Weed Technology 2007 21:310323

Status and Management of Grass-Weed Herbicide Resistance in Latin America

Bernal E. Valverde*
Twenty-one grass weeds have evolved resistance to herbicides in Latin America, particularly in rice, soybean, wheat, and orchards. Junglerice, the most widespread and economically important rice weed, evolved resistance to propanil, acetylcoenzyme A carboxylase (ACCase)-inhibitor herbicides, quinclorac, and imazapyr in Central America, Colombia, and Venezuela. Some junglerice populations are resistant to at least three herbicide modes of action. Other herbicide-resistant (HR) rice weeds are barnyardgrass and gulf cockspur to quinclorac in Brazil, and saramollagrass to ACCase-inhibitor herbicides in Colombia and bispyribac in Venezuela. Populations of weedy rice resistant to imidazolinones are now emerging, most likely originated from gene flow from imidazolinone-resistant rice. Saramollagrass also became resistant to nicosulfuron in corn in Venezuela. Eight species associated with soybean are resistant to ACCase-inhibitor herbicides in Brazil (alexandergrass, goosegrass, and southern crabgrass) and Bolivia (Louisiana cupgrass, itchgrass, sudangrass, and two common wild sorghum species). Four more ACCase-inhibitorresistant species (hedgehog dogtailgrass, wild oat, rigid ryegrass, and Italian ryegrass) are found in Chile infesting canola and wheat. ACCase-inhibitorresistant hood canarygrass, littleseed canarygrass, and wild oat are important in wheat in Mexico. Resistance to acetolactate synthase (ALS)-inhibitor herbicides has been reported in itchgrass, goosegrass, and Mexican grass. Italian ryegrass populations resistant to glyphosate have been found in Chile and Brazil. Glyphosate resistance has also evolved in goosegrass in Bolivia and johnsongrass in Argentina. In general, little is done to prevent resistance evolution. An exception is the stewardship programs aiming to prevent gene flow from imidazolinone-resistant rice to weedy rice. Once resistance evolves, HR populations are mostly managed by shifting to herbicides with different modes of action and, in some cases, by slightly modifying agronomic practices. Propanil formulations containing a synergist are used to manage propanil-resistant junglerice. Increased no-till agriculture and planting of glyphosate-resistant crops are likely to select more glyphosate-resistant weeds. Nomenclature: Bispyribac; glyphosate; imazapyr; nicosulfuron; propanil; quinclorac; alexandergrass, Brachiaria plantaginea (Link) A. S. Hitchc. BRAPL; barnyardgrass, Echinochloa crus-galli (L.) Beauv. ECHCG; common wild sorghum, Sorghum verticilliflorum (Steud.) Stapf. and S. saccharatum (L.) Moench; goosegrass, Eleusine indica (L.) Gaertn. ELEIN; gulf cockspur, Echinochloa crus-pavonis (Kunth) J. A. Schultes. ECHCV; hedgehog dogtailgrass, Cynosurus echinatus L. CYXEC; hood canarygrass, Phalaris paradoxa L. PHAPA; Italian ryegrass, Lolium multiflorum Lam. LOLMU; itchgrass, Rottboellia cochinchinensis (Lour.) W. D. Clayton. ROOEX; johnsongrass, Sorghum halepense (L.) Pers. SORHA; junglerice, Echinochloa colona (L.) Link ECHCO; littleseed canarygrass, Phalaris minor Retz. PHAMI; Louisiana cupgrass, Eriochloa punctata (L.) Desv. ex Hamilt. ERBPO; Mexican grass, Ixophorus unisetus (Presl) Schult. SETUN; rigid ryegrass, Lolium rigidum Gaudin. LOLRI; saramollagrass, Ischaemum rugosum Salisb. ISCRU; southern crabgrass, Digitaria ciliaris (Retz.) Koel. DIGSP; sudangrass, Sorghum sudanense (Piper) Stapf. SORSU; weedy rice, Oryza sativa L. ORYSA; wild oat, Avena fatua L. AVEFA; canola, Brassica napus L.; corn, Zea mays L.; rice, Oryza sativa L.; soybean, Glycine max (L.) Merr.; wheat, Triticum aestivum L. Key words: herbicide resistance management.

Twenty-one grass weeds have evolved resistance to herbicides in Latin America (Table 1). The most widely studied species and cases of herbicide resistance are those of junglerice and other Echinochloa species in Central America and northern South America, and acetolactate synthase (ALS, EC 4.1.3.18) and acetyl coenzyme-A carboxylase (ACCase, EC 6.4.1.2)-inhibitor resistance in several species in Brazil and Chile. Although there is no account of the economic impact of herbicide resistance in Latin America, some herbicideresistant (HR) weeds are agriculturally relevant. In Central America, an estimated 60% of the rice area is infested with propanil-resistant junglerice. Farmers still spend about US$18 million on propanil (regular formulations) applied under these conditions, mostly because of its low cost, crop selectivity, and good efficacy on other weeds. Alternative

DOI: 10.1614/WT-06-097.1 * Visiting Scientist, Faculty of Life Sciences, The University of Copenhagen, 13, Taastrup, DK-2630, Denmark. E-mail: bev@life.ku.dk Hojebakkegaard Alle and ideatrop@ice.co.cr

herbicides to control propanil-resistant junglerice, such as ACCase- and ALS-inhibitor herbicides, are more expensive and have already imposed enough selection pressure for some populations to evolve HR biotypes (Valverde and Itoh 2001). It is estimated that more than 2 million ha are infested with HR weeds in Brazil, resulting in crop damage of about US$80 million per year (Vidal et al. 2006a). About 100,000 ha of the 400,000 ha of wheat planted in Chile have HR grass weeds at varying infestation levels (N. Espinoza, personal communication). This article is an updated account of known cases of HR grasses in Latin America and covers aspects related to the selection of resistant populations, resistance mechanisms, and management options. For this purpose, I have covered information published in refereed international as well as regional and local journals, articles and reports in Spanish and Portuguese of more limited distribution, conference abstracts and extended summaries, and undergraduate and graduate theses. Additional information comes from conversations and written communication with Latin American researchers and from my own studies and experience. Where the information

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is available electronically, I have tried to provide reliable links to be easily accessed. Presentation of resistance cases is organized by herbicide mode of action.
Resistance to Photosystem-II Inhibitors

