Fundam. appl. NemalOl., 1997.20 (6), 521-539

Forum article

NEMATODES AND AGRICULTURE IN CONTINENTAL ARGENTINA.AN OVERVIEW

Marcelo E. DOUCET and Marîa M.A. DE DOUCET

Laboratorio de Nematologia, Centra de Zoologia Aplicada, Fant/tad de Cien.cias Exactas,Fisicas y Naturales, Universidad Nacional de Cordoba, Casilla df Correo 122, 5000 C6rdoba, Argentina.

Acceplecl for publication 5 November 1996.

Summary - In Argentina, soil nematodes constitute a diverse group of invertebrates. This widely distributed group incJudesmore than twO hundred currently valid species, among which the plant-parasitic and entomopathogenic nematodes are themost remarkable. The former includes species that cause damages to certain crops (mainly MeloicU:igyne spp, Nacobbus aberrans,Ditylenchus dipsaci, Tylenchulus semipenetrans, and Xiphinema index), the latter inc1udes various species of the Mermithidaefamily, and also the genera Steinernema and Helerorhabditis. There are few full-time nematologists in the country, and they workon taxonomy, distribution, host-parasite relationships, control, and different aspects of the biology of the major species. DuetO the importance of these organisms and the scarcity of information existing in Argentina about them, nematology can beconsidered a promising field for basic and applied research.

Résumé - Les nématodes et l'agriculture en Argentine. Un aperçu général - Les nématodes du sol représentent enArgentine un groupe très diversifiè. Ayant une vaste répartition géographique, il comprend actuellement plus de deux centsespèces, celles parasitant les plantes et les insectes étant considèrées comme les plus importantes. Les espèces du genre Me/oi­dogyne, ainsi que Nacobbus aberrans, Dùylenchus dipsaci, Tylenchulus semipenetrans et Xiphinema index représentent un réel dangerpour certaines cultures. Les parasites d'insectes comprennent plusieurs espèces de Mermithidae, ainsi que des représentantsdes genres Steinernema et Heterorhabditis. Les nématologistes travaillant en Argentine sont peu nombreux. Ils poursuivent desrecherches très diversifiées: taxinomie, répartition, relation hôte-parasite, résistance, contrôle, biologie des principales espèces.Etant donné l'importance de ces organismes et leur connaissance encore incomplète, la nématologie peut être considéréecomme ayant en Argentine un futur plein de promesses dans les domaines des recherches fondamentale et appliquée.

Key words: Agriculture, Argentina, nematology.

This article is an overview of the main features ofthe histary of nematalogy in Argentina and its currentstatus, with emphasis on its association with agricul­ture.

The information presented here cornes from a cri ti­cal analysis of the most relevant bibliographic infor­mation, from the personal experience of the authorsand from their interaction with other specialists in thecountry.

Mainland Argentina has an area of 2 791 810 km 2

and is characterised by a great diversity of climates(Chiozza & van Domselaar, 1958) (Fig. 1) and soils(Papadakis, 1964; Moscatelli, 1990). This diversityhas favoured the development of diversified agricul­ture, with both extensive and intensive farming sys­tems in different regions of the country (Fig. 2).Extensive farming is used for crops such as wheat,soybean, corn, sorghum, sunf1ower, rice, sugar cane,barley, rye, alfalfa, peanut, rape, tabacco, grapevine,sweet potato, and potato; intensive farming producesail kinds of vegetables, and mostly tomato, garlic, pep­per, beans, asparagus, and lentils. In addition, many

types of fruit trees are grown in wide areas throughoutthe country (Gomez Riera, 1992). Aromatic andorna mental plants also are important items in thevegetal production of Argentina.

Throughout its development, Argentine agriculturehas adapted ta the demand of international markets.Because of this, different crops have received priorityin different regions. Currently, soybean, wheat, corn,sunf10wer and potata are the most important speciesfrom the viewpoint of exports and domestic consump­tion (Cap el al., 1993a).

Within this context of marked agricultural diversity,there exist numerous species of soil nematodesbelonging to the major trophic categories (plant-para­sitic, free-living, mycophagous, predators, and ento­mopathogenic species).

Plant-parasitic nematoàes include sorne species thatcan cause serious damages tO several crops, while vari­ous entomopathogenic species could be used as bio­logical control agents against harmful insects. Recentresults (Doucet & Doucet, 1996) and older data as

Fundam. app!. NemolO/. J J 64-557 1/97/06 1 ~. Gauthier- Villars - ORSTOM 521

kf. E. Douce! & M. J'IIf. A. de Douce!

o 200

TYPES OF CLIMATE

tç{:~g/ . // SUI3TROPICAJ~ WITHOUT DRY SEASON

rTTTTm~SUI3TROPICAL WITH DRY SEASON

~TROPICAL OF THE MOUNTAINS

~ ARID OF THE ANDES AND PUNA

~TEMPERATE OF THE PAMPAS

~ OCEANIC TEMPERATE

t~===~ TRANSITIONAL TEMPERATE

~ AHID OF THE MOUNTAINS~ AND nOLSONS

b==~ SEMIARID

DJIIID PATAGONIAN ARID

~TEMPERATE OF THE MOUNTAINS

o COLD-WET OF THE PATAGONIAN ANDES

[[[[II] MAGALLl\NIAN

li.'>:] NIVAL

qOO Km

Fig. 1. Types of climale ill Argentina (adapledfrom Chiozza & van DorllSe/aar, 1958).

522 Fundam. appl. Nemarol.

ARGENTlNJ\REPUElLIC

f!

SUNFLOWF:R

NemalOdes in conlinelllai Argelllina

Fig. 2. Distriburion of the main crops in Argenlina.

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M. E. Doucel & M. M. A. de Doucel

presented in this article emphasise the great diversityof soil nematodes in Argentina.

Historical background

The first reference to soil nematodes in Argentina isrelated to entomopathogenic species. A German natu­ralist (Weyembergh, 1876) working in the NationalAcademy of Sciences (C6rdoba City) described underthe name of Gordius aeridiorum a parasite associated toone of the most important agricultural pest at thattime: the locust Schiscocerca paranensis (Burm). Later,the same investigator, with others also from Europe,reported the afore mentioned species and G. acrydii inlocusts from Buenos Aires and C6rdoba (\Veyen­bergh, 1878, 1879; Conil, 1878, 1879; BruneI',1898). The descriptions made at that time were notdetailed, but the data published by these authors onthe life-cycle and interactions of these parasites withtheir hosts make it obvious that these nematodesbelonged to the family Mermithidae (Berg, 1898).

Only fifteen years separate the first reference toentomopathogenic species and the discovery of aplant-parasitic nematode. At the beginning of the1890's, the Head of the Chemistry Office of theMunicipality of C6rdoba city - a chemist and natural­ist from Germany - informed the Head of theNational Department of Agriculture at Buenos Airesof the finding of "Helerodera radicicola de Greer' asso­ciated with grapevine, and reported sorne problemscaused by phylloxera. He also rnentioned that thispathogen had also been observed on roots of peachtrees (Doering, 1891) and he added: "we have foundthe linle animal together with the eggs in the cavitiesof the root-knot" (c1early a reference to the genusJ'vIeloidogyne) .

The subsequent deveJopment of nematology in thecountry followed different paths for plant-parasitic,entomopathogenic, and free-living soil nematodes.

Plant-parasitic nematodes, because of their closeassociation with economic losses to the agriculture,have received the most attention. We will nowdescribe the activities that took place in Argentinaduring the development of nematological researchconcerning each of the three types of nematodes.

Plant-parasitic nematodes

At the beginning to this cent ury, several articleswere published on Helerodera radicicola (now Meloido­gyne spp.) associated with tomato plants. Thosepapers are remarkable by the good information theyprovide on yield loss in the attacked plants, sorneaspects of the life cycle of the nematode and its rel a­tionships with sorne environmental factors, and themost convenient methods for reduction of the nema­tode damages (Lynch Arribalzaga, 1901; H uergo1902, 1903).

