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Potential of “Lure and Kill” in Long-Term Pest Management andEradication of Invasive Species

A. M. EL-SAYED,1,2 D. M. SUCKLING,1 J. A. BYERS,3 E. B. JANG,4 AND C. H. WEARING5

J. Econ. Entomol. 102(3): 815Ð835 (2009)

ABSTRACT “Lure and kill” technology has been used for several decades in pest management anderadication of invasive species. In lure and kill, the insect pest attracted by a semiochemical lure isnot “entrapped” at the source of the attractant as in mass trapping, but instead the insect is subjectedto a killing agent, which eliminates affected individuals from the population after a short period. Inpast decades, a growing scientiÞc literature has been published on this concept. This article providesthe Þrst review on the potential of lure and kill in long-term pest management and eradication ofinvasive species. We present a summary of lure and kill, either when used as a stand-alone controlmethodor incombinationwithothermethods.Wediscuss its efÞcacy incomparisonwithothercontrolmethods. Several case studies in which lure and kill has been used with the aims of long-term pestmanagement (e.g., pink bollworm, Egyptian cotton leafworm, codling moth, apple maggot, biting ßies,and bark beetles) or the eradication of invasive species (e.g., tephritid fruit ßies and boll weevils) areprovided. Subsequently, we identify essential knowledge required for successful lure and kill programsthat include lure competitiveness with natural odor source; lure density; lure formulation and releaserate; pest population density and risk of immigration; and biology and ecology of the target species.The risks associated with lure and kill, especially when used in the eradication programs, arehighlighted. We comment on the cost-effectiveness of this technology and its strengths and weak-nesses, and list key reasons for success and failure. We conclude that lure and kill can be highlyeffective in controlling small, low-density, isolated populations, and thus it has the potential to addvalue to long-term pest management. In the eradication of invasive species, lure and kill offers a majoradvantage in effectiveness by its being inverse density dependent and it provides some improvementsin efÞcacy over related control methods. However, the inclusion of insecticides or sterilants in lureand kill formulations presents a major obstacle to public acceptance.

KEY WORDS lure and kill, attract and kill, attracticides, semiochemicals, pheromones

Reducing the quantity of insecticide applied in theenvironment is a major objective that drives researchfor the discovery of new behavior-modifying chemi-cals (semiochemicals) and for investigation of theirpotential in pest management and eradication of in-vasive species. Semiochemicals are being used in pestmanagement either alone as in mass trapping or mat-ing disruption (Carde and Minks 1995, Suckling 2000,El-Sayed et al. 2006, Byers 2007) or in combinationwith insecticides, sterilants or insect pathogenstermed “lure and kill” (or “lure and sterilize” and “lureand infect”). Lure and kill typically uses semiochemi-cals and insecticides in a concentrated area at the luresource to provide pest control. The insect respondingto the semiochemical lure is not “entrapped” at the

source of the attractant by adhesive, water, or otherphysical device as in mass trapping, but instead theinsect is subjected to a killing or sterilizing agent,which effectively eliminates it from the populationafter a short time (Jones 1998). This tactic has beendescribed in the literature with different nomencla-tures, for example, lure and kill, attract and kill, maleannihilation, bait sprays, and attracticide. In somecases, the boundaries between mass trapping and lureand kill are further blurred, such as when traps areinsecticide treated. Success of the lure and kill ap-proach in pest management depends on 1) insectscontacting the insecticide either mixed with semio-chemical or applied adjacent to the lure, 2) adequatedosingwith the insecticidebefore leaving the lure, and3) the level of mortality or adverse behavior-modify-ing effects that are eventually detrimental to the insectpopulation. Usually, insects can be attracted to a pointsource either by chemical signals, visual cues, acousticcues, or combination of any of these signals and cues.

Attractants used in lure and kill can be either crudebaits or synthetic semiochemicals. Crude baits havebeen used extensively with crawling insects (e.g., ants

1 HortResearch, Canterbury Research Centre, Lincoln, 8152, NewZealand.

2 Corresponding author, e-mail: [email protected] 2US Arid-Land Agricultural Research Center, USDAÐARS, 21881

North Cardon Lane, Maricopa, AZ 85238.4 U.S. PaciÞc Basin Agricultural Research Center, USDAÐARS, P.O.

Box 4459, Hilo, HI 96720.5 674 Rolling Ridges Rd., RD 4, Timaru, New Zealand.

0022-0493/09/0815Ð0835$04.00/0 � 2009 Entomological Society of America

and cockroaches), whereas semiochemicals-basedlure and kill has been used mainly with ßying insects(e.g., Lepidoptera, Diptera, and Coleoptera). This ar-ticle focuses only on lure and kill that use semiochemi-cals, which include pheromones (e.g., sex phero-mones), kairomones (e.g., host volatiles), attractantswith a known behavioral function (e.g., host plant oroviposition odors), and attractants identiÞed throughthe screening of candidate chemicals with poorlyknownbehavioral functions(BerozaandGreen1963).Semiochemicals used in lure and kill should have sev-eral key attributes to be suitable: 1) the deployed luresreleasing the odor plumes are perceived by nearly alladult males or females or both in the treated area, 2)the odor plumes are able to attract males or females orboth more effectively than natural odor sources (e.g.,virgin or mated females in the case of sex pheromone)within the treated area, 3) the lures entice these adultinsects to make direct contact with an insecticidal (orsterilant) component where all or a very high per-centage are subsequently killed (or sterilized), and 4)treatment is done from the time of Þrst adult emer-gence to the time of last adult emergence in thetreated area (this may be seasonal or continuous). Aspart of a pest management program, the effectivenessof lure and kill may not need to be optimal, providedthe beneÞts of male and/or female removal result indamage reduction or crop yield increases that aregreater than the cost of the treatment. In pest man-agement, residual populations that remain after treat-ment can be tolerable if populations are kept under an“economic” threshold, but this is not the aim in pesteradication. Therefore, the key objective for successof this technology in an eradication program of inva-sive species is to lure and kill all adult insects in aspeciÞed area before they mate, disperse, and repro-duce.

Considerable data have been accumulated in thescientiÞc literature on the application of semiochemi-cals in pest management (El-Sayed 2008). In a re-cently published article, El-Sayed et al. (2006) pro-vided a review on the potential of mass trapping inlong-term pest management and eradication of inva-sive species. In the present article, we provide a com-plementary review on the potential of lure and killapproaches that use semiochemicals for these pur-poses. Reviewing the literature on lure and kill indi-cates this approach has been mainly used against ag-riculturally and medically important pests, and too alesser extent against forestry pests. We provide anoverview on the application of lure and kill in pestmanagement, and eradication of invasive species sup-ported by case studies, and we summarize the knowl-edge that is needed for successful lure and kill pro-grams. We discuss different methodologies used tomeasure the efÞcacy and risks associated with thisapproachandhighlight thecritical issuesaffecting lureand kill efÞcacy based on published data.Lure and Kill Formulation. Lure and kill technol-

ogy is patentable, and this has resulted in commercialdevelopment (Antilla et al. 1996, Charmillot et al.2000) accompanied by commercial trials, and market-

ing of a variety of formulations. Those products usingdroplets of paste or gel, applied by hand, are variouslynamed Attract and Kill, Sirene, Appeal, and most re-cently GF-120 and SPLAT. The pheromone/semio-chemical and insecticide are incorporated in a paste,gel, or wax at known concentrations during manufac-ture, and the user applies speciÞed numbers of drop-lets per hectare directly from the commercial hand-held applicator. These formulations dominate theliterature. There are also microencapsulated formu-lations (pheromone/semiochemical and insecticide)that are applied with a hand-held sprayer to providea known number of lures per hectare. This methodalso can accommodate other formulations of the in-secticide (e.g., emulsiÞable concentrate). A thirdhand-applied lure and kill formulation uses plasticsheets containing the pheromone that are cut intoindividual lures; these are stapled to the substrate andthen hand-sprayed with insecticide. The Ecogen No-mate hollow Þbers (also called Attract and Kill) can beapplied by air; the hollow Þbers, containing the pher-omone, are premixed with an adhesive containing theinsecticide before application.Lure and Kill as a Stand-Alone Control Method. In

evaluating the results of various lure and kill programs,several were initially considered to cause substantialreductions in the target pest population or damage.These included some testing paste/gel products asstand-alone treatments against pests at low pest den-sity (Suckling and Brockerhoff 1999, Charmillot et al.2000, Ebbinghaus et al. 2001, Ioriatti and Angeli 2002),although more detailed analysis was needed to con-Þrm the conclusions (see below). Lure and kill teststhat resulted in much less reduction of pest numbersor damage (Charmillot et al. 1996, Trematerra et al.1999, Angeli et al. 2000) were considered unlikely tobe promising and may indicate methods or situationsto avoid. Lure and kill programs that gave no evidenceof population or damage reduction (Moraal et al. 1993,Downham et al. 1995) were considered most likely toprovide information on the conditions under whichlure and kill should not be attempted; in these cases,the authors themselves concluded that lure and killwas not suitable for control of the target pest (Moraalet al. 1993, Downham et al. 1995), although later tech-nology improvements may change this view. In onecase, the authors also concluded that lure and kill wastoo expensive (Moraal et al. 1993); however, in regardto invasive species with high potential economic im-pact, the costÐbeneÞt equation may be quite different.Programs in which lure and kill alone was able tosubstantially reduce the target pest population wereconsidered of particular interest for assisting in erad-ication of invasive species; they also provided a betterindication of the effectiveness of the lure and killtechnology without complex interactions with othercontrol methods. A prime example is the applicationof “male annihilation” for tephritid fruit ßies (Cun-ningham 1989) that are considered quarantine pests inmany countries.