Propanil resistance in junglerice is the only confirmed case in Latin America of resistance to herbicides that inhibit the electron transport in photosystem II (PS II). Indeed, this was the first case of herbicide resistance in a grass species reported in this region. Resistance to propanil was found almost at the same time in the late 1980s in Costa Rica and Colombia (Fischer et al. 1993; Garro et al. 1991). It was later confirmed in neighboring Central American countries (Valverde et al. os-Espinoza 1999, Villa2000), Mexico (Valverde and Bolan Casarez 1998), and Venezuela (Ortiz et al. 2000) but not in Bolivia (Valverde and Itoh 2001). A review provides detailed information of the extent, spread, importance, and management of propanil-resistant junglerice in Central America (Valverde et al. 2000). Resistance evolved particularly in areas of continuous rice production, where propanil was applied (1) more than once per crop cycle, (2) usually alone or in mixture with herbicides not active on junglerice, (3) at increasing doses, and (4) after 20 or more yr of use; the herbicide had been introduced in Latin America in the early 1960s. Failure to control junglerice at normal propanil doses (1.9 to 2.2 kg/ha) prompted farmers to use higher doses (up to 5.7 kg/ha) and two to three applications during the cropping season. The persistent selection pressure by propanil applied at increasing doses is most likely responsible for the great variation in resistance levels observed among populations collected in the field; some populations have low levels of resistance, whereas others cannot be controlled with extremely high herbicide doses. It is still unclear if these differences in resistance levels are related to gene amplification or other multifactorial mechanisms. In the early 1980s, systemic graminicides that inhibit ACCase became commercially available for use in dicotyledonous crops. Soon after, it was found that some of these herbicides could be used with an acceptable selectivity level to control grasses in rice. Fenoxaprop, and to a lesser extent sethoxydim, became very popular in Costa Rica and other countries because they eliminated propanil-resistant junglerice and also allowed late postemergence control of other grasses. Overreliance on fenoxaprop soon resulted in complaints from farmers about its performance and to the discovery of fenoxaprop-resistant junglerice populations. Some of these populations were resistant to propanil and fenoxaprop, whereas others were resistant to just one of them (Caseley et al. 1997). Fenoxaprop-resistant populations from Costa Rica were cross-resistant to other ACCase herbicides, including fluazifop-P, quizalofop, cyhalofop, cycloxydim, and sethoxydim; some of these at the time had not yet been commercially introduced (Caseley et al. 1997; Knights 1995; Riches et al. 1996). Where the initial fenoxaprop-resistant populations were found, quinclorac was an option for control. Its limited weed spectrum and comparatively higher cost, however, prevented its widespread use and did not result in the selection of quinclorac-resistant junglerice in Costa Rica. In Colombia,

however, quinclorac-resistant junglerice populations have been found (Valverde and Itoh 2001). Faced with increasing resistance problems, farmers began to worry about their production schemes but were limited in their management options, especially crop rotation. Where rice is grown under irrigation with heavy investment in field leveling and construction of levees, rotation with row crops is seldom practiced because of the cost of rebuilding the infrastructure. Of several agronomic practices, the stale seedbed preparation was found to be an excellent option for managing HR junglerice. Typically junglerice germinates in flushes, at least two and often three (Valverde et al. 2000). By preparing the soil in advance or delaying seeding to allow killing the first cohorts with a total, nonselective herbicide (usually glyphosate) or with light tillage, rice seedlings become established in association with diminished junglerice densities. This practice was already adopted by farmers to control weedy rice in fields that were heavily infested. Junglerice plants then emerging with the rice crop or soon after could be controlled with preemergence or early-postemergence herbicides. Of several herbicides tested, pendimethalin proved to be excellent to control junglerice and as a suitable partner for mixing with propanil, even when propanil-resistant populations were present (Riches et al. 1997; Valverde et al. 2001). Farmers dealing with resistance refused to discard propanil because it remained active against other grasses and economically controlled broadleaf seedlings and emerging sedges. In addition to pendimethalin, several other herbicides also could be used to control junglerice, including preemergence (anilofos, butachlor, clomazone, molinate, pretilachlor, thiazopyr, and thiobencarb) and postemergence (azimsulfuron, bispyribac, ACCase inhibitors, pyribenzoxim, and quinclorac) herbicides. In contrast to the great majority of cases of resistance to PSII inhibitors, resistance to propanil in junglerice is not targetsite based. Propanil uptake and translocation were similar among susceptible and resistant plants (Leah et al. 1995), but major differences in metabolism were observed when both types were compared. In susceptible plants, parental propanil remained at higher concentrations and for longer periods than in resistant plants. The hydrolysis product 3,4-dichloroaniline and especially its conjugates were formed and accumulated more rapidly, respectively, in the resistant than in the susceptible plants. Rapid propanil hydrolysis in resistant plants is endowed by an increased arylacyl amidase (AAA, EC 3.1.1.a) activity (Leah et al. 1994, 1995). Propanil selectivity in rice is also due to hydrolysis by AAA (Frear and Still 1968; Still and Kuzirian 1967; Yih et al. 1968), thus junglerice mimicked rice in its physiological mechanism to degrade propanil. Similar results have been found in barnyardgrass resistant to propanil (Carey et al. 1997). In rice, propanil selectivity is lost when organophosphate or carbamate insecticides are sprayed soon before, together with, or soon after propanil. These insecticides inhibit AAA activity, allowing propanil to interfere with the PS II (Frear and Still 1968; Leah et al. 1995; Matsunaka 1968). Because the propanil resistance mechanism in junglerice was similar to that of rice selectivity, several insecticides of these two groups were assessed in mixture with propanil to determine if it was
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Table 1. Weedy grasses evolving resistance to herbicides in Latin America. Species Avena fatua Biotype Quino and Lautaro Origin Region IX, Chile Resistant toa DCF, CDF Also resistant tob HXP, CTD, STD, TKD Susceptible to IDF, FCZ, IZX, IZX + IZP IMB, IPT TKD, MSS + IDF, FCZ TKD CTD CTD, TPD Reference Espinoza and Zapata 2000; Espinoza and Nitsche 2002; Venegas et al. 2001a, 2001b lito et al. 2005 Cruz-Hipo Tafoya-Razo 2005 Tafoya-Razo 2005 Gazziero et al. 2000 Gazziero et al. 2004

o, Mexico El Baj o, Mexico El Baj Brachiaria plantaginea B-196 BRAPL-1.02 Mexicali, Mexico Mangueirinha, state, Parana Brazil Ponta Grossa, southern region state, of Parana Brazil Parana state, Brazil Chile Castro (R1), Ponta Grossa (R2), Carambe (R3) and Pira do Sul (R4) o ABC, Fundac a state, Parana Brazil state, Brazil Parana Echinochloa colona Central America, Mexico, Colombia, Venezuela Costa Rica, Nicaragua, Colombia

FNP (RI 5 4.0)c,d CDF CDF STD STD IDF, FCZ FZP, HXP, FNP ND

STD CDF STD, FZP

TPD FCZ TPD ND

Cynosurus echinatus Digitaria ciliaris

Golc alves Cortez et al. 2001 Espinoza et al. 2005b; N. Espinoza, personal communication pez-Ovejero et al. 2005 Lo

STD PPN

FNP

CHF, FZP, QZF, CDF, CXD, STD

FNP + CTD, BTD, CLY Several herbicides available. See Valverde et al. 2000 Several herbicides available. See Valverde et al. 2000

Buzatti 2001, 2002a, 2002b Fischer et al. 1993; Ortiz et al. 2000; Valverde et al. 2000 Caseley et al. 1997; Valverde et al. 2000; C. Fuentes, personal communication Zambrano and Espinoza 2004a Caseley et al. 1997; Valverde et al. 2000 B. E. Valverde, unpublished data Valverde and Itoh 2001 Melo et al. 2003a Ferreira et al. 2003 Noldin et al. 2005 Concenc o et al. 2003 Noldin et al. 2005 Valverde et al. 1993

Venezuela Costa Rica

FNP PPN, FNP, AZF IZP QCR QCR QCR QCR QCR QCR IZP STD (RI 5 18) GLY FZP HXP, CHF, CXD PPN, PBZ, BPB, GLY, GLU IZQ, IZT, CRM, SFT CHF, FNP, PQF, BTD BPB, CFD PPN, BPB, FNP CFD PPN, BPB, FNP Several herbicides available. See Valverde et al. 2000

