Tuta Resistance To Insecticides

Resistance of Tuta absoluta to insecticides
Because of the short generation time and the frequent applications of insecticide to manage T. absoluta
resistance to several insecticides has developed.
In 1999 signifcant resistance of T. absoluta to acephate and deltamethrin was reported
1
. In the same
year resistance to deltamethrin, lamba=cyhalothrin, mevinphos, metamidophos and esphenvalerate was
reported in Chile
2
.
In 2000 resistance to Cartap was reported in Brazil
3,4
. In 2001 resistance to abamectin was additionally
reported in Brazil
4,5
. In 2005 an Argentine study confrmed T. absoluta resistance in that country to
deltamethrin and abamectin as well as methamidophos
6
.
Among the widely used insecticides that are still effective are imidacloprid
7
and Bacillus thruingiensis.
References
1.
CASTELO BRANCO, M.; FRANA, F.H., MEDEIROS, M.A.; LEAL, J.G.T. Uso de inseticidas para o controle da
traa-do-tomateiro e traa-dascrucferas: um estudo de caso. Horticultura Brasileira, v. 19 n. 1, p. 60-63, maro, 2001.
2.
SALAZAR E.; ARAYA J. Tomato moth, Tuta absoluta (Meyrick) response to insecticides in Arica, Chile.
Agric. Tc. v.61 n.4 Chilln oct. 2001
3.
SIQUEIRA H. A. A. ,GUEDES R. N. C.; PICANCO M. C. Cartap resistance and synergism in populations of Tuta
absoluta (Lep., Gelechiidae). J. Appl. Ent. 124, 233-238. 2000
4.
SIQUEIRA H. A. A. ,GUEDES R. N. C.; PICANCO M. C . Insecticide resistance in populations of Tuta
absoluta (Lepidoptera: Gelechiidae). Agricultural and Forest Entomology. Volume 2, Issue 2, Pages 147-153. 2001
5.
SIQUEIRA H. A. A., GUEDES R. N. C, FRAGOSO D. B and MAGALHA L. C. Abamectin resistance and synergism
in Brazilian populations of Tuta absoluta (Meyrick) (Lepidoptera: Gelechiidae) International Journal of Pest Management.
47(4) 247-251. 2001
6.
LIETTII M., BOTTOII E., ALZOGARAY R. Insecticide resistance in Argentine populations of Tuta absoluta (Meyrick)
(Lepidoptera: Gelechiidae). Neotrop. Entomol. vol.34 no.1 Londrina Jan./Feb. 2005
7.
DANTE.M; GIMNEZ R. Effcacy of imidacloprid to control the tomato borer (Tuta absoluta meyrick).
IDESIA (Chile) V 26, N 1, p 65-67. 2008
60 Hortic. bras., v. 19, n. 1, mar. 2001.
E
m agosto de 1999, foi verificado que
no Ncleo Rural da Taquara (DF) a
produo de tomate e brassicas estava
seriamente comprometida devido aos
danos ocasionados pela traa-do-toma-
teiro (Tuta absoluta) e pela traa-das-
crucferas (Plutella xylostella). Diver-
sos tipos de inseticidas, com freqncia
que variava de semanal a diria, foram
utilizados na regio. A impossibilidade
de controle das pragas foi atribuda, pe-
los agricultores, possvel falsificao
dos produtos. No foi levantada a hi-
ptese de que a ineficincia dos produ-
tos poderia ser devida resistncia das
pragas aos inseticidas. Resistncia de
traa-do-tomateiro a cartap j foi obser-
vada no Brasil (Siqueira et al., 2.000) e
CASTELO BRANCO, M.; FRANA, F.H., MEDEIROS, M.A.; LEAL, J.G.T. Uso de inseticidas para o controle da traa-do-tomateiro e traa-das-
crucferas: um estudo de caso. Horticultura Brasileira, v. 19 n. 1, p. 60-63, maro, 2.001.
Uso de inseticidas para o controle da traa-do-tomateiro e traa-das-
crucferas: um estudo de caso.
Marina Castelo Branco
1
; Flix H. Frana
1
; Maria A. Medeiros
1
, Jos Guilherme T. Leal
2
1/
Embrapa Hortalias, C. Postal 218, 70.359-970, Braslia - D.F; E-mail: marina@cnph.embrapa.br
2/
EMATER-DF. Escritrio Local
Ncleo Rural da Taquara, s/n. 70.000-000 Braslia DF
RESUMO
Em agosto de 1999, produtores de tomate e brassicas da Ncleo
Rural da Taquara tiveram seus cultivos seriamente comprometidos
devido impossibilidade de controle da traa-do-tomateiro e da tra-
a-das-crucferas. Diversos inseticidas, alguns com o mesmo prin-
cpio ativo ou, pertencentes ao mesmo grupo qumico, eram aplica-
dos de uma a sete vezes por semana sem qualquer eficincia no
controle das pragas. Lavouras foram abandonadas em diferentes estdios
de desenvolvimento. A fim de definir uma estratgia de controle
que viabilizasse a produo de tomate e brassicas na regio, foi ava-
liado em laboratrio a eficincia da dose comercial de alguns inseti-
cidas usados no controle das duas pragas. Para isso, foram coletadas
duas populaes de traa-do-tomateiro e uma populao de traa-
das-crucferas. Para traa-do-tomateiro, cartap, abamectin, lufenuron,
acefate e deltametrina causaram respectivamente 100; 90; 67 e 0%
de mortalidade das larvas. Para traa-das-crucferas, B. thuringiensis,
abamectin, cartap, acefate and deltametrina causaram 100; 96; 86;
79 e 5% de mortalidade respectivamente. De acordo com estes re-
sultados foi recomendada a suspenso imediata do uso de piretrides
e organofosforados para o controle das duas pragas. Abamectin e
cartap foram recomendados para o controle da traa-do-tomateiro e
B. thuringiensis para o controle de traa-das-crucferas.
Palavras-chave: Brassica oleracea, Lycopersicon esculentum,
Tuta absoluta, Plutella xylostella, tomate, repolho, couve-flor,
controle qumico, resistncia a inseticida.
ABSTRACT
Use of insecticides for controlling the South American Tomato
Pinworm and the Diamondback Moth: a case study.
In August 1999, at the Ncleo Rural da Taquara, Federal
District, Brazil, tomato and brassica crops were severely damaged
by the South American Tomato Pinworm (Tuta absoluta) and the
Diamondback Moth (Plutella xylostella). During that time growers
related that they had been spraying insecticides one to seven times
per week without controlling the pests. In the fields it was observed
that there were crops with different ages and levels of chemical
residues which allowed the pests to multiplicate continuously. Then
it was decided that the first step to solve the problem would be to
evaluate the efficacy of the recommended field rate of some
insecticides in laboratory bioassays. Two Brazilian Tomato Pinworm
populations and one Diamondback Moth population were collected.
Cartap, abamectin, lufenuron, acephate and deltamethrin caused 100;
90; 67 and 0% of larval mortality to the South American Tomato
Pinworm, respectively. B. thuringiensis, abamectin, cartap, acephate
and deltamethrin caused 100; 96; 86; 79 and 5% of mortality to the
Diamondback Moth, respectively. According to laboratory results it
was recommended that the use of pyrethroid and organophosphorous
compounds must be suspended immediately. Abamectin and cartap
must be used to control the South American Tomato Pinworm and
B. thuringiensis must be employed to Diamondback Moth control.
Keywords: Brassica oleracea, Lycopersicon esculentum, Tuta
absoluta, Plutella xylostella, tomato, cabbage, cauliflower,
chemical control, insecticide resistance.
(Aceito para publicao em 04 de janeiro de 2.001).
resistncia de traa-das-crucferas a di-
versos inseticidas j foi observada em
vrias partes do mundo (Castelo Branco
& Gatehouse, 1997; Cameron & Walker,
1998; Baker, 1999; Kovaliski, 1999).
Observaes preliminares de tomate
do local constataram a presena de mi-
nas de traa-do-tomateiro em pratica-
mente todas as folhas e, em alguns ca-
sos, at 100% de frutos danificados. Em
lavouras de brassicas foram observados
furos de traa-das-crucferas em folhas
de repolho e couve-flor e, em um cultivo
de couve-flor, foram encontradas mais de
100 larvas/planta. O sistema de produ-
o destas culturas envolvia: plantio con-
tnuo e sucessivo de tomate e brassicas;
abandono de restos culturais nas reas de
cultivo; mistura de inseticidas; utilizao
em rotao de dois ou trs produtos dife-
rentes, em uma mesma semana, sem ob-
servao de critrios tcnicos.
Zhao et al. (1995), em ensaios reali-
zados na China, observaram que testes
de laboratrio onde se avaliava a eficin-
cia da dose comercial de inseticidas para
o controle da traa-das-crucferas eram
bons indicadores da eficincia dos inse-
ticidas em campo. A fim de determinar
quais os inseticidas ineficientes para o
controle da traa-das-crucferas e traa-
do-tomateiro no Ncleo Rural da
Taquara, testes de laboratrio foram rea-
lizados. De posse destes dados e das ob-
servaes de campo, recomendaes para
o manejo da cultura foram sugeridas.
MATERIAIS E MTODOS
1. Populaes coletadas
Foram coletados ovos, larvas e
pupas de duas populaes de traa-do-
tomateiro (Populaes 1 e 2) e de uma
populao de traa-das-crucferas (Po-
pulao 3) no Ncleo Rural da Taquara.
Os agricultores forneceram dados sobre
os inseticidas utilizados e freqncia de
aplicao, conforme segue:
1.1 Traa-do-tomateiro: abamectin,
Bacillus thuringiensis, chlorfluzuron,
ciflutrina, deltametrina, fenpropatrina,
lufenuron, metomil, permetrina e
triflumuron. As pulverizaes foram
realizadas com um inseticida ou com
mistura de produtos a cada 24 horas.
1.2 Traa-do-tomateiro: abamectin,
Bacillus thuringiensis, betaciflutrina,
ciflutrina, cartap, fenpropatrina,
lufenuron, metomil, permetrina,
triflumuron. As pulverizaes eram rea-
lizadas a cada trs dias, com um inseti-
cida ou com mistura de dois inseticidas
(piretride + lufenuron).
1.3 Traa-das-crucferas: abamectin,
chlorfluzuron, deltametrina, metamidofs
e outros inseticidas no identificados. As
pulverizaes eram feitas com interva-
lo que variavam de um a trs dias.
2. Bioensaios
2.1. Traa-do-tomateiro: Foram
utilizadas larvas de segundo e terceiro
estdio de traa-do-tomateiro provenien-
tes diretamente do campo. Os insetici-
das abamectin (9 g.i.a./ha), acefate (750
g.i.a./ha), cartap (625 g.i.a./ha),
deltametrina (10 g.i.a./ha) e lufenuron
(40 g.i.a./ha) foram diludos consideran-
do-se o volume de calda de 1.000 L/ha.
Fololos que no continham larvas de
traa-do-tomateiro foram imersos na
soluo de inseticida por 10 segundos
e, em seguida, colocados para secar a
temperatura ambiente. Aps estarem
secos, 10-15 larvas de traa-do-tomatei-
ro foram colocadas em trs fololos em
placa de Petri (15 cm de dimetro). Em
outro teste, fololos infestados com lar-
vas de traa-do-tomateiro foram imersos
na soluo de inseticida por 10 segun-
dos (10-15 larvas/repetio) e transfe-
ridos para placas de Petri. Foram utili-
zadas quatro repeties por tratamento.
Para todos os inseticidas, a mortali-
dade de larvas foi avaliada aps 24 h,
exceto para lufenuron, onde a mortali-
dade de larvas foi avaliada aps seis
dias. Para este ltimo inseticida foi ain-
da avaliado o nmero de adultos emer-
gidos. Para a populao 1 foram testa-
dos abamectin, acefate, deltametrina e
lufenuron. Para a populao 2 foram tes-
tados acefate, cartap e deltametrina. Os
produtos cuja mortalidade de larvas foi
superior a 90% foram considerados como
eficientes para o controle da praga.
Para a anlise estatstica foi utiliza-
do o esquema fatorial 5 x 2 e 4 x 2 [in-
seticidas x posio das larvas nas folhas
(sobre ou dentro das minas)] para as
populaes 1 e 2, respectivamente. Os
dados foram submetidos anlise de
varincia e ao teste DMS (p<0,05) para
a separao das mdias.
2.2.Traa-das-crucferas: Larvas e
pupas foram coletadas em cultivo de
couve-flor e criadas em laboratrio at
a emergncia de adultos. Os adultos fo-
ram liberados em gaiola contendo fo-
lhas de repolho para a obteno dos ovos
os quais foram mantidos em caixas pls-
ticas com folhas de repolho at que as
larvas se desenvolvessem at o segun-
do estdio, quando foram utilizadas no
bioensaio. Foi avaliada a eficincia dos
seguintes inseticidas: acefate (750 g i.a./
ha), abamectin (9 g i.a./ha), Bacillus
thuringiensis (18 g i.a./ha), cartap (300
g i.a./ha) e deltametrina (6 gi.a./ha). As
diluies foram realizadas consideran-
do-se um volume de calda de 400 l/ha.
Discos de folhas de repolho de 4 cm
de dimetro foram imersos na soluo
inseticida e secos temperatura ambien-
te, no laboratrio. Foram transferidos
para placas de Petri com 9 cm de di-
metro e sobre cada folha foram coloca-
das 15 larvas de traa-das-crucferas.
A mortalidade de larvas foi avalia-
da aps 48 h. Os dados foram submeti-
das anlise de varincia e foi utilizado
o teste DMS (p<0,05) para a separao
de mdias.
2.3. Inimigos naturais: Um total de
50 ovos de traa-do-tomateiro foram
coletados no campo em cada uma das
duas reas de tomate. Os ovos foram
individualizados em cpsulas de gelati-
na para a verificao de ocorrncia de
parasitides.
Larvas de traa-das-crucferas
coletadas no campo foram tambm se-
paradas e criadas at o estgio de pupa.
Quando as pupas foram obtidas (172 no
total), foram individualizadas em cp-
sulas de gelatina para a observao da
emergncia de parasitides ou adultos
da praga.
RESULTADOS E DISCUSSO
1. Traa-do-tomateiro: Para as duas
populaes de traa-do-tomateiro hou-
ve apenas efeito do inseticida na morta-
lidade das larvas. No houve efeito da
posio das larvas sobre os fololos (lar-
vas sobre os fololos ou no interior des-
tes) nem da interao inseticida x posi-
o das larvas. Este resultado diferen-
te do observado por Castelo Branco &
Frana (1993) onde, quando folhas de
tomate foram tratadas com cartap, a
mortalidade de larvas de traa-do-toma-
teiro no interior das minas foi significa-
tivamente menor do que a mortalidade
de larvas sobre as folhas. A causa desta
diferena no pde ser identificada, mas
possvel que o grau de suscetibilidade
das populaes ao inseticida de alguma
maneira interfira nos resultados.
As doses comerciais dos inseticidas
deltametrina e acefate causaram a mor-
talidade de menos de 2% das larvas das
populaes 1 e 2 (Tabelas 1 e 2).
Lufenuron ficou em uma posio
intermediria, causando entre 67 e 72%
de mortalidade das lagartas (Tabela 1).
No entanto, este produto, por ser um
regulador de crescimento, afeta tambm
a emergncia de adultos. Uma mdia de
13% dos adultos emergiram, quando as
larvas foram colocadas sobre as folhas.
J quando as larvas estavam dentro das
folhas, este percentual subiu para 26%.
Ainda que mais de 10% dos adultos da
traa-do-tomateiro tenham emergido,
inseticidas reguladores de crescimento
como lufenuron afetam a fertilidade de
fmeas (Frana & Castelo Branco,
1996), podendo contribuir para a redu-
o da populao da praga em campo.
Abamectin causou a mortalidade de
mais de 90% das larvas da populao 1
(Tabela 1) e cartap causou a mortalida-
de de todas as larvas da populao 2
(Tabela 2). Estes resultados indicam que
estes dois inseticidas so os produtos
Uso de inseticidas para o controle da traa-do-tomateiro e traa-das-crucferas: um estudo de caso.
mais eficientes para o controle da praga
na regio.
Dos 50 ovos da traa-do-tomateiro
coletados de cada populao, nenhum
parasitide emergiu. Este resultado pode
indicar a ausncia de parasitides na
regio ou a eliminao destes.
2. Traa-das-crucferas:
Deltametrina foi o produto menos eficien-
te, causando a mortalidade de menos de
6% das larvas (Tabela 3). Acefate e
cartap se situaram em uma posio in-
termediria com uma mortalidade varian-
do de 79 a 86% (Tabela 3).Abamectin e
Bacillus thuringiensis causaram morta-
lidade superior a 96% (Tabela 3). Estes
resultados indicaram uma boa eficcia
dos dois inseticidas para o controle da
praga. Abamectin no registratdo para
a cultura de brssicas, no tendo por-
tanto o seu uso recomendado.
Das 172 pupas de traa-das-
crucferas obtidas, apenas duas estavam
parasitadas. Uma por Apanteles sp. e a
outra por Oomyzus sokolowiskii. Entre
as pupas 87 originaram adultos e 83 no
emergiram. Esta baixa ocorrncia de
parasitides pode ser atribuda ao ele-
vado nmero de aplicaes de insetici-
da e ao uso de produtos extremamente
txicos como por exemplo metamidofs
e deltametrina (Talekar & Yang, 1991;
Kao & Tzeng, 1992). Este resultado di-
fere do observado por Frana &
Medeiros (1998) onde em uma avalia-
o de inseticidas em campo foi obser-
vada populao alta de parasitides (m-
dia > 4,0 adultos por planta) nas parce-
las tratadas com deltametrina, indican-
do a sobrevivncia destes no local do
experimento. Como nesta rea de culti-
vo foram utilizados diferentes tipos de
inseticida, no foi possvel a identifica-
o dos produtos que mais contriburam
para a reduo da populao dos

Tabela 1. Mortalidade de larvas de traa-do-tomateiro tratadas com diferentes inseticidas. Larvas sobre folhas ou no interior das minas.
Populao 1. Taquara, Embrapa Hortalias, 1999.
1/
nmero de larvas encontradas aps 24 h para todos os inseticidas a exceo de Match, onde o nmero de larvas o nmero de larvas
encontrado aps seis dias.
Mdias seguidas de mesma letra no diferem entre si pelo teste DMS (p> 0,05)