816 JOURNAL OF ECONOMIC ENTOMOLOGY Vol. 102, no. 3

Lure and Kill in Combination with Other Ap-proaches. There are few cases in which lure and killtechnology has been used in combination with generalapplication of insecticides (Hofer and Angst 1995,Antilla et al. 1996, Ioriatti and Angeli 2002). In twocases (Hofer and Angst 1995, Antilla et al. 1996), therewere major beneÞts from the inclusion of lure and killdue to major damage reduction and/or reduced levelsof insecticide sprays. In the third case, the use ofinsecticides was a confounding inßuence that com-plicated the interpretation of trial results (Ioriatti andAngeli 2002).Comparison of Lure and Kill with Other ControlMethods. Some authors compared lure and kill di-rectly in their trials with other control methods, orthey discussed such comparisons. For example, Char-millot et al. (2000) evaluated the efÞcacy of both masstrapping and lure and kill against codling moth andconcluded that lure and kill was more effective, a viewalso held for other pests based on likely efÞciency ofmale removal compared with traps (Suckling andBrockerhoff 1999). Lure and kill also was consideredas effective as conventional applications of insecti-cides, or better, against cotton bollworm (Hofer andAngst 1995) and codling moth (Olszak and Pluciennik1999). However, most comments were reserved forcomparing lure and kill with mating disruption. It wasespecially noted that lure and kill was more effectivethan mating disruption on small, hilly sites (Hofer andAngst 1995, Ioriatti and Angeli 2002, Charmillot et al.2000) and was less sensitive than mating disruption toproblems caused by the shape and size of treatedareas, by higher population densities, by the need forisolation, and by environmental factors such as wind(Charmillot et al. 2000). Some concluded simply thatlure and kill was more effective, especially on high-density populations (Hofer and Angst 1995), and mat-ing disruption was phased out of some trials (Antillaet al. 1996) in favor of lure and kill. The high cost ofpheromone for mating disruption was also a concern(Ioriatti and Angeli 2002). All these factors are im-portant in deciding the potential added value that lureand kill could bring to pest management and eradi-cation of invasive species, compared with mating dis-ruption or other approaches using semiochemicals.

Use of Lure and Kill in Long-Term PestManagement

Most of the lure and kill publications in the litera-ture refer to the use of this technology as part of pestmanagement programs. Although there were a fewcases of lure and kill being used in combination withother treatments, notably insecticides, it was normallya stand-alone treatment. Both situations are of interestfor determining its potential for long-term manage-ment. The use of lure and kill has become an integralpart of an areawide integrated pest management(IPM) approach to long-term management of fruitßies by using semiochemical-based “bait sprays” andmale annihilation (Vargas et al. 2003a,b; also see Te-

phritid Fruit Flies). Many of the examples in the casestudies that follow are summarized in Table 1.Case Studies. Pink Bollworm. The pink bollworm,Pectinophora gossypiella (Saunders), is a major eco-nomic pest of cotton, Gossypium hirsutum L., in allcotton-growing regions around the world. P. gossyp-iella was the Þrst species of Lepidoptera to be inves-tigated for the suitability of lure and kill to managethese pests (Hummel et al. 1973). Gossyplure [1:1mixture of (Z,Z)-7,11-hexadecadienyl acetate and(Z,E)-7,11-hexadecadienyl acetate] is the pink boll-worm sex pheromone that has been commerciallyused to control this moth since 1977 by mating dis-ruption (Gaston et al. 1977). The success of matingdisruption against pink bollworm encouraged re-searchers to investigate the addition of small amountof insecticides to kill male pink bollworm moths at-tracted to and contacting the pheromone sources(Butler and Las 1983, Beasley and Henneberry 1984).The Þrst trials investigating the potential of lure andkill approach against pink bollworm were conductedin California and Arizona in the early 1980s (Butlerand Las 1983, Beasley and Henneberry 1984). In thesetrials, gossyplurewasappliedaeriallyeither asNoMateÞbers or disruptant ßakes at a rate of 1.85Ð2.77 g/hawith and without permethrin added. Assessment wascarried out by monitoring male moth trap catch, andby crop damage (counting pink bollworm-infestedßowers and infested bolls). Both NoMate Þbers anddisruptant ßakes without the insecticide were highlyeffective in reducing male moth catches. However,the addition of permethrin to the disruptant ßakessigniÞcantly improved the effectiveness of control.The results of these trials indicated that a lure and killapproachwas feasiblewithpinkbollworm. In1987, theÞrst patent that describes a device for lure and kill(Ecogen hollow Þber) was approved in the UnitedStates for management of pink bollworm (Conlee andStaten 1986). Subsequently, an areawide trial for con-trol of pink bollworm program was conducted inParker Valley, AZ, over 6 yr by using the EcogenNoMate Þbers combined with permethrin. Lure andkill densities varied, and the gossyplure dose rangedfrom 25 to 35 g/ha, with two to four applications peryear. Assessment was achieved by larval counts inbolls. The authors concluded that the program pro-vided excellent control of pink bollworm, whichgreatly reduced the damage and the need for sprays,and with very low residual populations (Beasley andHenneberry 1984).

Another trial investigating the potential of lure andkill approach for management of pink bollworm wasconducted in Egypt in 1990 for two consecutive years(Hofer and Angst 1995). An isolated cotton Þeld of 14ha was treated with 7,000Ð8,000 lure and kill dropletsof 50 �l each (i.e., 0.6 g of gossyplure and 25.5 g ofcypermethrin per ha) that was applied four times perseason. Assessment was achieved by larval counts inthe bolls and crop yields. They concluded that theprogram resulted in reducing larval infestation andimproving yields.

June 2009 EL-SAYED ET AL.: “LURE AND KILL” WITH SEMIOCHEMICALS 817

Tab

le1

.L

ure

and

kill:

anal

ysis

offie

ldtr

ials

Tar

get

Refe

ren

ceL

ure

Lure

den

sity

Lure

dosa

ge;

inse

ctic

ide

dosa

ge

Tri

alperi

od

Resu

ltan

dco

ncl

usi

on

C.pomonella

Hofe

ran

dB

rass

el

(199

2)A

ttra

ctan

dK

ill

6Ð8

�10

0�

ldro

ps

per

tree

0.8

g/h

a;33

gfu

rath

ioca

rb/h

a/ap

plica

tion

1yr

Dam

age

very

low

but

sign

iÞca

nce

un

kn

ow

n;

“con

trol”

was

asi

ngle

hig

hly

infe

sted

tree

near

by

Iori

atti

and

An

geli

(200

2)Sir

en

eC

M1,

100Ð

2,00

0�

57m

gdro

ps

per

ha

0.10

Ð0.1

8g/h

a;4�

7g

perm

eth

rin

2yr

Con

trol

seem

ed

sim

ilar

toco

nven

tion

alin

sect

icid

es

but

tria

lsco

nfo

un

ded

by

imm

igra

tion

,pse

udore

plica

tion

,an

dsp

ray

inte

rven

tion

sE

bbin

gh

aus

et

al.(2

001)

Appeal

2,4,

6,00

0�

100

�l

dro

ps

per

ha

Not

stat

ed

2yr

Con

trol

at4Ð

6000

dro

ps

per

ha

equiv

alen

tto

IGR

spra

yin

g;very

low

resi

dual

pop

An

geli

et

al.(2

000)

Sir

en

eC

M1,

200Ð

2,00

0�

50�

ldro

ps

per

ha

0.08

Ð0.2

0g/h

a;3Ð

7.5

gperm

eth

rin

/ha

3yr

Con

trol

equiv

alen

tto

con

ven

tion

alin

sect

icid

eon

isola

ted

orc

har

dw

ith

low

den

sity

;le

sseff

ect

ive

ath

igh

er

den

sity

;re

sidual

pop

rem

ain

ed

Ch

arm

illo

tet

al.(1

996,

2000

)Sir

en

eC

M10

40Ð4

140

�50

or

100

�l

dro

ps

per

ha

0.08

Ð0.4

3g/h

a3Ð

16.2

gperm

eth

rin

/ha

3yr

Good

contr

olw

her

elo

win

itia

lpes

tden

sity

Ñat

end,ve

rylo

wre

sidual

pop;poor

contr

olw

her

ehig

hin

itia

lpes

tden

sity

Lose

let

al.(2

000)

Cas

tor

oil

bas

ed

(Appeal

?)7,

500

�10

0�

ldro

ps

per

ha

0.75

g30

gcy

�uth

rin

/h

a1

yr

Con

trol

equiv

alen

tto

IGR

inse

ctic

ides

and

very

low

resi

dual

pop;

use

dlo

wden

sity

CM

site

s;lu

redro

pden

sity

�5

per

tree

Ols

zak

and

Plu

cien

nik

(199

9)A

ppeal

1Ð3

dro

ps

per

tree

Not

stat

ed

1yr

Con

trol

equal

toor

bett

er

than

triß

um

uro

n;ta

rget

CM

den

sity

(un

treat

ed)

cause

ddam

age

of

5Ð6%

Paranthrenetabaniformis

Mora

alet

al.(1

993)

Sti

cky

ribbed

tube

�m

odel

fem

ale

30ri

bbed

tubes

per

ha

Cyperm

eth

rin

1yr

Fai

led

tore

duce

dam

age;

pro

bab

lydue

toim

mig

rati

on

of

mat

ed

fem

ales

on

tosm

all

plo

tsfr

om

old

heav

ily

infe

sted

trees

Con

tin

ued

on

follow

ing

pag

e


Tab

le1

.C

onti

nued

Tar

get

Refe

ren

ceL

ure

Lure

den

sity

Lure

dosa

ge;

inse

ctic

ide

dosa

ge

Tri

alperi

od

Resu

ltan

dco

ncl

usi

on

P.gossypiella

An

tilla

et

al.(1

996)

NoM

ate

Þber

�perm

eth

rin

Var

ious

25Ð3

7g/h

a6

yr

Inse

ctic

ides

overs

pra

yed;

exc

ellen

tco

ntr

ol

wit

hgre

atly

reduce

dsp

rays

and

dam

age,an

dvery

low

resi

dual

pop

Hofe

ran

dB

rass

el

(199

2)A

ttra

ctan

dK

ill

5Ð60

0?�

50�

ldro

ps

per

ha

0.6

g25

.5g

cyperm

eth

rin

1yr

Dam

age

on

eth

ird

of

“con

trol”

plo

t(c

on

ven

tion

ally

gro

wn

cott

on

)H

ofe

ran

dA

ngst

(199

5)Sir

en

e5,

000

�50

�l

dro

ps

per

ha

0.4

g15

.8g

cyperm

eth

rin

2yr

Reduce

dla

rval

infe

stat

ion

allo

wed

dela

yed

and

reduce

din

sect

icid

e;

com

merc

ialize

d19

94;

bett

er

than

inse

ctic

ide

pro

gra

mm

e(i

ncl

udin

gyie

ld)

S.littoralis

Dow

nh

amet

al.(1

995)

MC

(Mic

ro-e

nca

psu

late

d)

or

PV

C50

0sp

ray

poin

tsor

PV

Clu

res

per

ha

MC

0.5Ð

2.5

g0.

95Ð1

.9g

�-c

yh

aloth

rin

PV

C0.