Costa Rica Colombia E. crus-galli ETB-14 ECH12 E. crus-galli and E. cruspavonis E. crus-pavonis Eleusine indica Brazil Capa o do Lea o (RS), Brazil Santa Catarina, Brazil Several locations, Rio Grande do Sul, Brazil Santa Catarina, Brazil Alajuela, Costa Rica Mato Grosso, Brazil Santa Cruz, Bolivia Eriochloa punctata Monte Cristo Santa Cruz, Bolivia

EC13

CTD, FZP, HXP, QZF

Vidal et al. 2006b P. Franco, personal communication Franco et al. 1999

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Table 1. Continued. Species Ischaemum rugosum Biotype Colombia Origin Resistant toa FNP Also resistant tob Susceptible to Reference C. Fuentes, personal communication; O. Almario, personal communication Zambrano and Espinoza 2004b Delgado et al. 2007; C. Zambrano, personal communication Chaves et al. 1997; Valverde et al. 1993 Espinoza et al. 2003b; Espinoza and Nitsche 2002 rez Kogan and Pe rez and 2002; Pe Kogan 2001, 2003; M. Kogan, personal communication Espinoza et al. 2003a Espinoza et al. 2005a; N. Espinoza, personal communication Vargas et al. 2004, 2005

rico, Venezuela Gua Venezuela

BPB NCS FRS + IDF

Ixophorus unisetus Lolium multiflorum Cajon

Puntarenas and Guanacaste, Costa Rica Region IX, Chile

IZP DCF (RI 5 9.7)

IZT, IZQ CDF (RI 5 3.3), STD (RI 5 2.7)

CFR, SFT HXP (RI 5 1.8), CTD (RI 5 1.5), TKD (RI 5 1.8), IDF, FCZ, IZX, IZX + IZP

San Bernardo, Olivar, San Fernando

Region VI, Chile

GLY

Chile Vil-1 Vilcum (Region IX), Chile Vacaria-RS, Brazil

GLY (RI 5 67.5) GLY (RI 5 4.6) GLY

CTD (RI 5 9.5), IDF (RI 5 2.0) IDF (RI 5 4.0), FCZ (RI 5 5.5) Low level of resistance to DCF, FNP, and HXP CDF (RI . 8.4), DCF (RI . 8.3), CTD (RI 5 2.5), STD (RI 5 4.2), TKD (RI 5 3.3)

DCF DCF, CTD GLU, FZP, CTD, STD, PQT, PQT + DRN, ATZ + SMZ, TFL, and MLC IDF, FCZ

Lolium rigidum

n Mulche

Tapejara-RS, Brazil Region VIII, Chile

GLY HXP (RI . 8.4)

Galli et al. 2005 Espinoza et al. 2002; Espinoza and Zapata 2000 B. E. Valverde, unpublished data

Oryza sativa

Costa Rica

Phalaris minor

Yaqui Valley (Sonora), Mexico El Bajio, Mexico

IZP (through gene flow from resistant crop) FNP FNP DCF FNP FNP NCS FZP GLY

TKD

DCF, CDF CDF, CTL, TBT

Phalaris paradoxa

El Bajio, Mexico El Bajio, Mexico Yaqui Valley (Sonora), Mexico

CXD, CHF, FNP, and other graminicides CDF, CTL, TBT TKD DCF, CDF FRS + IDF HXP, STD, CTD CTD

Tamayo-Esquer and Martinez-Carrillo 2002 Morgado Gutierrez et al. 2001 Ruiz-Santaella et al. 2006 Morgado Gutierrez et al. 2001 Tamayo-Esquer and Martinez-Carrillo 2002 C. Zambrano, personal communication Avila et al. 2007 De la Vega et al. 2007, J. Delucchi, personal communication; M. O. Starke, personal communication Franco and Aguanta 2000

Rottboellia cochinchinensis Sorghum halepense

Venezuela Santa Cruz, Bolivia Salta, Argentina

Sorghum saccharatum

El Fuerte

Santa Cruz, Bolivia

CTD (RI 5 5.6)

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Table 1. Continued. Species Biotype Origin Santa Cruz, Bolivia Santa Cruz, Bolivia Resistant toa HXP (RI 5 14.9) HXP (RI 5 6.4), CTD (RI 5 6.0) Also resistant tob CTD, CDF, FNP, QZF Susceptible to BTD, GLY, CRM, IZT, IZX Reference Franco and Aguanta 2000 Franco and Aguanta 2000

Sorghum Marcial sudanense Sorghum Murillo verticilliflorum

a Herbicides: ATZ, atrazine; AZF, azimsulfuron; BPB, bispyribac; BTD, butroxydim; CDF, clodinafop; CFD, 2-[1-2-(4-chlorophenoxy)propoxyimino)-butyl]3-oxo5-thian-3-ylcyclohex-1-enol (proposed common name clefoxydim); CFR, chlorsulfuron; CHF, cyhalofop; CRM, chlorimuron; CTD, clethodim; CTL, chlortoluron; CXD, cycloxydim; DCF, diclofop; DRN, diuron; FCZ, flucarbazone; FNP, fenoxaprop; FRS, foramsulfuron; FZP, fluazifop; GLU, glufosinate; GLY, glyphosate; HXP, haloxyfop; IDF, iodosulfuron; IMB, imazamethabenz; IPT, isoproturon; IZP, imazapyr; IZQ, imazaquin; IZT, imazethapyr; IZX, imazamox; MLC, metolachlor; MSS, methyl 2-[[[[(4,6-dimethoxy-2-pyrimidinyl)amino]carbonyl]amino]sulfonyl]-4-[[(methylsufonyl) amino]methyl]benzoate (proposed common name mesosulfuron); MTS, metsulfuron; NCS, nicosulfuron; PBZ, pyribenzoxim; PPN, propanil; PQF, propaquizafop; PQT, paraquat; QCR, quinclorac; QZF, quizalofop; SFT, sulfometuron; SMZ, simazine; STD, sethoxydim; TBT, terbutryn; TFL, trifluralin; TKD, tralkoxydim; TPD, (EZ)-(RS)-2-{1-[(2E)-3-chloroallyloxyimino]propyl}-3hydroxy-5-perhydropyran-4-ylcyclohex-2-en-1-one (proposed common name tepraloxydim). b Because the mechanisms of resistance are not completely known, no attempt has been made to separately tabulate multiple herbicide resistance and nontarget-site cross-resistances. c Abbreviations: ND, not determined; RI, resistance index. d RIs are given only as a reference value when reported in the literature. RIs obtained in different studies or using different approaches are not comparable.