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From that time on, several species were sporadicallydetected in differem regions, and linked to problemsobserved in several crops.

As time went by, sorne people from national organi­zations - 1) Ministry of Agriculture; Instituto Nacio­nal de Tecnologia Agropecuaria (INTA): Estaci6nRegional Experimental Balcarce (Buenos Airesprovince), Roque Saenz Pena (Chaco province), SanRafael (Mendoza province); il) Universities (Univer­sidad de Buenos Aires, Universidad Nacional de C6r­doba, Universidad Nacional de Cuyo, UniversidadNacional de La Plata) ; provincial organizations ­Direcci6n de Sanidad Vegetal, Ministerio de AsuntosAgrarios, Buenos Aires province; Esraci6n Experi­mental AgI'o Industrial Obispo Colombres, Tucumanprovince ; and private organizations - Centro deInvestigaciones Nemato16gicas de Corrientesbecame interested in the studies of these organisms.Agriculture Engineers Burkart, Fresa and Marchion­ano and Dr. R. Gutierrez were sorne of the peoplewho out stood in sorne of those institutions for theirwork. Thus, "centres" of nematological activitiesemerged in the Buenos Aires, Mendoza and Tucumanprovinces; in these last IWO provinces, Engineers Edu­ardo Vega and Miguel Costilla developed importantapplied works related to management of plant-para­si tic nematodes associated to numerous crops. Thefirst articles defining the status of p!ant-parasiticnematodes in relation to agriculture in Argentina wereprepared in Buenos Aires and Mendoza (Moreno,1964; L6pez Crist6bal, 1965; Vega, 1971). Dr. Ama­lia Moreno worked in the Patologia Vegetal Institute(Centro de Investigaciones Agropecuarias, INTACastelar, Buenos Aires) where she analysed sarnplesfrom different sites in the country and obrained theinitial information about the distribution of differentspecies and their relationships with their hosts.

In 1980, a new "centre" of nematological activitieswas created at the National University of the city ofC6rdoba, under the direction of the authors of thisarticle. Both had studied nematology at "Station deRecherches sur les Nématodes" (INRA, Antibes,France). In C6rdoba, they engaged in the study of soilnematodes (plant-parasitic, entomopathogenic andfree-living species) and organised courses on Agricul­tural nematology. The first course - "Introduction tonematology" - in Argentina was presented in the Cen­tro by the present authors in July 1980.

Almost at the same time, another nematologist, Dr.E. Chaves, was being formed at the University ofGhent, Belgiurn, who later joined the UnidadIntegrada EEA Balcarce, Universidad Nacional deMar dei Plata, Buenos Aires.

Subsequently nematological activities started at theUniversidad Nacional de Cuyo, Mendoza province(control of plant-parasitic nematodes) and at the Uni-

Fundam. appl. j\lemawl.

versidades Nacionales of Buenos Aires and La Plata(Buenos Aires province) (survey of plant-parasiticnematodes and taxonomy-biology of entomopatho­genic nematodes, respecrively).

More recently, other young researchers from Argen­tina have done postgraduate studies in Belgium andthe USA, on taxonomy and the search for resistantcharacters.

Agriculturallosses caused by plant-parasiticnematodes

In general, agricultural losses caused by nematodesare not yet known with any degree of accuracy.

According to specialised agronomists, it is possibleto evaluate approximate yield reductions in sornezones or average production losses for specific cul­tures. For example, in the western part of the country,tomato and peach trees were affected until 1970 byMeloidogyne spp. that reduced the yields by 20 % and15 %, respectively. Meloidogyne spp. and Xiphinernaindex attacks on grapevine reduced yields by about5 %. The use of resistant varieties and nematicidesproducts have now reduced the losses to 2, 1, and3 %, respectively. On garlic, Ditylenchus dipsacicaused damages that affected 30 % of the productionbenveen 1965-1970; the selection of non contami­nared seeds and different methods of control havereduced the damages to less than 1 % of the produc­tion (Vega, pers. comm.).

Potato can be attacked by Meloidogyne spp. and/orNacobbus aberrans, depending on the regions. Themain problems occur for seed potato production. Spe­cies of Meloidogyne have been observed in the West(Vega el al., 1986; Doucet & Ponce de Leon, 1996a)affecting 40 % of the production, the North-West(Costilla, comm. pers.), and the South-East (Chaves& Torres, 1993; Doucet & Ponce de Leon, 1996a)affecting 30 % of the production (Vega, comm. pers.).

Nacobbus aberrans has been found so far only inTucuman and Buenos Aires areas (Costilla el al.,1978; Chaves & Torres, 1993). The current legisla­tion prohibits the use of infected tubercles for seeds;they are sold for food at lower prices.

These and other nematodes can cause a decrease ofthe production of several other crops. This situationhas been evident for sorne years in horticultural cropsunder hot-house conditions; the economical loss hasnot yet been evaluated with precision.

Current list of nematode species in the country

The number of nematologists specialised in popula­tion characterisation, and identification and descrip­tion of species has been and still is very low.

Unfortunately, sorne doubts have been expressed onpast identification of sorne species because the vari-

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Nematodes in continental Argentina

abiliry of the characters used was not taken intoaccoun t and detailed analyses of the populations werenot conducted. Sorne identifications were probablyinaccurate because those species have never beenfound again, although many samples were analysed.Furthermore, voucher specimens were almost neverkept in local institutions or deposired in known collec­tions.

As a consequence, it is difficult to recognise whichspecies are of quarantine importance for the country.In sorne cases, the commercialisation of sorne agricul­tural products is difficult on international markets.

The list of identified species is given in the Appen­dix.

Active fields of research

The srudies on plant-parasitic nematodes in Argen­tina can be divided into two main categories.

The first one is related to the acquisition of basicinformation: distribution, host range, sorne host-para­site relationships, identification and characterisationof populations of various species, and some aspects ofthe biology of new species. This kind of informationhas been obtained on the most common plant-para­sitic nematodes associated with the major crops of thecountry (Lapez Cristobal, 1946, 1965; Moreno,1958; La Red & Vega, 1968; Vega, 1971; Costillael al., 1976; Chaves, 1984; Doucet, 1986, 1989;Chaves & Torres, 1993). These activities are stillbeing developed, and expanded to different generaand species of nematodes.

The second category is related to the managementof problems caused by harmful nematodes.

From the very first projects on plant-parasiticnematodes in Argentina, the necessity of controllingnematode populations has been emphasised. The useof various chemical products has been proposed and ithas been recommended to use preventive measures toavoid spreading these pathogens (Doering, 1891;Lynch Arribalzaga, 1901; Huergo, 1902, 1903).Then, several methods of crop protection were imple­mented.

In different cases, resistant varieties were selected(Burkart, 1934, 1937; Galmarini, 1978; Pucci &Avila, 1978; SisJer & Casaurang, 1983; March el al.,1985; Costilla, 1985, 1986; Castellanos & Del Toro,1994), crop rotation was established (Vega, 1981;Costilla, 1991), and different chemicaJ methods wereused, as explained below under the major nematodesspecies. Several nematicides have been tested, includ­ing DD, DBCP, methyl bromide (Gargiulo & Moy­ano, 1948; Vega & Gatica, 1968, 1972; Del Toro &Vega, 1982), carbofuran (Costilla, 1973), and aldi­carb (Del Toro, 1988a). Because of their high cost,these are used generally in crops with high profitabi­lity.

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Experiments have started on the use of solarizationtechniques as an alternative control method.

Finally, surveys have recently been initiated tOdelimitate areas free of speciflc harmful nematOdes.This should make it possible tO commercialise agri­cultural products in accordance with internationalregulations (Del Toro el al, 1994a).