5g

1.45

g�-

cyh

aloth

rin

3yr

Bri

efsu

ppre

ssio

nof

mat

ing

but

no

reduct

ion

of

egg

mas

ses;

not

avi

able

contr

olte

chniq

ue

inth

ese

tria

ls;hig

hden

sity

targ

etpes

t;par

tial

mat

ing

dis

rupti

on

achie

ved

rath

erth

an“lu

rean

dkill”

E.kuehniella

Tre

mat

err

a(1

995)

Lam

inar

dis

pen

ser

plu

ssi

gn

stim

ulu

sO

ne

dis

pen

ser

every

220Ð

280

m3

2m

gof

sex

ph

ero

mon

eplu

s5

mg

of

cyperm

eth

rin

2Ð3

yr

Num

ber

of

moth

sca

ptu

red

intr

eat

ed

mill

was

sign

iÞca

ntl

ylo

wer

than

con

trol

mill;

itw

asn

ot

poss

ible

toelim

inat

eal

lin

fest

atio

nRhyacioniabuoliana

Sukovat

aet

al.(2

004)

Rh

vkil

(ric

inole

icac

idplu

spetr

ole

um

oil)

1,00

0Ð2,

000

dro

ple

tsper

ha

Eac

hdro

ple

tsis

0.05

g,

con

tain

s0.

25%

sex

ph

ero

mon

ean

d2%

perm

eth

rin

3yr

No

sign

iÞca

nce

dif

fere

nce

intr

apca

tch

betw

een

treat

ed

and

con

trol

plo

ts;h

ow

ever,

shoot

dam

age

was

reduce

dsi

gn

iÞca

ntl

yin

treat

ed

plo

tRhagoletispomonella

Bost

ania

nan

dR

acett

e(2

001)

Red

sph

ere

s,yellow

boar

ds

wit

hbuty

lh

exa

noat

e

Every

2Ð3

mal

on

gorc

har

dperi

ph

ery

6Ð12

%cy

perm

eth

rin

or

1.3Ð

1.7%

delt

ameth

rin

5yr

Apple

mag

got

acti

vit

yle

ssin

Quebec

solu

rean

dkill

asu

ccess

ful

alte

rnat

ive

toco

nven

tion

alin

sect

icid

euse

Con

tin

ued

on

follow

ing

pag

e


Tab

le1

.C

onti

nued

Tar

get

Refe

ren

ceL

ure

Lure

den

sity

Lure

dosa

ge;

inse

ctic

ide

dosa

ge

Tri

alperi

od

Resu

ltan

dco

ncl

usi

on

Glossina

spp.

Est

erh

uiz

en

et

al.(2

006)

Blu

ean

dbla

ckcl

oth

wit

hac

eto

ne,1-

oct

en

-3-o

l,an

d4-

meth

ylp

hen

ol

8Ð12

per

km

2E

ach

stat

ion

rele

ased

350

mg/

hac

etone,

5.7

mg/

h1-

oce

ten-3

-ol,

15.5

mg/

h4-

met

hyl

-phen

ol;

cloth

dip

ped

in0.

8%del

tam

ethri

n

1Ð2

yr

99%

reduct

ion

infe

mal

es

ofG.austeni

but

on

ly85

%re

duct

ion

inG.

brevipalpis

due

toit

sh

igh

er

mobilit

y

S.calcitrans

Meif

ert

et

al.(1

978)

William

str

aps

pla

ced

betw

een

poult

rym

anure

and

feedin

gca

ttle

1st

atio

nper

5ca

ttle

(5st

atio

ns

10m

apar

t)L

ive

catt

leodors

;2.

5g

perm

eth

rin

/m2

8d

Up

to90

%re

duct

ion

inst

able

ßy

pop

afte

r8

dof

treat

men

ts,ab

out

30%

of

pop

per

day

rem

oved

by

stat

ion

sM.domestica

Ch

apm

anet

al.(1

998b

)W

hit

eboar

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ith

(Z)-

9-tr

icose

ne

mix

ed

wit

hsu

gar

2.5

gof

40%

(Z)-

9-tr

icose

ne/b

ait;

org

anoph

osp

hat

eal

facr

on

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azam

eth

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stat

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not

est

imat

ed

but

con

cluded

that

lure

and

kill

wit

hfe

mal

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yph

ero

mon

eis

pro

mis

ing

D.frontalis

Couls

on

et

al.(1

973)

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nta

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induce

dn

atura

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mon

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dpin

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ee

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6per

tree

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gfr

on

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rein

pla

stic

via

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ps;

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acid

(syst

em

icar

sen

ate)

4m

oIn

sect

icid

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reat

ed

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had

sign

iÞca

ntl

yle

ssbro

od

Dendroctonusbrevicomis

Hal

let

al.(1

982)

exo-

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vic

om

in,fr

on

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)42

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ide

treat

ed

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2m

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mpoun

dper

tree;

1Ð4%

chlo

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ph

os

and

1Ð4%

carb

aryl

topoin

tof

run

off

1yr

4%C

hlo

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os

and

4%ca

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yl

were

eff

ect

ive

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tect

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pon

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Dendroctonusvalens

Hal

l(1

984)

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theti

cE

FM

induce

dat

tack

s(p

itch

)ofD.

brevicomis

attr

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toD.valens

795

attr

acti

ve

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treat

ed

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sect

icid

es

2m

g/d

EF

M(a

sab

ove);

1Ð4%

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rpyri

ph

os

and

carb

aryl

asab

ove,

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itro

thio

n,

0.1Ð

0.4%

perm

eth

rin

3yr

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ly4%

carb

aryl

or

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nit

roth

ion

cause

dsi

gn

iÞca

nt

reduct

ion

inat

tack

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plica

tion

sin

July

more

eff

ect

ive

than

inSept.

;tr

eat

men

tsth

atpro

tect

ed

pin

es

from

D.brevicomis

did

not

work

agai

nstD.valens

Ipstypographus

Dedek

et

al.(1

988)

Syn

theti

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gre

gat

ion

ph

ero

mon

e(cis

-verb

en

ol

and

meth

ylb

ute

nol)

2per

gro

up

of

4Ð6

trees,

15gro

ups

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mg/dcis-

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en

ol

and

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g/d

meth

ylb

ute

nol)

;2-

mm

-th

ick

by

20-c

m-

wid

eri

ng

of

pas

te(1

5%m

eth

amid

oph

os)

on

trun

k(1

g/c

mtr

un

kdia

m)

3m

oB

ark

bee

tle

atta

cked

trea

ted

tree

seq

ual

lyto

contr

ols

,in

dic

atin

gno

repel

lent

action

of

inse

ctic

ide;

no

bro

od

pro

duct

ion

inin

sect

icid

e-tr

eate

dtr

ees;

trea

tmen

tof

tree

saf

ter

win

dfa

llsal

soef

fect

ive

Con

tin

ued

on

follow

ing

pag

e


Tab

le1

.C

onti

nued

Tar

get

Refe

ren

ceL

ure

Lure

den

sity

Lure

dosa

ge;

inse

ctic

ide

dosa

ge

Tri

alperi

od

Resu

ltan

dco

ncl

usi

on

Dru

mon

tet

al.(1

992)

Ph

ero

pra

xlu

re(cis

-verb

en

ol

and

meth

ylb

ute

nol)

1per

tree,80

trees

amon

g16

site

sA

bout

asab

ove;25

gof

cyh

aloth

rin

/lit

er

(12

ml/

lite

rw

ater/

m2

bar

k)

3m

oT

rap

trees

caugh

t4,

600Ð

16,8

00beetl

es

or

2.3Ð

12.7

tim

es

atr

ap,

equiv

alen

tto

bro

od

from

1co

lon

ized

tree

S.multistriatus

Lan

ier

and

Jon

es

(198

5)M

ult

ilure

(dai

ly:25

�g

of

4-m

eth

yl-

3-h

epta

nol,

6�

gof

�-m

ult

istr

itin

,50

�g

of

�-c

ubeben

e)

14bai

ted

trees,

2co

ntr

ol

1M

ult

ilure

per

tree;

27.4

%ca

codylic

acid

toru

noff

inax

efr

ill

on

trun

k,so

me

caco

dylic-

treat

ed

spra

yed

wit

h0.

5%ch

lorp

yri

fos

5m

oL

ure

attr

acte

dbeetl

es

totr

eat

ed

and

un

treat

ed

trees

but

more

attr

acte

dto

treat

ed;n

o.

of

beetl

es

kille

dgre

atly

incr

eas

ed

by

caco

dylic-

treat

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and

inse

ctic

ide

spra

yed

wit

hch

lorp

yri

fos

P.scarabaeoides

Gon

zale

san

dC

ampos

(199

5)2-

(Chlo

roet

hyl)

phosp

honic

acid

treat

men

tpro

duci

ng

attr

acti

ve

eth

yle

ne

4Ð7

bai

ted

bar

rier

trees

next

toin

sect

icid

e-t

reat

ed

trees

12%

Eth

rel;

2.5%

(wt:

vol)

lam

bda

cyal

oth

rin

e-A

at5.

7lite

rsper

tree

3m

oIn

sect

den

sity

intr

eat

ed

plo

tsw

as11

Ð13%

that

of

con

trol

plo

tsaf

ter

treat

men

tbut

incr

eas

ed

to40

Ð60%

aten

dof

exp

3m

ola

ter

Bactroceraspp.