possible to control resistant plants (Leah et al. 1994; Valverde et al. 2001). Propanil efficacy was restored in resistant plants by the addition of low doses of the insecticides, further proving that the resistance mechanism was associated with the rapid hydrolysis of the herbicide. The principle of synergizing propanil to control propanil-resistant junglerice being established, further research allowed us to develop piperophos and anilofos, organophosphate herbicides registered in some countries, as propanil synergists for preemergence weed control in rice (Caseley et al. 1996; Valverde et al. 1997, 1999). In collaboration with the local development group of Rohm and Haas, a commercial formulation containing propanil plus piperophos (440 g/L plus 40 g/L, respectively) was registered for resistance management, first in Costa Rica and later in other Latin American countries (Valverde 1996, Valverde et al. 2000). The formulation was introduced and rapidly accepted by farmers. Within 4 yr, about 75% of the total rice area planted in Costa Rica was treated with the synergized propanil formulation (Figure 1). A couple of years later, anilofos was also introduced in Colombia and Central America as a synergist, but not in coformulation with propanil. Currently, there are several formulations of propanil

plus either synergist, which are still being used to control junglerice; in Central America, about 22% of the rice area is currently treated with these formulations (Table 2). The herbicide rice market in Latin America was quickly modified when bispyribac was introduced. This herbicide became widely used because of its broad spectrum, excellent selectivity, and efficacy on HR junglerice. Capture of the market by bispyribac severely reduced sales of synergized propanil and cyhalofop, as well as those of sulfonylurea herbicides. Within a couple of years after bispyribacs introduction, lack of control of Digitaria spp. and Leptochloa spp., two genera that are naturally tolerant to the herbicide, brought back some balance to the market and the use of cyhalofop, fenoxaprop, and clefoxydim resurged. The same situation occurred with propanil in Colombia, where the market was quite limited because propanil was considered inadequate for Echinochloa spp. control due to resistance. In 2005, however, propanil experienced increased sales, probably for controlling Digitaria spp. (O. Almario, personal communication). Similarly, propanil use in Venezuela decreased from 97% of the rice-planted area treated in 1996 to about 5% in 2005; grasses are now being controlled with cyhalofop, clefoxydim, and bispyribac (A. Ortiz, personal communication). Because there are no other commercially important PS-II inhibitors used in rice to control grasses, no additional cases of resistance to herbicides with this mode of action have evolved
Table 2. Current use (as percentage of rice planted area treated) of propanil formulations that contain either piperophos or anilofos as synergists of propanil for controlling propanil-resistant junglerice in Central America. Country Rice ha Guatemala Honduras El Salvador Nicaragua Costa Rica Panama Total 8,000 5,600 3,990 65,000 53,120 60,000 195,710 Treated % 45 65 23 8 25 28 22

Figure 1. Historical annual use (as percent of rice area treated) of the formulation of propanil containing piperophos as synergist for controlling propanil-resistant junglerice in Costa Rica.

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in this crop. There is suspicion of itchgrass evolving resistance to hexazinone and diuron in Costa Rica, but confirmation is pending.
Resistance to ACCase-Inhibitor Herbicides

In Latin America, 16 weedy grasses have been confirmed resistant to ACCase-inhibitor herbicides (Table 1), most of them in cereal crops. Chile and Bolivia report five ACCaseinhibitorresistant species each. Resistant Species in Chile. About 400,000 ha of wheat are planted in Chile (Food and Agriculture Organization [FAO] 2006), of which about 25% have resistant weeds at varying infestation levels (N. Espinoza, personal communication). Three of the most important grass weeds of cereal crops have evolved resistance and are the most agronomically important: Lolium spp. (about 75,000 ha), wild oat (40,000 ha), and hedgehog dogtailgrass (25,000 ha); more than one HR species can occur in a single field. In Chile, diclofop was introduced in the late 1970s, followed by clodinafop and tralkoxydim in the late 1980s. These herbicides were widely used in wheat. Beginning in the late 1980s, other ACCase-inhibitor herbicides also became prominent in canola and lupin (Lupinus angustifolius L.), including haloxyfop, fluazifop, quizalofop, and sethoxydim. Failure of these herbicides to control grass weeds became evident in the early 1990s, but resistance was not documented until the latter part of the decade (N. Espinoza, personal communication). Use of diclofop, and to a lesser degree clodinafop and haloxyfop, likely selected for grass-weed resistance. There were cases in which farmers who continuously used diclofop switched to clodinafop, only to find that this herbicide was also ineffective. Initially, resistance was more common to aryloxyphenoxypropionate (APP) herbicides. Later, however, resistance to tralkoxydim used in wheat and barley (Hordeum vulgare L.) and sethoxydim used in broadleaf crops became common, and these herbicides lost most of their market share. Interestingly, clethodim and the more recently introduced cyclohexanedione, trepaloxydim, are still quite effective against otherwise ACCase-inhibitorresistant biotypes (N. Espinoza, personal communication). At the beginning of the 1990s, a rigid ryegrass population n (Region VIII) was not controlled with from Mulche haloxyfop in canola. Bioassays confirmed resistance to this herbicide and cross-resistance to other ACCase-inhibitor herbicides (diclofop, clodinafop, tralkoxydim, clethodim, and sethoxydim), but not to iodosulfuron or flucarbazone (Espinoza et al. 2002; Espinoza and Zapata 2000). A similar case with the congener Italian ryegrass was found in 1997 in n (Region IX) after diclofop failed to control this weed in Cajo wheat (Espinoza et al. 2003b, Table 1). In this case, the biotype was cross-resistant to clodinafop and sethoxydim but not to haloxyfop, clethodim, or tralkoxydim. Wild oat resistant to diclofop and clodinafop was documented in Quino and Lautaro (Region IX). These biotypes are also resistant to other ACCase-inhibitor herbicides, but remain susceptible to iodosulfuron or flucarbazone (Espinoza and Nitsche 2002; Espinoza and Zapata 2000; Venegas et al.

