the host root tissues declining plants that were usually stunted. In (Inserra and Vovlas, 1977). In another study pathogenicity tests, obvious disease symptoms (cid del prado Vera and Maggenti, 1988), were observed as reduced and rotted roots (four colonies of juveniles and females of P. months after nematode inoculation) as well as hamicaudatus ciddel prado Vera & Maggenti, reduced plant growth and height plus reduced 1988 induced specialized feeding site in the rotted roots (eight months after nematode cortex of the roots of the Coast Redwood inoculation). Histological observations Sequoia sempervirens (D. Don) Endl indicated that P. shenzhenensis is an Brzeski et al. (1999) found the species P. endoparasitic pathogen of anthurium roots. arculatus Luc & Guiran, 1962 in a soil sample Whole nematode bodies were observed in the from olive tree nursery in the south of Spain. The outermost epidermal cells and root hairs, and population density varied from 0.03 to 2.49 of

250 Ghaderi______J. Crop Prot. (2019) Vol. 8 (3)

3 nematode per cm of soil. During the examination P. curvitatus van der Linde, 1938 of olive roots, the author found many females on suppressed plant height, stem sturdiness, root roots. As stated by Háněl (2000), the species P. length and to a little extent flower size on straeleni (de Coninck, 1931) Oostenbrink, 1960 is carnation at the initial inoculum level of 500 probably a typical species of soil fauna under individuals per pot. The severity of the damage silver birch Betula pendula Roth. culture, because increased with the corresponding increase of it occurs numerously in birch rhizosphere of wet the inoculum level. Plants affected by this as well as dry soils in South Bohemia. Čermák species were stunted, and their leaves turned and Renčo (2010) also noticed that P. Straeleni yellow. Floral stalks of such plants were weak was dominant species of plant-parasitic leading to poor quality of flowers. This study nematodes in the wet birch wood of Slovak and indicated that P. curvitatusis a potential threat the Czech Republic. to profitable cultivation of carnation in the Feeding habits of P. latescens (Raski, 1976) infested areas (Khanna and Jyot, 2002). Siddiqi, 1986 was well discussed in detail by The findings of Masdek et al. (2007) two separate studies (Troccoli et al., 2002; indicated that infestation by Paratylenchus sp. Inserra et al., 2003). These studies indicated is the most probable cause of yield decline of that mature females of P. latescens remain as pineapple on peat in Johor, India. Foliar sedentary ectoparasites attached by the stylet to analysis of the pineapple plants showed the surface of timber bamboo roots decreased content of potassium in the leaf. (Phyllostachys bambusoides Siebold &Zucc.) After the parent crop was sprayed with for their entire life. Troccoli et al. (2002) herbicide and burned later, the nematode indicated that slender females initiate root population decreased, but the population in the infection. These slender females remain root and soil still could affect the next crop. attached to the root surface by the stylet. Soil particles and cell debris accumulate around the Concluding remarks anterior portion of the female body outside the The pin nematodes of the genus Paratylenchus root. As females reach sexual maturity, they occur in the rhizosphere of many plants and become swollen and secrete a gelatinous feed on a wide range of host plants. matrix, which covers and protects their body. Paratylenchus species sometimes may produce The gelatinous matrix hardens around females, no specific symptoms in plants, but, it is