Entomopathogenic nematodes

After the discovery that marks the beginning ofArgentine nematOlogy (Weyembergh, 1876), morethan half a century passed before a second ento­mopathogenic nematode was detected: the mermithidAgamermis decaudaw found parasitizing specimens ofthe JocuSt Dichroplus arrogans Stal in Buenos Aires(Gutierrez, 1945).

At the beginnings of the 1970's, nematodes belong­ing to the Steinernematidae and Heterorhabditidaefamilies, natural enemies of Coleoptera of agronomi­cal importance (genus GraphognalUs), were reportedfor the first time in the humid pampa soils (Ahmad,1972).

The Jast 15 years have seen the beginning of sys­tematic studies of entomopathogenic nematOdes.Research concentra tes almost exclusively on speci­mens of the Mermithidae, Steinernematidae andHeterorhabditidae families. Only a few studies con­cern nematodes of the Thelastomatidae, Rhabditidaeand Diplogasteridae families (Miralles, 1982; Doucet& Doucet, ] 989, 1992; StOck & Camino, 1991;Stock, 1993c).

In general, rhe nematode families which are beingstudied are those that include species that are beingused, or may be used, as an alternative for chemicalcontrol of insect pests.

Active fields of research

Research has been developed on the following sub­jects:

TAXONOMIC STUDIES

These include the characterisation of local popula­tions of known species and also descriptions of newgenera and species in the Mermithidae (Doucet,1982, 1986a; Doucet & Poinar, 1984; Camino,1985a, b, c, 1987, 1988a, 1990, 1991, 1993; Camino& Poinar, 1989; Camino & StOck, 1989, 1994; Srock& Camino, 1992a,b), Steinernematidae (Doucer,1986b, 1995; Poinar el al., 1988; Doucet & Doucer,1990; Stock, 1993b), and Heterorhabdiridae families(Stock, 1993a; Doucer el al., 1996).

Several articles on species of the 1asr rwo familieshave emphasised the need ro consider rhe intraspecificvariabiliry of rhe characrers used (Doucet & Doucer,

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1986; Doucerelal., 1990,1991, 1992a,Doucerelal.,1996).

BIOLOGICAL STUDIES

The published srudies described rhe embryonicdevelopment and 1ife cycle of rhe mermirhids (Cami­no, 1986; Camino & Reboreda, 1994) and the lifecycle of hererorhabdirids (Doucer & Poinar, 1985;Doucer el al., 1996).

Srudies on hosr range and aggressiviry of Hele­rorhabdilis bacleriophora Poinar, 1975 have recentlysrarted (Doucer el al., 1992b; Doucer & Giayerto,1994; Doucer el al., 1996).

ECOLOGICAL STUOIES

Studies concern the geographic distribution ofmermithids, sreinernematids, and heterorhabditids(Doucet & Cagnolo, 1995a; Doucet & Bertolotti,1995; Doucet & Doucet, 1996) and the spa tial distri­bution patterns of Heterorhabditidae and Steinerma­tidae infective larvae in the soil (Doucet, 1992).

These studies continue in the Centro de EsrudiosParasitOI6gicos y Vectores (La Plata, Buenos Airesprovince), the Laboratorio de Nematologia of theCentro de Zoologia Aplicada, and the Catedra deParasitologia, Facultad de Ciencias Exactas, Fisicas yNaturales (Universidad Nacional de C6rdoba).

Free-living soi! nematodes

Most of the articles on free-living nematodes con­cern descriprions of species from various parts of thecountry. In particular, Cruznema lriparlilum has beenstudied in relation to its possible association with adisease of garlic (Gutierrez, 1949; Avila & Pucci,1980; Doucet & Ponce de Le6n, 1994) and its even­tuai behaviour as a facultative entOmopathogenic spe­cies (Doucet & Doucet, 1989; Doucet, 1994).Another interesting case is that of the free-living spe­cies DislOlabrellus veechi Anderson, 1983 which, inspecifie circumstances, becornes a facultative ento­mopathogenic species (Doucet & Doucet, 1992).

The Argentine Society of Nematology

In 1980, at the request of INTA, the ArgentineSociery of Nematology was created with the objectiveof coordinating nematological activities in the fieldand promoring conracts between the specialists in thecountry. Its first President was the senior author of thepresent article and one of the most important actionsof the Society was to create an awareness of the agri­cultural significance of soil nematodes, the damagesthey cause, and the possible ways to manage theseparasites. To this aim, courses were organised inseveral instirutions ail over the country for teachers,technicians, researchers and other professionals

Fundam. appl. Ne rn alOI.

involved in plant pathology. These courses proved rabe one of the best ways of promoting nemaralogy.Among those courses, one of the most remarkable wasthe one sponsored by ORSTOM (France) in 1986 atthe Centro de Zoologia Aplicada. The growing aware­ness of nemarade problems initiated by these coursesamong agriculture technicians and farmers resulted inthe passing by the national and provincial Ministriesof Agriculture of laws intended ra limit the spread ofnematodes harmful to agriculture. In this manner, notonly is agricultural production preserved but also theproducts obtained are clean and can be commercia­lised abroad.

Present situation

At the present, there are about ten researchersworking full time on nematology. The studies that arecarried out concern several subjects (survey, taxo­nomy, biology, management, search for resistancesources) and are principally related ra plant-parasiticand entomopathogenic nematodes. Occasionally, thefew specialised laboratories offer identification ser­vices for the species of importance for various cropsand act as consultants for organisations related toagriculture at the national, provincial and privatelevels.

Major genera and species of plant-parasiticnematodes reported in Argentina andthe problems they cause

From the beginning, attention has focused onharmful species in the genera Ditylenchus, MelOl'do­gyne, Nacobbus, Tj1lenchulus, and Xiphinema.

In this context, it is necessary to mention the occur­rence of the species Dùylenchus dipsaci, Meloidogynespp., Nacobbus aberrans, 'Tylenchulus semipenetrans andXiphinema index associated to cultivated alfalfa andgarlic, potatoes and several vegetables, citrus andgrapevine, respectively. Because their possible eco­nomic importance, other genera (Pratylenchus andHelicotylenchus) are also ta ken into account.

GENUS DITYLENCHUS

Two species are found in Argentina: D. imermediusand D. dipsaci.

Ditylenchus intermedius

Found in association with the tree Platanus sp.(Moreno & Turica, 1960), P. acen/alia (Brugnoni,1980), Citrus sp., and grass (Moreno, 1961). Theimpact of this nemarade on agriculture has not yetbeen evalua ted.

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Nematodes in cominellla/ Argentina

Ditylenchlls dipsaci

This species has created problems mostly on alfalfaand garlic. Ir was first recorded in the country associ­ated with damages on aJfalfa in the North-West(Anon., 1929). Several studies on D. dipsaci were car­ried out before the 1950's. The species was believed tobe widely distributed in the central and northern partsof the country and to be responsible for damages ofvariable importance (Baez, 1925; Marchionano et al.,1926; Anon., 1929; Blanchard, 1930; Burkart, 1943;Lopez Cristobal, 1946; Moreno, 1964). Observationsmade at that time revealed the presence of plantsimmune to this nematode in sorne fields in BuenosAires. Resistant varieties of alfalfa were then selectedfor the management of D. dipsaci (Burkart, 1934,1937; Ragonese & Marco, 1943; Miccio Peralta,1956; Godeck & Favret, 1965; Godeck & Stilinovic,1966). Since then, no more damages caused by thisplant-parasitic nematode have been detected in alfalfafields.