Tabela 2: Mortalidade de larvas de traa-do-tomateiro tratadas com diferentes inseticidas. Larvas sobre folhas ou no interior das minas.
1/
nmero de larvas encontradas aps 24 h para todos os inseticidas a exceo de Match, onde o nmero de larvas o nmero de larvas
encontrado aps seis dias.
M. Castelo Branco et al.
parasitides. Estudos que visem avaliar
a seletividade de alguns destes produ-
tos se fazem necessrios.
sabido que as doses recomenda-
das de qualquer inseticida so capazes
de matar um determinado percentual da
populao da praga, geralmente 95%,
independentemente da sua densidade
populacional (Knipling, 1979). Ento,
quando a densidade populacional bai-
xa, os produtos tendem a ser mais efi-
cientes do que quando a densidade
populacional mais elevada. No Ncleo
Rural da Taquara, o sistema de produ-
o de tomate e brassicas (plantios su-
cessivos e no eliminao de restos cul-
turais) e as condies ambientais (tempo
quente e seco) eram favorveis ao cres-
cimento descontrolado das populaes de
traa-das-crucferas e traa-do-tomatei-
ro (Frana et al., 1985; Haji et al., 1988;
Castelo Branco, 1992). Deste modo, ne-
nhum inseticida, mesmo os considerados
eficientes em testes de laboratrio, apre-
sentaram eficincia no campo.
Assim, para a viabilizao de lavou-
ras de tomate e brassicas na regio e
sobrevivncia de parasitides e preda-
dores que possam auxiliar na reduo
das populaes das pragas, so neces-
srias a implementao de medidas ra-
cionais de uso de inseticidas e outras
prticas de manejo da cultura que visem,
principalmente, reduzir as condies
favorveis ao crescimento populacional
dos insetos. So recomendadas as se-
guintes medidas:
a) pulverizaes semanais de inseti-
cidas;
b) eliminao de inseticidas perten-
centes a grupos qumicos considerados
ineficientes nos testes de laboratrio;
c) introduo de um esquema de rota-
o de inseticidas (Castelo Branco, 2.000);
c) uso de irrigao por asperso para
remoo de ovos e mortalidade de lar-
vas e pupas (Costa et al., 1998;
Junqueira et al., 1998);
d) destruio de restos culturais;
e) no utilizao de plantio
seqenciado de tomate ou brssicas.
As recomendaes aqui descritas
foram seguidas por um agricultor do
Ncleo Rural da Taquara e com isso foi
recuperada uma lavoura de tomate e um
plantio de couve-flor que j haviam sido
considerados perdidos.
AGRADECIMENTOS
A Hozanan P. Chaves pelo auxlio
nos trabalhos de campo e laboratrio.
Aos agricultores do Ncleo Rural da
Taquara pelas informaes prestadas.
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Workshop. Taiwan: AVRDC, 1992. p. 287-
296.
KNIPLING, E.F. The basic principles of insect
population suppression and management. Wa-
shington: USDA, 1979. 659 p.
SIQUEIRA, H.A.A.; GUEDES, R.N.C.;
PICANO, M.; OLIVEIRA, E.E. Cartap
resistance and synergism in populations of Tuta
absoluta (Lepidoptera: Gelechiidae). In.
INTERNATIONAL CONGRESS OF
ENTOMOLOGY, 21; BRAZILIAN
CONGRESS OF ENTOMOLOGY, 18., 2000.
Foz do Iguau, Abstracts. Londrina: SEB/
Embrapa Soja, 2000. v. 1, p. 353.
TALEKAR, N.S.; YANG, J.C. Characteristic of
parasitism of diamondback moth by two larval
parasites. Entomophaga, v. 36, p. 95-104,
1991.
ZHAO, J.Z.; WU, S.C.; ZHU, G.R. Bioassays with
recommended field concentrations of several
insecticides for resistance monitoring in
Plutella xylostella. Resistant Pest
Management, v. 7, n. 1, p. 13-14,1995.

Tabela 3 Mortalidade de larvas de traa-das-crucferas tratadas com diferentes inseticidas.
Uso de inseticidas para o controle da traa-do-tomateiro e traa-das-crucferas: um estudo de caso.
Agricultura Tcnica
Agric. Tc. v.61 n.4 Chilln oct. 2001
RESPUESTA DE LA POLILLA DEL TOMATE, Tuta absoluta
(Meyrick),
A INSECTICIDAS EN ARICA
1

Tomato moth, Tuta absoluta (Meyrick) response to
insecticides in Arica, Chile
Erika R. Salazar
2
y Jaime E. Araya
3

1
Recepcin de originales: 22 de noviembre de 1999.
2
Instituto de Investigaciones Agropecuarias, Centro Regional de Investigacin La
Platina, Casilla 439, Correo 3, Santiago, Chile.
3
Universidad de Chile, Facultad de Ciencias Agronmicas, Departamento de Sanidad
Vegetal, Casilla 1004, Santiago, Chile. E-mail: jaraya@uchile.cl
ABSTRACT
Larval susceptibility of Tuta absoluta (Meyrick) collected on tomato (Lycopersicon
esculentun Mill.) crops in Azapa, Arica (18 31 S lat, 70 11 W long), Chile, was
compared by toxicological tests with several commonly used insecticide doses applied
on two development stage larvae groups (stadia 1-2 and 3-4). To determine resistance
to insecticides, LD
50
, LD
90
, and regression slopes between probit mortality and log
dosage were calculated. Resistance to studied insecticides was verified, since LD
50
at
least doubled those in Ovalle and Quillota, locations where T. absoluta had the
greatest resistance in other study. Deltamethrin and mevinphos were the least and
most toxic compounds, respectively. Larvae of both development levels were equally
susceptible to deltamethrin, while larger larvae were more resistant to mevinphos than
smaller ones. Results with esfenvalerate and - cyhalothrin on large larvae, and
metamidophos on small larvae, were too variable and this caused a defective probit
analysis and resistance evaluation to these insecticides in such larval groups. Parasite
may have also developed resistance to insecticides; this may explain the resurgence of
populations in the Valley. Possible parasite resistance, which are present even with
insecticide treatments at extremely high dosages, could have affected accuracy in the
regressions obtained, particularly on large larvae.
Key words: Deltamethrin, esfenvalerate, - cyhalothrin, mevinphos, metamidophos.
RESUMEN
Se compar la susceptibilidad larvaria de la polilla del tomate, Tuta absoluta
(Meyrick), colectada en tomate (Lycopersicon esculentun Mill.) en Azapa, Arica (18
31 lat. Sur, 70 11 long. Oeste), mediante pruebas de toxicologa con varias dosis de
insecticidas de uso comn, aplicados sobre grupos de larvas de dos niveles de
desarrollo (estados 1-2 y 3-4). Para determinar la resistencia a los insecticidas se
calcularon las DL
50
, DL
90
y pendientes de las regresiones entre mortalidad (probit) y
dosis (log). Se verific la resistencia a los insecticidas estudiados, pues las DL
50
al
menos duplicaron aquellas en Ovalle y Quillota, localidades donde T. absoluta
present la mayor resistencia en otro estudio. Deltametrina y mevinfos fueron los
compuestos menos y ms txicos, respectivamente. Las larvas de ambos niveles de
desarrollo fueron igualmente susceptibles a deltametrina, mientras que las larvas
grandes fueron ms resistentes a mevinfos que las pequeas. Los resultados con
esfenvalerato y - cihalotrina sobre larvas grandes, y metamidofos en larvas
pequeas fueron muy variables, lo que impidi un buen ajuste probit y la evaluacin
de resistencia a estos compuestos en dichos grupos larvarios. El parsito tambin
puede haber desarrollado resistencia a insecticidas, lo que puede explicar que en el
Valle se hayan restablecido sus poblaciones. La posible resistencia de parsitos,
presentes incluso en tratamientos insecticidas a dosis extremadamente altas, podra
haber afectado la precisin de las regresiones obtenidas, especialmente sobre larvas
grandes.

Palabras clave: Deltametrina, esfenvalerato, - cihalotrina, mevinfos, metamidofos.
INTRODUCCIN
La polilla Tuta absoluta (=Scrobipalpuloides absoluta) (Meyrick) (Lepidoptera:
Gelechiidae) es clave en el cultivo del tomate (Lycopersicon esculentum Mill.) en
Chile (Gonzlez, 1989; Prado, 1991); en cada temporada, el cultivo requiere
aplicaciones frecuentes de insecticidas para evitar una reduccin drstica de la
produccin y calidad de los frutos (Vargas, 1970; Apablaza, 1984). Salazar y Araya
(1997) describieron la importancia de los daos y comprobaron el desarrollo de
resistencia a algunos de estos compuestos mencionado por diversos autores (Acua,
1970; Vargas, 1970; Campos, 1976; Moore, 1983) en varias localidades productoras.
Los insecticidas pueden aumentar los rendimientos al reducir los daos causados por
las plagas, pero su uso repetido puede seleccionar gradualmente insectos resistentes
(Lockwood et al., 1984; Brattsten, 1989; Metcalf, 1989).
Los cultivos de tomate en el Valle de Azapa reciben 15-17 aplicaciones de numerosos
insecticidas en la temporada para controlar a T. absoluta, muchas veces en mezclas y
con dosis mayores a las comerciales, lo que sugiere el desarrollo de resistencia. El
objetivo de este estudio fue evaluar en laboratorio, con una metodologa simple y de
bajo costo, la efectividad de algunos insecticidas de uso habitual en el control de la
polilla del tomate en el Valle de Azapa, utilizando larvas obtenidas en rastrojos de
tomate.
MATERIALES Y MTODOS
Las pruebas se hicieron con larvas colectadas en rastrojos de tomate en el Valle de
Azapa, en el Centro de Investigacin y Capacitacin Agrcola (CICA), San Miguel de
Azapa (18 31 lat Sur; 70 11 long Oeste), Universidad de Tarapac (a 12 km de
Arica), en 1995. Se evaluaron los piretroides deltametrina (Decis 2,5 EC),
esfenvalerato (Halmark 7% LE) y - cihalotrina (Karate 5% EC), y los fosforados
mevinfos (Phosdrin 24% LE) y metamidofos [Monitor 600 (g L
-1
) CS], por su
amplia utilizacin por productores de tomate (Salazar y Araya, 1997).
Las larvas se colectaron del follaje de rastrojos de tomate con pincel y se utilizaron
diariamente, manteniendo el excedente a baja temperatura (alrededor de 5C) para uso
en pruebas posteriores. En cada tratamiento se usaron larvas pequeas (estados 1-2,
de 1,0-2,5 mm de longitud) y grandes (estados 3-4, de 4,5-7,5 mm). Las larvas de
2,5-4,5 mm se descartaron, para separar claramente ambos tamaos larvarios.
En las pruebas se determin la mortalidad por contacto, asperjando con un aspersor
manual De Vilbiss 1 mL de solucin insecticida sobre grupos de 20 larvas
seleccionadas al azar, en placas Petri inclinadas 45 de la horizontal, con papel filtro
N1 en el fondo. Aspersiones sobre papel fotogrfico determinaron una gota similar a
la de la torre Potter utilizada por Salazar y Araya (1997) y el aspersor manual a 1 m
entre el aspersor y la placa Petri en este estudio. Se aplicaron al menos cinco dosis
crecientes por insecticida y tamao larvario, desde las dosis mnimas comerciales (0,4
mL L
-1
de mezcla insecticida para los piretroides, y 1 2 mL L
-1
para los fosforados
metamidofos y mevinfos, respectivamente), con tres repeticiones por tratamiento. Una
vez secas, las larvas tratadas se trasladaron a frascos con follaje fresco de tomate sin
insecticida y se mantuvieron a 182C. La mortalidad se evalu a las 48 h,
considerando muertas aquellas larvas sin motilidad al ser estimuladas con pincel.
Para verificar la accin de cada insecticida (Busvine, 1980), los resultados de
mortalidad se corrigieron por la frmula de Abbott (1925) y procesaron mediante
anlisis probit (Finney, 1971; Busvine, 1980), siguiendo la metodologa aplicada por
Salazar y Araya (1997), para calcular, utilizando el programa computacional POLO
PC, las DL
50
, DL
90
y pendientes de las regresiones entre mortalidad (probit) y dosis
(log). Se realizaron pruebas de Chi
2
para comprobar el ajuste entre las mortalidades
obtenidas y las esperadas, y corroborar el anlisis probit. Las DL
50
se analizaron
mediante anlisis de varianza; los promedios se separaron mediante pruebas de rango
mltiple de Duncan (1955). Las diferencias estadsticas entre las pendientes de las
regresiones se evaluaron mediante la prueba t de Student.
RESULTADOS Y DISCUSIN
Los resultados del anlisis por insecticida se presentan en los Cuadros 1-3. Al no
obtenerse mortalidad mayor al 90% con dosis razonables de insecticida, algunos DL
90

se estimaron con las ecuaciones obtenidas con el programa POLO PC.
El Cuadro 1 presenta las DL
50
, intervalos de confianza al 95% y pendientes
desviacin estndar de la regresin lineal de los insecticidas evaluados.

Cuadro 1. Respuesta de larvas de Tuta. absoluta de San Miguel de Azapa, Arica
1
.
Table 1. Tuta absoluta larvae response from San Miguel de Azapa, Arica
1
.
Insecticidas DL
50
Intervalo de
confianza al 95%
Pendiente desviacin estndar
Estados 1-2
Deltametrina 571,26 201,30 - 412,09 2,720,31
Esfenvalerato 46,12 _ 1,180,25
- cihalotrina 35,28 17,40 - 332,94 0,960,16
Mevinfos 21,45 - 3,110,42
Estados 3-4
Deltametrina 476,09 _ 2,390,46
Mevinfos 35,84 26,02 - 53,76 23,290,35
Metamidofos 24,21 _ 1,940,29
1
Las poblaciones no presentaron distribucin normal, lo que impidi su anlisis estadstico.
Las pruebas de Chi
2
indicaron que las rectas log (dosis) x mortalidad de esfenvalerato
y - cihalotrina para larvas grandes (estados 1-2), y metamidofos para larvas
pequeas (estados 3-4), no se ajustaron a poblaciones de distribucin normal de
tolerancia, lo que impidi su anlisis estadstico. Sin embargo, a pesar que la
metodologa en este ensayo es menos precisa que la que utiliza la torre Potter,
igualmente se constat la poca eficiencia y la posible resistencia a los insecticidas
estudiados, pues las DL
50
obtenidas al menos duplican aquellas determinadas para
Ovalle y Quillota, localidades con los mayores niveles de resistencia en el estudio de
Salazar y Araya (1997).
Deltametrina no caus diferencias significativas de susceptibilidad entre ambos
tamaos larvarios, aunque la mayor DL
50
numrica ocurri con las larvas pequeas
(Cuadro 2). Este resultado, diferente a los obtenidos por Salazar y Araya (1997) en
otras localidades, pudo deberse al diferente mtodo de aplicacin de los insecticidas.
En el tratamiento con mevinfos las larvas grandes fueron ms resistentes que las
pequeas.

Cuadro 2. DL
50
(mL L
-1
) de los dos grupos larvarios (estadios 1-2 y 3-4) de Tuta
absoluta de San Miguel de Azapa, Arica, tratados con cinco insecticidas
1
.
Table 2. LD
50
(mL L
-1
) of both larval groups (stadia 1-2 and 3-4) of Tuta absoluta
from San Miguel de Azapa, Arica, treated with five insecticides
1
.
Larvas Estados 1-2 Estados 3-4
Deltametrina 571,26 a 447,09 a
Esfenvalerato 46,12 -
-
cihalotrina
35,27 -
Mevinfos 21,45 b 35,84 a
Metamidofos - 24,21
1
Promedios en cada columna con letras iguales no son diferentes significativamente (P=0,05).
El signo - indica que no hubo un buen ajuste probit.
Para las larvas de San Miguel de Azapa, deltametrina y mevinfos fueron los
compuestos de menor y mayor toxicidad, respectivamente (Cuadro 3).

Cuadro 3. Resistencia relativa de las larvas de Tuta absoluta de San Miguel de
Azapa, Arica, a los insecticidas, en DL
50
mltiplos de la dosis comercial
1
.
Table 3. Tuta absoluta larvae from San Miguel de Azapa, Arica, relative resistance to
the insecticides, in LD
50
-folds of the commercial dosage
1
.
Ingrediente activo Estados 1-2 Estados 3-4
Deltametrina 1428,16 a 1117,72a
Esfenvalerato 115,30 b -
- cihalotrina 88,19 b -
Mevinfos 17,92 c 10,72 c
Metamidofos - 24,21 b
1
Promedios en cada columna con letras iguales no son diferentes significativamente (P=0,05).
El signo - indica que no hubo un buen ajuste probit.

Salazar y Araya (1997) encontraron factores de resistencia (FR) a deltametrina de 7,1
y 8,2 para larvas grandes y pequeas, respectivamente, de T. absoluta de Ovalle, y
adems confirmaron la resistencia a este piretroide en Quillota y Colina. Tambin
encontraron resistencia a esfenvalerato en Ovalle y Quillota, e incipiente en Colina,
con FR de 2,0 y 1,9 para larvas grandes y pequeas, respectivamente. Las mayores
DL
50
para - cihalotrina en larvas pequeas y grandes ocurrieron en Ovalle y
Quillota, respectivamente, con un menor nivel de resistencia en Colina. Sin embargo,
estos autores indicaron que podran haber subestimado los FR para deltametrina y
esfenvalerato, debido a las altas dosis requeridas en ambos casos para matar al 50%
de la poblacin (control) susceptible de Requinoa, la que ya tendra algn grado de
resistencia a estos insecticidas. Las poblaciones de Ovalle, Quillota y Colina fueron
menos susceptibles a - cihalotrina. Las de Ovalle, Quillota y Colina fueron menos
susceptibles para metamidofos que las de Requinoa. Para mevinfos, los mayores FR
ocurrieron en Ovalle (5,45 y 4,72 para larvas grandes y pequeas, respectivamente).
Para todas las localidades, mevinfos fue el insecticida de mayor toxicidad relativa y
por ende ms efectivo. Sin embargo, el uso de este insecticida fue prohibido en Chile
en 1995 por el Servicio Agrcola y Ganadero.