Downloaded from jcp.modares.ac.ir at 5:11 IRST on Sunday October 3rd 2021 males, newly hatched J2s, and eggs. Multiple supposed that large populations reduce the infections by four or five females packed absorption capacity of roots and promote root together in the same gelatinous matrix were death; probably they affect the general common. J2s leave the gelatinous matrix and physiology of the plant (Linford et al., 1949). If move to the soil and molt to initiate another a large number of Paratylenchus inhabit the cycle. Inserra et al., (2003) pointed out that subsoil zone, the plants show symptoms of vermiform females insert their long stylet into quick death (Jenkins, 1960; Mai et al., 1960; root tissues and remain attached to the root Adams and Eichenmuller, 1962; Corbett, 1966). surface, where they mature and swell. Generally, Paratylenchus is not considered Penetration of the nematode body to the root damaging on most crops unless it occurs in high tissue does not occur and thus, no anchorage numbers, more than 500 per 100cm3 of soil can be seen at the feeding site. The electron- (Talavera and Navas, 2002). Heavy dense deposit, probably produced by the root accumulation of Paratylenchus in the cell walls, appears to cement and thus anchor rhizosphere of plants and even in the roots is the stylet to the roots, allowing a sophisticated not necessarily conclusive of their parasitic life form of parasitism involving feeding site style. Interestingly, the researchers were often formation similar to that of other sedentary puzzled by the absence of a distinct correlation tylenchulids and cyst-forming nematodes. between the size of populations of

251 The damage potential of pin nematodes ______J. Crop Prot.

Paratylenchus and visible symptoms of the Adams, R. E. and Eichenmuler, J. J. 1962. plant condition, so, very large populations were Gracila cuscapitatus n. sp. from scarlet oak in often found in the rhizosphere of apparently West Virginia. Nematologica, 8: 87-92. healthy plants (Cobb, 1923; Linford et al., Allen, M. W. and Jensen, H. J. 1949; Reuver, 1959; Mcglohon et al., 1961; 1950.Cacopaurus epacris, new species Lownsbery et al., 1964). An important note in (Nematoda: Criconematidae), a nematode the pathogenicity of Paratylenchus which parasite of California black walnut roots. should be considered in making management Proceedings of the Helminthological Society decisions, is the ability of its species to increase of Washington, 17: 10-14. from low number to damaging levels during a Andrássy, I. 1985. Paratylenchus microdorus. short time (Jenkins and Taylor, 1956; Coursen C. I. H. descriptions of plant-parasitic and Jenkins, 1958; Rhoades and Linford, 1961; nematodes. Set 8, No. 107. Faulkner, 1964; Odihirin and Jinkins, 1965; Bally, W. and Reydon, G. A. 1931. De Townshend, 1972; Townshend et al., 1973; tegenwoordige st and van het vraagstuk van Brzeski, 1976). de wortelaaltjes in de koffiecultur. According to the literature, studies on the Archiefvoor de Koffiecultuur in Nederl and pathogenicity and damage to plants have not Indie, 5: 23-216. been well performed for pin nematodes (as a Bell, N. L. and Watson, R. N. 2001. group having ectoparasitic feeding behavior, Identification and host range assessment of and capable for causing damages) compared to Paratylenchus nanus (: sedentary or migratory endoparasites (e.g., root- Tylenchulidae) and Paratrichodorus minor knot nematodes, cyst nematodes, root-lesion (Triplonchida: Trichodoridae). Nematology, nematodes). However, the difficulties in 3: 483-490. working with these very small-sized nematodes Boag, B. 1974. Nematodes associated with are well known for plant pathologists. Much of forest and woodland trees in Scotland. the damage to crops is recorded for certain Annals of Applied Biology, 77: 41-50. widespread species including P. bukowinensis, Braun, A. L., Palmiter, D. H. and Keplinger, J. P. dianthus, P. hamatus, P. nanus, P. A. 1966. Nematodes found in apple orchards neoamblycephalus and P. projectus. This means in the Hudson Valley, 1956-1965. Plant our current knowledge on the damage potential Disease Reporter, 50: 783-786.