The crop that was the most affected by D. dipsaci isgarlic. The climatic and edaphic conditions of thewestern, central-southern and south-eastern regionscontributed to the fast increase of populationswherever infested seeds were used (Cucchi et ai.,1967; Urbieta Salvarredi et ai., 1971). D. dipsaci wasfirst found on garlic in 1965 when it provoked a totalloss of the garlic production in sorne areas of the prov­ince of Mendoza, the major producer of garlic in thecountry (Cucchi et al., 1967). Simultaneously, theclose association of this nematode with fungi to forma major pathologie complex was detecred (Avila &Pucci, 1980). This situation was at the origin ofdetailed studies on the behaviour of the nemarade(Urbieta Salvarredi, 1972), which resulted in theimplementation of several forms of control, includingthermotherapy (Del Toro & Mavrich, 1977), chemicalcontrol with synthe tic nematicides (Rivera & DelToro, 1982; Del Toro et al, 1988), and use of naturalproducts (Del Toro, 1988b).

Today, garlic is grown on about 15 000 ha dis tri­buted in various regions, mainly the Central-West(Burba, 1992). Given the specific requirements of theArgentine regulatory legislation, special care is takenfor the production of nemarade-free seeds, which isenough ta ensure an excellent production of garlicwithout nematode damages (Del Toro el al., 1994b).

It seems that the host range of D. dipsaci is wide inArgentina. This nemarade has been found in pastures(Burkart, 1935; Moreno & Fresa, 1969), wheat(Brugnoni, 1966), ornamental plants and weeds(Burkart, 1937; Moreno, 1956a; Doucet, 1992; Mar­eggiani & Russo, 1992), cotton (Mallo, 1961), andforest trees (Moreno & Turica, 1960; Brugnoni,1980).

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Dilylenchus spp.

Unidentified species of Dùylenchus have beenreported in association with sugar cane in the north­western part of Argentina (Costilla el al., 1976) andwith corton (Gutierrez, 1959).

GENUS PRA TYLENCHUS

This genus is widely distribured throughout thecountry. So far, fourteen species have been reportedin Argentina. The population densities of these spe­cies are generaly low, except in the North-West wherePralylenchus spp. cause damages to banana plantations(Costilla, 1973) and can affect the development andyield of corn (Costilla, 1992).

GENUS NACOBBUS

This genus was first found in the mountainousregion of the province ofTucuman (2000 m above sealevel), in soils of potata fields where it parasitises rootsof cultivated and wild potatoes, Cucurbita maxima,Beta spp., and sorne weeds (Costilla el al., 1977).

Subsequent studies showed that this nemarode iswidely distributed in the country (Chaves & Sisler,1980; Chaves, 1984; Costilla, 1985; Doucet, 1989)and that it can be associated with a large number ofplants, such as potataes, vegetables and weeds (Ojedael al., 1978; Agüero el al., 1984; Costilla, 1985; Dou­cet & Ponce de Leon, 1986; Ponce de Leon & Dou­cet, 1989; Doucet el al., 1992; Doucet, 1992).

The analysis of morphological characters of variouspopulations of Nacobbus from Argentina showed thatthey ail belong ta the species N. aberrans (Chaves,1984; Costilla, 1985; Doucet, 1989; Doucet & DiRienzo, 1991). However, preliminary observations ofthe behaviour of different populations in relation tadifferent hosts revealed significant differences (Cos­tilla, 1986). Subsequent studies based on morpho­metric characters (Doucet & Di l'Zienzo, 1991) andisoenzyme phenotypes (Doucet & Gardenal, 1992;Doucet el al., 1996) revealed important differencesbetween populations. This supports the hypothesis ofa complex of forms within the species N. aberrans(Jarala & Golden, 1977).

The major problems caused by this species arerelated to the cultivation of potaro (Ojeda el al., 1978;Costilla, 1985).

Besides forming knots in the roots, third and fourthstage juveniles settie below the lenticels of the tuberwhere they reach a state of anhydrobiosis. In thisstate, the nematades remain viable during storage andmay be dispersed by contaminated tuber seed (Cos­tilla, 1985). N. aberrans causes significant damages rocrops, seriously affecting production. Because of this,the legislation requires the use of non-contaminatedpotata seed.

528

Various strategies are used against this nematade,such as the use of nematicides (Costilla & Gomez,1981), the assessment of resistance of different culti­vars (Sisler & Casaurang, 1983), and crop rotation(Costilla, 1991).

A search for sources of resistance in Lycopersicon hasrecently been launched (Cap el al., 1993b).

GENUS HELICOTYLENCHUS

This genus is widely distributed in Argentina. Irincludes several populations, most of which have notbeen identified at the species level.

The most important species identified so far ­H. multicinaus - was found in the North-West onroots of banana trees, where it prod uces characteristiclesions (Costilla el al., 1979).

GLOBODERA ROSTOCHIENSIS

This nematode was first reported in Argentina inthe soil of a garlic field in the western part of thecountry (Moreno, 1956b). Later, it was mentioned inthe Andes, Province of Jujuy, parasitizing roots of wildpotaroes (Brücher, 1960, 1961). Years later, it wasreported in the soil of a potato field in another moun­rainous region in the West (Virsoo, 1967).

Since then, the numerous analyses carried out bydifferent laborarories on soil samples and tubers frommany different regions ail had negative results(Chaves, 1993). Work on cyst nematades from Argen­tina suggests that this species is not present in thecountry (Chaves, 1987, 1993; Chaves & Torres,1993) .

Because of the first four bibliographic references,sorne countries hinder exports of certain agriculturalproducts and demand regularory certifications. How­ever, there are no objective proofs of the occurrence ofG. roslOchiensis in the country.

For the last two years, surveys have been in progressin order to demonstrate that specific areas of garlic,potata and onion production are free of G. rosw­chiensis (Del Toro el al., 1994a).

GENUS MELOIDOGYNE

As mentioned above, the discovery of a species ofthis genus marks the beginning of the study of plant­parasitic nematodes in Argentina. Since then, speciesof the genus Meloidogyne have been found on roots ofvarious plants: vegetables, potaroes, robacco, soybean,corton, ornamental plants, fruit trees, forest trees, andweeds.

Reviews of the existing information on the genushave recently been published (Doucet & Pinochet,1992; Doucet, 1993; Doucet & Ponce de Leon,1996b).

Nine species have been detected: M. acrita,iVI. arenaria (race 2), M. chitwoodi, M. cruciani)M. decalineata) M. hapla, M. incognita (races l, 2,

Fundam. appl. NemalOl.

and 3), M. javanica, M. ouersoni, in addition to sornepopulations reported as Meloidogyne spp. whose spe­cifie identity is unknown.

One hundred plant species (belonging ta 33 fami­lies) can be parasitised by the eight species of Meloi­dogyne. M. incognila and M. arenaria are the most fre­quently found species, and they parasitise 56 % and27 % of plants, respectively (Doucet, 1993). Culti­vated as weil as wild plants (native and imroduced)can be attacked by these nematodes, but mostly weedsand vegetables.

Although the genus is widely distributed and has avery wide range of hosts, populations, and damages,appear to be limited to sorne relatively small areas.

Major damages are observed in vegetables (particu­larly tomato and pepper), potata, grapevine, andsoybeans. They are managed by conventional nemati­cides controlling soil populations, crop rotations, andresistant varieties.

Potatoes present a particular challenge because thenematodes parasitise the tubers and can thus be trans­ported to new sites when contaminated tubers areused for seed. Because of the high reproductivepotential of these nematodes and their polyphagousbehaviour, planting pieces of infested tubers will cre­ate a serious problem to soils and future crops(Chaves & Torres, 1993; Doucet & Ponce de Leon,1996a). The legislation prohibits the use of infestedpotata seed, which helps preventing the dispersal ofthese nematades.