Cun

nin

gh

aman

dSuda

(198

6)M

eth

yle

ugen

ol

6st

atio

ns

(1st

atio

nper

10h

a)U

nsp

eci

Þed

amt

of

meth

yle

ugen

ol

�5%

nal

ed;25

gof

25%

mal

ath

ion

per

stat

ion

4m

o�

99%

reduct

ion

of

mal

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ies,

but

on

ly48

%re

duct

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uit

infe

stat

ion

,pro

bab

lydue

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mig

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on

of

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es

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mat

ed

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hfe

mal

es

wh

ow

ere

not

caugh

tB.dorsalis

Ste

iner

et

al.(1

965)

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yl

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sqm

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aled

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aria

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Bactroceraoleae

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um

aset

al.(2

002)

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mon

ium

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ssy

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deca

ne)

15,0

00Ð2

0,00

0tr

aps

per

300

ha

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ameth

rin

15m

g(A

I)/t

rap

4yr

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er

trap

captu

res

and

fruit

infe

stat

ion

levels

com

par

ed

wit

hst

andar

dbai

tsp

rays

on

lyin

the

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B.cucurbitae

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nin

gh

aman

dSte

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(197

2)M

eth

yl

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plu

sn

aled

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ns

per

2m

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luti

on

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nal

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ßy

pop,lo

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resi

dual

life

of

lure

-pois

on

Ceratitscapitata

Nav

arro

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et

al.(2

008)

Bio

lure

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PA

lure

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re10

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over

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s


Egyptian Cotton Leafworm. The Egyptian cottonleafworm, Spodoptera littoralis (Boisduval), is consid-ered a key pest of cotton in the Nile delta of Egypt.Trials examining the potential of lure and kill approachto control S. littoraliswere investigated over 3 yr in thelate 1980s in upper Egypt (McVeigh and Bettany 1986,Downham et al. 1995). In these trials, 1 mg per lure ofa binary mixture of the two pheromone compounds,(Z,E)-9,11-tetradecadienyl acetate and (E,E)-10,12-tetradecadienyl acetate, in a ratio 99:1, respectively,was used to lure male S. littoralis. Lure and kill wasapplied using two different formulations: sprayablemicroencapsulated formulation containing a mixtureof sex pheromone and �-cyhalothrin, or polyvinyl-chloride pheromone formulation stapled to the un-derside of cotton leaves with �-cyhalothrin appliedover it. Both formulations were distributed to give adensity of 500 point sources per ha. Assessments of theeffectiveness of lure and kill approach were madeusing tethered females, male catch, and egg masscounts. Although lure and kill caused a signiÞcantreduction in mating of the tethered females and trapcatch, this was not corroborated with a reduction inegg masses compared with control plots. The results ofthese trials indicate that the efÞcacy of lure and killseemed to be poor and short-lived for controllingEgyptian cotton leafworm. In the same trial, lure andkill was compared with mating disruption (also 500point sources per ha), and similar degrees of matingsuppression were obtained with both methods. There-fore, it was concluded that the mating suppressionobserved with lure and kill was due to disruption ofmating communication rather than lethal contactswith insecticide incorporated into the pheromonesources. The failure of these preliminary lure and killtrials to provide adequate control of S. littoralis couldbe attributed to many factors, including a suboptimalpheromone blend used to attract males because fe-males produce other minor compounds (Campion etal. 1980) that were not included in the lure. Therelease rate of the pheromone from the point sourceswas not measured in these trials, and it is known thatoptimum release rate can be very important to achievefalse trail following to the point of contact with thesource (i.e., effective attracticide). Also, the density ofthe attracticide sources employed in these trials couldhave been below that needed to provide adequatecontacts with lure and kill formulations. Other factorsthat might have contributed to the failure of thesetrials could be the high density of the target pestand/or the partial but incomplete mating disruptionachieved by pheromone sources rather than lure andkill. Since these preliminary trials, no attempts havebeen made to improve the efÞcacy of lure and killapproaches to control S. littoralis. In spite of thesedisappointing results, the potential of the lure and killapproach against this important pest has not been fullyexplored as yet, because much basic information in-cluding blend composition, optimum release rate, op-timum lure density, formulation persistence, mortalityfrom insecticide, and sublethal effects on behavior andreproduction, is still lacking.

Codling Moth. The success of lure and kill againstseveral lepidopterous pests encouraged researchers totry this approach against the codling moth, Cydiapomonella (L.) (Charmillot et al. 2000, Ebbinghaus etal. 2001). In general, most of the lure and kill trials gavecontrol that was similar to/or better than treatmentwith insecticides or insect growth regulators (IGRs)(Olszak and Pluciennik 1999, Angeli et al. 2000, Loselet al. 2000), whereas in a few trials lure and kill did notprovided an acceptable control level (Hofer and Bras-sel 1992). In all these trials, only the primary sexpheromone compound of codling moth [(E,E)-8,10-dodecadienol] was combined with various insecti-cides (i.e., furathiocarb, permethrin, cypermethrin,and cyßuthrin) in various formulations (Charmillot etal. 2000, Ebbinghaus et al. 2001).

In a 3-yr trial conducted in an apple (Malus spp.)orchard in Switzerland between 1995 and 1997 (Char-millot et al. 1996, 2000), a gel formulation that con-tained 0.16% codlemone and 6% permethrin was ap-plied at densities of 1,000Ð4,000 droplets per ha, whichcorresponded to 0.08Ð0.43 g of codlemone and 3Ð16.2g of permethrin per ha. The droplets were applied twotimes per season with 5Ð7-wk intervals, and successwas assessed by female mating and counting fruit dam-age caused by codling moth. This trial achieved goodcontrol in areas where low initial pest density wasrecorded, and poor control in a plot with high initialpest density. A small lure and kill trial using a similarformulation was conducted in the southern Okanaganvalley, British Columbia, to investigate the response ofmale codling moth to lure and kill formulations(Krupke et al. 2002). The number of droplets perhectare is an important factor that determined thesuccess of this technology against codling moth inthese trials because a higher droplet density resultedin a higher frequency of males contacting the lure andkill formulation. However, another important factor isa high ratio of the lure and kill droplets to callingfemales so that males are more likely to contact thedroplets than females. It was found that male codlingmoth exhibited autotomy of thoracic legs when ex-posed to sublethal doses of insecticides which im-paired their ability to mate. However, some male cod-ling moths were caught in traps baited with callingfemales, indicating that these males were able to lo-cate and ßy toward calling females in the treated plot.This might be due to either males locating a callingfemale due to suboptimal attractiveness of the drop-lets under Þeld conditions, or that males ßew to thelure and kill droplets but they did not actually contactthe droplets.

Another trial that used a caster oil-based formula-tion containing 0.1% codlemone and 4% cyßuthrin wasconducted in apple orchards in Poland between 1998and 1999 (Ebbinghaus et al. 2001). The lure and killdroplets were applied as 100-�l droplets at densities of2,000, 4,000, and 6000 droplets per ha. The dropletswere applied two times per season with 6-wk intervals,and the success of the trial was assessed by countingfruit damage caused by codling moth larvae. It wasconcluded that the lure and kill approach provided a


good control at densities of 4,000Ð6,000 droplets perha, which was equivalent to control by spraying withIGRs (Ebbinghaus et al. 2001). In a subsequent studyby the same group, it was determined that the spatialdistribution of the lure and kill formulation was animportant factor for effective control, and the verticalposition of the droplets was more important than thehorizontal position because male codling moths arepredominately active in upper parts of the treecrowns. Environmental degradation of the formula-tion also can lead to reduction of the attractiveness ofthe pheromone and the knockdown effect of insecti-cides. The population density of codling moth also isan important factor in determining the success of thelure and kill against codling moth, and the approach ismore effective at low population density than at highpopulation density. The population density interactswith the density of the droplets per ha in determiningthe efÞcacy of lure and kill against codling moth, withmore than three droplets/tree required for efÞcientcontrol.Stored-Product Moths. Lure and kill approaches

have been tested for control of stored-products pests,mainly the Mediterranean ßour moth, Ephestia kueh-niella (Zeller), and the Indianmeal moth, Plodia in-terpunctella (Hubner), in ßour mills and warehouses.Trematerra and Capizzi (1991) investigated the po-tential of the lure and kill approach against E. kueh-niella. They used laminar pheromone dispensersbaited with 2 mg of the sex pheromone [(Z,E)-9,12-tetradecadienyl acetate and (Z,E)-9,12-tetradecadi-enol in 6:1 ratio] and 5 mg of cypermethrin at a densityof one dispenser every 220Ð280 m3. Under these cir-cumstances lure and kill was effective in maintainingthe population level of E. kuehniella in the ßour millbelow the economic threshold. Trematerra andCapizzi (1991) demonstrated the importance of visualstimuli in increasing the efÞcacy of the lure and killformulation against E. kuehniella when higher num-bers of males were attracted to silhouette subtriangu-lar forms resembling the female of this species. An-other trial investigating the potential of lure and kill tocontrol E. kuehniella was conducted in Italy for twoconsecutive years (1992Ð1993) in a 16,000-m3 ßourmill. In this trial, 5 mg of cypermethrin was applied tolaminated dispensers containing 2 mg of the sex pher-omone that released 13 �g of sex pheromone per day.Assessment was achieved by recording number ofadult males caught in pheromone baited funnel traps.The lure and kill formulations were combined with avisual stimulus to form a “sign stimulus.” A similar ßourmill in the same area was used as control. The presenceof lure and kill formulations led to a signiÞcant re-duction in the number of males caught in pheromone-baited traps throughout the mill and caused a signif-icant decrease in the E. kuehniella population. Theauthors highlighted the need to control outdoor pop-ulations to manage the risk of immigration and thusreinfestation.Arecent studyaimedat investigating thepotential of lure and kill to control a similar species,the Indianmeal moth, in small warehouse rooms in-

dicates that this approach is effective only when thepopulation level was low (Nansen and Phillips 2004).Apple Maggot. Lure and kill is a promising method

to control the apple maggot, Rhagoletis pomonella(Walsh), in eastern United States and Quebec, Can-ada. Fein et al. (1982) identiÞed apple volatiles at-tractive to apple maggot ßies (Reissig et al. 1982,1985). Yellow colors mimic apple foliage that is at-tractive to immature ßies, whereas red attracts sexu-ally mature ßies that mate and oviposit on matureapples (Bostanian and Racette 2001). Odor-baited redspheres (representing apples) coated with adhesivehave been used for apple maggot control in IPM(Prokopy et al. 1990). Red spheres mimicking appleswere coated with sucrose solutions or Þlled with su-crose and gelatinized corn ßour and insecticide thatkilled any attracted ßies (Hu et al. 2000). Yellowboards and red spheres were sprayed with cyper-methrin and deltamethrin in kerosene with butyl hex-anoate at 2Ð3-m intervals along the periphery of anapple orchard to reduce ßy infestations (Bostanianand Racette 2001). For the treatment to be consideredeffective with a high percentage of uninjured fruit, nomore than 13 ßies should be caught per four trapsbaited as above on the plot periphery, which was 1.6times the action threshold.Biting Flies. The treatment of large tracts of land

with insecticide to reduce insect vectors of disease hasmany potential problems. Among them are high costs,chemical resistance, nontarget insect mortality, and alack of public acceptance of widespread sprayings(Day and Sjogren 1994). Dipteran vectors such asmosquitoes and biting ßies are often attracted to car-bon dioxide, lactic acid, and octenol, associated withvertebrate hosts, as well as various ovipositional stim-ulants (DeFoliart and Morris 1967, Acree et al. 1968,Gillies 1980, Adeyeye and Butler 1991, Kline et al.1991).