En todas las localidades evaluadas por Salazar y Araya (1997), las larvas 3-4 fueron
menos susceptibles a los insecticidas estudiados, en general con DL
50
1,2-2,5 veces
mayores que para larvas de menor desarrollo (estados 1-2). Estos resultados
confirman otros informados en la literatura, pues la susceptibilidad a un producto
qumico disminuye con el aumento del tamao larvario (Busvine, 1980).
En la comparacin de Salazar y Araya (1997) de la toxicidad relativa de los
insecticidas evaluados, mevinfos y deltametrina fueron los compuestos ms y menos
txicos, respectivamente, sobre todas las poblaciones de T. absoluta. De los
piretroides evaluados para todas las localidades y tamaos larvarios, - cihalotrina
fue el ms txico. En Ovalle, mevinfos y - cihalotrina fueron el fosforado y el
piretroide ms txicos. Metamidofos fue ms txico que - cihalotrina sobre larvas
grandes, mientras que ambos insecticidas no se diferenciaron estadsticamente en
larvas pequeas. En Quillota, mevinfos y - cihalotrina fueron los insecticidas ms
txicos. Los fosforados fueron ms txicos que los piretroides en larvas grandes,
mientras que mevinfos fue ms txico que - cihalotrina sobre larvas pequeas.
Metamidofos fue el tercer insecticida en toxicidad. En Colina, mevinfos y -
cihalotrina fueron tambin los insecticidas ms txicos; se observ adems, un
aumento en la toxicidad de esfenvalerato sobre los resultados para Quillota y Ovalle,
aunque sin diferencia estadstica con metamidofos. En Requinoa, al igual que en las
otras localidades, mevinfos y - cihalotrina fueron los compuestos ms txicos.
Ambos insecticidas presentaron resultados similares, con DL
50
cercanos a las dosis
comerciales recomendadas. Destacaron adems, las DL
50
para deltametrina sobre los
estados 3-4 (29,85 veces) y 1-2 (12,34 veces), lo que evidenci una alta resistencia
para deltametrina en esta localidad.
Si los resultados se comparan con los de Salazar y Araya (1997), se puede concluir
que la poblacin de Azapa ha desarrollado alta resistencia a los insecticidas
estudiados. Esta hiptesis debe ser comprobada, sin embargo, utilizando una
metodologa de precisin similar a la de ese estudio, e incluyendo una poblacin
control susceptible.
Durante la evaluacin de mortalidad se observaron algunas larvas grandes con alguna
motilidad al estimulo con pincel, pero con tegumento marrn oscuro, por lo que se
consideraron muertas. En otros recuentos se encontraron pupas de parsitos adosadas
al cuerpo de las larvas de T. absoluta, revelando que aquellas larvas oscuras
consideradas muertas estaban parasitadas. Se criaron algunas pupas, las que sin
embargo se deshidrataron y no produjeron parasitoides adultos para su identificacin.
El parsito tambin puede haber desarrollado resistencia a insecticidas, lo que ha
permitido que en el Valle de Azapa se restablezcan sus poblaciones (Vargas, 1970).
La posible resistencia de parsitos larvarios es lgica al encontrarse larvas y/o pupas
en sectores tratados con insecticidas en concentraciones extremadamente altas. Este
factor podra haber afectado la precisin de las regresiones en los tratamientos sobre
larvas grandes.
Una forma razonable de manejar la resistencia a los plaguicidas son programas de
manejo integrado que reduzcan la frecuencia e intensidad de la seleccin y apliquen
un mejor control natural y cultural, y el uso de variedades resistentes. En conjunto,
estas medidas pueden eliminar gran parte de los individuos seleccionados antes que
produzcan una progenie resistente a insecticidas (Metcalf, 1980). Para el manejo
integrado es vital disminuir la presin de seleccin, aplicar insecticidas con menor
frecuencia, evitar el uso de compuestos persistentes en el ambiente, formulaciones de
lenta liberacin, compuestos con presin de seleccin en varios estados de desarrollo
del insecto, e incorporar mtodos alternativos biolgicos y culturales (Metcalf, 1980,
1989). Para manejar los insecticidas se deben analizar las poblaciones para conocer la
susceptibilidad original y detectar temprano el desarrollo de resistencia, de manera de
extender la vida til de un insecticida hasta que la respuesta de la poblacin indique la
necesidad de cambiarlo. La secuencia de insecticidas alternativos debe limitar los
compuestos con resistencia simple y cruzada (e.g., dimetoato y piretroides) y el uso de
mezclas insecticidas. Se debe considerar el umbral econmico, utilizar mtodos
adecuados y aplicacin oportuna, usar plaguicidas inocuos para los enemigos
naturales, y productos selectivos en vez de compuestos txicos de amplio espectro.
Aunque la metodologa en este trabajo logr resultados muy variables y fue menos
precisa que la utilizada por Salazar y Araya (1997) con una torre Potter en laboratorio,
nuestros resultados comprobaron la ineficacia de los insecticidas ms utilizados contra
T. absoluta en el Valle de Azapa, verificando el deterioro ambiental y productivo
causado por la intervencin con insecticidas de amplio espectro en ambientes poco
diversificados y ecolgicamente inestables. Se debe estudiar la incorporacin de
nuevos compuestos insecticidas con diversos modos de accin, de manera de evitar el
desarrollo de resistencia y/o regenerar el sistema productivo de tomates en el extremo
norte de Chile, con el objeto de proteger esta importante actividad agrcola, fuente de
ingresos para numerosos productores durante todo el ao, y evitar repetir situaciones
de crisis productivas de tipo regional como las descritas por Metcalf (1989).
CONCLUSIONES
Aunque la metodologa utilizada permiti una medicin rpida de la prdida de
efectividad de los insecticidas ms utilizados contra la polilla del tomate en el Valle
de Azapa, algunos resultados fueron muy variables, lo que impidi un buen ajuste
probit y la evaluacin consiguiente de resistencia a estos compuestos. Estos estudios
pueden afinarse mediante el uso de una torre de precisin tipo Potter y la comparacin
con una poblacin susceptible. Tambin se debe estudiar el desarrollo de resistencia a
insecticidas en los enemigos naturales de T. absoluta, con miras a su posible
utilizacin en programas de control integrado de esta plaga.
AGRADECIMIENTOS
Los autores agradecen la valiosa colaboracin del Ingeniero Agrnomo Sr. Juan
Machuca del Servicio Agrcola y Ganadero, Arica, y de los entomlogos Sres. Hctor
Vargas y Dante Bobadilla, CICA, San Miguel de Azapa, Universidad de Tarapac.
LITERATURA CITADA
Abbott, W.S. 1925. A method for computing the effectiveness of an insecticide. J.
Econ. Entomol. 18:265-267. [ Links ]
Acua, J. 1970. Control qumico de la polilla del tomate Gnorimoschema absoluta
(Meyr.). IDESIA, Universidad del Norte (Chile) 1:75-110. [ Links ]
Apablaza, J. 1984. Incidencia de insectos y moluscos plagas en siete hortalizas
cultivadas en las regiones V y Metropolitana, Chile. Ciencia e Investigacin Agraria
11:27-34. [ Links ]
Brattsten, L.B. 1989. Insecticide resistance: Research and management. Pestic. Sci.
26: 329-332. [ Links ]
Busvine, J.R. 1980. Recommended methods for measurement of pest resistance to
pesticides. 132 p. FAO, Rome, Italy. [ Links ]
Campos, R. 1976. Control qumico del "Minador de las hojas y tallos de la papa"
(Scrobipalpula absoluta M.) en el Valle de Caete. Rev. Peruana Entomol. 19:102-
106. [ Links ]
Duncan, D.B. 1955. Multiple range and multiple F tests. Biometrics 11:1-42.
[ Links ]
Finney, D.J. 1971. Probit analysis. 33 p. 3 ed. Cambridge Univ. Press, London,
England. [ Links ]
Gonzalez, R.H. 1989. Insectos y caros de importancia agrcola y cuarentenaria en
Chile. 310 p. Ograma, Santiago, Chile. [ Links ]
Lockwood, J.A., T.C. Sparks, and R.N. Story. 1984. Evolution of insect resistance to
insecticide: A reevaluation of the roles of physiology and behavior. Bull. Entomol.
Soc. Amer. 30:41-51. [ Links ]
Metcalf, R.L. 1980. Changing role of insecticides in crop protection. Ann. Rev.
Entomol. 25:219-256. [ Links ]
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[ Links ]
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Management 29:231-238. [ Links ]
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en Chile. 207 p. Serie Bol. Tcnico N 169. Instituto de Investigaciones
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tomate. Simiente 67:8-22. [ Links ]
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Insecticide resistance in populations of Tuta absoluta (Lepidoptera: Gelechiidae)
Herbert lvaro A. Siqueira, Raul Narciso C. Guedes and Marcelo C. Picano
Departamento de Biologia Animal, Universidade Federal de Viosa, Viosa, MG 36571 000, Brazil
Correspondence: Raul Narciso C. Guedes. Tel: + 55 31 899 2006; fax: + 55 31 899 2537; e-mail: guedes@mail.ufv.br
KEYWORDS
Abamectin cartap methamidophos permethrin tomato leafminer
ABSTRACT
Abstract
1 Control failures of insecticides used against the tomato leafminer Tuta absoluta (Meyrick) (Lepidoptera: Gelechiidae) in Brazil
led to the investigation of the possible occurrence of resistance of this insect pest to abamectin, cartap, methamidophos and
permethrin.
2 The insect populations were collected from seven sites in the states of Minas Gerais, Rio de Janeiro, and So Paulo. These
populations were subjected to concentrationmortality bioassays using insecticide-impregnated filter papers.
3 We were unable to obtain a single population which provided a susceptibility standard for all insecticides tested. Therefore,
the resistance levels were estimated in relation to the most susceptible population to each insecticide. Resistance to abamectin
and cartap were observed in all populations when compared with the susceptible standard population, with resistance ratios
ranging from 5.2- to 9.4-fold and from 2.2- to 21.9-fold for abamectin and cartap, respectively. Resistance to permethrin was
observed in five populations with resistance ratios ranging from 1.9- to 6.6-fold, whereas resistance to methamidophos was
observed in four populations with resistance ratios ranging from 2.6- to 4.2-fold.
4 The long period and high frequency of use of these insecticides against this insect pest suggest that the evolution of
insecticide resistance on them has been relatively slow. Alternatively, the phenomenon might be widespread among Brazilian
populations of T. absoluta making the finding of suitable standard susceptible populations difficult and leading to an
underestimation of the insecticide resistance levels in this pest.
5 Higher levels of resistance to abamectin, cartap and permethrin are correlated with greater use of these compounds by
growers. This finding suggests that local variation in insecticide use was an important cause of variation in susceptibility.
Accepted: 14 April 2000;
DIGITAL OBJECT IDENTIFIER (DOI)
10.1046/j.1461-9563.2000.00062.x About DOI
Introduction
The tomato leafminer Tuta absoluta (Meyrick) (Lepidoptera: Gelechiidae) is an oligophagous insect which attacks solanaceous
crops, especially tomato, where it is one of its main pests ( Picano et al. 1998 ). This insect was originally described in Peru,
but it is widespread throughout South America ( Povolny 1975; Carballo et al. 1981 ; Muszinski et al. 1982 ; Souza & Reis
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Published Online: 24 Dec 2001
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Insecticide resistance in populations of Tuta absoluta (Lepidoptera: Gelechiidae)
Herbert lvaro A. Siqueira, Raul Narciso C. Guedes and Marcelo C. Picano
Departamento de Biologia Animal, Universidade Federal de Viosa, Viosa, MG 36571 000, Brazil
Correspondence: Raul Narciso C. Guedes. Tel: + 55 31 899 2006; fax: + 55 31 899 2537; e-mail: guedes@mail.ufv.br
KEYWORDS
Abamectin cartap methamidophos permethrin tomato leafminer
ABSTRACT
Abstract
1 Control failures of insecticides used against the tomato leafminer Tuta absoluta (Meyrick) (Lepidoptera: Gelechiidae) in Brazil
led to the investigation of the possible occurrence of resistance of this insect pest to abamectin, cartap, methamidophos and
permethrin.
2 The insect populations were collected from seven sites in the states of Minas Gerais, Rio de Janeiro, and So Paulo. These
populations were subjected to concentrationmortality bioassays using insecticide-impregnated filter papers.
3 We were unable to obtain a single population which provided a susceptibility standard for all insecticides tested. Therefore,
the resistance levels were estimated in relation to the most susceptible population to each insecticide. Resistance to abamectin
and cartap were observed in all populations when compared with the susceptible standard population, with resistance ratios
ranging from 5.2- to 9.4-fold and from 2.2- to 21.9-fold for abamectin and cartap, respectively. Resistance to permethrin was
observed in five populations with resistance ratios ranging from 1.9- to 6.6-fold, whereas resistance to methamidophos was
observed in four populations with resistance ratios ranging from 2.6- to 4.2-fold.
4 The long period and high frequency of use of these insecticides against this insect pest suggest that the evolution of
insecticide resistance on them has been relatively slow. Alternatively, the phenomenon might be widespread among Brazilian
populations of T. absoluta making the finding of suitable standard susceptible populations difficult and leading to an
underestimation of the insecticide resistance levels in this pest.
5 Higher levels of resistance to abamectin, cartap and permethrin are correlated with greater use of these compounds by
growers. This finding suggests that local variation in insecticide use was an important cause of variation in susceptibility.
Accepted: 14 April 2000;
DIGITAL OBJECT IDENTIFIER (DOI)
10.1046/j.1461-9563.2000.00062.x About DOI
Introduction
The tomato leafminer Tuta absoluta (Meyrick) (Lepidoptera: Gelechiidae) is an oligophagous insect which attacks solanaceous
crops, especially tomato, where it is one of its main pests ( Picano et al. 1998 ). This insect was originally described in Peru,
but it is widespread throughout South America ( Povolny 1975; Carballo et al. 1981 ; Muszinski et al. 1982 ; Souza & Reis
1 de 7 1/11/2010 2:48 PM
1986). It was initially reported in Brazil between September 1979 and October 1980 at Morretes county, state of Paran, from
where it spread throughout the country, and has been considered a serious problem for tomato production since then (
Muszinski et al. 1982 ; Souza & Reis 1986; Frana 1993; Guedes et al. 1994 ; Picano et al. 1995 ).
The larval injury to the tomato plant leads to direct yield loss. The leafminers attack the tomato plants in all of their
developmental stages, damaging the stems, apices, flowers and fruits besides mining their leaves ( Souza et al. 1992 ; Miranda
et al. 1998 ). This kind of damage resembles those of other Gelechiidae pests of solanaceous crops, such as Keiferia
lycopersicella (Wals.) in Central and North America and Phthorimaea operculella (Zell.) in the Americas, Europe, Africa and
Asia ( Sannino & Nicodemo 1979; Juvick et al. 1982 ; Abbas et al. 1993 ; Ebora et al. 1994 ; Pouey et al. 1994 ). When in high
densities, T. absoluta is able to cause significant production losses in tomato crops ( Souza et al. 1992 ; Picano et al. 1998 ).
The main control method for T. absoluta is the use of insecticides, but some of the compounds recommended for its control are
apparently not providing the desired effect ( Castelo Branco & Frana 1992; Guedes et al. 1994 ). It has been hypothesized that
excessive insecticide applications commonly applied to the tomato crop during a single cultivation period, sometimes up to 36
sprays, could have led to the evolution of resistant populations, besides eliminating their natural enemies, and leading to
additional occupational hazards ( Castelo Branco & Frana 1992; Gonalves et al. 1994 ; Picano et al. 1995 ).
Permethrin and cartap were initially the only insecticides for use against T. absoluta, leading to their large scale utilization in
Brazil for a long period ( Souza & Reis 1986). The use of abamectin and methamidophos is more recent (i.e. early 1990s) and
abamectin, in mixture with mineral oil, shows high efficiency against the pest ( Guedes et al. 1995 ; Castelo Branco et al. 1996 ;
Picano et al. 1998 ). However, some investigators pointed out that the use of mineral oil in insecticide mixtures can increase
problems of insecticide resistance in insect pest populations ( Castelo Branco et al. 1996 ; Picano et al. 1996 ).
Insecticide resistance has been reported all over the world to almost every group of insecticides used against insect pests (
Lockwood et al. 1984 ; Georghiou 1986; Kay & Collins 1987). Therefore, it is necessary to develop management tactics to delay
or even prevent the evolution of insecticide resistance in insect pest populations, and the detection and monitoring of such
phenomena is of key importance to achieve this ( Dennehy et al. 1983 ; Tabashnik & Roush 1990). Owing to control failures of
insecticides used against T. absoluta and the report of resistant populations of this pest in Chile ( Moore 1983; Salazar & Araya
1997), the present study was carried out to detect the existence of Brazilian populations of T. absoluta resistant to the main
insecticides used against it and to quantify that resistance and its relationship with insecticide use in seven sites in Brazil.
Materials and methods
Five populations of T. absoluta from the state of Minas Gerais, one population from the state of Rio de Janeiro, and another
from So Paulo ( Fig. 1; Table 1) were used in this study. These last two populations were obtained from laboratory colonies,
established from local populations, with intended use as potential standard susceptible populations. For each of these sites,
information on insecticide use was obtained from appropriate growers or State extension personnel. Colonies of T. absoluta
were established from at least 200 larvae obtained from heavily infested leaves from each sampling site. The individual
populations were reared on tomato plants of variety Santa Clara, without insecticide exposure, enclosed in cages and
maintained in a greenhouse.
Fig. 1 Sites of origin of the populations of tomato leafminer (T.
absoluta) in the Brazilian states of Minas Gerais, Rio de Janeiro and
So Paulo. The numbers correspond to locations indicated in
Table 1.
[Normal View ]
Table 1 Origin and year of collection of populations of the tomato leafminer (T. absoluta).
Code number County State Place Month and year collected
1 Araguari Minas Gerais Field August 1998
2 Guiricema Minas Gerais Field September 1998
3 Lavras Minas Gerais Field August 1998
4 Uberlndia Minas Gerais Field April 1998
5 Viosa Minas Gerais Field October 1997
6 So Joo da Barra Rio de Janeiro Laboratory August 1997
7 Paulnia So Paulo Laboratory August 1998

The four technical grade insecticides used were abamectin (Novartis Biocincias, So Paulo, SP), cartap (Iharabrs, Sorocaba,
SP), methamidophos (Bayer, So Paulo, SP), and permethrin (Zeneca Agrcola, Holambra, SP). All of these insecticides are
currently recommended for controlling T. absoluta in Brazil.
2 de 7 1/11/2010 2:48 PM
Insecticide bioassays were carried out using insecticide-impregnated filter paper (9 cm diameter) placed in Petri dishes (9 cm
diameter 1.5 cm height). For each bioassay, six to seven different insecticide concentrations were applied; control treatments
(acetone for abamectin and permethrin, ethanol for methamidophos, and distilled water for cartap) were included. Three
replicates with 20 second instar larvae of T. absoluta were used at each insecticide concentration. For each replicate, we used
a filter paper impregnated with 1 mL of insecticide dissolved in the appropriate solvent (i.e. acetone, ethanol or distilled water).
Insecticide concentrations were calculated as g a.i./cm
2
of treated surface. Insects were counted as dead if they were unable
to walk. Concentrationmortality data were subjected to probit analysis (Proc Probit, Sas Institute 1997).
Correlation analysis (Proc CORR, Sas Institute 1997) was used to test the association between the insecticide use and its
resistance ratio across the sites studied. No association between insecticide use and resistance ratio could result from poor
estimation of insecticide use, resistance ratio, or both, or lack of a causal relationship between insecticide use and resistance
ratio. In addition, multiple regression analysis (Proc REG, Sas Institute 1997) was used to determine whether the use of the
other insecticides contributed to variation in resistance ratio of a particular insecticide. The dependent variable (i.e. the
resistance ratio of a particular insecticide) was used in the multiple regression models to test the four independent variables (i.e.
the use of each of the four insecticides under investigation).
Results
The results of
2
test (
2
and P values) used to measure how well the data of each concentrationresponse curve fit the
assumptions of the probit model are presented on Tables 25. As values (responses) predicted by the probit model did not differ
significantly from values actually observed in the bioassays (low
2
-values and P > 0.05), the probit model was suitable for the
concentrationresponse analyses.
Table 2 Comparative susceptibility of populations of tomato leafminer (T. absoluta) to permethrin.
Population n Slope SEM
LC
50
(95% CL)
g a.i./cm
2
LC
90
(95% CL)
g a.i./cm
2
Resistance
ratio *
2
Prob.
Uberlndia 421 0.51 0.04 47.8 (39.259.0) 319 (221532) 4.06 0.54
Viosa 362 0.63 0.07 71.5 (58.785.9) 389 (275669) 1.50 6.37 0.17
Paulnia 360 0.70 0.07 89.3 (76.6103) 305 (237447) 1.87 0.26 0.99
Lavras 422 0.56 0.09 142 (108219) 575 (3252194) 2.97 9.57 0.09
So Joo da Barra 360 0.58 0.08 158 (117290) 528 (2894475) 3.31 7.80 0.10
Araguari 420 0.80 0.07 187 (165212) 600 (469875) 3.90 5.63 0.34
Guiricema 365 0.86 0.07 316 (273361) 1088 (8451628) 6.61 2.59 0.63
* LC 50 resistant/LC50 susceptible.