1'YLENCHULUS SEMIPENETRANS

7j;lenchulus semipenelrans was found for the first timein the North-East of the country on roots of citrustrees (Fresa, 1943). Later, it was found associatedwith different varieties of citrus in several regions,mainly the North-West and the North-East (Marchio­natto, 1945, 1946, 1947; Schultz, 1945; Gutierrez,1947; Lapez Cristobal, 1965; Moreno, 1969). In aHcases, the nematade was linked ta a decrease in citrusyields. The distribution of this nematade is very Iim­ited. The problems it causes are managed by resistantgraft (Doucet & Ponce de Leon, 1992; Costilla, pers.comm.).

FAMILY LONGIDORIDAE

So far, three genera have been detected: Xiphinema,Xiphidorus, and Paraxiphidorus. The first genus has alow diversity; only seven species are present in Argen­tina. On the contrary, ail the known species of the sec­ond genus are represented in the country to theexception of X. uruguayensis. The third genus includesonly the type species (Coomans & Chaves, 1995).

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Nematodes in continental Argenrina

Regarding the relative importance of these genera, itis necessary to mention the following species:

Xiphinema index

Xiphinema index was firs! found in the West ongrapevines infested with grape-vine fan leaf virus andgrape yellow mosaic virus (Feldman & Pontis, 1963,1964). This species is widely distributed in the wine­growing region of the Province of Mendoza (La Red& Vega, 1968; Cucchi el al., 1971; Vega, 1971, 1978).Ir was also found in grapevine plantations in the cen­tral part of the country - without any virus-relateddiseases - and on citrus roots in the North-East (Luc& Doucet, 1990).

The damages caused by this species on grapevineswere controlled by nematicides (La Red & Vega,1970).

Xiphinema americanum sensu law

X. americanum sensu law is the species of the genusXiphinerna the most frequently found in the country.It is associated with a great variety of crops (tabacco,citrus, sugar cane, grapevine, barley, tomato, severalfruit trees) as weil as wild plants. Ir is widely distrib­uted (Luc & Doucet, 1990). It is not known whetherthe populations of this species can transmit patho­genic viruses.

Main genera and species of entomopathogenicnematodes recorded in Argentina

FAMILY MERMITHIDAE

From the results of the studies made so far, it can besaid that the Mermithidae family is characterised by agreat generic and specifie diversity in the country. Fif­teen genera have been observed: three terres trial andtwelve aquatic genera. Two of them were found forthe first time in Argentina: Dilremarnermis Camino &Poinar, 1989 and Divisispiculimermis Doucet, 1986.The best represented genera are Mesomermis and Gas­tromermis, both including mostly species parasites ofsimulids.

So far, Slrelkovimermis spiculalUs - a parasite of mos­quitaes - is the only species srudied in the laborataryto obtain information on its physiology and modalitiesof infection (Camino, 1988b,c; Camino & Garcia,1991,1992).

Among terres trial mermithids, the most remarkableare the genus Hexamermis including the largest num­ber of species and the species Agamermis decaudatabecause of its large geographic distribution (Caminoel al., 1986; Doucet & Cagnolo, 1995b).

FAMILY HETERORHABDITIDAE

The genus Heterorhabdilis is represented by the spe­cies H. argeminiensis and H. baeteriophora. Only one

529

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population of the first species has been observed(Stock, 1993a) whereas several populations of thesecond species have been found in various locations inthe country (Doucet & Bertolotti, 1995; 1996).

In the laboratory, two H. bacleriophora populationsfrom C6rdoba have shown promises as efficient bio­logica! control agents against Lepidoptera andColeoptera (Scherma & Rodriguez Mosquera, 1989;Doucet & Giayetto, 1994).

FAMILY STEINERNEMATIDAE

So far, six species of the genus Sleinernema areknown in Argentina. S. felriae was found in the SantaFe province (Stock, 19936), S. scaprerici in BuenosAires (Stock, 1992), and S. carpocapsae, S. glaseri,S. rara, and S. rilleri were found in C6rdoba.

Except for S. glaseri, which was observed in moun­tain forest soil, the Sleinernerna species were found incultivated soils.

Two species, S. carpocapsae and S. rara couldbecome good control agents of harmful Homoptera,Lepidoptera and Coleoptera (Doucet, 1981).

Conclusion

Our knowledge of the soil nemarodes that arepresent in the country is still insufficient. Because ofthe great diversity of conditions, it is reasonable tosuppose that there is a marked diversity of species. Itmay be necessary to confirm the previous identifi­cation of some of the species by means of detailedstudies.

Concerning the species of plant-parasitic nema­rodes, it remains necessary to assess more accuratelythe impact they have on the crops they attack.Because of a lack of symproms, the attack of thesepathogens sometimes goes unnoticed or is ascribed toother agents.

In those cases in which the harmful associationnematode-plant is confirmed, it is essential to obtainmore information on the biology of these species. Theknowledge of the type of behaviour and the character­isation of the different populations of particular spe­cies (particularly Meloidogyne spp. and Nacobbusaberrans) will allow the development of strategiesaimed at ensuring plant production and preservinggood soil conditions.

In Argentina, research remains aimed at the studyof the major species of plant-parasitic nematodesassociated with potato and soybean and with vegeta­bles in general, and roward the development of differ­ent types of control measures.

In relation ro entomopathogenic nemarodes, theobjective of the investigations is to increase our know­ledge on the species present, on the aggressivity ofdifferent populations, and to evaluate their potentialsas biological control agents.

530

Acknowiedgements

The authors thank the Consejo de Investigaciones Cienti­ficas y Técnicas (CONICET), the Consejo de Investiga­ciones Cientificas y Tecnologicas de la Provincia deC6rdoba (CONICOR), the Secretaria de Ciencia y Técnicaof the Universidad Nacional de C6rdoba and the Comisi6nAdministradora dei Fondo de Promoci6n de la TecnologiaAgropecuaria (CAFPTA) for their fmancial support thatmade it possible to obtain part of the information reviewedhere. They are also grateful to the members of the: SociedadArgentina de Nematologia, Biblioteca dei Ministerio deAgricultura de la Naci6n and Archivo Historico de laMunicipalidad de C6rdoba, who collaborated in the searchand compilation of bibliographie references. They alsothank Dr. M. Luc for his valuable suggestions and com­ments.

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DOUCET, M. E. & DE PONCE DE LEON, E. L. (1986). Che­nopodium album L.: eficiente hospedador de Nacobbusaberrans (Thome, (1935) Thorne & Allen, 1944 y Meloi­dogyne javanica (Treub, 1885) Chitwood, 1949 en la Pro­vincia de C6rdoba. Inftivo Inveslig. agropec., 437-440: 36­43.

DOUCET, M. E. & DE PONCE DE LEON, E. L. (1992). Tylen­chulus semipenelrans Cobb, 1913 (Nemata: Tylenchida) ysu asociaci6n con Citrus auramium L. y Citrus reshrli Hort.ex Tan. Agriscientia, 9: 113-116.

DOUCET, M. E. & DE PONCE DE LEON, E. L. (1994). Aso­ciaci6n de Cruznema lripartir:um con plantulas clor6ticasde ajo. NemalOl. medit., 22: 149-151.

DOUCET, M. E. & DE PONCE DE LEON, E. L. (l996a).Meloidogyne spp. (Nematoda): una seria amenaza para lapapa Solanum wberosum L. RevLa Investig. agropec., 26:45-51.

DOUCET, M. E. & DE PONCE DE LEON, E. L. (1996b).Alteraciones histol6gicas ocasionadas por Meloidogyneincognita (Kofoid & White, 1919) Chitwood, 1949 enraices de plantas omamentales florales. RevLa Investig.agropec., 26: 83-91.

DOUCET, M. E. & DE PONCE DE LEON, E. L. & COSTILLA,M. A. (1992). Histopatologia en CapsiCLIm annuum L.inducida por Nacobbus aberrans (Thorne, 1935) Thome &Allen, 1944. RevLa Investig. agropec., 23: 69-76.