The density of trapping stations is critical to suc-cessful control. Removal trapping of the tsetse ßy,Glossina spp., populations that vector trypanosomiasis(sleeping sickness) was done in West Africa in theearly 1940s (Morris and Morris 1949). It was observedthat baited traps could reduce local populations by upto 70% but had little effect on overall populations inthe region. Small numbers of baited traps (Þve toseven traps per acre) had little effect until the densityreached �20 traps per acre when successful controlwas achieved. Biconical traps impregnated with 400mg of deltamethrin and spaced at 300-m intervals overa large area caused a 96.6% reduction in Glossinapalpalis (Robineau-Desvoidy) populations and inter-rupted their breeding cycle (Kupper et al. 1985).However, the baited traps with insecticide were aheadof their time and phased out in the 1950s, being re-placed by larger scale insecticide treatments. In 1978,use of baited traps with insecticide was rediscovered(Vale et al. 1985, Laveissiere 1988, Day and Sjogren1994). Recently, Esterhuizen et al. (2006) treated a35-km2 area in Zululand, South Africa, with eight to 12lure and kill stations per km2. Cloth targets were madeof blue and black cloth dipped in 0.8% deltamethrin


and baited with acetone (350 mg/ha), 1-octene-3-ol(5.7 mg/h), and 4-methylphenol (15.5 mg/h). Theyreported a 99% reduction in G. austeni Newstead fe-males after 13 mo of treatment and up to 85% for G.brevipalpis Newstead. It was concluded that the G.brevipalpis were less affected by the program due totheir higher ßight mobility.

The stable ßy, Stomoxys calcitrans (L.), is easilydisturbed and ßies often between vertebrate hostswhile feeding, thus transmitting equine infectionssuch as anemia, anthrax, and trypanosomes (Day andSjogren 1994). The Williams trap (Williams 1973) wasused by Rugg (1982) to reduce populations of S. cal-citrans at the Taronga Zoo in Sydney, Australia, by 79%after only 7.5 d of trapping. When Williams trapscontained a pyrethroid compound, permethrin (2.5g/m2), and they were placed between the ßy source(poultry manure from a poultry house) and the “lure”of cattle feeding nearby, populations were reducedeven more, by up to 90% in a week (Meifert et al.1978). They calculated that 30% of the adults wereremoved each day when one station per Þve domesticanimals was used.

The house ßy,Musca domestica L., although it doesnot bite, has been involved in the spread of numerousdiseases including salmonella, diphtheria, tuberculo-sis, hepatitis, and amoebic dysentery (Hanley et al.2004). High populations of house ßy are associatedwith livestock feeding lots and garbage landÞll siteswhere up to 1,500 ßies can be produced per m2 oflandÞll waste. Commonly, synthetic insecticides aresprayed at the site to control house ßies, but this canpose a health risk and is not very effective due tomultiple insecticide resistance (Chapman et al. 1993).Target sites baited with house ßy sex pheromone (Z)-9-tricosene mixed with sugar and insecticide wereused to reduce house ßy outbreaks at poultry units(Chapman et al. 1998a, 1998b). The advantages arereduced exposure of humans to insecticide and lesspotential for insecticide resistance due to ingestioncompared with cuticular contact. In these studies,males are primarily attracted and killed, so improvedbaits for females would likely enhance the success ofsuch programs.Bark Beetles. “Trap trees” have been felled in Eu-

ropean forests for several centuries to concentrate theattracted bark beetles in an attempt to lower theirpopulations and spare desired standing trees from at-tack (Bakke and Riege 1982). In the wide sense, traptrees lure and kill beetles attracted to the natural odorsof the dying trees and pheromones of attacking insects(Byers 2004), until the trees are removed by the for-ester. Thus, it was a logical step to treat these trap treeswith insecticide or to bait a healthy tree with barkbeetle pheromone to create a trap tree that also couldbe treated with insecticide. In United States, Coulsonet al. (1973) used a synthetic aggregation pheromonemixture (Frontalure) to attract southern pine beetles,Dendroctonus frontalisZimmermann, tocacodylic acidpoisoned pines. Hall et al. (1982) sprayed 420 pon-derosa pines, Pinus ponderosa Dougl., with the insec-ticides carbaryl or chloropyrifos and then baited with

western pine beetle, Dendroctonus brevicomis Le-Conte, synthetic pheromone components. All but onetree received arriving beetles that bored through thebark, but most were eventually overcome by the in-secticide and the trees survived. However, more pred-ators, Temnochila chlorodia (Mannerheim), attractedto the lure were killed by the insecticides than inbaited control trees. In a similar test, Hall (1984) Þrstbaited ponderosa pines withD.brevicomispheromonecomponents to induce resinous attacks that thencaused attraction of the red turpentine beetle, Den-droctonus valens LeConte, but if these trees had beensprayed with several insecticides (carbaryl, chloropy-riphos, fenitrothion, and permethrin), the trees wereprotected from colonization by D. valens. Lanier andJones (1985) used trap trees of American elm baitedwith Scolytusmultistriatus (Marsham) synthetic pher-omone (Multilure) to attract these beetles that vectorDutch elm disease. Some of the trees were sprayedwith chlorpyrifos that killed arriving beetles and pre-vented the elms from being colonized.

In Europe, Norway spruce trees baited with syn-thetic pheromone components of the bark beetle Ipstypographus (L.) (cis-verbenol and 2-methyl-3-buten-2-ol) were treated with a band of 32P-labelled meth-amidophos insecticide that penetrated the ascendingsap as a systemic that did not prevent beetle entry ofthe tree but signiÞcantly inhibited brood production(Dedek and Pape 1988, Dedek et al. 1988). Trap treesbaited with I. typographus pheromone and treatedwith pyrethroid insecticide (cyhalothrin) killed at-tacking beetles of I. typographus (Drumont et al.1992). The pyrethroid insecticides also were recom-mended for control of the ambrosia beetle Trypoden-dron lineatum (Olivier) that enters the sapwood. Theyestimated that a lure on a trap tree attracts and kills upto 14 times more beetles than a lure in a trap alone, oralmost as many beetles as is produced by a colonizedtree (2,300Ð17,000 beetles). Olive trees when treatedby 2-(chloroethyl)phosphonic acid release a primaryattractant, ethylene, that attracts the olive beetle, Ph-loeotribus scarabaeoides (Bernard), and when com-bined with insecticide lambda cyalothrine-A sprays,caused a signiÞcant reduction in attack density andpopulation level in the treated area (Gonzalez andCampos 1995). Pena et al. (1998) also released ethyl-ene from dispensers placed on olive logs to attract P.scarabaeoides where they were killed by cyper-methrin, resulting in no colonization.

Use of Lure and Kill in Eradication of InvasiveSpecies

Despite the use of lure and kill in eradication effortswith orders such as Diptera and Coleoptera, no evi-dence could be found that lure and kill has been usedfor pest eradication or even as part of an eradicationprogram for any lepidopterous pest. This is probablybecause the technology is relatively new against mothpests. However, it should not be assumed that lure andkill is unsuitable for use in moth eradication. Thepractice of lure and kill has been used for many years


in eradication programs against other pests, the mostimportant being for containment or eradication offruit ßies. This programs demonstrate the validity ofthe application principles potentially to eradicate lep-idopteran pests.Case Studies. Tephritid Fruit Flies. Case studies on

the tephritid fruit ßies are an interesting contrast tothose reported for other species. Due to their statureas key regulatory pests and the inability to determineinfestation easily (larvae are internal feeders), te-phritid fruit ßies have garnered the attention of agri-cultural states such as California, Florida, and Texas inthe United States and countries such as New Zealandand Japan where the ßies are not established. Effortsto eradicate fruit ßies have largely overshadowedmethods for their control and only recently has IPMbeen developed to replace insecticide cover spraysused for decades to control these pests. Successfuleradication of fruit ßies has followed the developmentof attractants for use in detection and delimitation, aswell as the lure and kill concept discussed herein.Froggatt (1909) suggested that male fruit ßies could be“annihilated” by attraction to kerosene, whereas in1912 Howlett (1915) showed that citronella oil (whichcontains methyl eugenol), was attractive to tephritidßies, probably mainly Bactrocera dorsalis (Hendel),the oriental fruit ßy (Bateman et al. 1966a). In 1931,traps baited with kerosene were used in a program tocontrol fruit ßies, and in 1935 terpinyl acetate was usedto attempt control of another fruit ßy,Ceratitis(�Pter-andrus) rosa Karsch (Karsch), in South Africa. Thiswas followed by the use of proteinaceous food baits(Gow 1954) in the United States and other parts of theworld, which were the basis for the protein bait spraysdeveloped 50 yr later with GF-120 (Prokopy et al.2003, Mangan et al. 2006). Food odor attractants in-cluding proteinaceous “bait” have been used exten-sively for fruit ßy control (Jang and Light 1996). Var-ious members of the family Tephritidae, whichincludes the genera Bactrocera, Anastrepha, Rhagole-tis, andCeratitis, are attracted to odors from hyrolyzedprotein baits. The Mediterranean fruit ßy, Ceratitiscapitata (Wiedemann), was suppressed by hydro-lyzed protein baits mixed with malathion, phloxine B,or spinosad (Peck and McQuate 2000; Vargas et al.2003a, 2003b). Katsoyannos and Papadopoulos (2004)found that yellow plastic spheres baited with foodattractants ammonium acetate, putrescine, and trim-ethylamine captured C. capitata, in this case morefemales were attracted. In Pakistan, simple woodenblocks soaked with insecticide and lure attracted andkilled 4 times more male fruit ßies (B. dorsalis) thanany commercial traps, and which were also more ex-pensive (Stonehouse et al. 2002).