2001a, 2001b). A biotype of hedgehog dogtailgrass, an important weed in wheat, barley, oat (Avena sativa L.), lupin, and canola, was found resistant to both ACCase- and ALSinhibitor herbicides (Espinoza et al. 2005b). Finally, there is suspicion that Vulpia sp. (Espinoza and Nitsche 2002) and little quakinggrass (Briza minor L.) have also evolved resistance to ACCase-inhibitor herbicides, but confirmation tests have not been conducted yet. In all of these cases, the mechanism of resistance is unknown. Failure to control important grasses with ACCase-inhibitor herbicides provided the opportunity for postemergence ALS herbicides active against grasses to capture part of the market. Iodosulfuron was introduced in 2001. Even before its commercial release, cross-resistance to this herbicide was observed in some ACCase-inhibitorresistant Lolium spp. biotypes (N. Espinoza, personal communication). With increased use of iodosulfuron, resistance to this herbicide among grass populations has also increased. Iodosulfuron, however, still controls most ACCase-inhibitorresistant biotypes. These cross-resistance patterns impose the need for further efficacy tests (using known HR biotypes) before alternative herbicides are registered and commercialized. Management of HR grass weeds in Chile encounters the typical limitations for crop and mode-of-action rotations. In oat, the only option for grass control is ACCase-inhibitor herbicides. In wheat, in addition to ACCase-inhibitor herbicides, postemergence ALS herbicides can be used. Preemergence grass killers for wheat production include triasulfuron (ALS inhibitor), prosulfocarb (lipid synthesis inhibitor), and diuron (PS-II inhibitor). In barley, herbicide rotations include ACCase inhibitors, prosulfocarb and diuron, and ACCase-inhibitor herbicides and simazine in lupin. In canola, ACCase-inhibitor herbicides can be rotated with trifluralin and metazachlor (N. Espinoza, personal communication). Emerging ACCase-Resistance Problems in Bolivia. Louisiana cupgrass, three weedy Sorghum spp., and itchgrass evolved resistance to ACCase-inhibitor herbicides in Bolivia. Louisiana cupgrass infests several crops in the Santa Cruz department of Bolivia, including corn, soybean, rice, and cotton (Gossypium hirsutum L.). Several Louisiana cupgrass populations from soybean fields were collected and bioassayed for fluazifop resistance. Putative HR populations selected by fluazifop for at least 5 yr were confirmed as resistant. A fluazifop-resistant biotype (Monte Cristo) was further characterized. It exhibited cross-resistance to haloxyfop, cyhalofop, and cycloxydim, but was susceptible to herbicides with different modes of action including propanil, pyribenzoxim, bispyribac, glyphosate, and glufosinate (Franco et al. 1999). Three Sorghum spp. are weedy in Santa Cruz, particularly in corn and soybean. Of these sudangrass is the most frequent, followed by two common wild sorghums. Two sudangrass biotypes were clearly resistant to haloxyfop (Toborocchi, resistance index [RI] 5 13.2 and Marcial, RI 5 14.9), but only one of them (Marcial) was cross-resistant to clethodim. The latter was also cross-resistant to quizalofop, fenoxaprop, and clodinafop, but not to butroxydim. ALS-inhibitor graminicides and glyphosate control this biotype (Franco and Aguanta 2000; P. Franco, personal communication). A
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common wild sorghum biotype was cross-resistant to haloxyfop and clethodim, and another (S. saccharatum) was resistant to clethodim, but not to haloxyfop (Franco and Aguanta 2000; P. Franco, personal communication). No further characterization of Sorghum biotypes has been conducted in recent years, but there are indications that resistance is becoming more widespread. Under the same production systems, populations of itchgrass have also been selected for ACCase-inhibitor resistance mainly by the selection pressure imposed by fluazifop. In addition to being resistant to fluazifop, the best-characterized biotypes so far are also resistant to haloxyfop, sethoxydim, and clethodim. HR and susceptible biotypes did not differ in the rate of [14C]sethoxydim foliar uptake, translocation, or metabolism. The concentrations of sethoxydim required to inhibit in vitro ACCase activity by 50% (I50) were substantially higher (about 11 times) for HR biotypes compared to the reference biotype, indicating that the HR itchgrass biotypes have an ACCase that is relatively insensitive to the graminicides (Avila et al. 2007). ACCase-Resistant Grass Weeds in Mexico. Two Phalaris spp., littleseed and hood canarygrass, evolved resistance to ACCase-inhibitor herbicides in wheat in Mexico. Information about these species is scattered, and to my knowledge, research results have not been published in any refereed journal. Wheat is grown in Mexico in irrigated land mainly in the northwestern and northern states, predominantly during the autumn/winter cycle. It is also planted in central Mexico in spring/summer in about 15% of the 600,000 ha dedicated to the crop (FAO 2006). The Northwest accounts for 56% of o (Guanajuato) for 22%. Barley is total production and El Baj also planted in both regions for a total of about 385,000 ha (FAO 2006). The main producing states of wheat and barley n, are Sonora, Baja California, Guanajuato, Sinaloa, Michoaca Chihuahua, Tlaxcala, Jalisco, and Hidalgo. Littleseed and hood canarygrass are present in about 40% and 30% of the wheat area, respectively ( J. A. Tafoya, personal communication). After receiving complaints from farmers about lack of control of both species with graminicides, preliminary studies found that biotypes collected in the Yaqui Valley (southern Sonora) were indeed resistant to the selecting herbicide, fenoxaprop. Although these studies included only two discrete herbicide doses, equivalent to the commercially recommended dose and 23, the biotypes apparently had cross-resistance to tralkoxydim but not to diclofop or clodinafop (TamayoEsquer and Martinez-Carrillo 2002). In the semiarid, subto, ACCase-inhibitorresistant Phalaris ropical region of El Baj spp. increased from about 300 ha in 1998 to 2,000 ha in 2003 (Tafoya Razo 2003) and currently affects 6,000 ha ( J. A. Tafoya, personal communication). Fenoxaprop-resistant biotypes of both Phalaris spp. collected in this area were controlled with postemergence mixtures of clodinafop, chlortoluron, and terbutryn (Morgado Gutierrez et al. 2001). Very little is known about the ACCase-inhibitor resistance mechanism in Phalaris. Unpublished preliminary studies indicate that ACCase-inhibitor resistance is likely due to enhanced herbicide metabolism ( J. A. Tafoya, personal communication), but Ruiz-Santaella et al. (2006) reported that a diclofop-resistant biotype of hood canarygrass (also cross-resistant to cycloxydim, cyhalofop, fenoxaprop, and 316 N
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other graminicides) had a diclofop-insensitive ACCase derived from the substitution of Gly by Ser at position 2096. This is a new mutation conferring resistance not previously reported in other species. Wild oat is also an important weed in cereal production in Mexico. It is present in about 40% of the Mexican wheat area, but fewer chemicals are aimed at its control. In the northwestern region (Mexicali Valley), about 3,000 ha are infested with fenoxaprop-resistant wild oat. Biotypes from this region are resistant to both ACCase-inhibitor herbicide families and frequently are cross-resistant to ALS-inhibitor herbicides (metsulfuron + iodosulfuron, flucarbazone, and imazethapyr), probably as a result of enhanced herbicide o but metabolism. HR populations are also found in El Baj only in about 500 ha ( J. A. Tafoya, personal communication). A fenoxaprop-resistant biotype from this area (RI 5 4.0 based on a rapid test) was susceptible to imazamethabenz and lito et al. 2005). Populations from El isoproturon (Cruz-Hipo o are typically resistant to fenoxaprop and other APP Baj herbicides, but not to cyclohexanediones (CHD); resistance is endowed by a target-site mutation ( J. A. Tafoya, personal o where resistance is communication). In the areas of El Baj a problem, farms that rotate crops are not affected by HR weeds. In some farms covering about 3,000 ha, farmers are using stale seedbed soil preparation and preplant irrigation to eliminate the first flush of emerging HR grasses with glyphosate or paraquat. In another 30,000 ha farmers rely on herbicide rotations and mixtures to control weedy grasses. In the Northwest, 100,000 ha are managed by combining mechanical and chemical control, a system that farmers are trying to implement in other regions ( J. A. Tafoya, personal communication). Grass Weeds Resistant to ACCase Herbicides in other Latin American Countries. Three ACCase-inhibitorresistant grasses have been found in Brazil. Alexandergrass is one of the most important grass weeds in soybean production in Brazil. Alexandergrass biotypes exhibit different cross-resistance patterns. A sethoxydim-resistant biotype from Parana state was susceptible to the CHD herbicides, clethodim and tepraloxydim (Gazziero et al. 2004). Another biotype from the same state was resistant to tepraloxydim and sethoxydim. Absorption and translocation of both herbicides were similar in resistant and susceptible biotypes. The I50 value for the inhibition of ACCase by sethoxydim was 1.0 to 2.5 mM and 600 mM for the susceptible and resistant biotypes, respectively; those for tepraloxydim were 0.25 to 0.5 mM and 4 mM for the resistant and susceptible biotypes, respectively. It was concluded that ACCase-inhibitor resistance of this alexandergrass biotype is due to an alteration of the target enzyme (Golc alves Cortez et al. 2001). In the same pez-Ovejero et al. (2005) production systems in Parana, Lo reported ACCase-inhibitorresistant populations of southern crabgrass. Populations were resistant to fluazifop, sethoxydim, and tepraloxydim (Table 1). Plants of this species emerging after harvesting a bean (Phaseolus vulgaris L.) crop in which graminicides failed to control it were treated with increasing doses of fenoxaprop plus clethodim (formulated mixture), sethoxydim, butroxydim, or glyphosate. All herbicides effectively controlled the southern crabgrass population,