DOUCET, M. M. A. DE (1981). Neoap/eClana carpocapsaeWeiser (Steinemematidae) su posible utiJizaci6n comoagente en el control de Zulia emrerriana (Berg) (Cercopi­dae). 4lh Jorn. fùosan. Arg. Cordoba, 19-21 AuguSI 1981:149-150. [Abstr.]

DOUCET, M. M. A. DE (1982). Una nueva especie de Gas­lromermis Micoletzky, 1923 (Nematoda: Mermithidae)parasito de Simulium wolffhengeli (Enderlein) (Diptera:Simulidae). Revr.a M useo A rg. Cienc. nal. "BernardinoRivadavia", 2: 11-17.

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Nemacodes in continerual A rgentina

DOUCET, M. M. A. DE (1986a). Divisispiculimermis mirus n.gen., n. sp., (Nematoda: Mermithidae) parasitizingmidges in C6rdoba (Argentina).]. NemaLOl., 18: 247-251.

DOUCET, M. M. A. DE (1986b). A new species of Neoaplec­lana Steiner, 1929 (Nematoda: Steinernematidae) fromC6rdoba, Argentina. Revue NémaLOI., 9: 317-323.

DOUCET, M. M. A. DE (1992). Distribuci6n de nematodosentom6fagos en suelos de C6rdoba, Argentina. 6lh Congr.latinoam. Fitopal. & 6lh Congr. nac. Soc. Esp. Fitopal. Tar­remolinos, Espar/a, 8-12 Apnl1992: 112. [Abstr.]

DOUCET, M. M. A. DE (1995). Caracterizacion de unapoblacion de Sleinernema carpocapsae (Nematoda: Stein­emematidae) aislada en Cordoba, Republica Argentina.NemaLOI. medit., 23: 181-189.

DOUCET, M. M. A. DE & BERTOLOTTI, M. A. (1995). Newrecords of entomogenous nematodes (Steinernematidae;Heterorhabditidae) in C6rdoba, Argentina. Congr. inl.Nem acol. 1rop. , Rio Queme, BraS/l, 4-9 June 1995: 74.[Abstr.]

DOUCET, M. M. A. DE & BERTOLOTTI, M. A. (1996). Unanueva poblaci6n de Heterorhabdùis bacteriophora Poinar,1975 de Argentina. Caracterizacion y accion sobre elhuésped. Nem acol. medit., 24: 169-174.

DOUCET, M. M. A. DE & CAGNOLO, S. (1995a). Diversityof mermithids in Cordoba, Argentina. Congr. im. Nema­col. lrop., Rio Queme, Brasil, 4-9 June 1995: 75. [Abstr.]

DOUCET, M. M. A. DE & CAGNOLO, S. (1995b). Agamer­mis decaudaw Cobb, Steiner y Christie, 1923 (Nematoda)parasito de is6podos y acrididos de C6rdoba, Argentina.9lh Jorn. fitosan. Arg., Mendoza, 13- 15 November 1995:110. [Abstr.]

DOUCET, M. M. A. DE, CAGNOLO, S. R. & BERTOLOTTI,M. A. (1996). New Heterorhabditis bacleriophora Poinar,1975 from Argentina. Variability and description of her­maphrodites from second and third generation. Fundam.appl. NemalOl., 19: 415-420.

DOUCET, M. M. A. DE, CARO, K., MIRANDA, M. &BERTOLOTTI, M. A. (1996). Efficacy of Heterorhabditisbacleriophora (strain OU) in relation to temperature, con­centration and origin of the infective juvenile. Nemalro­pica, 26: 129-133.

DOUCET, M. M. A. DE & DOUCET, M. E. (1986). Nuevosdaros para el conocimiento de Heterorhabditis bacteriophoraPoinar, 1975. Revta Invesllg. agropec., 21: 1-10.

DOUCET, M. M. A. DE & DOUCET, M. E. (1989).Cruznema lripanill/m (Linstow, 1906) Zu!lini, 1982: unnuevo nematodo entomofago facultativo. 7lh Jorn. fiLOsan.Arg., Salta, 5-8June 1989: 1 p. [Abstr.]

DOUCET, M. M. A. DE & DOUCET, M. E. (1990). Descrip­tion of Sleinernema rilleri n. sp. (Nemaroda: Steinemema­tidae) with a key to the species of the genus. NemaLOlogica,36: 257-265.

DOUCET, M. M. A. DE & DOUCET, M. E. & BERTOLOTTI,M. A. (1991). Efecto de la temperatura de cria e inten­sidad de infestaci6n sobre los caracteres morfométricos deHelerorhabditis baCleriophora Poinar, 1975. Analisis devariabilidad. Nemalropica, 21: 37-49.

DOUCET, M. M. A. DE, DOUCET, M. E. & DI RIENZO, J.(1990). Analisis de variabilidad de los caracteres morfo-

533

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métricos de Helerorhabditis bacleriophora Poinar, 1975.Nemalropica, 20: 4-5.

DOUCET, M. M. A. DE, DOUCET, M. E. & DI RIENZO, ].(1992a). Diferencias entre las larvas infectantes de Het.e­rorhabditis bacœriophora Poinar, 1975 (Nemata: Hete­rorhabditidae) segùn su generaci6n de origen. Nemalrop­ica, 22: 121-122.

DOUCET, M. M. A. DE, DOUCET, M. E. & NIENSTEDT,K. M. (1992b). Diferencias inter e intraespecificas en lacapacidad infectiva de poblaciones de Het.erorhabditis ySœinernema aislados en Argentina. NernaLropica, 22: 237­242.

DOUCET, M. M. A. DE & GIAYETTO, A. (1994). Gama dehuéspedes y especificidad de Heœrorhabditis bact.eriophoraPoinar, 1975 (Nematoda: Heterorhabditidae). NemalOl.medù., 22: 171-178.

DOUCET, M. M. A. DE & POINAR, G. O., JR. (1984). Gas­lrOlIIermlS kolleonis n. sp. (Nematoda: Mermithidae) a par­asite of midge (Chironomus sp., Chironomidae) fromArgentina. J NemalOl., 16: 252-255.

DOUCET, M. M. A. DE & POINAR, G. O., JR. (1985). Estu­dia dei ciclo de vida de una poblaci6n de HeœrorhabdiLissp. proveniente de Rio Cuarto, Provincia de C6rdoba.Revla Univ. nac. Rio Cuano, 5: 253-258.

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Appendix

Lisl of sail nemalOdes reporœd in continental A rgemina (doublful records are marked with an asterish (*j).

Class AdenophoreaSubclass EnopliaOrder Enoplida

Family Tripylidae

TrischislOma monohysleroides Altherr, 1963TrischislOma monohystera (de Man, 1880) SchuurmansStekhoven, J951Tripyla sp.Tobrilus maerospiculum Altherr, 1963Tobrilus papil/icaudatus Altherr, 1963Tobrilus sewsus Altherr, 1963Tobrilus sp.