A unique lure and kill technology termed male an-nihilation or MAT was developed for fruit ßies in thegenus Bactrocera by using a natural product methyleugenol (4-allyl-1,2-diemethoxybenzenecarboxylate)(Cunningham and Suda 1986, Cunningham 1989).This chemical was so attractive to male fruit ßies thatit was used to eradicate new infestations in many partsof the world including Rota (oriental fruit ßy; Steiner

et al. 1965), Japan (melon ßy; Kuba et al. 1996), Aus-tralia (papaya fruit ßy, Bactrocera papayae Drew &Hancock; Cantrell et al. 2002) and on numerous oc-casions against oriental fruit ßy in California. Anothercompound called cuelure [4-(p-acetoxyphenyl)-2-butanone)], developed by Beroza et al. (1960) wasfound to be attractive to males of the Bactrocera com-plex as well but is not able to eradicate fruit ßies as astand-alone technology. Beroza et al. (1960) tried an-alogs of a known male lure, anisylacetone, and foundseveral para-substituted derivatives of 4-phenylbu-tanone were attractive to melon ßies, Bactrocera(�Dacus) cucurbitae (Coquillett). Cunningham andSteiner (1972) used cue-lure soaked on Þberboardblocks and released 9,000 male B. cucurbitae that hadbeen dyed different colors in the treated and un-treated areas. They found that 96% fewer ßies wererecaptured in the treated plot despite its having morebaited traps, whereas marked ßies were recaptured upto 3 mo after being released in the check plot only.Cue-lure was the most active compound for B. cucur-bitae and a range of other species in the genus and wasused with an insecticide to destroy �50% of maleQueensland fruit ßy, Bactrocera tryoni (Frogatt), in atown in Australia (Bateman et al. 1966a). However,they concluded that although the insecticide killed allßies that came to the lure, a stronger lure was needed,or a higher density of trap stations. In a larger test,protein baits with insecticide, attractive to both maleand female Queensland fruit ßy, were compared withmale lures with insecticide and the combination inseveral towns in Australia. Bateman et al. (1966b)found that sometimes the protein baits were effectivebut that combination baits (protein � insecticides)were the most effective in preventing fruit infestation.

Several attractants have been identiÞed to attractmale Mediterranean fruit ßies, including trimedlureand ceralure (Jang and Light 1996; Jang et al. 2001,2003, 2005). However, these have not been found to beattractive enough or tested sufÞciently as a standalonecontrol technology and currently must be used withother techniques such as the sterile insect technique.Navarro-Llopis et al. (2008) evaluated several trapÐfoodÐlure combinations for attract and kill of Medi-terranean fruit ßy and found speciÞc traps and lurespotentially useful for areawide control of this pest inSpain. California currently uses a combination of baitsprays, and sterile insects as the primary methods toeradicate introductions of Mediterranean fruit ßy(CDFA 1999).

One concern is that nontarget and beneÞcial insectscan be attracted to the baits and killed by insecticides.Michaud (2003) tested two fruit ßy bait/insecticides,Nu-Lure/malathion and GF-120/spinosad for theirtoxicity to coccinellid, lacewing, and ßower bug spe-cies. Coccinellids [Ollav-nigrum(Mulsant) andScym-nus spp.] and the insidious ßower bug,Orius insidiosus(Say), did not succumb to Nu-Lure/malathion. How-ever, Nu-Lure was attractive to some syrphid ßies thatwere then killed. Both Nu-Lure and GF-120 causedmortality of two parasitoid wasps. In another study,Uchida et al. (2003, 2007) baited traps with cue-lure


and methyl eugenol and found many insect specieswere attracted, although most of the attraction ofnontarget insects may have been to odors of decayinginsects in the traps.

Olive fruit ßies, Bactrocera oleae (Gmelin), are ofgreat economic importance in the Mediterranean re-gions of Europe, and as such, attempts to control oliveßy have been well reported in the literature. Likeother Bactrocera, early control measures used widespectrum organophosphate insecticides as coversprays, which in many cases resulted in serious effectson the nontarget fauna (Feron and DÕAquilar 1962).Later, attractive proteinaceous baits were developedwith insecticides as early “attract and kill” formula-tions (Orphanidis et al. 1958). The identiÞcation of theolive ßy pheromone led to tests of pheromone andfood attractant combinations with various toxicants(Broumas et al. 1985, Haniotakis et al. 1986) Olive ßywas controlled by Ecotraps (Rovesti 1997) that com-bine ammonia-releasing salts as food attractants witha sex pheromone (a spiroacetal) and deltamethrininsecticide at one trap per tree. Broumas et al. (1985)conducted a 4-yr study in which they compared theammonium carbonate/pheromone lure with delta-methrin on a 300-ha orchard and found both trapcapture and fruit infestation was lower compared withorchards in which a bait spray only was used. Anareawide pest management system was constructed inItaly that predicts whether an area is suitable for lureand kill (or mass trapping) if its active infestation doesnot exceed 30% on 80% of farms by the third week ofOctober (Petacchi et al. 2003).

Lureandkillmethodshavebeen testedonanumberof other fruit ßy species. The Mexican fruit ßy, Anas-trepha ludens (Loew), and the Caribbean fruit ßy,Anastrepha suspensa(Loew), are serious pests of citrusand other fruits and may invade southern Texas andoccasionally California and Florida (Nilakhe et al.1991, Epsky et al. 1993). McPhail traps baited withtorula yeast or other proteinaceous baits have beenused for the last century (Robacker and WarÞeld 1993,Thomas et al. 2001). However, during the last decades,synthetic food-odor lures such as trimethyl amine,ammonium acetate and putrescine have become moreprevalent because these components are more selec-tively attractive to the fruit ßies (Heath et al. 2004).McPhail and Multi-Lure traps catch more than“Mitchell” killing stations with these lures, but theMitchell station is less expensive per unit (Holler et al.2006). The Mitchell station uses permethrin insecti-cide that kills ßies �30 min after contact (Holler et al.2006).Boll Weevil. The boll weevil, Anthonomus grandis

grandis (Boheman), entered the United States fromMexico in 1892 and soon afterward caused seriouseconomic damage to cotton (Ridgway et al. 1990,Smith 1998). Insecticides such as DDT controlled theboll weevil from 1945 until it became resistant to allchlorinated hydrocarbon insecticides by 1960. Eradi-cation of the boll weevil from the southwest was ac-complishedbyacombinationofcultural control, pher-omone trapping, and insecticide application in

response to pheromone trap catches. Mass trappingboll weevils began in 1968 and indicated that low-density populations could be reduced further, but theprobability of success declined as population densityincreased (Hardee 1982, Ridgway et al. 1990). At thehighest density of traps (14 per ha), it was estimatedthat 92% of the nonoutbreak population of emergingweevils could be trapped. Mitchell et al. (1976) de-termined that baited pheromone traps at 10 traps peracre captured 76% of the overwintering weevils and�96% of the late-emerging population. Lloyd et al.(1981) reported that three to four traps per acre cap-tured 80Ð90% of the females. Knipling (1979), usingpopulation models with expected capture rates, sug-gested that populations could be suppressed to verylow levels with as few as four traps per acre. Thesestudies merely trapped weevils but demonstrated theeffectiveness of the lure. More recently, Villavaso et al.(1998) compared pheromone-baited traps to baitedsticks with adhesive to determine the relative attrac-tiveness. They found that three times more boll wee-vils contacted the bait sticks than the pheromonetraps. All weevils that contacted the sticks that hadinsecticide were killed. Thus, bait sticks with insecti-cide (malathion) should be about three times moreeffective than baited traps.

Essential Knowledge for Successful Lure and Kill

Lure Competitiveness with Natural Odor Source(Including Insecticide–Lure Interactions). Differentkinds of research and development have been under-taken to ensure that lure and kill formulations doindeed lure and kill the majority of adult insects of thetarget species (Brockerhoff and Suckling 1999, Loselet al. 2000, Poullot et al. 2001, Evenden and McLaugh-lin 2004). The addition of insecticide to the phero-mone adds a new layer of complexity to the manyfactors inßuencing efÞcacy. Key among these factorsare semiochemical blend, semiochemical dose (andassociated Þeld longevity, including possible blendchanges), lure formulation, lure density, insecticidechoice, and insecticide dose (and associated Þeld lon-gevity). Other factors that affect the success of lureand kill include lure/trap placement, lure/trap height,and lure/trap design/size. There are complex inter-actions between the pheromone/semiochemical, theinsecticide, and insect behavior which need to beunderstood if high kill rates are to be achieved. Thepresence of the insecticide or other formulation com-ponents (such as gel, oil, or adhesive) must not com-promise the ability of the semiochemicals to causeinsects to land and contact the toxic substrate. Forexample, the ability of a sex pheromone lure to com-pete with wild females is fundamental to the successof lure and kill technology in Lepidoptera (e.g.,Krupke et al. 2002). Lure and kill has been able toexploit existing knowledge of sex pheromone compo-sition or other behaviorally active semiochemicals ofthe target insects. Nevertheless, the pheromone isbeing used to inßuence male behavior, and research isnecessary in this context to conÞrm its greater attrac-


tiveness than virgin females and its ability to inducemales to contact the lures. Research needs to directlycompare lure attractiveness with virgin females tooptimize pheromone blend, release rate, and dose(Downham et al. 1995, Brockerhoff and Suckling 1999,Poullot et al. 2001), all of which may need reÞnements,particularly to increase male-lure contact (Downhamet al. 1995) and insecticidal efÞcacy (Brockerhoff andSuckling 1999, Losel et al. 2000). This kind of researchalso assists in determining the period of attractivenessand effectiveness of the lures and the required fre-quency of lure replacement, which may vary from amatter of days (Downham et al. 1995) to severalmonths depending on the formulation, pheromonecomposition, and insecticide used (Brockerhoff andSuckling 1999, Suckling and Brockerhoff 1999). BothÞeld and wind tunnel research has been valuable inthis regard (Poullot et al. 2001). The constant releaseof pheromone by the lures gives them an advantageover virgin females which “call” during restricted pe-riods of the day; this is particularly useful becausemany males are active earlier, both daily and season-ally, when the lures do not have to compete with wildfemales (Losel et al. 2000).Lure Density. At a given pest population density,

the efÞcacy of lure and kill increases as the number ofpoint sources per ha increases (Downham et al. 1995,Suckling and Brockerhoff 1999, Krupke et al. 2002);however, this is up to some upper limit. For examplewith sex pheromone, lure density per ha must besufÞcient to compete with the numbers of wild fe-males calling in the Þeld, and this is most readilyachieved where population densities are low (see be-low). Compared with many traps in mass trapping,high densities of lure and kill point sources can bedeployed more easily and with lower labor costs; trialsof paste or gel formulations have been used at up to7,500 droplets per ha (Losel et al. 2000) and the Eco-gen Nomate hollow Þbers (aerial applications) (An-tilla et al. 1996) seem to have been used at even higherdensities. However, care is required because higherdensities of pheromone point sources may carry agreater risk of interference between them (Sucklingand Brockerhoff 1999), causing some mating disrup-tion of the target pest in the case of sex pheromone(Downham et al. 1995), and failure to obtain sufÞcientlure and kill (Downham et al. 1995). In this context,understanding the behavior of insects close to andcontacting the lures is critical. It is possible that in-creasing the density of lure and kill dispensers can leadto an increase in pest immigration into the treatedarea. This will result in an increase in the populationdensity of the target pests and can lead to failure of thelure and kill program. Therefore, understanding thecorrelation between density of applications and insectimmigration is critical.Lure Formulation. Lure formulation has been