Table 3 Comparative susceptibility of populations of tomato leafminer (T. absoluta) to abamectin.
LC
50
(95% CL)
g a.i./cm
2
LC
90
(95% CL)
g a.i./cm
2
Resistance
ratio *
2
Prob.
Uberlndia 360 0.80 0.10 0.97 (0.871.09) 2.40 (1.993.14) 4.34 0.36
Paulnia 361 0.76 0.12 5.00 (4.215.81) 18.2 (14.525.1) 5.16 3.30 0.51
Guiricema 360 0.67 0.06 5.87 (4.906.96) 21.9 (16.832.3) 6.04 2.80 0.59
So Joo da Barra 420 0.45 0.03 5.96 (4.517.52) 48.7 (35.176.7) 6.14 1.19 0.95
Viosa 421 0.49 0.04 6.76 (5.288.37) 47.1 (34.871.2) 6.97 1.93 0.86
Lavras 360 0.67 0.08 8.31 (7.109.87) 34.7 (25.156.6) 8.57 0.48 0.98
Araguari 360 0.63 0.09 9.09 (7.6810.92) 36.0 (25.365.0) 9.37 6.79 0.15
* LC
50
resistant/LC
50
susceptible.

Table 4 Comparative susceptibility of populations of tomato leafminer (T. absoluta) to methamidophos.
3 de 7 1/11/2010 2:48 PM
Insecticide bioassays were carried out using insecticide-impregnated filter paper (9 cm diameter) placed in Petri dishes (9 cm
diameter 1.5 cm height). For each bioassay, six to seven different insecticide concentrations were applied; control treatments
(acetone for abamectin and permethrin, ethanol for methamidophos, and distilled water for cartap) were included. Three
replicates with 20 second instar larvae of T. absoluta were used at each insecticide concentration. For each replicate, we used
a filter paper impregnated with 1 mL of insecticide dissolved in the appropriate solvent (i.e. acetone, ethanol or distilled water).
Insecticide concentrations were calculated as g a.i./cm
2
of treated surface. Insects were counted as dead if they were unable
to walk. Concentrationmortality data were subjected to probit analysis (Proc Probit, Sas Institute 1997).
Correlation analysis (Proc CORR, Sas Institute 1997) was used to test the association between the insecticide use and its
resistance ratio across the sites studied. No association between insecticide use and resistance ratio could result from poor
estimation of insecticide use, resistance ratio, or both, or lack of a causal relationship between insecticide use and resistance
ratio. In addition, multiple regression analysis (Proc REG, Sas Institute 1997) was used to determine whether the use of the
other insecticides contributed to variation in resistance ratio of a particular insecticide. The dependent variable (i.e. the
resistance ratio of a particular insecticide) was used in the multiple regression models to test the four independent variables (i.e.
the use of each of the four insecticides under investigation).
Results
The results of
2
test (
2
and P values) used to measure how well the data of each concentrationresponse curve fit the
assumptions of the probit model are presented on Tables 25. As values (responses) predicted by the probit model did not differ
significantly from values actually observed in the bioassays (low
2
-values and P > 0.05), the probit model was suitable for the
concentrationresponse analyses.
Table 2 Comparative susceptibility of populations of tomato leafminer (T. absoluta) to permethrin.
LC
50
(95% CL)
g a.i./cm
2
LC
90
(95% CL)
g a.i./cm
2
Resistance
ratio *
2
Prob.
Uberlndia 421 0.51 0.04 47.8 (39.259.0) 319 (221532) 4.06 0.54
Viosa 362 0.63 0.07 71.5 (58.785.9) 389 (275669) 1.50 6.37 0.17
Paulnia 360 0.70 0.07 89.3 (76.6103) 305 (237447) 1.87 0.26 0.99
Lavras 422 0.56 0.09 142 (108219) 575 (3252194) 2.97 9.57 0.09
So Joo da Barra 360 0.58 0.08 158 (117290) 528 (2894475) 3.31 7.80 0.10
Araguari 420 0.80 0.07 187 (165212) 600 (469875) 3.90 5.63 0.34
Guiricema 365 0.86 0.07 316 (273361) 1088 (8451628) 6.61 2.59 0.63

Table 3 Comparative susceptibility of populations of tomato leafminer (T. absoluta) to abamectin.
LC
50
(95% CL)
g a.i./cm
2
LC
90
(95% CL)
g a.i./cm
2
Resistance
ratio *
2
Prob.
Uberlndia 360 0.80 0.10 0.97 (0.871.09) 2.40 (1.993.14) 4.34 0.36
Paulnia 361 0.76 0.12 5.00 (4.215.81) 18.2 (14.525.1) 5.16 3.30 0.51
Guiricema 360 0.67 0.06 5.87 (4.906.96) 21.9 (16.832.3) 6.04 2.80 0.59
So Joo da Barra 420 0.45 0.03 5.96 (4.517.52) 48.7 (35.176.7) 6.14 1.19 0.95
Viosa 421 0.49 0.04 6.76 (5.288.37) 47.1 (34.871.2) 6.97 1.93 0.86
Lavras 360 0.67 0.08 8.31 (7.109.87) 34.7 (25.156.6) 8.57 0.48 0.98
Araguari 360 0.63 0.09 9.09 (7.6810.92) 36.0 (25.365.0) 9.37 6.79 0.15
* LC
50
resistant/LC
50
susceptible.

Table 4 Comparative susceptibility of populations of tomato leafminer (T. absoluta) to methamidophos.
3 de 7 1/11/2010 2:48 PM
LC
50
(95% CL)
g a.i./cm
2
LC
90
(95% CL)
g a.i./cm
2
Resistance
ratio *
2
Prob.
So Joo da Barra 362 0.67 0.06 59.9 (50.970.8) 243 (184357) 7.75 0.10
Viosa 401 0.62 0.07 62.3 (53.972.1) 192 (154256) 1.04 8.06 0.15
Lavras 420 0.67 0.05 79.3 (53.499.0) 239.6 (204305) 1.32 7.78 0.17
Uberlndia 440 0.80 0.11 155 (137175) 530 (420744) 2.59 8.10 0.23
Araguari 343 2.17 0.42 191 (179201) 279 (263303) 3.19 5.34 0.25
Paulnia 400 1.16 0.17 225 (202246) 473 (421556) 3.76 5.66 0.34
Guiricema 400 1.23 0.16 252 (228275) 535 (473637) 4.22 7.51 0.19

Table 5 Comparative susceptibility of populations of tomato leafminer (T. absoluta) to cartap.
LC
50
(95% CL)
g a.i./cm
2
LC
90
(95% CL)
g a.i./cm
2
Resistance
ratio *
2
Prob.
Paulnia 363 1.90 0.21 0.44 (0.410.47) 0.74 (0.670.84) 5.34 0.25
Guiricema 363 2.24 0.52 0.97 (0.921.03) 1.60 (1.392.19) 2.25 4.68 0.32
Lavras 420 1.90 0.20 1.87 (1.761.98) 3.15 (2.893.55) 4.22 5.06 0.41
Uberlndia 400 1.04 0.13 4.57 (4.135.04) 10.4 (9.0812.5) 10.4 4.30 0.51
Viosa 420 0.71 0.10 4.81 (4.225.47) 15.5 (12.520.7) 10.9 5.44 0.36
So Joo da Barra 420 0.77 0.04 7.16 (6.348.18) 20.9 (16.828.1) 16.2 6.66 0.25
Araguari 341 1.01 0.11 9.68 (8.4710.87) 25.3 (21.332.4) 21.9 6.86 0.14
* LC
50
resistant/LC
50
susceptible.