Family Ironidae

ironus filiformis Altherr, 1963

Order Mononchida

Family Cobbonchidae

Cobbonchus regulus Altherr, 1963

Family Mononchidae

Clarkus papillatus (Bastian, 1865) Jairajpuri, 1970Coomansus ger/ache! (de Man, 1904) Jairajpuri & Khan,1977Mononchus aqualù;us Coetzee, 1968Mononchus bel/us Andnissy, 1985MonOr/chus sp.Prionchulus muscorum (Du;ardin, 1845) Wu & Hoeppli,1929

Family Mylonchulidae

Mylonchulus brachyuris (Bütschli, 1873) Altherr, 1953Mylonchulus hawaiiensis (Cassidy, 1931) Andrâssy, J958Mylonchulus minor (Cobb, 1893) Andrâssy, 1958Mylonchulus parabrachyurus (Thome, 1924) Andrâssy,1958Mylonchulus sigmalUrus (Cobb, 1917) Alrherr, 1953Mylonchulus sp.Sporonchulus ibitiensis (Carvalho, 1951) Andrâssy, 1958

Family IotOnchidae

IOlOnchus arenicola Altherr, 1963IOlOnchus geminis Heyns & Lagerwey, 1965IOfOnchus monhyslera (Cobb, 1917) Jairajpuri, 1970

536

Order Mermithida

Family Mermithidae

Agamermis decaudata Cobb, Steiner & Christie, 1926Amphimermis bonariensis Miralles & Camino, 1983Balhymermis simuliae Camino, 1993Dùremamermis simuliae Camino & Poinar, 1989Divisispiculimermis rnirus Doucet, 1986Gaslromermis cordobensis Camino, 1991Gaslromermis doloresi Camino, 1993Gaslromermis fidelis Doucet, 1982Gasll'omermis kol/eonis Doucet & Poinar, 1984Gaslromermis vaginiferous Camino, 1985Hexamermis coch/earius StOck & Camino, 1992Hexamermis hortensis Cami no & StOck, 1989Hexamermis macroswma Camino & StOck, 1994Hexamermis ovislriata StOck & Camino, 1992Hydromermis sp.Hydromermis doloresi Camino, 1993Isomermis sp.Isomermis sierrensis Camino, 1994Limnomermis bonariensis Camino, 1989Limnomermis sp.Mermis sp.Mesomermis crassivaginae Camino, 1985Mesomermis dissimilis Camino, 1985Ivlesomermis delponteiana Camino, 1990Mesomermis nonensis Camino, 1991Mesomermis ochrae Cami no, 1985Ivlesomermis subandina Camino, 1985Mesomermis sp.OClOmyomermis albicans Camino, 1985OClOmyomermis arecoensis Camino, 1991OClOmyomermis bonariensis Camino, 1988OClOmyomermis longispiculae Camino, 1992Pseudomermis sp.Slrelkovimermis spiculalus Poinar & Camino, 1986Slrelkovimermis sp.

Order Dory1aimida

Family Actinolaimidae

AClinolaimus sp.

Family Aporcelaimidae Heyns, 1965

Aporcelaimel/us obscurus (Thome & Swanger, 1936)Heyns, 1965

Fundam. appl. Nemawl.

Aporcelaimellus sp.Aporcelaimus sp.

Family Belondiridae

Dorylaimellus moneicolus Clark, 1963Dorylaimellus virginianus Cobb, 1913

Family Charcharolaimidae

Carcharolaimus fOl"mosus Lordello, 1957Carcharolaimus rocundicauda (de Man, 1880) Thome,1939Carcharolaimus sp.

Family LeptOnchidae

LeplOnchus gramdosus Cobb, 1920LeplOnchus sp.Tylencholaimellus montanus Thome, 1939

Family Longidoridae

Paraxiphidorus michelluci Coomans & Chaves, 1995Paraxiphidorus sp.Xiphidorus achalae Luc & Doucer, 1984Xiphidorus balcarceanus Chaves & Coomans, 1984Xiphidorus tucumanensis Chaves & Coomans, 1984Xiphidorus saladillensis Chaves & Coomans, 1984Xlphidoms yepesara Monreiro, 1976Xlphidorus sp.Xiphinema americanum Cobb, 1913Xlphinema diversicaudacum (Micoletzky, 1927) Thome,1939Xlphinema index Thome & Allen, 1950Xlphinema krugi Lordello, 1955Xlphinema sewriae Luc, 1958Xlphinema rivesi Dalmasso, 1969Xiphinema surinamense Loof & Maas, 1972Xlphinema sp.

Family Mydonomidae

Dorylaimoides elongalUs Husain & Khan, 1968

Family Nordiidae

Pungemus monhystera Thome & Swanger, 1936

Family Dorylaimidae

Dorylaimus sp.Mesodorylaimus aberrans Loof, 1969Mesodorylaimus adalberti Andrassy, 1963Mesodorylaimus argentinus Alrherr, 1963Mesodorylaimus demi Loof, 1969Mesodorylaimus meridianlls Andrassy, 1963Mesodorylaimus pseudobastiani Loof, 1969Mesodorylaimus puellae Andrassy, 1963Mesodorylaimus szekessyi Andrassy, 1960Mesodorylaimus sp.

Family Nygolaimidae

Nygolaimus sp.

Family Qudsianemaridae

Eudorylaimus monohystera (de Man, 1880) Andrassy, 1959

Vol. 20, nO 6 - 1997

NemalOdes in continenwl Argentina

Eudorylaimus obcusicalldacus (Basrian, 1865) Andrassy,1959Eudorylaimus profeslUs Andrassy, 1963Eudorylaimus quadramphidius Andrassy, 1963Eudorylaimus sp.Labronema sp.

Order Triplonchida

Family Diphrerophoridae

Dzphterophora communis de Man, 1880Diphterophora sp.

Family Trichodoridae

Paratrichodorus minor (Colbran, 1956) Siddiqi, 1974Paratrichodorus sp.Trichodorus obscurus Allen, 1957Trichodorus sp.

SubcIass ChrornadoriaOrder Araeolairnida

Family Plecridae

AnapleclUs sp.Pleetus (Pleetoides) pazagonieus de Man, 1904Pleetus sp.

Order Chrornadorida

Family Cyatholaimidae

Odomolaimus aquacicus \Xl Schneider, 1937

Order Monhysterida

Family Monhysreridae

Monhystera lepidura Andrassy, 1963Monhystera sp.

Class SecernenteaSubcIass RhabditidaOrder Rhabditida

Family Cephalobidae

Acrobeles sp.Acrobeles emmatus Shahina & De Ley, 1997Acrobeloides sp.Cephalobus sp.Cervidellus sp.Eucephalobus sp.Nothacrobeles lunensis Shahina & De Ley, 1997Zeldia sp.

Family Rhabdiridae

Cephaloboides sp.Cruznema crzpartùum (LiRstOw, 1906) Zullini, 1982Cruznema sp.DislOlabrellus veechi Anderson, 1983Oxycerca brevispina (de Man, 1895) Andrassy, 1983Pellioduis pellio (Schneider, 1866) Timm, 1960

537

M. E. Doucet & M. M. A. de Doucet

Rhabdùis spp.

Family Heterorhabditidae

Heterorhabdùis argeminiensis S rock, 1993Heœrorhabdùis baCleriophora Poinar, 1975Heterorhabdùis sp.

Family Steinemematidae

Sœinernema carpocapsae (\X1eiser, 1955) Poinar, 1990Sœinernema felliae (Filipjev, 1934) Poinar, 1990Steinernema glaseri (Steiner, 1929) Poinar, 1990Steinernema rartlm (Doucet, 1986) Poinar, 1990Steinernema rilleri Doucet & Doucet, 1990Steinernema scapterisCl Nguyen & Smart, 1991Steinenzema sp.

Family Panagrolaimidae

Panagrolaimus sp.

Order Ascaridida

Family The1asromatidae

Leidynema appendieulawm (Leidy, 1890) Chitwood, 1932Pseudonymus sp.ThelaslOma desselae Adamson, 1985

Order Diplogasterida

Family Diplogastridae

Micoletzkya vidalae Srock, 1993

Order Aphelenchida

Family Aphelenchidae

Aphelenchus avenae Bastian, 1865Aphelenchus sp.

Family Aphelenchoididae

Aphelenchoides bicaudaws (lmamura, 1931) Filipjev &Schuurmans Stekhoven, 1941Aphelenchoides blaslOphthorus Franklin, 1952Aphelenchoides fragariae (Ritzema Bos, 1890) Christie,1932Aphelenchoides parietinus (Bastian, 1865) Steiner, 1932Aphelenchoides riizemabosi (Schwartz, 1911) Steiner, 1932Aphelenchoides sp.Laimaphelenchus cocuccii Doucet, 1992Laimaphelenchus sp.Seimlra sp.