especially at the highest doses tested, except sethoxydim, which did not cause any damage (Buzatti 2001, 2002a, 2002b). The most recent species becoming resistant to graminicides in Brazil is goosegrass. HR plants were discovered in 2003 in soybean fields in Mato Grosso treated with these herbicides at least once a year for almost 15 yr. Plants were resistant to sethoxydim (RI 5 18) and crossresistant to cyhalofop, fenoxaprop, propaquizafop, and butroxydim, but not to clethodim, fluazifop, haloxyfop, or quizalofop (Vidal et al. 2006b). Preliminary studies indicate that the response of the HR plants to ACCase-inhibitor herbicides is influenced by light, temperature, and insecticides, which may be an indirect indication that resistance could be related to enhanced herbicide metabolism (R. Vidal, personal communication). As previously mentioned, junglerice populations evolved resistance to fenoxaprop and other ACCase-inhibitor herbicides in rice production systems in Central America and in Colombia; saramollagrass also became resistant to fenoxaprop in Colombia (O. Almario, personal communication; C. Fuentes, personal communication). In these countries, the use of fenoxaprop dropped drastically, and cyhalofop and clefoxydim, the most recent graminicides commercially used in rice, show increasing failures in controlling Echinochloa spp. and saramollagrass. The latter herbicides remain the most important postemergence grass killers controlling Digitaria spp., Leptochloa spp., and goosegrass (O. Almario, personal communication). Rapid tests were developed for field screening of fenoxaprop-resistant junglerice and saramollagrass, and a kit is locally available for verification of the resistance status of Ospina 2001). Four these species (Fuentes et al. 2003; Girata junglerice biotypes recently collected in Portuguesa state, Venezuela, were resistant to fenoxaprop, with RI between 3.5 and 4.5 (Zambrano and Espinoza 2004a).
Resistance to ALS-Inhibitor Herbicides

used frequently and at high doses to maintain areas around chicken houses weed-free. Under this extreme selection pressure, a particular population of goosegrass evolved resistance to the imidazolinone. Based on whole plant bioassays, it was determined that the resistant population was cross-resistant to imazethapyr, imazaquin, sulfometuron, and chlorimuron (Valverde et al. 1993). ALS-HerbicideResistant Rice Grass Weeds. Saramollagrass rico, populations collected in rice fields in the state of Gua Venezuela were confirmed resistant to bispyribac (Zambrano and Espinoza 2004b, Table 1). These populations had been exposed to bispyribac in rice for about 4 yr. We also have recently identified populations of junglerice that are resistant to imazapyr in Costa Rica. These populations were found at two different rice farms, one in the rain-fed area in the Central Pacific and another in the irrigated district of Guanacaste (Northern Pacific region). In both cases, populations came from fields that had been planted for at least three cropping seasons with an imidazolinone-resistant (ClearfieldH) rice variety. A commercial mixture of imazapyr plus imazapic was used at least twice during the cropping season to control weedy rice and other weeds, including junglerice. Whole plant bioassays confirmed resistance. Cross-resistance patterns are being investigated (B. E. Valverde, unpublished data). Weedy rice populations resistant to imazapyr have now been identified in Costa Rica in fields where Clearfield rice had been planted two or three times sequentially (B. E. Valverde, unpublished data). Laboratory studies have confirmed that resistance in these populations was conferred by gene flow from imidazolinone-resistant rice to the weedy types (K. B. Holm and B. E. Valverde, unpublished data). There is also preliminary experimental greenhouse evidence of barnyardgrass populations evolving resistance to bensulfuron in rice fields in Chile. Because this herbicide is usually applied in combination with molinate, the problem has not become evident in the field (M. Kogan, personal communication). Other ALS-HerbicideResistant Grasses. Some biotypes of Phalaris spp. that evolved resistance to ACCase-inhibitor herbicides, and of Lolium spp. resistant to glyphosate, are also cross-resistant to ALS-inhibitor herbicides. These biotypes are described in the respective sections of this paper. In addition, itchgrass evolved resistance to nicosulfuron in corn in Venezuela. Of 13 populations sampled in Portuguesa state that had been exposed to nicosulfuron applications for 8 yr, 11 exhibited varying degrees of resistance (Delgado et al. 2007). Several of these populations are also resistant to the mixture of foramsulfuron and iodosulfuron (C. Zambrano, personal communication). Farmers from the same region also complain that nicosulfuron no longer controls johnsongrass and common wild sorghum, but confirmation studies are pending (A. Ortiz, personal communication).
Resistance to Glyphosate

Imazapyr-Resistant Mexican Grass and Goosegrass in Costa Rica. Although there are several cases of ALSinhibitor-resistant broadleaf weeds in Latin America, only a few grasses have evolved resistance to this group of herbicides. Mexican grass evolved resistance to imazapyr in Costa Rica. This was the first case worldwide of a grass weed becoming resistant to an imidazolinone herbicide (Valverde et al. 1993). HR populations were found along irrigation canals and ditch banks in rice and sugarcane (Saccharum officinarum L.) fields in the northwestern region of Costa Rica. In all cases, populations had been treated with imazapyr for at least 5 yr. HR plants were cross-resistant to other imidazolinones (imazethapyr and imazaquin), but not to the grass-active sulfonylureas, chlorsulfuron, and sulfometuron (Chaves et al. 1997). The mechanism of resistance was determined to be an altered ALS enzyme, but the mutation responsible for the enzyme insensitivity is unknown. Because Mexican grass rarely infests sugarcane or irrigated rice fields, resistance never posed any major threat to rice or sugarcane production, except for the increased cost of switching to alternative, low-persistence herbicides. At about the same time, a second case of imazapyr resistance was found. At a poultry farm, imazapyr was being