There was significant variation in the susceptibility among the insect populations studied for the four insecticides investigated
based on the criterion of failure of 95% CL at the LC50s to overlap. We were unable to identify a general standard susceptible
population. Therefore, the resistance ratios were taken by comparison with the population most susceptible to each insecticide.
The population from Uberlndia was the most susceptible one for permethrin and abamectin ( Tables 2 and 3), whereas So
Joo da Barra was the most susceptible to methamidophos ( Table 4) and Paulnia was most susceptible to cartap ( Table 5).
LC
50
s for permethrin were significantly greater for five of the populations of T. absoluta relative to the population from
Uberlndia ( Table 2). LC
50
for the population from Viosa resembled that of Uberlndia and Paulnia, differing from the
remaining ones. Permethrin resistance ratios ranged from 1.5- to 6.6-fold among the populations of the leafminer studied. LC
50
s
for abamectin were significantly greater for all insect populations when compared with the population from Uberlndia, with
resistance ratios ranging from 5.2- to 9.4-fold ( Table 3).
Resistance to methamidophos was observed in only four insect populations when compared with the population from So Joo
da Barra, the most susceptible one to this insecticide ( Table 4). These four populations were from Araguari, Guiricema,
Paulnia and Uberlndia, and their resistance ratios to methamidophos ranged from 2.6- to 4.2-fold. By contrast, the populations
from Lavras and Viosa were susceptible to this insecticide. Resistance to cartap was observed in all of the populations studied
in comparison with the most susceptible one ( Table 5), as was observed for abamectin, but with a different susceptibility
standard. The cartap resistance ratios ranged from 2.3- to 21.9-fold among the T. absoluta populations.
Slopes of the concentrationmortality curves for permethrin and abamectin were relatively similar among the different insect
populations. However, there was a greater variation in slopes for the insecticides methamidophos and cartap, with some
populations showing slopes twice as great as those of some others. These high slopes of concentrationmortality curves
probably reflect a higher homogeneity of response to these insecticides in these populations ( Finney 1971).
Insecticide use varied among sites and the total number of sprays per cultivation cycle per site ranged from seven to 22, the
number of applications in Uberlndia and Araguari, respectively. Overall number of sprays per cultivation cycle per site for
abamectin (mean = 4.3, range = 08) and cartap (mean = 4.7, range = 08) were greater than for methamidophos (mean = 3.0,
range = 010) and permethrin (mean = 1.4, range = 03). Variation in use of each insecticide was significantly correlated with
the variation in resistance ratio for the same insecticide ( Fig. 2), except for methamidophos (r= 0.36, P= 0.42). Multiple
regression analyses showed that the use of other insecticides did not contribute to the resistance ratio of any particular
insecticide. Only the use of the insecticide to which the resistance has been observed provided good regression models, with
4 de 7 1/11/2010 2:48 PM
the exception of methamidophos, for which we were unable to obtain any significant model (dferror = 5, F= 9.88, P= 0.02 and
r
2
= 0.66 for abamectin resistance; df
error
= 5, F= 17.08, P= 0.01 and r
2
= 0.77 for cartap resistance; df
error
= 5, F= 4.23, P=
0.10 and r
2
= 0.46 for methamidophos resistance; dferror = 5, F= 9.13, P= 0.03 and r
2
= 0.65 for permethrin resistance).
Fig. 2 Correlations between number of sprays per cultivation cycle and
resistance ratio of T. absoluta for the same insecticide across seven
sites.
[Normal View ]
Discussion
Among the insecticides investigated here, only resistance to methamidophos had been previously recorded in Chilean
populations of T. absoluta ( Salazar & Araya 1997). Resistance of T. absoluta to abamectin, cartap and permethrin is reported
here for the first time. The resistance levels observed in Brazilian populations of the tomato leafminer are relatively low
(< 10-fold), but they are possible causes of the control failures with these insecticides ( Souza et al. 1992 ).
There were significant differences in the resistance levels among different populations for each insecticide. Such variability in
resistance levels indicates the occurrence of differential selection pressures, genetic diversity in the resistance mechanisms
among the insect populations, or both ( Kerns & Gaylor 1992). The response to abamectin was similar among the resistant
insect populations, but there was greater variability of response among populations for the other insecticides. This may be
caused by the different level of use of these compounds in the different areas from where the populations were obtained. The
resistance levels were lower for permethrin and methamidophos, and greater for abamectin and cartap, which may be due to a
higher selection pressure provided by the more intensive use of abamectin and cartap ( Souza et al. 1992 ; Picano et al. 1995
), or the lack of good susceptible standard populations, especially for the insecticides methamidophos and permethrin,
underestimating the resistance to these two compounds.
The persistence of an insecticide on a plant leads to the continuous selection of resistant individuals, which may contribute to a
faster resistance evolution ( Roush 1989). The lower resistance levels to permethrin in comparison with cartap, compounds
used for about the same length of time in Brazil, might be due to the lower persistence of the former, leading to a lower selection
pressure for resistance on the leafminer populations ( Souza & Reis 1986; Guedes et al. 1994 ). In addition, permethrin is
mainly used during the tomato harvest, due to its short pre-harvest interval, unlike cartap, which may have favoured a stronger
selection pressure for cartap resistance ( Guedes et al. 1994 ). Despite the low permethrin resistance levels in Brazilian
populations of T. absoluta observed in our study, the more resistant populations of this insect were obtained from areas of high
frequency of insecticide application and where there were reports of chemical control failures, as in the counties of Araguari and
Guiricema ( Souza et al. 1992 ; Picano et al. 1995 ).
The populations of T. absoluta showed low levels of resistance to methamidophos, similar to those observed for permethrin.
This may be due to the relatively lower efficiency of this compound in controlling T. absoluta compared to abamectin and cartap,
for example, as reported by Souza et al. 1992 ) . This is probably one of the reasons for its lesser use in comparison with the
other compounds, as well as its high toxicity to mammals ( Ware 1994). The resistance levels to methamidophos reported here
for Brazilian populations of T. absoluta resemble those for Chilean populations ( Salazar & Araya 1997).
Abamectin resistance has never been reported in T. absoluta before, despite suspicions of the occurrence of this phenomenon
in different populations of this pest as a result of its widespread use, usually in mixture with mineral oil ( Guedes et al. 1995 ;
Castelo Branco et al. 1996 ; Castelo Branco et al. 1996 ). The use of mineral oil in mixture with abamectin apparently increases
the persistence of the insecticidal effects on the plant ,exerting a higher selection pressure on the pest population ( Castelo
Branco et al. 1996 ). The use of the insecticide alone does not seem to produce strong selection pressure for resistance
because of the low doses used in the field and its fast degradation in the environment without bioaccumulation ( Clark et al.
1994 ). The low levels of resistance to abamectin observed in our study suggest that the importance of mineral oil in the
evolution of abamectin resistance in T. absoluta is probably overestimated. The low variability in response to this insecticide
among different populations of the pest insect is probably a reflection of its widespread use throughout Brazil.
Cartap resistance has not been the object of much attention. The only studies on the subject are those reported by Liu et al.
(1982) and Cheng 1988) with the diamondback moth Plutella xylostella. Our results indicate resistance to cartap in Brazilian
5 de 7 1/11/2010 2:48 PM
the exception of methamidophos, for which we were unable to obtain any significant model (dferror = 5, F= 9.88, P= 0.02 and
r
2
= 0.66 for abamectin resistance; df
error
= 5, F= 17.08, P= 0.01 and r
2
= 0.77 for cartap resistance; df
error
= 5, F= 4.23, P=
0.10 and r
2
= 0.46 for methamidophos resistance; dferror = 5, F= 9.13, P= 0.03 and r
2
= 0.65 for permethrin resistance).
Fig. 2 Correlations between number of sprays per cultivation cycle and
resistance ratio of T. absoluta for the same insecticide across seven
sites.
[Normal View ]
Discussion
Among the insecticides investigated here, only resistance to methamidophos had been previously recorded in Chilean
populations of T. absoluta ( Salazar & Araya 1997). Resistance of T. absoluta to abamectin, cartap and permethrin is reported
here for the first time. The resistance levels observed in Brazilian populations of the tomato leafminer are relatively low
(< 10-fold), but they are possible causes of the control failures with these insecticides ( Souza et al. 1992 ).
There were significant differences in the resistance levels among different populations for each insecticide. Such variability in
resistance levels indicates the occurrence of differential selection pressures, genetic diversity in the resistance mechanisms
among the insect populations, or both ( Kerns & Gaylor 1992). The response to abamectin was similar among the resistant
insect populations, but there was greater variability of response among populations for the other insecticides. This may be
caused by the different level of use of these compounds in the different areas from where the populations were obtained. The
resistance levels were lower for permethrin and methamidophos, and greater for abamectin and cartap, which may be due to a
higher selection pressure provided by the more intensive use of abamectin and cartap ( Souza et al. 1992 ; Picano et al. 1995
), or the lack of good susceptible standard populations, especially for the insecticides methamidophos and permethrin,
underestimating the resistance to these two compounds.
The persistence of an insecticide on a plant leads to the continuous selection of resistant individuals, which may contribute to a
faster resistance evolution ( Roush 1989). The lower resistance levels to permethrin in comparison with cartap, compounds
used for about the same length of time in Brazil, might be due to the lower persistence of the former, leading to a lower selection
pressure for resistance on the leafminer populations ( Souza & Reis 1986; Guedes et al. 1994 ). In addition, permethrin is
mainly used during the tomato harvest, due to its short pre-harvest interval, unlike cartap, which may have favoured a stronger
selection pressure for cartap resistance ( Guedes et al. 1994 ). Despite the low permethrin resistance levels in Brazilian
populations of T. absoluta observed in our study, the more resistant populations of this insect were obtained from areas of high
frequency of insecticide application and where there were reports of chemical control failures, as in the counties of Araguari and
Guiricema ( Souza et al. 1992 ; Picano et al. 1995 ).
The populations of T. absoluta showed low levels of resistance to methamidophos, similar to those observed for permethrin.
This may be due to the relatively lower efficiency of this compound in controlling T. absoluta compared to abamectin and cartap,
for example, as reported by Souza et al. 1992 ) . This is probably one of the reasons for its lesser use in comparison with the
other compounds, as well as its high toxicity to mammals ( Ware 1994). The resistance levels to methamidophos reported here
for Brazilian populations of T. absoluta resemble those for Chilean populations ( Salazar & Araya 1997).
Abamectin resistance has never been reported in T. absoluta before, despite suspicions of the occurrence of this phenomenon
in different populations of this pest as a result of its widespread use, usually in mixture with mineral oil ( Guedes et al. 1995 ;
Castelo Branco et al. 1996 ; Castelo Branco et al. 1996 ). The use of mineral oil in mixture with abamectin apparently increases
the persistence of the insecticidal effects on the plant ,exerting a higher selection pressure on the pest population ( Castelo
Branco et al. 1996 ). The use of the insecticide alone does not seem to produce strong selection pressure for resistance
because of the low doses used in the field and its fast degradation in the environment without bioaccumulation ( Clark et al.
1994 ). The low levels of resistance to abamectin observed in our study suggest that the importance of mineral oil in the
evolution of abamectin resistance in T. absoluta is probably overestimated. The low variability in response to this insecticide
among different populations of the pest insect is probably a reflection of its widespread use throughout Brazil.
Cartap resistance has not been the object of much attention. The only studies on the subject are those reported by Liu et al.
(1982) and Cheng 1988) with the diamondback moth Plutella xylostella. Our results indicate resistance to cartap in Brazilian
5 de 7 1/11/2010 2:48 PM
populations of T. absoluta. The observed cartap resistance levels in T. absoluta reached moderate levels (> 10-fold and <
100-fold) in four of the populations studied (i.e. Uberlndia, Viosa, So Joo da Barra and Araguari), with the highest level
observed in the population from Araguari (22-fold). Such resistance levels are a probable explanation for the control failures with
cartap against T. absoluta reported by Frana 1993) . However, Castelo Branco et al. 1996 ) suggested that the low efficiency
of cartap against T. absoluta was probably due to a poor application technology. However, our results do not provide support
for this contention. Nevertheless, poor application may be a contributing factor to the low efficiency of cartap in controlling
resistant populations of T. absoluta in some areas. The low to moderate levels of resistance to cartap and the long period of
use of this insecticide in Brazil suggest that the resistance to cartap has a slow rate of evolution in populations of T. absoluta.
The significant positive correlations between frequency of application and resistance ratio suggest that the variation in
susceptibility of T. absoluta populations was caused by local variation in insecticide use. The only exception to this observation
was the insecticide methamidophos, whose resistance ratios were not significantly correlated with its use, probably due to a
poor estimation of this insecticide use in So Joo da Barra, where it is commonly used (10 sprays per cultivation cycle). The
population collected in this site was the most susceptible to this insecticide. After eliminating these data from the correlation
analysis (for methamidophos), there was a significant and positive correlation between frequency of methamidophos application
and its resistance ratio across the remaining sites (r= 0.88, P= 0.02). The results obtained with the multiple regression did not
suggest any pattern of cross-resistance among the insecticides studied.
In summary, our data indicate that populations of T. absoluta from Brazil show resistance to abamectin, cartap, methamidophos
and permethrin, confirming the suspicions raised by other investigators (e.g. Souza et al. 1992 ; Frana 1993; Guedes et al.
1994 ). The resistance levels ranged from low (< 10-fold) to moderate (> 10-fold and < 100-fold) levels for the main insecticides
used for a long time, which suggests a low rate of evolution of this phenomenon in T. absoluta, but also explains some of the
control failures observed with the use of these compounds against this insect. Alternatively, the phenomenon might be
widespread among Brazilian populations of T. absoluta, making finding suitable standard susceptible populations difficult and
leading to an underestimation of the insecticide resistance levels in this insect. This is one of the few reported cases of
resistance to cartap in insect pests reaching moderate levels in some populations. This finding opposes claims of poor
application technology as the sole explanation of control failures with the use of this insecticide against T. absoluta. The
correlations between insecticide use and resistance ratio across seven sites suggest that local variation in insecticide use was
an important cause of variation in susceptibility. There was no evidence of cross-selection among the insecticides studied.
Acknowledgements
We would like to express our gratitude to Nilton C. Picinato (DuPont) and Norma E. Pereira (Universidade Estadual do Norte
Fluminense) for providing the T. absoluta populations from Paulnia (SP) and So Joo da Barra (RJ), respectively; and to the
agrochemical companies Iharabrs, Novartis Biocincias, Zeneca Agrcola and Bayer for providing the technical grade
insecticides used in this study. The information about insecticide use provided by growers and State extension personnel was
greatly appreciated. We also would like to thank A. D. Watt and three anonymous referees for their valuable suggestions.
Financial support was provided by FAPEMIG, CNPq and CAPES and is acknowledged here.
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Sannino, L. & Nicodemo, F. 1979 Su una insolita infestazione di Phthorimea operculella Zell. (Lepidoptera: Gelechiidae) al
tabacco nel salernitano . Annales Nel Istituto Superior Nel Tabacco Scafati, 5, 125 135.
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Guedes, R.N.C., Picano, M.C., Guedes, N.M.P., Madeira, N.R. 1995 Sinergismo do leo mineral sobre a toxicidade de
inseticidas para Scrobipalpuloides absoluta (Lepidoptera: Gelechiidae) . Pesquisa Agropecuria Brasileira, 30, 313 318.
Guedes, R.N.C., Picano, M.C., Matioli, A.L., Rocha, D.M. 1994 Efeito de inseticidas e sistemas de conduo do tomateiro no
controle de Scrobipalpuloides absoluta (Meyrick) (Lepidoptera: Gelechiidae) . Anais Da Sociedade Entomolgica Do Brasil,
23, 321 325.
Juvick, J.A., Berlinger, M.J., Bem-David, T., Rudich, J. 1982 Resistance among accessions of genera Lycopersicon and