Family Ektaphelenchidae

Ektaphelenchus tenuidens (Thorne, 1935) Rühm, 1956

Order Ty1enchida

Family Tylenchidae

Atylenchus sp.Arylenchus decalinealUs Cobb, 1913Boleodorus sp.

538

Cephalenchus sp.Coslenchus sp.Filenchus sp.Lelenchus sp.Psilenchus bahiablancae Doucet, 1996Psilenchus hilarus Siddiqi, 1963Psilenchus praœnsis Doucet, 1996Psilenchus sp.Tetylenchus sp.Tylenchus sp.Tylenchus davanei Bastian, 1985Tylenchus ftlzformis IVl.icoletzky, 1922

Family Anguinidae

Anguina agrostis (Steinbuch, 1799) Filipjev, 1936Anguina sp.Dùylenchus dipsaci (Kühn, 1857) Fiiipjev, 1936Dùylenchus imermedius (de Man, 1880) Filipjev, 1936Dùylel1chus sp.Orrina phyl/obia (Thome, 1934) Thome, 1961

Family Beiono1aimidae

Belol1olaimus gracilis Steiner, 1949 *Belol1olaimus brevicaudaws (Doucet, 1983) Fortuner &Luc, 1987Tetylenchus sp.Triversus feslOnaws (Doucet, 1985) Fortuner & Luc, 1987Tylenchorhynchlls acti Hooper, 1959Tylenchorhynchus annulalUs (Cassidy, 1930) Golden, 1971Tylenchorhynchus capùalUs Allen, 1955Tylenchorhynchus cylindricus Cobb, 1913Tylenchol'hynchus dubius (Bütschli, 1837) Filipjev, 1936Tylenchorhynchlls sp.

Family Dolichodoridae

Dolichodorus aquatulis Doucet, 1985Dolichodorlls heterocephalus Cobb, 1914 *Dolichodorlls longicaudatus Doucet, 1981Neodolichodorus leiocephalus Doucet, 1981

Family Prarylenchidae

Pralylenchus agilis Thome & Malek, 1968Pratylenchus brachyurus (Godfrey, 1929) Filipjev &Schuurmans Stekhoven, 1941Pratylenchus delaarei Luc, 1958Pratylenchus goodeyi Oostenbrink, 1953Pratylenchus hexincisus Taylor & Jenkins, 1957Pratylenchlls microstylus Bajaj & Bahatti, 1984Pratylenchus neglectus (Rensch, 1924) Filipjev & Schuur­mans Stekhoven, 1941Pratylenchlls penetrans (Cobb, 1917) Filipjev & Schuur­mans Srekhoven, 1941Pratylenchus praterlSis Meyl, 1954Pratylenchus pseudopratensis Seinhorst, 1968Pratylenchus scribneri Steiner, in Sherbakoff & Stanley,1943Pratylenchus thornei Sher & Allen, 1953Pratylenchus vulnus Allen & Jensen, 1951Pratylenchus zeae Graham, 1951

Fundam. appl. NemalOl.

Pralylenchus sp.Radopholus similis (Cobb, 1893) Thome, 1949Nacobbus aberrans (Thome, 1935) Thome & Allen, 1944

Family Hoplolaimidae

Aorolaimus longislylus (Doucet, 1980) Fortuner, 1987Aorolaimus perscilus (Doucet, 1980) Fortuner, 1987Aorolaimus conicori (Doucet, 1984) Fortuner, 1987Aorolaimus brevicaudaLUS (Doucet, 1984) Fortuner, 1987Aorolaimus lriticeus (Doucet, 1984) Fortuner, 1987Aorolaimus sp.Helicolylenchus nannus Steiner, 1945Helicolylenchus digonicus Perry, in Perry, Darling &Thome, 1959Helicolylenchus multicincLUs (Cobb, 1893) Golden, 1956Helicolylenchus pseudorobusLUS (Steiner, 1914) Golden,1956H elicorylench us sp.Hoplolaimus galealUs (Cobb, 1913) Thome, 1935Hoplolaimus sp.Rorylenchus buxophilus Golden, 1956ROlylenchus goodeyi Loof & Oosrenbrink, 1958Rorylenchus robuslUs (de Man, 1876) Filip;ev, 1936Rorylenchus ul1lformis (Thome, 1949) Loof & Oosren­brink, 1958Rorylenchus sp.

Family Heteroderidae

CaclOdera amaranlhi (Sroyanov, 1972) Kra Il' & Krall',1978CaclOdera sp.Globodera roslOchiensis (Wollenweber, 1923) Behrens,1975 *Globodera labacum labacum (Lownsberry & Lownsberry,1954) Behrens, 1975Globodera sp.Helerodera schachlii Schmidt, 1871 *Helerodera galeopsidis Goffart, 1936Helerodera spp.J\![eloidogyne acrita Chitwood, 1949Meloidogyne arenaria (Neal, 1889) Chitwood, 1949Meloidogyne chitwoodi Golden, O'Bannon, Santo & FinJey,1980Meloidogyne cruàani Garcia-Martinez, Taylor & Smart,1982Meloidogyne decalineata Whitehead, 1968Meloidogyne hapla Chitwood, 1949Meloidogyne incognita (Kofoid & White, 1919) Chitwood,1949Meloidogyne javanica (Treub, 1885) Chitwood, 1949Meloidogyne naasi Franklin, 1965Meloidogyne ollersoni (Thome, 1969) Franklin, 1971

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NemalOdes in continental Argentina

Meloidogyne sp.

Family Criconematidae

Criconema dupliciveslitum (Andrassy, 1963) Raski & Luc,1985Criconema sp.Nfesocriconema curvaLUm (Raski, 1952) Loof & De Grisse,1989Mesocriconema douceti (Siddiqi, 1986) Ebsary, 1991Mesocriconema ornaLUm (Raski, 1958) Loof & De Grisse,1989Nfesocriconema peruenslforme (De Grisse, 1967) Loof & DeGrisse, 1989Mesocriconema rilleri (Doucet, 1980) Loof & De Grisse,1989Mesocriconema sphaerocephala (Taylor, 1936) Loof & DeGrisse, 1989Mesocriconema talense (Chaves, 1983) Loof & De Grisse,1989Mesocriconema xenoplax (Raski, 1952) Loof & De Grisse,1989Mesocriconema sp.NOlhocriconema mUlabile (Taylor, 1936) De Grisse &Loof,1965NOlhocriconema sp.Caloosia paradoxa (Luc, 1958) Brzeski, 1974Hemicycliophora arenaria Raski, 1958Hemicycliophorafilicauda Doucet, 1982Hemicycliophora fragilis Doucet, 1982Hemicycliophora gracilis Thome, 1955Hemicycliophora penelrans Thome, 1955Hemicycliophora poranga Monteiro & Lordello, 1978Hemicycliophora rara Doucet, 1983Hemicycliophora rionegrensis Doucet, 1982Hemicycliophora ripa Van den Berg, 1981Hemicycliophora 1enUiSlriata Doucet, 1982Hemicycliophora lhienemanni (Schneider, 1925) Loos,1948Hemicycliophora zuckermani Brzeski, 1963Hemicycliophora sp.Ogma afternaLUm (Doucet, 1986) Raski & Luc, 1987Ogma comahuense Brugni & Chaves, 1994Ogma muftisquamaLUm (Kirjanova, 1948) Mehta & Raski,1971

Family Tylenchulidae

Gracilacus colina Huang & Raski, 1986Paralylenchus neoamblycephalus Geraert, 1965Paralylenchus sp.Tylenchulus semipenelrans Cobb, 1913

Family Allantonematidae

Metaparasitylenchus sp.

539