largely the preserve of commercial companies, butseveral programs have added their own design fea-tures, such as to improve contact between insect andlure/toxin (McVeigh and Bettany 1986), even to thepoint of adding visual female or ßower models to

induce landing behavior (Miller et al. 1990, Moraal etal. 1993). As with mass trapping, an understanding ofoptimum lure height and placement has contributedto efÞcacy of lure and kill. The insecticide used in lureand kill must be chosen carefully, particularly to avoidsuch problems as repellency that could prevent lurecontact. The preferred compounds are mainly fast-acting pyrethroids (e.g., cyßuthrin, �-cyhalothrin,cypermethrin, furathiocarb, and permethrin). Al-though pyrethroids are known to be repellent in somesituations and to some insects, lure and kill researchhas been able to develop formulations without thisproblem (De Souza et al. 1992, Haynes et al. 1996,Brockerhoff and Suckling 1999). Recent research hasfocused on obtaining a better understanding of theway insects obtain and respond to both lethal andsublethal doses of insecticide because this is funda-mental to the success of lure and kill. These studiesinclude the effects of insecticides on insect behavior,required time of contact, rapidity of kill, sublethaleffects on mating ability of insects, and the longevityof insecticide efÞcacy (De Souza et al. 1992, Hayneset al. 1996, Suckling and Brockerhoff 1999, Poullot etal. 2001). All these factors contribute to the selectionof insecticide and semiochemical dose to determinethe frequency of lure replacement (i.e., the number ofapplications required). Lure and kill formulationsprobably require longerperiodsofefÞcacy thanwouldbroadcast conventional insecticide spraying (in partbecause it prevents the next generation as the mech-anismofcontrol) andsomepyrethroids, suchascyper-methrin (Ioriatti and Angeli 2002), are sufÞcientlypersistent to offer this beneÞt.

Unlike mass trapping, sublethal effects of lure andkill can be an important component of efÞcacy be-cause the insecticidal contact may reduce the abilityof males to respond to and mate with females, even ifmales are not killed outright (Suckling and Brocker-hoff 1999). Insects that have acquired a sublethal dos-age may be more vulnerable to natural enemies. Al-though lower, sublethal dosages may be worrisomedue to risks of survivors developing resistance, if thedose is high enough to reduce the ability of theindividual to protect itself against natural enemies,then the risk of developing resistance may actuallybe very low.

Careful experimentation has been able to separatethe effects of lure competition and insecticidal com-ponents in lure and kill efÞcacy against some pests(Brockerhoff and Suckling 1999) and shown thatagainst other pests, the insecticidal component is thekey and not semiochemical disruption (Charmillot etal. 1996). Without this crucial insecticidal effect, lureand kill becomes less effective and no more thanpartial mating disruption (Downham et al. 1995).Suckling and Brockerhoff (1999) used exclusion cagesplaced over lure and kill droplets to show that 500point sources of pheromone reduced trap catch in theplots by point source competition, but the catch wasreduced about half as much when the droplets wereexposed, thereby allowing mortality from contact withthe droplets.


Population Density of Target Pest and Risk of Im-migration. There are some indications that lure andkill is more effective than mass trapping or matingdisruption when attempting control at higher popu-lation densities, for example, initial trials against somehigher density pest species has shown promise (Suck-ling and Brockerhoff 1999). However, the majority oftrials conÞrm the critical importance of populationdensity (Downham et al. 1995, Angeli et al. 2000, Loselet al. 2000, Ebbinghaus et al. 2001, Krupke et al. 2002)and the inverse density dependence of lure and kill(Krupke et al. 2002). There are frequent examples oflure and kill failing to control pest populations at highdensity (e.g., Trematerra et al. 1999, Angeli et al. 2000,Charmillot et al. 2000). However, lure and kill is assusceptible as mass trapping and mating disruption tothe negative inßuence of immigration, and the liter-ature regularly identiÞes the need to use lure and killfor the control of isolated smaller populations (Moraalet al. 1993, Downham et al. 1995, Angeli et al. 2000,Charmillot et al. 2000). This will not necessarily over-come the problem of attempting to control at high pestdensity (Trematerra et al. 1999), although a higherdensity of lures may help compensate in controllinghigher densities. The potential for lure and kill tech-nology in the eradication of invasive species lies intreatment of “populations” that are initially at lowdensity, in complete coverage of the population basedon delimitation surveys, and when there is a low riskof immigration.Biology and Ecology of Target Species.With fewer

pest species having been targeted by lure and killcompared with mass trapping or mating disruption,there is less information in the literature on the in-ßuence of pest biology and ecology on efÞcacy. It isreasonable toassumethat the inßuenceswill be similarfor the three methods, and this is supported by limitedevidence to date. For example, several authors stressthe importance of initiating control early in the season(Hofer and Angst 1995) before the onset of male mothßight (Charmillot et al. 2000), and ensuring that lureand kill is maintained throughout each generation(Ebbinghaus et al. 2001). These recommendations areto exploit protandry and would beneÞt from univolt-inism. The risk to effective lure and kill posed byimmigration is aggravated by mobile adult males andfemales (Moraal et al. 1993). With tephritid fruit ßies,early studies on melon ßy ecology showed that melonßies preferred to “roost” on hedgerows and boardersoutside of melon and cucurbit Þelds (Nishida and Bess1957), and females moved into the Þeld primarily tolay eggs on host fruit. As a result recent use of thisinformation has been successfully implemented in anareawide melon ßy pest management program in Ha-waii where border foliage attractive to the ßies arepreferentially planted and bait sprays applied to theborders at regular interval resulting in population re-duction in the Þelds (Vargas et al. 2003a).Measuring Efficacy of Control. The efÞcacy of lure

and kill against moths is routinely measured by mon-itoring the males of the target pest with pheromonetraps to conÞrm population decline. This is usually

supported by records of pest damage or infestation(Moraal et al. 1993, Downham et al. 1995, Trematerraet al. 1999), which is preferable to merely monitoringpopulations of males but much more time consuming.These methods provide an overall assessment that canbe compared with untreated “control” plots but do notdistinguish the effects of lure and kill from other mor-tality factors affecting the population. In particular, itis important to determine the impact of lure and killon mating (Krupke et al. 2002). The proportion oftethered or caged virgin females which mate(McVeigh and Bettany 1986, Downham et al. 1995,Charmillot et al. 2000)hasbeenused toprovideamoredirect measure of lure and kill efÞcacy, and to guidethe choice of lure density (Charmillot et al. 1996). Thismethod assumes that the females used are equal inattractiveness to wild virgin moths. ModiÞcations ofthis approach include recording the proportion ofmonitoring traps (containing either caged females orpheromone) that fail to attract a male (Suckling andBrockerhoff 1999, Krupke et al. 2002), because thesetraps can be considered as equivalent to females thathave not mated. The mated status of males, which canbe determined in some species of Lepidoptera bydissection (Evenden et al. 2003), may be a usefultechnique for assessing the proportion of virgin malescaught in the monitoring traps. MarkÐrecapture ofmales is another technique that has been used in Þeldcages and in the Þeld to measure the efÞcacy of lureand kill and to assist with determining an appropriatelure density (Krupke et al. 2002, Brockerhoff andSuckling 1999).Lure and Kill: Risks.The insecticidal component of

lure and kill could be perceived by the public as a riskif this technology were to be used in urban environ-ments, such as in the eradication of invasive species.Concern over the use of the broad-spectrum organo-phosphate insecticide “naled” as the active ingredientin “minugel,” a lure and kill formulation applied wellabove ground level to telephone poles, against Bac-trocera fruit ßies in urban California is a case in point.For years, a mixture of methyl eugenol and naled 5%mixed with the thickening agent minugel has beenapplied to urban infestations of oriental fruit ßy. Whenapplied at a rate of 600 spots per sq mile onto tele-phone poles, this technique has historically been usedsuccessfully for eradication of small outbreaks in Cal-ifornia (CDFA 1993). However, increasing concernover organophosphate insecticides has spurned re-search into alternatives. These “perceived risks,” how-ever, must be weighed against the beneÞt, which inthis case can be avoiding signiÞcant reductions intrade, increased use of other insecticides or perma-nent establishment of alien invasive species. The pres-ence of numerous insecticidal point sources in anurban environment can be expected to elicit at leastsome public opposition, even though the candidateinsecticides have low mammalian toxicity, are used invery low quantities per ha compared with conven-tional spraying (Suckling and Brockerhoff 1999,Trematera et al. 1999, Losel et al. 2000), and mostformulations have a low risk of lethality to nontarget


organisms. In some cases, natural enemies can behighly attracted to semiochemicals used in lure andkill formulations, and therefore the effect of lure andkill on nontarget species has to carefully be investi-gated to minimize this undesirable effect on nontargetspecies. The aerial application of the hollow Þberformulation of lure and kill (Antilla et al. 1996) wouldbe totally unacceptable to the general public, partic-ularly in an urban environment. EfÞcacy of all the lureand kill formulations is closely dependent on the num-ber of point sources per ha, and, even with paste or gelformulations, optimal densities would likely be in1,000 sources rather than 100 sources per ha. In ad-dition, the concentration of insecticide in the lurescould be expected to be �5Ð6%, and the droplets ofsome formulations are susceptible to rain splash thatspreads them to some extent after application. If thedroplets are not totally inaccessible, children or otherscontacting them, although at minimal risk, may havean allergic response, or a belief that this is happening,which could create severe negative public relations.Risk of lure and kill failure could come from variousother aspects of methodology, such as an inadequateodor blend or insufÞcient lure density for the targetpopulation density. Many of these risks can be mini-mized by careful research and development. A modelof mass trapping using knowledge of the lureÕs effec-tive attraction radius can be used to investigate lureand kill efÞcacy in regard to lure density versus pestdensities (Byers 2007).