Solanum to four of the main insect pests of tomato in Israel . Phytoparasitica, 10, 145 156.
Kay, I.R. & Collins, P.J. 1987 The problem of resistance to insecticides in tropical insect pests. Insect Science and its
Applications, 8, 715 721.
Kerns, D.L. & Gaylor, M.J. 1992 Insecticide resistance in field populations of the cotton aphid (Homoptera: aphididae). Journal
of Economic Entomology, 85, 1 8.
Liu, M.Y., Tzeng, Y.J., Sun, C.N. 1982 Insecticide resistance in the diamondback moth. Journal of Economic Entomology, 75,
153 155.
Lockwood, J.A., Sparks, T.C., Story, R.N. 1984 Evolution of insect resistance to insecticides: a reevaluation of the roles of
physiology and behavior. Bulletin of the Entomological Society of America, 30, 41 57.
Miranda, M.M.M., Picano, M., Zanuncio, J.C., Guedes, R.N.C. 1998 Ecological life table of Tuta absoluta (Meyrick)
(Lepidoptera: Gelechiidae) . Biocontrol Science and Technology, 8, 597 606.
Moore, J.E. 1983 Control of tomato leafminer (Scrobipalpula absoluta) in Bolvia . Tropical Pest Management, 29, 231 238.
Muszinski, T., Lavendowski, I.M., Maschio, L.M.A. 1982 Constatao de Scrobipalpula absoluta (Meyrick) (Lepidoptera:
Gelechiidae), como praga do tomateiro (Lycopersicon esculentum Mill.), no litoral do Paran. Anais Da Sociedade
Picano, M.C., Guedes, R.N.C., Leite, G.L.D., Fontes, P.C.R., Silva, E.A. 1995 Incidncia de Scrobipalpuloides absoluta
(Meyrick) (Lepidoptera: Gelechiidae) em tomateiro sob diferentes sistemas de tutoramento e controle qumico de pragas .
Horticultura Brasileira, 13, 180 183.
Picano, M., Leite, G.L.D., Guedes, R.N.C., Silva, E.A. 1998 Yield loss in trellised tomato affected by insecticidal sprays and
plant spacing. Crop Protection, 17, 447 452.
Picano, M.C., Silva, E.A., Lbo, A.P., Leite, G.L.D. 1996 Adio de leo mineral a inseticidas no controle de Tuta absoluta
(Meyrick) (Lepidoptera: Gelechiidae) e Helicoverpa zea (Bod.) (Lepidoptera: Noctuidae) em tomateiro . Anais Da Sociedade
Pouey, G.F., Chirinos, D.T., Rivero, G. 1994 Notas sobre Keiferia lycopersicella (Walsingham), (Lepidoptera: Gelechiidae), en
Venezuela . Boletin de Entomologia Venezoelana, 9, 203 206.
Povolny, D. 1975 On three neotropical species of Gnorimoschemini (Lepidoptera: Gelechiidae) mining Solanaceae . Acta
Universitatis Agriculturae, 23, 379 393.
Roush, R.T. 1989 Designing resistance management programs: how can you choose? Pesticide Science, 26, 423 441.
Salazar, E.S. & Araya, J.E. 1997 Deteccin de resistencia a insecticidas em la polilla del tomate. Simiente, 67, 8 22.
Sannino, L. & Nicodemo, F. 1979 Su una insolita infestazione di Phthorimea operculella Zell. (Lepidoptera: Gelechiidae) al
tabacco nel salernitano . Annales Nel Istituto Superior Nel Tabacco Scafati, 5, 125 135.
SAS Institute 1997 SAS User's Guide: Statistics, Version 6.12. SAS Institute, Cary, NC, USA.
Souza, J.C. & Reis, P.R. 1986 Controle da traa-do-tomateiro em Minas Gerais. Pesquisa Agropecuria Brasileira, 21, 343
354.
Souza, J.C., Reis, P.R., Salgado, L.O. 1992. Traa Do Tomateiro: Histrico, Reconhecimento, Biologia, Prejuzos E Controle.
EPAMIG, Belo Horizonte, MG, Brazil.
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Chapman & Hall. New York.
Ware, G.W. 1994. The Pesticide Book. 4th edn. Thomson, Fresno, CA.
Accepted 14 April 2000
7 de 7 1/11/2010 2:48 PM
Abamectin resistance and synergism in Brazilian populations of Tuta absoluta (Meyrick)
(Lepidoptera: Gelechiidae)
(Keywords: tomato leafminer, Brazil, insecticide resistance, diethyl maleate, piperonyl butoxide, triphenylphosphate)
H. A. A. SIQUEIRA, R. N. C. GUEDES*, D. B. FRAGOSO and L. C. MAGALHA
ES
Departamento de Biologia Animal, Universidade Federal de Vic osa, Vic osa, MG 36571-000, Brazil
Abstract. Failures in the control of the tomato leafminer Tuta absoluta
(Meyrick) by means of abamectin in Brazil, and a recent report of
abamectin resistance in Brazilian populations of this pest species, led to
the investigation of the possible involvement of detoxification enzymes
using insecticide synergists. Resistance to abamectin was observed in
all populations when compared with the standard susceptible popula-
tion, with resistance ratios ranging from 5.2- to 9.4-fold. Piperonyl
butoxide was the most efficient synergist with abamectin synergism
ratios ranging from 3.0- to 5.3-fold and providing significant resistance
suppression, but complete suppression of abamectin resistance was
only obtained in one population of T. absoluta. Triphenylphosphate was
an abamectin synergist which was not as efficient as piperonyl butoxide,
but it provided complete suppression of abamectin resistance in four of
the six resistant populations studied, suggesting a major involvement of
esterases as an abamectin resistance mechanismin these populations.
The importance of cytochrome P450, inhibited by piperonyl butoxide,
seems secondary to esterases. Diethyl maleate also synergized
abamectin in nearly all populations, but provided only partial suppres-
sion of abamectin resistance in the leafminer populations studied.
Therefore, glutathione-S-transferases seem to be of minor importance
as an abamectin resistance mechanism in Brazilian populations of T.
absoluta.
1. Introduction
Brazil is the eighth greatest producer of tomatoes in the
world, with 60,000 ha of the crop grown annually, and an
average productivity of 44 t.ha
-1
(FAO, 1994). Tomato produc-
tion for fresh consumption is the most important source of
income for small producers in several regions of the country.
However, this vegetable crop is often severely damaged by
pests in the tropics, and pesticides are intensively used to
control insects and pathogens in Brazil (Gravena, 1991;
Makishima, 1991; Picanc o et al., 1998). This situation substan-
tially increases the crop production cost and leads to human
health and environmental problems in addition to selecting for
resistance to pesticides. The tomato leafminer, Tuta (=Scrobi-
palpuloides) absoluta (Meyrick) (Lepidoptera: Gelechiidae), is
considered to be one of the most important pests of tomato
production in Brazil (Souza and Reis, 1986; Souza et al., 1992).
It is an oligophagous insect, distributed throughout the Neo-
tropical region, attacking solanaceous plants. It is of major
importance in Venezuela, Colombia, Chile, Equator, Bolivia,
Argentina, Peru (Povolny, 1975), and Uruguay (Carballo et al.,
1981). Its occurrence has been reported even in Japan (Nakano
and Paulo, 1983).
It has been a serious problemto tomato production in Brazil
since its introduction and rapid spread in the 1980s. The
damage caused by T. absoluta resembles that caused by other
Gelechiidae such as Keiferia lycopersicella (Wals.) in Central
and North America, and Phthorimaea operculella (Zell.) in the
Americas, Europe, Africa and Asia (Sannino and Nicodemo,
1979; Juvick et al., 1982; Abbas et al., 1993; Ebora et al., 1994;
Pouey et al., 1994). At high densities it can cause severe yield
losses (Souza et al., 1992; Picanc o et al., 1998).
Chemical control has been the primary control strategy for
this insect since the early 1980s (Souza and Reis, 1986), with
some farmers still making up to 36 insecticide applications
during the crop cycle (Picanc o et al., 1995). Producers have
observed insecticide failures in controlling this pest and
researchers have shown reduced insecticide activity in the field
(Souza et al., 1992; Franc a, 1993; Guedes et al., 1994),
suggesting the development of resistant populations to com-
pounds used against T. absoluta (Gonc alves et al., 1994).
Abamectin was first used around 1990 due to control failures
with other insecticides (Castelo Branco, 1990). Studies on
abamectin resistance in T. absoluta populations have not yet
been carried out, despite reports of resistance in this species to
other insecticides in Chile (Salazar and Araya, 1997). However,
studies on abamectin resistance have been carried out in
resistant strains of house flies (Musca domestica L.) (Roush and
Wright, 1986), German cockroaches (Blattella germanica (L.))
(Cochran, 1990, 1994), Colorado potato beetle (Leptinotarsa
decemlineata (Say)) (Argentine and Clark, 1990; Argentine et
al., 1992), and spider mites (Tetranichus urticae Koch and T.
mcdanuli McGregor) (Campos et al., 1995; Beers et al., 1998).
No cross-resistance (i.e., resistance to two or more insecticides
due to the same mechamism) to abamectin was detected in
these studies (Clark et al., 1995). Among lepidopteran pests,
only the diamondback moth, Plutella xylostella, has been
subjected to more intense investigation of abamectin resistance
(Abro et al., 1988; Iqbal and Wright, 1997). Resistance
mechanisms to abamectin have been studied in a few species,
where several major resistance mechanisms and some minor
factors have been implicated (Clark et al., 1995).
The aim of this study was to survey abamectin
resistance in Brazilian populations of T. absoluta and to
assess the possible involvement of detoxification enzymes in
International Journal of Pest Management
ISSN 0967-0874 print/ISSN 1366-5863 online 2001 Taylor & Francis Ltd
http://www.tandf.co.uk/journals
DOI: 10.1080/09670870110044634
*To whom correspondence should be addressed. Fax: +55 (31)3899-2537; e-mail: guedes@mail.ufv.br
INTERNATIONAL JOURNAL OF PEST MANAGEMENT, 2001, 47(4) 247251
abamectin resistance using insecticide synergists in `in vivo
assays.
2. Material and methods
Five populations of T. absoluta from the State of Minas
Gerais, one from Sao Paulo State and another from Rio de
Janeiro State, were used in this study (Table 1). A field
population collected from Uberlandia County, State of Minas
Gerais, was used as an abamectin susceptible standard
population because it was obtained from a site where this
insecticide had not been used and it was reported as a
population susceptible to this insecticide in a previous investiga-
tion (Siqueira, 1999). Colonies of T. absoluta were established
from at least 200 larvae obtained from heavily infested leaves
fromeach sampling site. The individual populations were reared
on tomato plants of variety Santa Clara, without insecticide
exposure, enclosed in cages and maintained under greenhouse
conditions at a temperature and daylength varying from 22 to
288C and from10 to 14 h respectively, during the study period.
The colonies were maintained for two generations in the
laboratory before starting the bioassays.
In vivo bioassays, using insecticide-impregnated filter paper
(9 cm diameter) placed in Petri dishes (9 cm diameter61.5 cm
height), were carried out. Technical grade abamectin (Novartis
Biociencias, Sao Paulo, SP) was used in the investigation. The
three synergists used were diethyl maleate, piperonyl butoxide
and triphenylphosphate (Aldrich Chemical Co., Milwauke, WI,
USA). Diethyl maleate is an inhibitor of glutathione-S-trans-
ferases, while piperonyl butoxide and triphenylphosphate are
inhibitors of cytochrome P450-dependent monooxygenases and
esterase respectively (Raffa and Priester, 1985; Bernard and
Philogene, 1993).
Three replicates with 20 second instar larvae of T. absoluta
were used at each insecticide concentration. For each concen-
trationmortality regression line, six to seven different insecticide
concentrations were applied; a control treatment of acetone
alone was included and used for correcting mortality in the other
treatments. On each replicate of each concentration we used a
filter paper impregnated with 1 ml of insecticide dissolved in
acetone. Insecticide concentrations were calculated as mg a.i./
cm
2
of treated surface. Insects were counted as dead if they
were unable to walk. The synergist bioassays followed the same
methodology described above using a insecticide:synergist
proportion of 1:10. The synergists were applied on filter paper
with dried insecticide residues at the specified concentration,
where 1 ml of the synergist solution proportional to each
insecticide concentration was applied. No mortality was ob-
served in preliminary assays with the synergists used alone at
the highest concentrations used in this investigation when in
mixture with abamectin. Concentrationmortality data were
subjected to probit analysis (PROC PROBIT, SAS Institute,
1987).
3. Results
The concentrationmortality regression lines were obtained
for all combinations of insecticide or insecticide +synergist and
every insect population under study (tables 25). There was
significant variation on the susceptibility among the insect
populations studied in response to abamectin. The resistance
ratios were calculated by dividing the LC
50
of the most
susceptible population (Uberlandia) to this insecticide by the
LC
50
of the other insect populations (table 2). The LC
50
of the
susceptible population was significantly smaller than those of
the other populations based on the criterion of failure of 95% CL
to overlap (table 2). The resistance ratio among the resistance
populations ranged from 5.2- to 9.4-fold. Slopes of the
concentrationmortality curves for abamectin were relatively
similar among the different insect populations, showing similar
homogeneity of response for all populations.
Piperonyl butoxide synergized abamectin for all populations,
based on the criterion cited above, except for the Uberlandia
population which was slightly antagonized. Resistance suppres-
sion was complete for the Paulnia population, and nearly
complete for the remaining populations (table 3). Diethyl maleate
H. A. A. Siqueira et al. 248
Table 1. Origin and year of collection of leafminer populations ( T.
absoluta)
County State Local
Month/
Collection year
Araguari Minas Gerais Field 08/1998
Guiricema Minas Gerais Field 09/1998
Lavras Minas Gerais Field 08/1998
Uberlandia Minas Gerais Field 04/1998
Vic osa Minas Gerais Field 10/1997
Sao Joao da Barra Rio de Janeiro Greenhouse 08/1997
Paulnia Sao Paulo Greenhouse 08/1998
Table 2. Relative susceptibility of populations of tomato leafminer (Tuta absoluta) to abamectin
Population n Slope+SEM
LC
50
(95% CL)
(mg a.i./cm
2
)
LC
90
(95% CL)
(mg a.i./cm
2
)
Resistance
ratio
a
w
2
P
Uberlandia 360 0.80+0.10 0.97 (0.87 1.09) 2.40 (1.99 3.14) 4.34 0.36 NS
Paulnia 361 0.76+0.12 5.00 (4.21 5.81) 18.22 (14.51 25.09) 5.16 3.30 0.51 NS
Guiricema 360 0.67+0.06 5.87 (4.90 6.96) 21.92 (16.78 32.34) 6.04 2.80 0.59 NS
Sao Joao da Barra 420 0.45+0.03 5.96 (4.51 7.52) 48.74 (35.13 76.67) 6.14 1.19 0.95 NS
Vic osa 421 0.49+0.04 6.76 (5.28 8.37) 47.15 (34.80 71.20) 6.97 1.93 0.86 NS
Lavras 360 0.67+0.08 8.31 (7.10 9.87) 34.66 (25.14 56.57) 8.57 0.48 0.98 NS
Araguari 360 0.63+0.09 9.09 (7.68 10.92) 35.97 (25.33 65.02) 9.37 6.79 0.15 NS
a
LC
50
resistant/LC
50
susceptible.
NS, not significant.
synergized significantly the populations from Araguari, Lavras,
Sao Joao da Barra and Vic osa; however, it had no effect on the
populations from Paulnia and Guiricema and even antagonized
abamectin in the Uberlandia population at its LC50. Resistance
suppression was smaller than with piperonyl butoxide for the
Araguari, Lavras, Sao Joao da Barra and Vic osa populations
(table 4). There was no resistance suppression for the Paulnia
and Guiricema populations. Triphenylphosphate synergized all
populations, except the one from Uberlandia, in which a slight
antagonismto abamectin was observed (table 5). This synergist
completely suppressed abamectin resistance in four populations
and nearly did so in the remaining two among the six tested
populations.
Synergising insecticides usually results in steeper concen-
trationresponse curves if the population is heterogeneous for
resistance. This was generally the case in our study, which is
expected for field-collected populations. However there were a
fewexceptions, mainly for triphenylphosphate, where no change
in slope was observed probably reflecting a higher homogeneity
of response to this synergist in the populations studied.
4. Discussion
T. absoluta resistance to abamectin was reported by
Siqueira (1999). The occurrence of this phenomenon in different
populations of this pest was suspected as a result of its
widespread use, usually in a mixture with mineral oil (Guedes et
al., 1995; Castelo Branco and Franc a, 1996; Castelo Branco et
al., 1996). The lowlevels of resistance to abamectin observed in
our study suggest that the importance of mineral oil in the
Abamectin resistance and synergism in T. absoluta 249
Table 3. Relative susceptibility of populations of tomato leafminer (Tuta absoluta) to abamectin+ piperonyl butoxide (1:10)
LC
50
(95% CL)
(mg a.i./cm
2
)
Resistance
ratio
a
Synergism
ratio
b
w
2
P
Uberlandia 360 1.25+0.11 1.30 (1.19 1.42) 0.75 2.51 0.64 NS
Paulnia 340 1.04+0.21 1.24 (1.04 1.40) 0.95 4.03 5.47 0.32 NS
Guiricema 360 1.75+0.19 1.94 (1.65 2.24) 1.49 3.03 8.74 0.07 NS
Sao Joao da Barra 360 1.22+0.23 1.68 (1.53 1.84) 1.29 3.55 4.68 0.24 NS
Vic osa 360 0.89+0.08 2.04 (1.77 2.31) 1.57 3.31 5.53 0.24 NS
Lavras 340 1.40+0.20 1.58 (1.44 1.72) 1.22 5.26 3.18 0.53 NS
Araguari 361 0.61+ 0.07 2.70 (2.08 3.30) 2.08 3.37 3.21 0.52 NS
a
Resistance Ratio (= LC
50
synergized resistant /LC
50
synergized susceptible).
b
Synergism Ratio (= LC
50
unsynergized abamectin/ LC
50
synergized abamectin).
Table 4. Relative susceptibility of populations of tomato leafminer (Tuta absoluta) to abamectin + diethyl maleate (1:10)
LC
50
(95% CL)
(mg a.i./cm
2
)
Resistance
ratio
a
Synergism
ratio
b
w
2
P
Uberlandia 366 1.47+0.20 1.84 (1.68 1.99) 0.53 4.49 0.34 NS
Paulnia 367 0.83+0.04 4.03 (3.45 4.63) 2.19 1.24 5.12 0.28 NS
Guiricema 400 0.55+0.08 4.84 (3.97 6.07) 2.63 1.21 8.89 0.11 NS
Vic osa 360 0.89+0.10 2.52 (2.18 2.86) 1.37 2.68 4.56 0.34 NS
Lavras 360 1.34+0.17 2.19 (2.01 2.39) 1.19 3.79 5.22 0.27 NS
Araguari 363 0.89+0.08 6.30 (5.43 7.13) 3.42 1.44 3.80 0.43 NS
a
50
50
b
50
50
Table 5. Relative susceptibility of populations of tomato leafminer (Tuta absoluta) to abamectin + triphenylphosphate (1:10)
LC
50
(95% CL)
(mg a.i./cm
2
)
Resistance
ratio
a
Synergism
ratio
b
w
2
P
Uberlandia 343 0.89+0.30 1.95 (1.67 2.25) 0.50 6.70 0.15 NS
Paulnia 361 0.57+0.08 2.18 (1.65 2.70) 1.12 2.29 7.05 0.76 NS
Guiricema 360 0.71+0.08 4.06 (3.37 4.77) 2.08 1.45 1.30 0.86 NS
Vic osa 360 0.64+0.08 2.17 (1.71 2.61) 1.11 3.12 6.16 0.19 NS
Lavras 366 1.29+0.19 2.21 (2.02 2.42) 1.13 3.76 5.09 0.28 NS
Araguari 362 0.67+0.05 5.06 (4.12 6.03) 2.60 1.80 2.82 0.59 NS
a
50
50
b
50
50
evolution of abamectin resistance in T. absoluta is probably
overestimated, even though it does provide support for the
contention that abamectin resistance is one of the causes of
control failures of this insecticide observed in field. The intensive
use of abamectin in T. absoluta control programmes seems not