Downloaded from jcp.modares.ac.ir at 5:11 IRST on Sunday October 3rd 2021 of many species in the genus is very limited. Braun, A. L. and Lownsbery, B. F. 1975. The pin However, more detailed studies encompassing nematode, Paratylenchus neoamblycephalus, several species of the genus are required to on Myrobalan plum and other hosts. Journal of clarify the exact importance and roles of this Nematology, 7: 336-343. group of nematodes in agroecosystems. On the Brzeski, M. W. 1971. [Pathogenicity of other hand, understanding the feeding behavior Paratylenchus bukowinensis on cabbage]. and corresponding mechanisms, the host plant Zezzytyproblemowe Postepów Nauk resistance and nematode virulence are essential Rolniczych, 121: 113-119. to have better insights into the pathogenicity of Brzeski, M. W. 1975. [Pathogenicity of the pin nematodes. Paratylenchus bukowinensis Micol. on celery]. Roczniki Nauk Rolniczych Seria, E References 5: 23-29. Brzeski, M. W. 1976. Paratylenchus Abivardi, C. 1970. Occurrence of bukowinensis. C. I. H. descriptions of plant- Paratylenchus hamatus on citrus in Iran and parasitic nematodes. Set 6, N 79. 2 pp. its sensitivity to two nematicides under Brzeski, M. W. 1991. Decline of Paratylenchus laboratory conditions. Plant Disease bukowinensis under non-host crops. Revue Reporter, 54: 1085-1088. de Nematologie, 14: 636-637.

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Brzeski, M. W. 1998.Nematodes of Tylenchina Trichodorus christei. Plant Disease in Pol and and temperate Europe. Warszawa, Reporter, 42: 456-460. Pol and, Muzeum i Instytut Zoologii PAN. Eroshenko, A. C. and Volkova, T. B. 2005. Brzeski, M. W., Háněl, L., Nico, A. I. and Plant nematodes of the Far East of Russia: Castillo, P. (1999) Paratylenchinae: The orders Tylenchida and Aphelenchida Redescription of Paratylenchus arculatus (Nematodyrastenity Dal’nego Vostoka Luc & de Guiran, 1962, a new senior Rossii). Vladivostok, USSR, Dalnauka. synonym of P. nainianus Edward & Misra, Faulkner, L. R. 1964. Pathogenicity and 1963 (Nematoda: Tylenchulidae). population dynamics of Paratylenchus Nematology, 1: 375-380. hamatus on Mentha spp. Phytopathology, Brzeski, M. W., Zepp, A. L. and D’Errico, F. P. 54: 344-348. 1975. Development stages of Paratylenchus Fisher, J. M. 1965. Studies on Paratylenchus bukowinensis Micoletzky. Roczniki Nauk nanus I. effects of variation in environment Rolniczych Seria, E 5: 145-151. on several morphometric characters of Čermák, V. and Renčo, M. 2010. The family adults. Nematologica, 11: 269-279. Paratylenchidae Thorne, 1949 in the Fisher, J. M. 1967. Effect of temperature and rhizosphere of grass and woody species in host on Paratylenchus neoamblycephalus Europe: a review of the literature. and effect of the nematode on the host. Helminthologia, 47: 139-146. Australian Journal of Agricultural Research, Cid Del Prado Vera, I. and Maggenti, A. R. 18: 921-929. 1988. Description of Gracilacus French, A. M., Lownsbery, B. F., Ayoub, S. M., hamicaudata sp. n. (Nemata: Weiner, A. C. and Gholl, N. E. 1964. Criconematidae) with biological and Pythiaceous fungi and plant-parasitic histopathological observations. Revue de nematodes in California pear orchards. II. Nématologie, 11: 29-33. Incidence and distribution of parasitic Ciobanu, M., Geraert, E. and Popovici, I. 2003. nematodes in orchard soils. Hilgardia, 35: The genera Paratylenchus Micoletzky, 1922 603-610. and Gracilacus Raski, 1962 in Romania Gaur, H. S. 1988. Dissemination and mode of (Nematoda: Tylenchulidae). Nematologia survival of nematodes in dust storms. Indian Mediterranea, 31: 55-59. Journal of Nematology, 18: 94-98.