Only a few weed species have evolved resistance to glyphosate worldwide. Most of them are broadleaf weeds:
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Palmer amaranth (Amaranthus palmeri S. Wats.), common ragweed (Ambrosia artemisiifolia L.), hairy fleabane [Conyza bonariensis (L.) Cronq.], horseweed [C. canadensis (L.) Cronq.], and buckhorn plantain (Plantago lanceolata L.); the only grasses presently reported in the International Survey of Herbicide Resistant Weeds (Heap 2006) were goosegrass in Malaysia, Italian ryegrass in Chile, Brazil, and the United States, and rigid ryegrass in Australia, the United States, and South Africa. In addition to glyphosate-resistant Italian ryegrass in Chile and Brazil, Latin America contributed three cases not yet registered in the survey, including a new HR broadleaf weed, ragweed parthenium (Parthenium hysterophorus L.) (Rosario Socorro and Fuentes 2005), and two grass species, johnsongrass and goosegrass (Table 1), which are considered in this section. Glyphosate-Resistant Italian Ryegrass. Italian ryegrass, the first glyphosate-resistant species in Latin America, was rez 2002; Pe rez and Kogan reported in 2001 (Kogan and Pe 2001, 2003). Resistant populations were found in grape orchards at two locations (San Bernardo and Olivar) in Region VI (central area) of Chile. The sites were being sprayed an average of three times per year with glyphosate (1.08 to 1.44 kg ae/ha) in combination with MCPA (1.5 kg ae/ha) for improved broadleaf weed control during the previous 8 to 10 yr. Based on whole plant bioassays, the San Bernardo and Olivar populations were two- and fourrez and fold, respectively, more resistant to glyphosate (Pe Kogan 2003) than the corresponding reference population. A third biotype (San Fernando) with similar characteristics was also found in the same region (M. Kogan, personal communication). In southern Chile (Region IX), yet another n) was documented in chemical resistant biotype (Vilcu fallows. This biotype evolved resistance after being exposed to glyphosate 12 times during 1989 to 2001, prior to the n planting of wheat or oat (Espinoza et al. 2005a). The Vilcu biotype, which was treated in 2001 with the ALS-inhibitor herbicide iodosulfuron is also cross-resistant to both iodosulfuron and flucarbazone (Table 1). Glyphosate resistance is an emerging problem in Chile, and thus far does not have the economic importance of ACCase-inhibitor resistance (N. Espinoza, personal communication). In central Chile, paraquat is more effective in controlling glyphosate-resistant Italian ryegrass than ACCase-inhibitor herbicides or amitrole (M. Kogan, personal communication). Italian ryegrass is also well adapted to the southern region of Brazil that includes Rio Grande do Sul (RS), Santa , where it is used in no-till agriculture as Catarina, and Parana a winter cover crop, as a cover crop or mulch in orchards, and as forage. In the first two situations, plants are desiccated with herbicides, mostly glyphosate (Galli et al. 2005). Italian ryegrass is also an important weed in soybean, wheat, and corn. Additionally, it is the most common grass weed in apple orchards in which three to four glyphosate applications (at 720 or 1,080 g ae/ha) are made yearly (Vargas et al. 2004, 2005). Glyphosate-resistant Italian ryegrass populations have been found at a few locations in RS in annual crops and orchards (Roman et al. 2004). One of them was documented at a field (Tapejara municipality) where glyphosate (1.08 kg ae/ha) failed to control Italian ryegrass at an early 318 N
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growth stage. Interestingly, the seed planted at this site came o Bonito (RS) that belonged to the same from a field in Capa farmer, where glyphosate resistance was also confirmed (Galli et al. 2005), but the common origin has not been ascertained by molecular studies. In general, herbicides with alternative modes of action can be effectively used to control the HR biotypes (Christoffoleti et al. 2005; Vargas et al. 2004, 2005). Thus, farmers now rely on systemic graminicides to control glyphosate-resistant Italian ryegrass (Vargas et al. 2005). Field and greenhouse studies with a resistant biotype collected in an area of soybeanwheat rotations in Lagoa Vermelha (RS) indicated that higher glyphosate doses are required for adequate control, as the resistant plants become older and some of the affected plants are able to resprout (Christoffoleti et al. 2005). HR populations can be controlled with glyphosate in mixture with ACCase-inhibitor herbicides (clethodim, tepraloxydim, sethoxydim, clethodim + fenoxaprop, and haloxyfop), and with mixtures of paraquat and diuron (Christoffoleti et al. 2005). But there are also indications of decreased response to some of these herbicides. Glyphosate GR50 values (dose required to reduce growth by 50% of an untreated control) for a resistant biotype found in an apple orchard and a reference susceptible biotype were 4,835 g ae/ha and 290 g ae/ha, respectively (Vargas et al. 2005). Although ACCase-inhibitor herbicides had never been used in the field to control the glyphosate-resistant biotype, it was slightly less affected by diclofop, fenoxaprop, and haloxyfop when compared to the susceptible biotype, especially at low doses. Fluazifop affected both the resistant and susceptible biotypes in the same manner. This is similar to the observations by Powles et al. (1998), who found that a glyphosate-resistant Italian ryegrass biotype from Australia was cross-resistant to diclofop but susceptible to fluazifop and sethoxydim. In analyzing the productivity of the resistant biotype, Vargas et al. (2005) found that it accumulated more aboveground biomass but fewer tillers, inflorescences, and seed, and it matured later than the susceptible biotype. The studies, however, were not conducted under competitive conditions to indicate fitness differences. The mechanism of resistance to glyphosate in the Latin American Italian ryegrass populations has not been elucidated yet. Studies comparing resistant and susceptible biotypes from Region VI of Chile determined that differential absorption, translocation, or allocation of glyphosate within the plant is rez et al. 2004). not the primary resistance mechanism (Pe Leaves and leaf segments of a glyphosate-resistant Italian ryegrass biotype from Oregon exposed to glyphosate accumulated more shikimic acid than those of the susceptible biotype. Amplification and sequencing of the enolpyruvylshikimate-3-phosphate synthase (EPSPS, EC 2.5.1.10) gene did not identify any amino acid change responsible for rez-Jones et al. 2005). In the congeneric species resistance (Pe rigid ryegrass, different mechanisms may be responsible for glyphosate resistance, depending on the biotype. In a biotype from Australia (NLR68/70), resistance appears to be associated with differences in translocation. The resistant and susceptible biotypes exhibit similar levels of expression of the EPSPS enzyme, which is equally sensitive to inhibition by

glyphosate in both populations. Additionally, plant metabolism of glyphosate does not contribute to resistance. Although resistant and susceptible plants both absorb glyphosate, the herbicide accumulates in the roots of susceptible plants, whereas it accumulates in the leaf tips of the resistant plants. (Lorraine-Colwill et al. 2002). Resistance in the NLR70 biotype is conferred primarily by a single nuclear gene and inherited as an incompletely dominant trait (Lorraine-Colwill et al. 2001). Similar to NLR70, three additional Australian HR biotypes (NLR71, NLR72, and NLR76) translocated glyphosate applied to a mature leaf more readily to the tip of the treated leaf than the susceptible one. Susceptible plants, however, translocated twice as much glyphosate to the stem meristems than resistant plants (Wakelin et al. 2004). In another Australian biotype, however, altered uptake, translocation, or metabolism of glyphosate was not responsible for resistance (Feng et al. 1999). Further studies with this biotype (48118a) indicated that resistance could be related to elevated activity of the EPSPS enzyme and messenger RNA levels, but the specifics are still unknown (Baerson et al. 2002). Glyphosate Resistance in Argentina. There has been much expectation regarding the possible selection of glyphosateresistant populations in Argentina, where no-till production and growing of glyphosate-resistant (Roundup ReadyH) crops, particularly soybean, corn, and cotton, are extensive. An estimated 128 million L of formulated glyphosate were used in Argentina in 2004. About 51 million L were applied to glyphosate-resistant crops; the rest was used during chemical fallows or as preplant treatments, particularly in no-till agriculture that covers 16 million ha ( J. Delucchi, personal communication). Published information only illustrates floristic changes in relation to glyphosate use as well as to tillage system, but not resistance evolution (Tuesca et al. 2001; Vitta et al. 2004). Minimum tillage favors prevalence of annual grass weeds (Tuesca et al. 2001). In a 5yr study, Puricelli et al. (2005) documented changes in weed species composition at sites where glyphosate was used regularly in different crop sequences that included wheat soybean, soybean monoculture, and soybeancorn under conventional and zero tillage. Glyphosate altered the local weed flora to a much greater extent than cropping system. Glyphosate suppressed highly competitive weeds, but increased the abundance of late-emerging weeds such as pellitory (Parietaria debilis Forst.) and field violet (Viola arvensis Murr.). Both species escape application of the herbicide because of their late emergence. Glyphosateresistant weeds were not found in this study. A survey conducted in the main sunflower (Helianthus annuus L.) -growing areas of Buenos Aires province did not detect cases of glyphosate resistance even though more than half of the farmers apply glyphosate at least three times per cropping season under no-till production compared to one application in conventional tillage. Changes in the floristic composition of the weed communities were also documented with new weed species such as Euphorbia spp. and hairy fleabane becoming more prevalent (Istilart 2005). The situation has changed recently, however, as farmers began to complain about poor control of johnsongrass by glyphosate in fields under no-till production where glypho-