Overall Evaluation of Lure and Kill

Lure and Kill: Success and Failure. Analysis of themain lure and kill programs and the associated re-search and development provide a guide to the keyreasons for success and failure. There are several ex-amples provided in the literature of the successful useof lure and kill in pest control, i.e., it provided a majorreduction in pest population or damage. Examinationof these cases shows that the key reasons for successcan be summarized as follows: 1) low-density targetpopulation (Angeli et al. 2000, Charmillot et al. 2000,Losel et al. 2000, Ebbinghaus et al. 2001); 2) isolatedtarget population (no or minimal immigration)(Downham et al. 1995, Angeli et al. 2000, Charmillotet al. 2000); 3) a lure competitive with wild females(Angeli et al. 2000, Charmillot et al. 2000, Ebbinghauset al. 2001); 4) high lure density (relative to pestdensity) (Angeli et al. 2000, Charmillot et al. 2000,Losel et al. 2000); 5) optimal lure placement (Char-millot et al. 2000, Losel et al. 2000, Ebbinghaus et al.2001); and 6) lure deployment before male emergenceand throughout ßight period (Charmillot et al. 2000,Ebbinghaus et al. 2001).

However, examination of the cases in which lureand kill failed to provide a major reduction in pestpopulation or damage indicate key reasons for failureof lure and kill which can be summarized as follows:1) too high density of target population (Downham etal. 1995, Trematerra et al. 1999, Angeli et al. 2000,Charmillot et al. 2000); 2) target population not iso-

lated (high risk of immigration, high pest mobility)(Moraal et al. 1993, Trematerra et al. 1999); 3) inad-equate pheromone lure that was not competitive withfemales (Downham et al. 1995); 4) insufÞcient densityof point sources in relation to target pest density(Moraal et al. 1993, Downham et al. 1995, Trematerraet al. 1999); and 5) lure and kill formulations appliedafter damage done (Charmillot et al. 2000).Lure andKill: Strengths andWeaknesses. Strengths.

The inverse density dependence of lure and kill (i.e.,its efÞcacy improves as target density declines) con-fers major advantages on the use of this technology ineradication of invasive species. Cost-effectiveness im-proves as the pest density declines and this countersthe usual problem of facing rising costs for removingincreasingly rare individuals as eradication continuesin the case of invasive species (Myers et al. 1998). Lureand kill is, therefore, highly effective against low-density populations (Suckling and Brockerhoff 1999,Charmillot et al. 2000, Ioriatti and Angeli 2002, Krupkeet al. 2002), particularly if they are isolated from im-migration. Today, toxic synthetic lures can be pro-duced which are highly competitive with wild virginfemales; this is due to the production of improvedpheromone blends and dosages, and partly becausethe pheromone is constantly released from the lures,even when females are not calling (Krupke et al.2002). The combination of lure and insecticide en-ables the use of very small amounts of insecticide pergiven area (Brockerhoff and Suckling 1999, Sucklingand Brockerhoff 1999, Losel et al. 2000). Moreover,the major formulations are very durable (Losel et al.2000) enabling the insecticide component to be twiceas persistent as conventionally applied insecticide(Hofer and Angst 1995) and therefore requiring fewerapplications. The target insects are killed by the brief-est contact with the insecticide (Losel et al. 2000,Poullot et al. 2001), during which adequate uptake ofthe toxin can occur (Losel et al. 2000). Much lesspheromone is used per ha than in mating disruption(Suckling and Brockerhoff 1999, Trematerra et al.1999, Charmillot et al. 2000), and this should bringanother major cost saving; this is because the primarymode of action is insecticidal. Unlike insecticide treat-ments and mating disruption which are sensitive totopography and environmental conditions, lure andkill technology can be used in a variety of conditions,including small, irregular, and hilly sites (Charmillot etal. 2000, Ioriatti and Angeli 2002). The pheromone isable to penetrate complex environments difÞcult toreach with spray coverage and this enables attractionand kill of males from remote, protected or cryptichabitats. In addition, lure and kill products do notcause spray drift (Suckling and Brockerhoff 1999),which occurs with conventional insecticides. Almostall formulations also can be applied in ways that avoidinsecticide being deposited on crops that will be har-vested as food (Suckling and Brockerhoff 1999, Ioriattiand Angeli 2002). The method of insecticide use pro-vides for high selectivity (Suckling and Brockerhoff1999, Charmillot et al. 2000, Losel et al. 2000, Ioriattiand Angeli 2002), with minimal impact on nontarget


organisms. This includes a high level of safety to work-ers and the public. No expensive special equipment isrequired to apply most lure and kill formulations. Theycan be deployed by a minimally trained work force(Olszak and Pluciennik 1999, Losel et al. 2000).Weaknesses. The major weaknesses of lure and kill

are the reciprocal of its strengths. The method hasdecreased efÞcacy at high pest density (Downham etal. 1995, Charmillot et al. 2000, Ioriatti and Angeli2002), due to competition from greater numbers ofcalling wild females (Losel et al. 2000) and becausemales in this situation use many other cues than pher-omone to locate their mates. Just like mass trapping,lure and kill is susceptible to immigration of the targetpest into the treated area (Moraal et al. 1993, Char-millot et al. 2000, Ioriatti and Angeli 2002), particularlyby ßying mated females. The extreme dependence oflure and kill on a highly competitive lure and rapid-acting insecticide can be seen as a major weaknessbecause of the many factors which must be optimizedfor successÑlure blend, dose, distribution, and place-ment, insecticide formulation, and dose. Research anddevelopment is essential to address these issues. It hasbeen pointed out that there are cost savings from lureand kill due to the small quantities of pheromone andinsecticide required per hectare. However, its abilityto compete with wild females is dependent on thenumber of pheromone point sources per hectare. Thelabor involved in deployment of the lures is signiÞcantand becomes a trade-off between efÞcacy and cost(Losel et al. 2000),particularly athigherpestdensities.This is a further reason to focus the potential use of thistechnology on low density, isolated populations.Lure and Kill: Cost-Effectiveness. A full costÐben-

eÞt analysis of lure and kill is beyond the scope of thisreview and is recognized as fraught with difÞculties inthe context of eradication (Myers et al. 2000). Sharovand Liebhold (1998) have shown that there are soundeconomic reasons for eradicating small incipient pop-ulations ahead of the main front of spreading gypsymoth populations. The important issue for the costÐeffectiveness of lure and kill is in its increasing efÞcacyas pest populations decline. This results in, Þrst, inef-fectiveness and too high cost to control high densitytargets (including intolerably high distribution/place-ment costs); and second, high and increasing effec-tiveness against low and falling populations. Thismakes the method suitable for the Þnal stages of aneradication program and overcomes the usual majorobstacle in eradication of rapidly rising costs for re-moval of rarer and rarer insects. Concentrating effortson the eradication of low-density populations alsoassists in limiting trap costs because few moths will becaught and trap design is unlikely to need to cope withtrap saturation problems. Mating disruption uses“large” quantities of pheromone per ha and this is amajor cost. Lure and kill uses much less pheromoneand this reduced cost has been identiÞed as an im-portant issue for the cost-effectiveness of the tech-nology (see above), while deployment costs of lureand kill are reported to remain similar to mating dis-ruption.

New Developments. The majority of lure and killmethods developed in the past 10 yr have used thefemale sex pheromones of the target pest species asattractants. However, it has been recognized that lureand kill could be improved if female attractants couldbe found, because this would greatly enhance efÞcacyby removing virgin and mated females, particularly ifthis could be added to male removal. Kairomones,which are the odors of the hosts or prey of insects, mayalso be attractive to both males and females. Recentresearch has identiÞed kairomones of codling mothwhich are being investigated for lure and kill (Pottingand Knight 2001). Also, ßoral volatiles that attractnoctuid moths (e.g., El-Sayed et al. 2008) can enhancethe efÞcacy of lure and kill for noctuid pests. Anothernew development for control of lepidopterous pests is“lure and sterilize” (Charmillot et al. 2002). In thistrial, the target pest is codling moth, and in this tech-nique the pheromone (codlemone at 0.12Ð0.35 g/haper application) attracts males to contact a chemo-sterilant, fenoxycarb, at 5% (4.6Ð13.2 g fenoxycarb/haper application). The result is autosterilization of themales that can then disperse and, in some cases, matewith virgin females which remain sterile. Five years oftrials have been undertaken, using 1,540Ð4,400 by 50�l drops/ha in two to three applications per season(i.e., 198Ð526 g formulated paste per ha per year), andplacing one third of lures in the lower parts of appleand pear trees and two thirds in the upper parts. Goodpopulation reduction (overwintering larvae) wasachieved in the Þrst 2 yr, but lowering the pasteamounts to �200 g/ha allowed population increase.Although autosterilization is predicted to approxi-mately double the efÞcacy of lure and kill with insec-ticides (Potting and Knight 2001, Charmillot et al.2002), this has still to be demonstrated because therewas a high residual population at the end of these trialsin 2000. Autosterilization could be considered for pestmanagement and eradication of invasive species as analternative to lure and kill. However, an effective che-mosterilant would have to be found and chemoster-ilant chemicals would likely create the same publicconcerns as insecticides.

Conclusions

Lure and kill has the potential to add value in bothlong-term pest management of many economicallyimportant pests and in the eradication of invasivespecies by being instrumental in control or eradicationof small, low-density, isolated populations either in-side the main distribution area of a pest or during theÞnal stages of eradication. We have identiÞed keyfactors that can contribute to success of lure and killas follows: low-density target population; isolated tar-get population; a lure competitive with wild females;high lure density relative to pest density; optimal lureplacement; and lure deployment before adult emer-gence and throughout ßight period are conditions thatfavor a successful lure and kill program. The key stepsin developing a successful lure and kill program underthe constraints of population and economic conditions


are illustrated in a ßow diagram (Fig. 1). Lure and killmay offer some improvements in efÞcacy over othercontrol methods using semiochemicals, but the inclu-sion of insecticides or sterilant in lure and kill formu-lations may present major obstacles to public accep-tance in urban areas, where new incursions are oftendetected Þrst. This could be the main reason that othercontrol methods (i.e., mass trapping or mating disrup-tion) have been preferred in eradication programs.

Acknowledgments

This work was supported by New Zealand Ministry ofAgriculture and Forestry (Biosecurity New Zealand), andthe Foundation for Research, Science and Technology(CO2X0501, Better Border Biosecurity; www.b3nz.org andInsecticide Risk Reduction in Horticulture, CO 6X0301).

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Received 9 March 2008; accepted 24 June 2008.


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