Downloaded from jcp.modares.ac.ir at 5:11 IRST on Sunday October 3rd 2021 Cobb, N. A. 1923. Notes on Paratylenchus, a Geraert, E. 1965. The genus Paratylenchus. genus of names. Journal of the Washington Nematologica, 11: 301-334. Academy and Sciences, 13: 254-257. Ghaderi, R. and Karegar, A. 2013. Some Corbett, D. C. M. 1966. Central African species of Paratylenchus (Nematoda: nematodes. II. Paratylenchus crenatus n. sp. Tylenchulidae) from Iran. Iranian Journal of (Nematoda: Criconematidae) from Malawi. Plant Pathology, 49: 137-156 (49-52) [In Nematologica, 12: 101-104. Persian with English abstract]. Corbett, D. C. M. 1978. Migratory soil Ghaderi, R., Geraert, E. and Karegar, A. 2016. Tylenchida. In: Southey, J. F. (Ed.). Plant The Tylenchulidae of the world, Nematology. Her Majestys Stationery identification of the family Tylenchulidae Office, London, pp. 188-206. (Nematoda: Tylenchida). Ghent, Belgium, Coursen, B. W. and Jenkins,W. R. 1958. Host- Academia Press. parasite relationships of the pin nematode Háněl, L. 2000. Soil nematodes (Nematoda) of Paratylenchus projectus on tobacco and tall alder, birch and oak forests in South and fesque. Phytopathology, 48: 460. Western Bohemia, Czech Republic. Coursen, B. W., Rohde, R. H. and Jenkins, W. ČasopisNárodnihoMuzeaŘadaPřírodovědné, R. 1958. Additions to the host lists of the 169: 107-117. nematodes Paratylenchus projectus and Inserra, R. N., Achor, D., Duncan, L. W. and

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Downloaded from jcp.modares.ac.ir at 5:11 IRST on Sunday October 3rd 2021 flora and fauna associated with roots. Soil Problemy Parazitologii, Kiev, pp: 230-231. Science, 53: 93-103. Nesterov, P. I. and Lisetskaya, L. F. 1965. [On Linford, M. B., Oliveira, J. M. and Ishii, M. the nematode fauna of some soils of the 1949.Paratylenchus minutus n. sp. a Moldavian SSR]. Parazity Zhivotnykh I nematode parasitic on roots. Pacific Science, Rastenii, Kishinev 1: 3-16. 3: 111-119. Nombela, G., Navas, A. and Bello, A. 1999. Lisetskaya, L. F. 1968. [Nematode fauna of Nematodes as bioindicators of dry pasture ethereal oil plants of Moldavia]. Material recovery after temporary rye cultivation. Natures Konferenz Po Parazitologie Sofia, Soil Biology and Biochemistry, 31: 535-541. pp. 47-48. Odihirin, R. A. and Jenkins, W. R. 1965. Host- Lisetskaya, L. F. 1969. [Nematodes of main parasite relationship of Impatiens balsamina ethereal oil plants of Moldavia]. Abstract of and certain nematodes. Phytopathology, 55: Candidates Dissertation, Kishinev, 1-19. 763-766. Loof, P. A. A. 1975. Paratylenchus projectus. Pennacchio, F., Derrico, F. P. and Tremblay, E. C.I.H. descriptions of plant-parasitic 1985. Spatial distribution pattern and nematodes. Set 5, No. 71. sequential sampling plan for Paratylenchus Lownsbery, B. F., Mitchell, J. T. and Paracer, dianthus. Nematologia Mediterranea, 13: S. M. 1964. Pythiaceous fungi and plant 137-146.