sate-resistant soybeans are grown. Monsanto began studying a potential case of a glyphosate-resistant johnsongrass population in northern Argentina in 2004 after a farmer from the Salta province submitted a glyphosate performance inquiry. Preliminary findings of field and greenhouse trials had indicated a lack of control with glyphosate at the label dose. However, results from more recent greenhouse studies with seedlings from the suspect biotype did not indicate a lack of control with glyphosate at the label rate. Therefore, additional research is being to fully elucidate the preliminary findings, both in the field and conducted in the greenhouse ( J. Delucchi, personal communication). A johnsongrass biotype from the same region was confirmed resistant to glyphosate by De la Vega et al. (2007). In fields where glyphosate-resistant soybean is rotated with bean, johnsongrass control is achieved with selective graminicides. Monsanto is also evaluating mixtures of glyphosate and grass herbicides for use in glyphosate-resistant soybean. The tank mixture of glyphosate and clethodim seems to be quite effective (M. O. Starke, personal communication). In a recent field visit to Salta, we corroborated the existence of spreading resistant johnsongrass, mostly distributed as clumps within soybean fields in vast areas around the province and at a single n province (B. E. Valverde and J. location in the Tucuma Gressel, unpublished observations). Putative Glyphosate-Resistant Goosegrass in Bolivia. There is also concern that goosegrass evolved resistance to glyphosate in Bolivia. Preliminary bioassays with populations collected in the Santa Cruz soybean area confirm failure of glyphosate at the recommended commercial dose to control some populations of this weed. Plants treated in full-tillering are able to resprout and produce seed (P. Franco, personal communication). Further work is underway to corroborate resistance and determine the corresponding RI.
Resistance to Quinclorac

Three Echinochloa spp., junglerice in Colombia (Valverde and Itoh 2001) and barnyardgrass and gulf cockspur in Brazil, became resistant to quinclorac in rice. Very limited research characterizing HR biotypes has been conducted, and most of it refers to the Brazilian cases. HR biotypes have been found in RS, a state that produces about 50% of the Brazilian rice in over one million ha of flooded fields. Quinclorac has been widely used in the state; by 2001, about 70% of the rice area was treated with this herbicide and HR populations were identified at several locations (Andres et al. 2002c). Barnyardgrass seeds were collected in ria do rice fields in four municipal districts (Santa Vito o, Camaqua , and Mostardas) where quinPalmar, Jaguara clorac failed to control the weed after its continued use for about 10 yr. Plants of a reference susceptible biotype (Cachoeirinha) never exposed to quinclorac were killed at the lowest dose (188 g/ha) tested in a greenhouse bioassay experiment. The four resistant biotypes survived quinclorac at increasing doses up to 1.5 kg/ha; maximum fresh weight reduction at the highest dose was only 20%, thus a GR50 value could not be calculated (Menezes et al. 2001).
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Between 84% and 94% of seedlings emerging from soil samples collected at the same sites survived quinclorac at a high dose (750 g/ha), indicating that the populations were already highly enriched with resistant individuals (Menezes et al. 2001). Similar responses to quinclorac have been obtained with other populations (Andres et al. 2002b, Ferreira et al. 2003). In a quick bioassay test, Echinochloa spp. seeds were germinated on filter paper moistened with quinclorac dilutions. Emerging seedlings of susceptible biotypes were severely affected by quinclorac at about 35 mg/L, whereas resistant seedlings did not reach a 50% growth reduction at the highest concentration (1,875 mg/ L) tested (Melo et al. 2003b). A biotype of each of the two Echinochloa spp. from Santa Catarina state was only slightly affected by quinclorac at 6 kg/ha, equivalent to 16 times the recommended dose (Noldin et al. 2005). Biotypes characterized so far can be controlled in rice with alternative herbicides. These include propanil, ALS-inhibitor herbicides (bispyribac), ACCase inhibitors (clefloxydim, cyhalofop, fenoxaprop), clomazone, and pendimethalin (Andres et al. 2002a, Concenc o et al. 2003; Ferreira et al. 2003; Menezes et al. 2001; Noldin et al. 2005; Schmidt et al. 2003).
Multiple Herbicide Resistance

were found ( J. A. Tafoya, personal communication). Further studies are required to establish the true multiple resistance of some of these cases, as they could also be explained by non target-site (intergroup) cross-resistances.
Acknowledgments

I would like to thank my colleagues and friends from Latin America and elsewhere who provided me with reprints of their work, shared information and unpublished results, and made useful comments on an early version of the manuscript: Leonardo Paniagua, Israel Garita, and Milton Pineda (Costa Rica); J. Antonio Tafoya (Mexico); Alvaro Salive, Octavio Almario, and Cilia Fuentes (Colombia); Aida Ortiz and Castor Zambrano (Venezuela); Jose Alberto Noldin, Dionisio Gazziero, Pedro Christoffoleti, Germani Concenc o, Walter Buzatti, and Ribas Vidal (Brazil); Julio Delucchi (Argentina); Michelle O. Starke (United States); Nelson Espinoza and Marcelo Kogan (Chile); and Pablo Franco (Bolivia).
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Multiple herbicide resistance, i.e., the ability of a biotype to withstand herbicides from different chemical classes by expressing more than one resistance mechanism, in agronomically important weeds substantially limits the chemical options to control them, and can increase production costs. As described in previous sections, a few grass weeds in Latin America exhibit resistance to more than one herbicide or chemical family with different modes of action, probably by accumulating more than a single resistance mechanism. The first grass species with multiple resistance in this region was junglerice, which in addition to being resistant to propanil and fenoxaprop (with cross-resistance to other ACCaseinhibitor herbicides), was also resistant to azimsulfuron (Valverde and Itoh 2001). An Italian ryegrass biotype from Chile confirmed resistant to glyphosate (RI 5 68) was also resistant to clethodim (RI 5 10), but not to diclofop. It also exhibited a low level (RI 5 2.0) of resistance to the ALS-inhibitor herbicide iodosulfuron (Espinoza et al. 2003a). Another biotype (Vil1) treated 12 times with glyphosate between 1989 and 2001 was also confirmed resistant to glyphosate, with crossresistance to the ALS-inhibitor herbicides iodosulfuron and flucarbazone. This second biotype remained susceptible to the ACCase-inhibitor herbicides diclofop and clethodim (Espinoza et al. 2005a). There is also indication that Mexican Phalaris spp. and wild oat resistant to ACCase-inhibitor herbicides have already evolved resistance to alternative herbicides. A wild oat biotype from Valle de Mexicali could not be controlled with flucarbazone or the mixture of mesosulfuron and iodosulfuron o where (Tafoya-Razo 2005). In 2 out of 10 sites in El Baj wheat was rotated with alfalfa (Medicago sativa L.) to help control HR Phalaris spp., plants resisting imazethapyr also 320 N
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Received June 5, 2006, and approved September 19, 2006.

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