to have been enough to select highly resistant strains. This is
probably due to the lowdose of abamectin used in the field and
its fast degradation in the environment without bioaccumulation
which do not seem to provide strong selection pressure for
resistance (Clark et al., 1995).
Besides using enzymes to maintain their normal home-
ostasis, insects may also use them for protection against
xenobiotics. Among the known insecticide resistance mechan-
isms, it is not surprising that enhanced activity of detoxification
enzymes is one of the most common mechanisms of resistance
to insecticides (Scott, 1990a). Insect detoxification enzymes are
important resistance mechanisms and synergists are helpful in
providing preliminary evidence of their involvement as resis-
tance mechanisms (Brindley and Selim, 1984; Scott, 1990b;
Bernard and Philogene, 1993; Ishaaya, 1993). In this study,
abamectin toxicity was enhanced by the synergists diethyl
maleate, piperonyl butoxide and triphenyl phosphate which
respectively inhibit the detoxification enzymes glutathione-S-
transferases, cytochrome P450-dependent monooxygenases,
and esterases (Raffa and Priester, 1985; Bernard and Philo-
gene, 1993), providing some interesting information regarding
abamectin resistance mechanisms in this insect-species.
Piperonyl butoxide nearly completely suppressed the resis-
tance to abamectin in all populations, except for the Paulnia
population of T. absoluta where the suppression was complete.
The synergism to abamectin by piperonyl butoxide suggests an
enhanced cytochrome P450-dependent monooxygenase activity
in this insect species as an abamectin resistance mechanism.
This effect by piperonyl butoxide has been observed in other
arthropod species. The involvement of monooxygenases in
abamectin resistance was observed in Colorado potato beetle,
house flies, two-spotted spider mites, Spodoptera littoralis and
Helicoverpa armigera (Scott, 1989; Christie and Wright, 1990;
Argentine et al., 1992; Campos et al., 1996).
Gluthatione-S-transferases are also probably involved in
abamectin resistance in T. absoluta. Our results indicate partial
suppression of abamectin resistance by diethyl maleate which
inhibit gluthatione-S-transferases. However this seems to be a
mechanism of minor importance. Argentine et al. (1992) found
that gluthatione-S-transferases were not important as a resis-
tance mechanism in L. decemlineata and carboxylesterases
possibly play an important role in abamectin resistance in this
species. Despite the occurrence of partial suppression by diethyl
maleate in the populations from Paulnia and Guiricema, the
synergistic effect was not significant. Triphenylphosphate and
piperonyl butoxide are stronger synergists to abamectin than
diethyl maleate suggesting enhanced detoxification by es-
terases and cytochrome P450-dependent monooxygenases in
populations of T. absoluta resistant to abamectin.
Triphenylphosphate completely suppressed abamectin re-
sistance in four T. absoluta populations (from Lavras, Paulnia,
Sao Joao da Barra, and Vic osa) and provided partial suppres-
sion in two populations (from Araguari and Guiricema). These
results suggest a major involvement of esterases as an
abamectin resistance mechanism in Brazilian populations of T.
absoluta, with cytochrome P450-dependent monooxygenases
probably playing a secondary role in addition to gluthatione-S-
transferases as a mechanism of minor importance. The
coexistence of different resistance mechanisms in the same
insect populations suggests an oligo or polygenic basis for
abamectin resistance in these populations.
The use of synergists in insecticide resistance management
programmes has been frequently suggested (e.g. Oppenoorth,
1985; Guedes, 1991; Denholmand Rowland, 1992; Bernard and
Philogene, 1993). However they may not be useful for managing
abamectin resistance in Brazilian populations of T. absoluta,
because of the likely occurrence of multiple resistance mechan-
isms. Nonetheless, synergists can be important tools for
managing T. absoluta populations lacking insecticide cross-
resistance. Resistance to abamectin is polyfactorial in both L.
decemlineata and M. domestica, which has resulted in several
major resistance mechanisms (e.g. excretion, oxidative meta-
bolism, penetration) and minor factors (e.g. altered target site,
conjugation, esteratic hydrolysis/sequestration) associated with
the phenomenon (Clark et al., 1995). The resistance to
abamectin in T. absoluta populations may be oligo- or even
polyfactorial, but further investigations are necessary to better
assess this possibility.
Acknowledgements
We would like to express our gratitude Nilton C. Picinato
(DuPont) and Norma E. Pereira (Universidade Estadual do
Norte Fluminense) for providing the T. absoluta population from
Paulnia (SP) and Sao Joao da Barra (RJ) respectively; and
Novartis Biociencias for providing the technical grade abamectin
used in this study. Financial support was provided by FAPEMIG,
CNPq and CAPES and is acknowledged here.
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Abamectin resistance and synergism in T. absoluta 251
January - February 2005 Neotropical Entomology 34(1) 113
CROP PROTECTION
Insecticide Resistance in Argentine Populations of Tuta absoluta (Meyrick)
(Lepidoptera: Gelechiidae)
MARCELA M.M. LIETTI
1
, EDUARDO BOTTO
2
AND RAL A. ALZOGARAY
3
1
Ctedra de Zoologa Agrcola, Facultad de Ciencias Agrarias, Universidad Nacional de Rosario (U.N.R.)
C.C. 14, (S2125ZAA) Zavalla, Santa Fe, Argentina. E-mail: mlietti@fcagr.unr.edu.ar
2
Insectario de Investigaciones para Lucha Biolgica, IMYZA-CNIA, INTA. C.C. 25, (1712)
Castelar, Buenos Aires, Argentina
3
Centro de Investigaciones de Plagas e Insecticidas (CIPEIN-CITEFA/CONICET)
J. B. de La Salle 4397, (B1603ALO) Villa Martelli, Buenos Aires, Argentina
E-mail: ralzogaray@hotmail.com
Neotropical Entomology 34(1):113-119 (2005)
RESUMO - A traa-do-tomateiro, Tuta absoluta (Meyrick), uma das pragas chaves no tomateiro na
Argentina. O controle qumico tem sido o principal mtodo de controle empregado a partir da sua
disperso nos anos 70. Contudo, tem-se observado uma reduo na eficcia de alguns dos inseticidas
recomendados a partir da dcada de 80. O objetivo deste trabalho foi estudar a toxicidade de trs
inseticidas amplamente usados no controle qumico de T. absoluta (abamectina, deltametrina e
metamidofs) em larvas de uma populao susceptvel de laboratrio (CASTELAR) e duas populaes
colectadas em casa de vegetao (ROSARIO e BELLA VISTA). Inseticidas foram diludos em acetona
e aplicados topicamente na regio dorsal mediana do abdome de larvas no segundo dia do quarto
estgio larval. Para cada inseticida estimou-se o LD
50
e calculou-se o Nvel de Resistncia (NR = LD
50
de
cada populao de casa de vegetao/LD
50
populao de laboratrio). As populaes de ROSARIO e
BELLA VISTA mostraram os seguintes NRs: > 68.38 para deltametrina; 2.48 e 3.49 para abamectina,
respetivamente; e 0.79 e 0.86 para metamidofs, respetivamente. A resistncia a deltametrina observada
em ROSARIO pode ser resultante da alta presso seletiva exercida pelos piretrides nessa localidade.
A resistncia incipiente a abamectina detectada em BELLA VISTA pode ter sido causado pelo uso
freqente do inseticida nessa localidade ou pode estar associada variao natural.
PALAVRAS-CHAVE: Traa-do-tomateiro, deltametrina, abamectina, metamidofs, resistncia a inseticidas
ABSTRACT - The tomato leafminer, Tuta absoluta (Meyrick), is one of the key pests of tomato in
Argentina. Since its dispersal in the 1970s, chemical control has been the main method of controlling it.
However, reduced efficacy of some of the recommended insecticides has been observed since the
1980s. The aim of this work was to study the toxicity of three insecticides widely used in chemical
control of T. absoluta (abamectin, deltamethrin and methamidophos) on larvae from a laboratory
susceptible population (CASTELAR) and two greenhouse populations (ROSARIO and BELLA VISTA).
Insecticides were dissolved in acetone and topically applied to the mid-dorsal abdominal region of two-
day old 4
th
instar larvae. LD
50
values were estimated and the Resistance Ratio (RR) for each insecticide
was calculated (RR = LD
50
value of each greenhouse population/LD
50
value of the susceptible
population). ROSARIO and BELLA VISTA populations showed the following RRs values: > 68.38 for
deltamethrin; 2.48 and 3.49 for abamectin, respectively; and 0.79 and 0.86 for metamidophos, respectively.
Deltamethrin resistance observed in ROSARIO could be due to the high selective pressure exerted by
pyrethroids in this location. Deltamethrin resistance in BELLA VISTA is more difficult to explain,
because pyrethroids were scarcely used in the greenhouse where the insects were sampled. The incipient
abamectin resistance detected in the BELLA VISTA population could result from the frequent use of
this insecticide in this location, although natural variation can not be discarded.
KEY WORDS: Tomato leafminer, deltamethrin, abamectin, metamidophos, insecticide resistance
Resistncia a Inseticidas em Populaes Argentinas de Tuta absoluta (Meyrick) (Lepidoptera: Gelechiidae)
114 Insecticide Resistance in Argentine Populations of Tuta absoluta (Meyrick)... Lietti et al.
Tomato crop is the second horticultural crop in Argentina,
with a harvested area of 24,000 ha and an average yield of
30,833 kg/ha (FAO 1996). Seventy per cent of the production
destination is for consumption in natura and the rest is
industrialized (Gmez Riera 1992). Tomato produced under
greenhouses as well as in outdoor areas for consumption in
natura brings the highest gross financial return to farmers in
the Litoral Region (Buenos Aires, Corrientes and Santa Fe
provinces) (Stoppani & Rodrguez 1992).
The tomato leafminer, Tuta absoluta (Meyrick), is a
neotropical oligophagous insect, which attacks solanaceous
crops. Since the 1960s it has become one of the key pests of
tomato crops in many South American countries (Souza et
al. 1983, Larran 1986, IAN 1994). A fruit importation from
Chile may have introduced it to Mendoza province
(Argentina) in April 1964 (Bahamondes & Mallea 1969);
dissemination to other tomato production regions occurred
through fruit commercialization (Benavent et al. 1978, Cceres
1992, Riquelme 1993).
Larvae can damage tomato plants during all growth stages,
producing large galleries in their leaves, burrowing stalks,
apical buds, green and ripe fruits (Cceres 1992, IAN 1994). It
can cause important yield losses in different production
regions and under diverse production systems (Benavent et
al. 1978, Cceres 1992).
Since its introduction, chemical control has been the main
method of control used against T. absoluta in all production
regions in Argentina. Horticultural growers have tried to
decrease its injure applying insecticides two times a week
during a single cultivation period. Effective chemical control
was difficult to achieve because of the mine-feeding
behaviour of larvae, lack of a threshold action, and deficient
spraying technology.
Initially, the only insecticides used against tomato
leafminer in Argentina were organophosphates, which were
gradually replaced by pyrethroids during the 1970s. In the
early 1980s, cartap, alternated with pyrethroids, and
thiocyclam were introduced with the former showing excellent
efficacy at that moment. During the 1990s, insecticides with
novel sites of action such as abamectin, acylurea insect
growth regulators, spinosad, tebufenozide and chlorfenapyr
were introduced (Galarza & Larroque 1984, Polack 1999,
Cceres 2000).
Reports of insecticide resistance development in
populations of T. absoluta were scarce. A decrease in the
control efficiency of organophosphorous insecticides was
observed in Bolivia and Chile, which could be satisfactorily
controlled by pyrethroids (Moore 1983, Larran 1986).
Recently, the existence of resistance to organophosphates
and pyrethroids in Chile (Salazar & Araya 1997, 2001) and to
abamectin, cartap, methamidophos and permethrin in Brasil
(Siqueira et al. 2000, 2001) were reported.
The suspicion about the development of resistance in
Argentine populations of T. absoluta has been present since
the 1980s, when growers and agronomic advisers observed
that some of the compounds recommended for its control
were loosing its effectiveness in the field. Apparently, the
loss in effectiveness has not occurred with the same intensity
and to the same compounds in all tomato producing regions.
Nevertheless, the susceptibility of different populations to
distinct active ingredients has not yet been determined.
Owing to a decreasing activity of some insecticides used
against T. absoluta and the report of resistant populations
in Chile and Brasil, it is necessary to detect and quantify
the resistance to the main insecticides used for its control
in Argentina.
The objective of this work was to evaluate the toxicity
of abamectin, deltamethrin and methamidophos to a
laboratory-susceptible and two greenhouse populations of
T. absoluta, in order to establish if insecticide resistance
has developed in the last ones. The three insecticides used
in this study are currently registered and widely used by
farmers as chemical control of T. absoluta and other tomato
pests in Argentina (CASAFE 2001).
Material and Methods
Biological Material. A susceptible population of T. absoluta
(CASTELAR population), reared since 1993 without
exposure to insecticides, was provided by the Insectario de
Investigaciones para Lucha Biolgica, IMIZA, CNIA-INTA.
(Castelar, Buenos Aires Province).
Greenhouse individuals of T. absoluta were collected
from tomato crops cultivated in the Mdulo Experimental
de Nuevas Tecnologas (Rosario, Santa Fe Province)
(ROSARIO population) and in the Experimental Station,
INTA (Bella Vista, Corrientes Province) (BELLA VISTA
population). Both populations came from experimental
crops grown under greenhouses and exposed to intense
chemical treatments to maintain them free of pests. The
greenhouses are made of wood and covered by plastic.
The sides of them are lifted, whenever required, for
regulating the temperature inside it. Information about
insecticides used in each site during the current cultivation
period was obtained. Individuals from both the susceptible
and the greenhouse populations were separately reared
on greenhouse organically-grown tomato plants
(Lycopersicon esculentum Mill) cv. Presto (Petoseed )
in a controlled environment room at 25 2C and under a
photoperiod of 14:10 L:D. The experimental work was done
on two-day old 4
th
larval stage.
Chemicals. The insecticides used in this study were
abamectin 94% (Chemotecnica, Argentina), deltamethrin >
99% (Roussell-Uclaf, France) and methamidophos 68%
(Bayer, Argentina). Acetone pro-analysis (Merck,
Argentina) was used as solvent.
Bioassay. Insecticides were topically applied to the mid-
dorsal abdominal region of the larvae using a micro syringe
provided with a dispenser. Each insect received 0.2 l of a
solution of insecticide in acetone. Control groups were
topically treated with acetone alone. Five to seven doses of
each compound and 15 to 20 larvae for each dose were used
to estimate the Lethal Dose 50% (LD
50
) values. Three to
five independent replicates for each bioassay were done.
Data of the different replicates were pooled when the
confidence limits of their respective LD
50
overlapped.
After treatment, the larvae were individually placed in 3 cm
3
plastic vials (13 x 35 mm) in a controlled environment room at 27
2C and under a photoperiod of 14:10 (L:D). Mortality was
recorded 24h after treatment under stereoscopic microscope (10x).
Larvae were considered as dead when they were not able to
move back to ventral position after being placed on their dorsum.
The LD
50
values were calculated using the Probit method
(Lichtfield & Wilcoxon 1949). In all cases, differences between
values were considered significant (P < 0.05) if the respective
95% confidence limits did not overlap.
Resistance Ratios (RRs) values were calculated by
dividing the LD
50
value of each greenhouse population by
the LD
50
value of the susceptible population. Confidence
limits of RRs were calculated according to Robertson &
Preisler (1992). RRs were considered significantly different
from 1 (P < 0.05) when their 95% confidence limits did not
include 1.
Results
Fig. 1 shows the dose-response relationship for abamectin
and metamidophos in an insecticide susceptible
(CASTELAR) and two greenhouse populations (BELLA
VISTA and ROSARIO) of T. absoluta. The DL
50
and RRs
values are shown in Table 1. The LD
50
value of deltamethrin
was 0.35 g/larva for the CASTELAR population. The LD
50
values of deltamethrin could not be calculated for the BELLA
VISTA and ROSARIO populations because the highest dose
applied (24 mg/larva, which is near the solubility limit of the
insecticide) caused only 31.9 and 18.3% of mortality,
respectively. The RRs values were > 68.4 in both cases,
indicating a high resistance to deltamethrin.
The LD
50
values of methamidophos were 0.81, 0.70 and
0.65 g/larva for the CASTELAR, BELLA VISTA and
ROSARIO populations, respectively. No significant
differences among those values were observed (P > 0.05).
The RRs values were not significantly different from 1 (P >
0.05). Hence, no resistance to this insecticide was observed
in the greenhouse populations.
The LD
50
values of abamectin were 0.16, 0.57 and 0.41 g/
larva for the CASTELAR, BELLA VISTA and ROSARIO
populations, respectively. The BELLA VISTA LD
50
value was
significantly higher than the CASTELAR one (P < 0.05), but
there was no significant difference between the ROSARIO
and CASTELAR values (P > 0.05). BELLA VISTA and
ROSARIO RRs were 3.49 and 2.48, respectively. Both values
differed significantly from 1 (P < 0.05). These results indicated
a slight resistance to abamectin in the BELLA VISTA
population.
Discussion
There are very few studies of insecticide resistance in T.
absoluta. Salazar & Araya (1997) reported resistance to
deltamethrin, metamidophos, esfenvalerate, lambda-
cyhalothrin and mevinphos in Chilean populations of this
pest. Studying different larval stages from several localities,
they found RRs values ranging from 2.2 to 8.2 for deltamethrin,
from 1.6 to 3.9 for metamidophos, from 1.9 to 12.6 for
Figure 1. Dose-response regressions for metamidophos
and abamectin topically applied on T. absoluta 4
th
larval stage
from a laboratory (CASTELAR) and two greenhouse
populations (ROSARIO and BELLA VISTA).
dose (g/insect)
0,1 1,0 10,0
m
o
r
t
a
l
i
t
y