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Philis, J. 2003. Controlling parasitic nematodes Workshop. National Sunflower Association, in an established vineyard in Cyprus. Bismarck, ND, pp: 24-25. Nematologia Mediterranea, 31: 61-63. Sohlenius, B., Boström, S. and S and or, A. Raski, D. J. 1975. Revision of the genus 1987. Long-term dynamics of nematode Paratylenchus Micoletzky, 1922 and communities in arable soil under four descriptions of new species. Part II of three cropping systems. Journal of Applied parts. Journal of Nematology, 7: 274-295. Ecology, 24: 131-144. Raski, D. J. and Lider, L. A. 1959. Nematodes Sohlenius, B., Boström, S. and Viketoft, M. in grape production. California Agriculture, 2011. Effects of plant species and plant 13: 13-15. diversity on soil nematodes-a field Reuver, I. 1959. Untersuchungenüber experiment on grassland run for seven years. Paratylenchus amblycephalus n. sp. Nematology, 13: 115-131. (Nematoda, Criconematidae). Nematologica, Solov’eva, G. I. 1975.Parasitic nematodes of 4: 3-15. woody and herbaceous plants, a review of Rhoades, H. L. and Linford, M. B. 1961. the genus Paratylenchus Micoletzky, 1922 Molting of pre-adult nematodes of the genus (Nematoda: Criconematidae). New Delhi, Paratylenchus stimulated by root diffusates. India, Amerind Publishing. Science, 130 (1960): 1476-1477. Steiner, G. 1924. On some plant parasitic Rich, A. E. 1955. The occurrence and control names and related forms. Paratylenchus of Paratylenchus hamatus on celery in nanus Cobb infesting the roots of Zinnea New Hampshire. Plant Disease Reporter, elegans. Journal of Agricultural Research, 39: 307-308. 28: 1064-1065. Rössner,J. 1969. Phytoparasitäre nematodes in Talavera, M. and Navas, A. 2002. Incidence of Forstpflanzgärten. Zeitschrift Fur plant-parasitic nematodes in natural and Angewandte Zoologie, 56: 1-64. semi-natural mountain grassland the host Ruehle, J. L. 1967.Distribution of plant- status of some common grass species. parasitic nematodes associated with forest Nematology,4: 541-552. trees of the world. Southeast Forestry Thorne, G. 1943.Cacopaurus pestis n. g., n. spec. Experimental Station Asheville North (Nematoda: Criconematidae), a destructive Carolina, Forest Service, USDA. parasite of the walnut, Juglans regia. Linn.

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forage crops in eastern Canada. Canadian Weischcr, B. 1957. Neuere Gesichtspunktezur Plant Disease Survey, 53: 131-136. Frage der biologie und ökologie der w and Townshend, J. L. and Potter, J. W. 1976. erndenwurzel nematodes. Nematologica, 2 Evaluation of forage legumes, grasses, and (Supplementary): 406-412. cereals as hosts of forage nematodes. Weischer, B. 1960. Der einfluss des bodens auf Nematologica, 22: 196-201. die verbreitungpflanzen-parasitärer Troccoli, A., Vovlas, N. and Inserra, R. N. 2002. nematodes in Rebanlagen. Mitteilungenaus Parasitism of timber bamboo roots by der Biologischen Bundesanstaltfür L and- Gracilacuslatescens Raski, 1976 and morpho- und. Forstwirtschaft, 99: 51-59. biological notes on mature and immature life Weischer, B. 1964. Über die stages. Nematropica, 32: 87-102. Beziehungenzwischen Befallszahl und Viketoft, M. 2008. Effects of six grassland plant Schäden Bei Pflanzen Parasitären Nematodes. species on soil nematodes: a glasshouse Mitteilungenaus der Biologischen experiment. Soil Biology and Biochemistry, Bundesanstaltfür L and-und. Forstwirtschaft, 40: 906-915. 111: 32-42. Viketoft, M., Palmborg, C., Sohlenius, B., Willis, C. B., Townshend, J. L., Anderson, R. Huss-Danell, K. and Bengtsson, J.2005. V., Kimpinski, J., Mulvey, R. H., Potter, J. Plant species effects on soil nematode W., Santerre, J. and Wu, L. Y. 1976. Species communities in experimental grassland s. of plant-parasitic nematodes associated with Applied Soil Ecology, 30: 90-103. forage crops in eastern Canada. Plant Wang, K., Li, Y., Xie, H., Wu, W. J. and Xu, Disease Reporter 60: 207-210. C.L. 2016. Pin nematode slow decline of Wood, F. H. 1973. Biology and host range of Anthurium and raeanum, a new disease Paratylenchus projectus Jenkins, 1956 caused by the pin nematode (Nematoda: Criconematidae) from a sub- Paratylenchus shenzhenensis. Plant alpine tussock grassland. New Zeal and Disease, 100: 940-945. Journal of Agricultural Research, 16: 381-384. Wasilewska, L. 1997. The relationship between Wu, L. Y. and Hawn, E. J. 1975. Paratylenchus the diversity of soil nematode communities neoprojectus n. sp. (Paratylenchinae: and the plant species richness of meadows. Nematoda) from alfalfa fields in Alberta, Ekologia Polska, 45: 719-732. Canada. Canadian Journal of Zoology, 53:

Downloaded from jcp.modares.ac.ir at 5:11 IRST on Sunday October 3rd 2021 Webster, G. R. and Hawn, E. J. 1973. 1841-1843. Distribution of Paratylenchus projectus in Yeates, G. W. 1984. Variation in soil nematode central and northern Alberta. Canadian Plant diversity under pasture with soil and year. Disease Survey, 53: 175-177. Soil Biology and Biochemistry, 16: 95-102. Webster, G. R., Orchard, W. R. and Hawn, E. J. Yeates, G. W. and Lee, W. G. 1997. Burning in 1972. Paratylenchus projectus in alfalfa a New Zeal and snow-tussock grassland: fields of central and northern Alberta. effects on vegetation and soil fauna. New Canadian Plant Disease Survey, 52: 75-76. Zeal and Journal of Ecology, 21: 73-79.

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ﭘﺘﺎﻧﺴﯿﻞ ﺑﯿﻤﺎرﯾﺰاﯾﯽ ﻧﻤﺎﺗﺪﻫﺎي ﺳﻨﺠﺎﻗﯽ (Paratylenchus spp. (Nematoda: Tylenchulidae در ﮔﯿﺎﻫﺎن

رﺿﺎ ﻗﺎدري

ﺑﺨﺶ ﮔﯿﺎهﭘﺰﺷﮑﯽ، داﻧﺸﮑﺪه ﮐﺸﺎورزي، داﻧﺸﮕﺎه ﺷﯿﺮاز، ﺷﯿﺮاز، اﯾﺮان. ﭘﺴﺖ اﻟﮑﺘﺮوﻧﯿﮑﯽ ﻧﻮﯾﺴﻨﺪه ﻣﺴﺌﻮل ﻣﮑﺎﺗﺒﻪ: [email protected] درﯾﺎﻓﺖ: 11 ﺷﻬﺮﯾﻮر 1397؛ ﭘﺬﯾﺮش: 7 اﺳﻔﻨﺪ 1397