(
%
)
10
20
30
40
50
60
70
80
90
95
98
99
CASTELAR (1)
BELLA VISTA (3)
ROSARIO (2)
2
3
1
metamidophos
dose (ng/insect)
0,01 0,10 1,00 10,00
m
o
r
t
a
l
i
t
y

(
%
)
10
20
30
40
50
60
70
80
90
95
98
CASTELAR (1)
BELLA VISTA (3)
ROSARIO (2)
2 3
1
abamectin
Metamidophos
CASTELAR (1)
ROSARIO (2)
BELLA VISTA (3)
Abamectin
M
o
r
t
a
l
i
t
y

(
%
)
M
o
r
t
a
l
i
t
y

(
%
)
Dose (ng/insect)
1 3
2
1
3
2
99
98
95
90
80
70
60
50
40
30
20
98
95
90
80
70
60
50
40
30
20
0.1 1.0 10.0
0,01 0,10 1,00 10,00
esfenvalerate, from 1.8 to 11.5 for lambda-cyhalothrin, and
from 1.9 to 5.5 for mevinphos. Later, Salazar & Araya (2001)
found higher RRs values for the same insecticides in other
populations of T. absoluta.
Resistance to abamectin, cartap, methamidophos and
permethrin was reported in several Brazilian populations of
T. absoluta (Siqueira et al. 2000). The RRs values varied from
5.2 to 9.4 for abamectin, from 2.2 to 21.9 for cartap, from 2.6 to
4.2 for metamidophos, and from 1.9 to 6.6 for permethrin. In
other study, Siqueira et al. (2001) studied the toxicity of
abamectin, with and without synergists, to six abamectin
resistant populations of T. absoluta. A complex result was
CASTELAR (1)
ROSARIO (2)
BELLA VISTA (3)
Dose ( g/insect)
Table 2. Insecticides applied to control the ROSARIO
population of T. absoluta before the insects were collected
for this study. Mdulo Experimental de Nuevas Tecnologas,
Rosario, Santa Fe.
Table 1. Susceptibility of Argentine populations of the tomato leafminer, T. absoluta, to insecticides.
1
95% CL = 95% confidence limits;
2
Resistance ratio = LD
50
greenhouse population/LD
50
laboratory population;
3
NS = No significant;
4
Significantly different from the laboratory-susceptible population (CASTELAR) (P < 0.05);
5
Significantly different from 1 (P < 0.05).
obtained: piperonyl butoxide (an inhibitor of the mixed
function microsomal oxidases activity MFMO)
suppressed the abamectin resistance completely in only one
population and partially in the rest; triphenylphosphate (an
inhibitor of the esterase activity) suppressed completely the
abamectin resistance in four populations; diethyl maleate (an
inhibitor of the glutathion S-transferase activity) suppressed
partially the resistance in nearly all populations. These results
suggested to the authors that the resistance to abamectin in
T. absoluta populations may be oligo or even polyfactorial,
and that several genes should be involved.
In this work, the existence of insecticide resistance in
Argentine populations of T. absoluta has been experimentally
demonstrated for the first time. Deltamethrin resistance was
observed in the two populations studied (ROSARIO and
BELLA VISTA), and a weak resistance to abamectin was
observed just in one of them (BELLA VISTA).
Deltamethrin is an insecticide widely used in the Horticultural
Belt of Rosario city (where the ROSARIO population was
collected). We found a high level of resistance (> 68.4) to
deltamethrin in individuals from the ROSARIO population. In
75% of cases, pyrethroids (deltamethrin or lambda-cyhalothrin)
were sprayed before our sampling (Table 2). The deltamethrin
resistance observed could be due to the high selective pressure
exerted by pyrethroids on this population.
Abamectin was introduced for tomato leafminer control
in the Horticultural Belt of Rosario city in the 1990s to
overcome the decrease of cartap efficacy. The ROSARIO
population received no application of abamectin, neither
before the insects collection nor in recent previous years
(Table 2). We found no significant differences between the
LD
50
values of this insecticide evaluated on the ROSARIO
and CASTELAR populations.
Metamidophos is a broad spectrum systemic insecticide
used by the farmers in the Horticultural Belt of Rosario city
only during the vegetative growth stage of tomato because of
its pre-harvest interval of 10 days. The lack of resistance to
metamidophos in the ROSARIO population could be due to
the scarce use of this compound in the sampling site (Table 2).
Deltamethrin was ineffective to control the tomato
leafminer in the Experimental Station of Bella Vista city as
resulted from efficacy assays carried out in the period 1981
to 1987 (Cceres 1992), when this pest began to compromise
tomato production in this location. However, deltamethrin is
used by farmers to control several tomato and pepper pest
species (9.5% of the total number of sprays per cultivation
cycle) (S. Cceres, personal communication). The total number
of sprays received by the BELLA VISTA population before
Insecticide Population n
Slope
SE
LD
50
(95% CL)
1
g a.i./larva
2
Resistance Ratio
2
(95% CL)
1
Deltamethrin CASTELAR 479 0.88 0.13 0.35
(0.22 0.55)
3.06
NS
3
--
ROSARIO 115 -- > 24.00 -- > 68.38
BELLA VISTA 130 -- > 24.00 -- > 68.38
Methamidophos CASTELAR 230 4.42 0.45 0.81
(0.65 0.96)
2.12
NS
--
ROSARIO 222 1.85 0.44 0.65
(0.21 1.06)
1.06
NS
0.79
(0.40 1.60)
BELLA VISTA 146 4.16 0.45 0.70
(0.55 0.85)
0.71
NS
0.86
(0.65 1.5)
g a.i./larva
Abamectin CASTELAR 348 0.97 0.12 0.16
(0.09 0.27)
4.83
NS
--
ROSARIO 324 1.35 0.19 0.41
(0.23 0.62)
0.98
NS
2.48
5
(1.22 5.02)
BELLA VISTA 188 1.43 0.29 0.57
4
(0.28 0.92)
1.47
NS
3.49
5
(1.64 7.42)
Months
(year 2000)
Insecticide
Number of
applications
Commercial
name
June to
October
Deltamethrin 7 Decis 5
July to
November
Lamdba-
cyhalotrin
5 Karate
July Methamidophos 1 Tamaron
August Cartap 1 Padam 50 SP
September
and October
Imidacloprid 2 Confidor
Table 3. Insecticides applied to control the BELLA VISTA
population of T. absoluta before the insects were collected
for this study. Experimental Station, INTA, Bella Vista,
Corrientes.
the insect collection was 11, with seven applications of
abamectin (63.6%), three of imidacloprid (27.3%) and one of
deltamethrin (9%) (Table 3). During the two previous years,
deltamethrin was not used in the greenhouse where this
population was collected, but it was applied in other
greenhouses of the same Experimental Station.
In the BELLA VISTA population, a high deltamethrin
resistance (RR > 68.4) was observed, although this population
received a relatively low number of pyrethroids applications.
This result is more difficult to explain. Deltamethrin was used
in other greenhouses in the Horticultural Belt of Bella Vista,
so resistance could be due to migration. An incipient
abamectin resistance was observed in BELLA VISTA (see
below), so an alternative explanation is that deltamethrin
resistance is due to cross-resistance after abamectin selective
pressure. Pyrethroids and abamectin have different sites of
action, however, a common metabolic detoxification
mechanism could confer resistance to both insecticide
groups. For instance, abamectin resistance in the Colorado
potato beetle, Leptinotarsa decemlineata (Say), largely
resulted from increased MFMO activity (Clark et al. 1994).
The involvement of MFMO in the resistance to deltamethrin
was also confirmed in Cidia pomonella (L.) after in vitro
metabolism studies (Sauphanor et al. 1997). Abro et al. (1988)
reported a cypermethrin resistant population of Plutella
xylostella (L.) which was also resistant to abamectin,
indicating that cross-resistance between pyretroids and the
latter is possible. The simultaneous application of piperonyl
butoxide was followed by partial reversion of the resistance
to deltamethrin in C. pomonella (Sauphanor et al. 1997).
Piperonyl butoxide also synergized abamectin in resistant
individuals of P. xylostella (Abro et al. 1988) and, as discussed
above, T. absoluta (Siqueira et al. 2001). These results suggest
that MFMO activity could be a mechanism of resistance to
both pyrethroids and abamectin.
Abamectin, in combination with mineral oil, was
introduced for chemical control of the tomato leafminer in
Bella Vista in 1994 and it is still effective in the field (Cceres
2000). Abamectin has been used in the greenhouses of the
Experimental Station of Bella Vista before collecting the
BELLA VISTA population and during the last two years, and
a weak resistance to this compound (RR = 3.5) was observed.
It is possible that resistance is arising due to the selective
pressure exerted by the application of abamectin. Other
possibility is that the values observed are due to natural
variation (this can not be confirmed because in Argentina
there is not a baseline of insecticide toxicity to T. absoluta).
Methamidophos was not used for chemical control of the
tomato leafminer in the Bella Vista Experimental Station, mainly
due to the relatively low efficacy of this compound in
comparison with abamectin (Cceres 1992). Nevertheless,
metamidophos is used by growers to control several tomato
and pepper pest species (26.2% of the total number of sprays
per cultivation cycle on tomato and pepper) (S. Cceres,
personal communication). The lack of usage in the sampling
site would explain that no resistance was observed for this
compound in the BELLA VISTA population (Table 3).
In the Litoral Region of Argentina, protected tomato
production offers a favorable environment for the
development of T. absoluta from May to August, providing
it with food and shelter. Moreover, since populations develop
faster in greenhouses than outdoors, the population level
and the consequent damage abruptly increase since early
October, which leads to early use of insecticides under
protected cultivation. The tomato leafminer has several
biological traits that favor resistance development. It has
many generations per year (more than five), which overlap in
protected crops, it has a relatively high fecundity (an average
of 40 to 55 eggs per female) and it survives in the soil at the
pupa stage (Botto 2000). The larvae and pupae are transported
form one region to another in infested containers and fruits,
allowing the mixing of individuals subjected to different
chemical regimens.
Although a rotation of different active ingredients was
done during each cultivation cycle in the two populations
studied, it was not aimed to manage resistance. The lack of
insecticide resistance management tactics favors resistance
development. The occurrence of resistant individuals under
greenhouses constitutes a potential danger. Taking into
account that the sides of the greenhouses are lifted whenever
required, for regulating the temperature inside them, the
resistant individuals might migrate outdoors in spring,
colonizing outdoor tomato crops. When insecticides are
applied, the resistant individuals have a biological advantage
in contrast with susceptible ones. This would contribute to
the development of resistance in outdoor tomato crops.
Resistance development is a consequence of insecticide
applications on an insect population and it must be
managed. Resistance management should be a component
of integrated pest management, which seeks to minimize
pesticide usage through the application of alternative
tactics such as cultural control and conservation of natural
control through selective insecticides. Monitoring the
susceptibility of different populations exposed to distinct
active ingredients is essential. The data obtained by means
of a monitoring plan should be used to design resistance
managing strategies, for example, the alternation of
insecticides with different modes of action and
detoxification, or the diminishing of conventional insecticide
use by employing alternative control tools. Several pests,
such as thrips and white flies, develop on tomato crops and
are sometimes controlled by the same compounds used
against the tomato leafminer. For this reason, insecticide
resistance management of T. absoluta should be considered
in the context of key-pest arthropod complexes of crops.
Month
(year 2000)
Insecticide
Number of
applications
Commercial
name
April Deltamethrin 1 Decis 5
May to
September
Abamectin 7 Vertimec
July to
September
Imidacloprid 3 Confidor
Acknowledgments
To the Centro de Investigaciones de Plagas e Insecticidas
for providing the technical grade insecticides deltamethrin
and abamectin. To Bayer Argentina for donating the technical
grade insecticide metamidophos. To Ing. Agr. Sara Cceres
(E.E.A.INTA Bella Vista) for sending the T. absoluta
population from Bella Vista. To the two anonymous reviewers
for their helpful comments.
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Sauphanor, B., A. Cuany, J.C. Bouvier, V. Brosse, M. Amichot,
& J.B. Berg. 1997. Mechanism of resistance to
deltamethrin in Cydia pomonella (L.) (Lepidoptera:
Tortricidae). Pest. Biochem. Physiol. 58: 109-117.
Siqueira, H.A. de, R.N. Guedes, D.B. Fragoso & L.C.
Magalhes. 2001. Abamectin resistance and synergism
in brazilian populations of Tuta absoluta (Meyrick)
(Lepidoptera: Gelechiidae). Int. J. Pest Manag. 47: 247-
251.
Siqueira, H.A. de, R.N. Guedes & M.C. Picano. 2000.
Insecticide resistance in populations of Tuta absoluta
(Lepidoptera:Gelechiidae). Agric. Forest Entomol. 2: 147-
153.
Souza, J.C., P.R. Reis, A. de Pdua Nacif, J.M. Gomes &
L.O. Salgado. 1983. Controle da traa-do-tomateiro.
Histrico, reconhecimento, biologa, prejuzos e controle.
Belo Horizonte, Empresa de Pesquisa Agropecuria de
Minas Gerais, 15p.
Stoppani, M.I. & J.P. Rodrguez. 1992. El tomate
(Lycopersicon esculentum Mill.) y su cultivo bajo
proteccin en la regin litoral sur del ro Paran
(Repblica Argentina). Informe Tcnico 61. Estacin
Experimental Agropecuaria San Pedro, INTA, 47p.
Received 07/X/04. Accepted 07/X/04.
65 Efecto del imidacloprid en el control de la polilla del tomate (Tuta absoluta Meyrick) Volumen 26, N 1, Pginas 65-72
IDESIA (Chile) Enero - Abril 2008
1
Ctedra de Zoologa Agrcola, Facultad de Agronoma, Universidad de Buenos Aires. Av. San Martn 4453, Buenos Aires
(CP C1417DSE), Argentina. E-mail: collavin@agro.uba.ar
2
Ctedra de Teraputica Vegetal, Facultad de Agronoma, Universidad de Buenos Aires. dem anterior. E-mail: rgimenez@
agro.uba.ar
Fecha de Recepcin: 06 Marzo 2006
Fecha de Aceptacin: 22 Octubre 2007
EFECTO DEL IMIDACLOPRID EN EL CONTROL DE LA
POLILLA DEL TOMATE (TUTA ABSOLUTA MEYRICK)
EFFICACY OF IMIDACLOPRID TO CONTROL THE TOMATO BORER
(TUTA ABSOLUTA MEYRICK)
Marcelo Dante Collavino
1
; Rosana Alejandra Gimnez
2
RESUMEN
La grave incidencia de Tuta absoluta en la produccin de tomate, el alto uso de plaguicidas para su control, el riesgo de contami-
nacin del ambiente y la generacin de resistencia hacen que sea muy importante encontrar formas alternativas de control eficiente
de esta plaga. Con el fin de evaluar la eficacia del imidacloprid en el control de la polilla del tomate, en distintas dosis y formas
de aplicacin se desarroll este ensayo en condiciones de invernculo.
Para ello se prepararon dos diluciones del insecticida, que fueron aplicadas de dos formas: por riego y por inmersin de la raz
del plantn con su pan de tierra.
Para evaluar el dao se realiz el clculo del porcentaje de fololos daados por planta y el nmero de lesiones por hoja en dis-
tintos momentos desde la aplicacin. El diseo fue factorial, completamente aleatorizado y con nueve repeticiones, aplicndose
ANOVA. Se comprob que hay diferencias significativas entre las diluciones y tambin entre las dos formas de aplicacin (riego
e inmersin). Siendo los tratamientos que mejor controlaron a la plaga: dilucin al 3,5% aplicada por riego y dilucin al 7%
aplicada por inmersin.
Palabras clave: Tuta absoluta, imidacloprid, riego, inmersin.
ABSTRACT
The incidence of Tuta absoluta in tomato production, the use of pesticides for its control, the risk of contamination to the envi-
ronment and the creation of a generation of resistance, makes it very important to find alternative ways to efficiently control the
tomato borer. This test was developed under greenhouse conditions to assess the effectiveness of the imidacloprid to the control of
this pest at different rates and application forms. Two dilutions of the insecticide were prepared and applied in two different ways:
by watering and by the immersion of the root of the plants into the soil bed. To evaluate the damage, calculations were carried
out on the percentage of the leaves damaged on each plant and on the number of lesions on each leaf at different moments of the
application. The design was factorial, completely randomized and with 9 replications, and an ANOVA was carried out. Significant
differences were confirmed among the dilutions and also among the two application forms (watering and immersion). The better
treatments were: dilution of 3,5% applied by watering and dilution of 7% applied by immersion.
Key words: Tuta absoluta, imidacloprid, watering, immersion.
INTRODUCCIN
En el cultivo de tomate el tipo de produccin ms
tecnificado es el realizado bajo invernculo, donde
el problema de la polilla del tomate (T. absoluta)
es mucho ms grave que a campo (Ripa, 1981).
Dentro del invernculo es menor la incidencia de
las bajas temperaturas, producindose condiciones
muy favorables para incrementar la incidencia de
esta plaga, que acorta su ciclo de vida y produce
muchas generaciones superpuestas a lo largo del
ciclo de cultivo.
IDESIA (Chile) Volumen 26, N 1, Enero-Abril, 2008 66
T. absoluta (Lepidoptera; Gelechiidae) pro-
duce daos muy importantes al cultivo del tomate
en casi todas las zonas de produccin hortcola de
Argentina (INTA, 1991). Las larvas lesionan el
follaje haciendo galeras dentro de las hojas que
pueden llegar a ocupar gran parte del rea foliar.
Tambin penetran en el fruto construyendo tneles
que al llenarse de excrementos causan la pudricin
y cada de los mismos (Betancourt, 1995). De esta
manera se pierde rendimiento y tambin disminuye
la calidad de los frutos obtenidos.
Los graves daos que esta plaga produce han
hecho que los productores aplicaran insecticidas
muy frecuentemente, generando resistencia a la
mayora de los insecticidas utilizados, especialmente
hacia los fosforados (Larran, 1986). En Argentina,
actualmente, estn registrados numerosos productos
para el control de esta plaga: ciflutrina, ciperme-
trina, deltametrina, fenvalerato, lambdacihalotrina
y permetrina (piretroides), acefato, clorpirifs,
fenitrotin, metamidofs, piridafentin y triazofs
(organofosforados), abamectn y cartap (microbio-
lgicos), clorfluazurn, lufenurn, metoxifenocide,
tebufenozide, teflubenzurn y triflumurn (IGR),
spinosad (Naturalyte) y tiocicln hidrogenoxalato
(nereistoxina) (CASAFE, 2001). Mientras que el
imidacloprid se encuentra registrado para aplicar en
el cultivo de tomate exclusivamente para controlar
moscas blancas (Bemisia tabaci, Trialeurodes spp. y
Aleurothripsus spp.) y minador del tomate (Faustinus
cubae) y trips (Frankliniella schultzei).
Otros investigadores han realizado estudios
para establecer la eficacia de insecticidas (Reis et
al., 1998; Lima y Machado, 1996) en aplicaciones
foliares para controlar T. absoluta.
El imidacloprid se caracteriza por tener un
modo de accin sistmico y de contacto, amplio
espectro de accin y posibilidad de ser absorbido
tanto por follaje como por races. Adems, posee
un mecanismo de accin diferente al de la mayora
de los insecticidas, por lo que no se han registrado
casos de resistencia en las plagas. Pertenece al
grupo de las nitroguanidinas y posee un efecto
residual de varias semanas y una baja toxicidad
para mamferos, con una dosis letal media (DL 50)
oral aguda en rata 24 h de 450 mg/kg (CASAFE,
2001). Adems, Epperlein et al. (1997) mostraron
que este insecticida aplicado al suelo no produjo
efectos adversos sobre artrpodos predadores de la
superficie del suelo.
Todas estas caractersticas hacen que sea posible
su aplicacin mediante mtodos no convenciona-
les, haciendo tratamientos menos contaminantes
y ms econmicos, como seran las aplicaciones
localizadas, evitando la pulverizacin foliar en
cobertura total.
As, con el fin de encontrar formas alternativas
de control eficiente de la polilla del tomate, evalua-
mos la aplicacin de imidacloprid mediante riego
y por inmersin de races al trasplante.
MATERIALES Y MTODOS
El insecticida con que se realiz este trabajo es
el formulado comercial Confidor SL100 de Bayer,
cuyo ingrediente activo es el imidacloprid.
El cultivo fue realizado en un invernculo de esta
Facultad, con tomate platense comn y ejemplares
de T. absoluta criados en cmaras.
La cra de T. absoluta se realiz en jaulas con
luz artificial, sobre plantines de tomate cultivados
en macetas plsticas.
La siembra de tomate fue realizada en alm-
cigos tipo speedling, con sustrato previamente
desinfectado con sulfato neutro de oxiquinoleina.
Estos almcigos se regaron peridicamente con
solucin fertilizante N-P-K (15-15-15).
Se utilizaron 45 macetas plsticas de 25 cm
de dimetro, para realizar all el trasplante de los
plantines.
Se prepararon dos diluciones del insecticida,
una al 3,5% y otra al 7%. Ambas fueron aplicadas
de dos formas distintas:
1) Riego de las macetas con regadera, un da antes
del trasplante. El riego se realiz hasta saturar
el suelo.
2) Inmersin de la raz del plantn con su pan de
tierra, durante 10 seg. Una vez escurrida, el
plantn se trasplantaba a la maceta.
Quedan as determinados cinco tratamientos:
Testigo (sin insecticida);
Dilucin al 3,5% aplicada por inmersin;
Dilucin al 3,5% aplicada por riego;
Dilucin al 7% aplicada por inmersin;
Dilucin al 7% aplicada por riego.
Luego de realizar la conduccin y la poda de
los brotes axilares de las plantas, se liberaron en
el interior del invernculo aproximadamente 100
ejemplares de T. absoluta (adultos y pupas), tapan-
67 Efecto del imidacloprid en el control de la polilla del tomate (Tuta absoluta Meyrick)
do con tul y papel todos las aberturas por donde
pudieran escapar las polillas.
Para evaluar el dao se realiz el clculo del
porcentaje de fololos daados por planta y el
nmero de lesiones por hoja, a los 27, 34, 41 y 48
das desde la aplicacin del producto.
Se implement un diseo estadstico factorial con
un tratamiento adicional, completamente aleatorio
y con nueve repeticiones por tratamiento; luego los
datos fueron analizados por medio de anlisis de
variancia ANOVA al 5 y 1 %.
RESULTADOS Y DISCUSIN
Como consecuencia de la aplicacin del imi-
dacloprid se advierte, en primera instancia, que las
plantas tratadas no sufren el ataque de la polilla, o lo
sufren en menor medida que aquellas a las cuales
no se les aplic dicho insecticida.
A continuacin se presentan los datos generados
en las observaciones semanales realizadas.
Como puede observarse en la Figura 1, el por-
centaje de fololos daados por planta se incrementa
Tabla 1
Promedios registrados a 27 das de la aplicacin
Testigo 3,5% riego 3,5% inmersin 7% riego 7% inmersin
Porcentaje de fololos daados
por planta
34,88 4,13 16,48 0,00 2,89
Nmero de lesiones por hoja 3,95 0,49 2,25 0,00 0,24
Tabla 2
por planta
44,60 10,44 26,75 3,20 11,26
Tabla 3
por planta
55,47 29,66 45,00 10,40 26,33
Tabla 4
por planta
63,48 31,01 48,43 13,47 28,68
a lo largo del tiempo. Los distintos tratamientos se
diferencian del testigo en mayor o menor medida
y en todos ellos se evidencia un dao de menor
magnitud que en ste.
En la Figura 2 se observa que a los 34 das
de la aplicacin hay 3 tratamientos con daos
que permanecen por debajo del umbral de dao
econmico (UDE) sugerido por Ariso (1997),
quien considera que para realizar una aplicacin
de insecticida en tomate cultivado en invernculo
se deben observar en promedio 3 lesiones por hoja.
As, en este ensayo (Figura 2) los tratamientos
realizados alcanzaron este umbral de dao en dis-
tintos perodos desde la aplicacin: el testigo en la
primera observacin (27 das) ya haba superado
el umbral mencionado; en los tratamientos 3,5%
inmersin, 7% inmersin, 3,5% riego alcanza el
mismo a los 28, 35 y 37 das, respectivamente,
mientras que el tratamiento 7% riego no alcanz
este UDE; sin embargo, en este ltimo tratamiento,
no se puede considerar que fue un largo perodo
de proteccin de las plantas, ya que se produjo
fitotoxicidad en el tomate.
Con anlisis estadstico se verific la existencia
de diferencias altamente significativas entre ambas
diluciones y tambin entre las dos formas de aplica-
cin (riego e inmersin) tanto para los resultados de
lesiones por hoja (Tablas 5 a 8) como para porcentaje
de fololos daados (Tablas 9 a 12).
No existe interaccin AB; Diferencias altamente
significativas entre diluciones; Diferencias altamente
significativas entre aplicaciones.
La dilucin al 7% aplicada por riego se descart
del anlisis por haber producido en todas las plantas
0
10
20
30
40
50
60
70
27 das 34 das 41 das 48 das
Das desde la aplicacin
P
o
r
c
e
n
t
a
j
e

d
e

f
o
l
o
l
o
s

d
a
a
d
o
s
Testigo 3,5% riego 3,5% inmersin
7% riego 7% inmersin
Figura 1. Evolucin del dao.
Tabla 5
ANOVA. Lesiones por hoja a los 27 das de la aplicacin.
Fuentes de
Variacin
SC GL CM F F 0.05 F 0,01
Dilucin 137.181 2 68.591 42.332 ** 3.185 5.070
Aplicacin 9.019 1 9.019 5.567 * 4.035 7.180
Interaccin AB 5.266 1 5.266 3.250 4.035 7.180
Error 79.394 49 1.620
Total 230.861 53 4.356
No existe interaccin AB; Diferencias altamente significativas entre diluciones; Diferencias significativas entre aplicaciones.
0
3
6
9
12
15
27 das 34 das 41 das 48 das
Das desde la aplicacin
L
e
s
i
o
n
e
s

p
o
r

h
o
j
a
3,5% riego 3,5% inmersin Testigo 7% riego 7% inmersin
Figura 2. Lesiones por hoja a travs del tiempo.
Tabla 6
ANOVA. Lesiones por hoja a los 34 das de la aplicacin
Fuentes de
Variacin
SC GL CM F F 0,05 F 0,01
Dilucin 431.402 2 215.701 50.353 ** 3.185 5.070
Aplicacin 70.492 1 70.492 16.456 ** 4.035 7.180
Interaccin AB 5.397 1 5.397 1.260 4.035 7.180
Error 209.904 49 4.284
Total 717.195 53 13.532
No existe interaccin AB; Diferencias altamente significativas entre diluciones; Diferencias altamente significativas entre
aplicaciones.
Tabla 7
Fuentes de
Variacin
SC GL CM F F 0,05 F 0,01
Dilucin 559.424 2 279.712 51.202 ** 3.185 5.070
Aplicacin 197.589 1 197.589 36.169 ** 4.035 7.180
Interaccin AB 0.902 1 0.902 0.165 4.035 7.180
Error 267.681 49 5.463
Total 1025.597 53 19.351
aplicaciones.
Tabla 8
Fuentes de
Variacin
SC GL CM F F 0,05 F 0,01
Dilucin 824.602 2 412.301 21.110 ** 3.185 5.070
Aplicacin 297.374 1 297.374 15.225 ** 4.035 7.180
Interaccin AB 0.421 1 0.421 0.022 4.035 7.180
Error 957.043 49 19.531
Total 2079.440 53 39.235
aplicaciones.
Tabla 9
ANOVA. Porcentaje de fololos daados a los 27 das de la aplicacin
Fuentes de
Variacin
SC GL CM F F 0,05 F 0,01
Dilucin 10789.821 2 5394.911 28.533 ** 3.185 5.070
Aplicacin 529.786 1 529.786 2.802 4.035 7.180
Interaccin AB 205.504 1 205.504 1.087 4.035 7.180
Error 9264.782 49 189.077
Total 20789.893 53 392.262
No existe interaccin AB; Diferencias altamente significativas entre diluciones; No hay diferencias entre aplicaciones.
Tabla 10
Fuentes de
Variacin
SC GL CM F F 0,05 F 0,01
Dilucin 13211.075 2 6605.537 36.066 ** 3.185 5.070
Aplicacin 1337.561 1 1337.561 7.303 ** 4.035 7.180
Interaccin AB 153.031 1 153.031 0.836 4.035 7.180
Error 8974.321 49 183.149
Total 23675.986 53 446.717
aplicaciones.
Tabla 11
ANOVA Porcentaje de fololos daados a los 41 das de la aplicacin
Fuentes de
Variacin
SC GL CM F F 0,05 F 0,01
Dilucin 12391.490 2 6195.745 35.632 ** 3.185 5.070
Aplicacin 2199.270 1 2199.270 12.648 ** 4.035 7.180
Interaccin AB 0.811 1 0.811 0.005 4.035 7.180
Error 3520.291 49 173.883
Total 2311.862 53 436.073
Tabla 12
Fuentes de
Variacin
SC GL CM F F 0,05 F 0,01
Dilucin 16266.865 2 8133.433 49.073 ** 3.185 5.070
Aplicacin 2396.430 1 2396.430 14.459 ** 4.035 7.180
Interaccin AB 11.001 1 11.001 0.066 4.035 7.180
Error 8121.290 49 165.741
Total 26795.586 53 505.577
aplicaciones.
tratadas una marcada disminucin del crecimiento
y prdida de hojas.
Los tratamientos que mejor controlaron a la
plaga fueron (figura 1):
Dilucin al 3,5 % aplicada por riego.
Dilucin al 7% aplicada por inmersin.
Como se aprecia en la Figura 2, la aplicacin por
inmersin de la dilucin al 7% present niveles de
dao inferiores al UDE hasta los 37 das despus del
tratamiento y la aplicacin por riego de la dilucin
al 3% hasta los 35 das.
A travs de un ensayo, bajo condiciones de
invernculo, no se puede establecer con certeza
cul es el tratamiento adecuado para controlar la
polilla del tomate en condiciones de campo, ya que
los resultados tienen sustento slo en este mbito
de produccin.
Finalmente, en cultivos protegidos de tomate,
teniendo en cuenta aspectos econmicos del con-
trol de plagas, sera recomendable la aplicacin de
imidacloprid a la raz de las plantas jvenes por
inmersin (al 7% en agua) durante el transplante,
ya que es el tratamiento que requiere en total una
menor cantidad de producto y tiene una alta eficacia
de control.
CONCLUSIONES
En condiciones de invernculo la aplicacin de
la dilucin al 7% aplicada mediante riego result
fitotxica para el tomate platense comn.
Las aplicaciones de la dilucin al 3,5% apli-
cada por riego y de la dilucin al 7% aplicada por
inmersin resultaron ser las ms adecuadas para el
control de la plaga, por su eficacia y periodo post-
tratamiento con daos menores al UDE.
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ARISO, E. (1997). Comunicacin personal, Departamento
Tcnico de Bayer.
BETANCOURT, C. M. y SCATONI I. B. (1995). Description
of the development stages of the tomato borer,
Scrobipalpuloides absoluta (Mayrick) (Lep., Gelechiidae).
Boletn de investigacin, Facultad de Agronoma, Universidad
de la Repblica. 1995, 45: 1-14. Uruguay.
CASAFE (Cmara de Sanidad Agropecuaria y Fertilizantes).
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EPPERLEIN, K.; SCHMIDT, H. W.; SCHWALBE, R. 1997.
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INTA Centro Regional Cuyo (1991). Manual de cultivo de
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LARRAN, P. (1986). Eficacia de insecticidas y frecuencia
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