ﭼﮑﯿﺪه: ﺟﻨﺲ Paratylenchus در ﺗﻌﺮﯾﻒ ﮔﺴﺘﺮده ﺧﻮد ﺷﺎﻣﻞ ﻧﻤﺎﺗﺪﻫﺎي ﻣﺘﻌﻠﻖ ﺑﻪ ﺟﻨﺲﻫﺎي Paratylenchus ( ﮔﻮﻧﻪﻫﺎي داراي اﺳﺘﺎﯾﻠﺖ ﺑﻪﻃﻮل 10 اﻟﯽ 40 ﻣﯿﮑﺮوﻣﺘﺮ)، Gracilacus ( ﮔﻮﻧﻪﻫﺎي داراي اﺳﺘﺎﯾﻠﺖ ﺑﻪﻃﻮل 40 اﻟﯽ 120 ﻣﯿﮑﺮوﻣﺘﺮ)، Gracilpaurus ( ﮔﻮﻧﻪﻫﺎي داراي ﺑﺮﺟﺴﺘﮕﯽﻫﺎي ﮐﻮﺗﯿﮑﻮﻟﯽ) و Paratylenchoides ( ﮔﻮﻧﻪﻫﺎي داراي ﺷﺒﮑﻪ ﮐﻮﺗﯿﮑﻮﻟﯽ ﺳﺮ رﺷﺪﯾﺎﻓﺘﻪ) اﺳﺖ. ﺗﻌﺪادي از ﮔﻮﻧﻪﻫﺎي داراي اﺳﺘﺎﯾﻠﺖ ﺑﻠﻨﺪ، از ﺑﺎﻓﺖ ﮐﻮرﺗﮑﺲ رﯾﺸﻪﻫﺎي ﻣﯿﺰﺑﺎنﻫﺎي ﭼﻮﺑﯽ ﺗﻐﺬﯾﻪ ﮐﺮده و ﻣﺘﻮرم ﻣﯽﺷﻮﻧﺪ، اﻣﺎ اﻏﻠﺐ ﮔﻮﻧﻪﻫﺎ اﻧﮕﻞ ﺳﻄﺤﯽ ﺳﺎﮐﻦ رﯾﺸﻪﻫﺎي ﮔﯿﺎﻫﺎن ﻣﯽﺑﺎﺷﻨﺪ. ﺑﻪﻋﺒﺎرت دﯾﮕﺮ، ﮔﻮﻧﻪﻫﺎي داراي اﺳﺘﺎﯾﻠﺖ ﮐﻮﺗﺎهﺗﺮ از 40 ﻣﯿﮑﺮوﻣﺘﺮ ﻣﻌﻤﻮﻻً از ﺳﻠﻮلﻫﺎي اﭘﯿﺪرﻣﯽ ﺗﻐﺬﯾﻪ ﻣﯽﮐﻨﻨﺪ اﻣﺎ ﮔﻮﻧﻪﻫﺎي داراي اﺳﺘﺎﯾﻠﺖ ﺑﻠﻨﺪﺗﺮ ﻗﺎدرﻧﺪ ﺑﺪون وارد ﺷﺪن ﺑﻪ رﯾﺸﻪ، از ﺑﺎﻓﺖ ﮐﻮرﺗﮑﺲ آن ﺗﻐﺬﯾﻪ ﻧﻤﺎﯾﻨﺪ. ﺑﻪﻃﻮر ﮐﻠﯽ ﻧﻤﺎﺗﺪﻫﺎي ﺳﻨﺠﺎﻗﯽ ﺟﻨﺲ Paratylenchus اﻧﮕﻞﻫﺎي ﮔﯿﺎﻫﺎن ﻋﺎﻟﯽ ﺑﻮده و ﺑﯿﺶﺗﺮﯾﻦ ﻓﺮاواﻧﯽ را در ﻓﺮار ﯾﺸﻪي درﺧﺘﺎن و ﮔﯿﺎﻫﺎن ﭼﻨﺪﺳﺎﻟﻪ دارﻧﺪ. ﻣﻘﺎﻟﻪ ﻣﺮوري ﺣﺎﺿﺮ ﺑﯿﻤﺎرﯾﺰاﯾﯽ و ﭘﺘﺎﻧﺴﯿﻞ ﺧﺴﺎرت ﻧﻤﺎﺗﺪﻫﺎي ﺳﻨﺠﺎﻗﯽ را در ﮔﯿﺎﻫﺎن ﻣﻮرد ﺑﺤﺚ ﻗﺮار داده اﺳﺖ.

واژﮔﺎن ﮐﻠﯿﺪي: Gracilacus، ﺧﺴﺎرت، ﺑﯿﻤﺎرﯾﺰاﯾﯽ، ﮔﯿﺎﻫﺎن ﭼﻨﺪﺳﺎﻟﻪ، درﺧﺘﺎن، ﻧﻤﺎﺗﺪﻫﺎي ﺳﻨﺠﺎﻗﯽ، ﺟﻤﻌﯿﺖ Downloaded from jcp.modares.ac.ir at 5:11 IRST on Sunday October 3rd 2021

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