grape berry moths in western european
DESCRIPTION
This chapter addresses the topic of pest biology and management for cluster-feeding Lepidoptera of European origin, from the viewpoint of applied entomologists working in Europe and North America. We describe the main biological, morpho- logical, and behavioral features of the Palaearctic moths harmful to grapes. Five species of Lepidoptera feed on grape clusters in Europe. Lobesia botrana (Denis & Schiffermüller) and Eupoecilia ambiguella (Hübner) are key pests that require spe- cific control measures. Argyrotaenia ljungiana (Thunberg), Cryptoblabes gnidiella (Millière) and Ephestia parasitella unicolorella Staudinger are occasionally harm- ful to vineyards. As the five species are similar in size and occur almost in the same ecological niche, morphological features allowing species identification are pro- vided in Table 14.1. For comparison, a description of the North American grape berry moth Paralobesia viteana (Clemens) is provided by Isaacs et al.TRANSCRIPT
339N.J. Bostanian et al. (eds.), Arthropod Management in Vineyards: Pests, Approaches, and Future Directions, DOI 10.1007/978-94-007-4032-7_14, © Springer Science+Business Media B.V. 2012
14.1 Introduction
This chapter addresses the topic of pest biology and management for cluster-feeding Lepidoptera of European origin, from the viewpoint of applied entomologists working in Europe and North America. We describe the main biological, morpho-logical, and behavioral features of the Palaearctic moths harmful to grapes. Five species of Lepidoptera feed on grape clusters in Europe. Lobesia botrana (Denis & Schiffermüller) and Eupoecilia ambiguella (Hübner) are key pests that require spe-cifi c control measures. Argyrotaenia ljungiana (Thunberg), Cryptoblabes gnidiella (Millière) and Ephestia parasitella unicolorella Staudinger are occasionally harm-ful to vineyards. As the fi ve species are similar in size and occur almost in the same ecological niche, morphological features allowing species identifi cation are pro-vided in Table 14.1 . For comparison, a description of the North American grape berry moth Paralobesia viteana (Clemens) is provided by Isaacs et al. (Chap. 15 ).
C. Ioriatti (*) Centre for Technology Transfer, FEM-IASMA , 38010 San Michele all’Adige , TN , Italy e-mail: [email protected]
A. Lucchi Department of CDSL, Section of Agricultural Entomology , University of Pisa , Via del Borghetto 80 , 56124 Pisa , Italy e-mail: [email protected]
L. G. Varela UC Cooperative Extension, Division of Agriculture and Natural Resources , University of California , 133 Aviation Blvd., Suite 109 , Santa Rosa , CA 95403-2894 , USA e-mail: [email protected]
Chapter 14 Grape Berry Moths in Western European Vineyards and Their Recent Movement into the New World
Claudio Ioriatti , Andrea Lucchi , and Lucia G. Varela
Tabl
e 14
.1
Mai
n m
orph
olog
ical
fea
ture
s of
the
prin
cipa
l gra
pe c
lust
er-f
eedi
ng m
oth
pest
s in
Eur
opea
n vi
neya
rds
Spec
ies
Fam
ily,
sub
fam
ily
Adu
lt E
gg
Firs
t ins
tar
larv
a Fi
fth
inst
ar la
rva
Pupa
Lobe
sia
botr
ana
Tortr
icid
ae,
Ole
thre
utin
ae
Win
gspa
n 11
–13
mm
. 0.
6 ×
0.7
mm
1
mm
10
mm
4–
6 m
m
Fore
win
gs ta
n-cr
eam
, mot
tled
with
gra
y-bl
ue, b
row
n, a
nd
blac
k bl
otch
es. H
indw
ings
gr
ay w
ith a
fri
nged
bor
der.
Win
gs h
eld
in a
bel
l sha
pe
over
the
abdo
men
at r
est
Slig
htly
elli
ptic
al,
yello
w s
traw
at
layi
ng, g
radu
ally
tu
rns
to tr
ansp
ar-
ent l
ight
gra
y w
ith
brig
ht ir
ides
cent
re
fl ect
ions
Cre
amy
whi
te
with
a
blac
k he
ad
Tan
to y
ello
wis
h br
own
head
an
d pr
otho
raci
c sh
ield
, re
ar e
dge
of th
e pr
otho
raci
c sh
ield
has
a
dark
er b
row
n to
bla
ck
bord
er. W
hite
tube
rcle
s at
the
base
of
the
body
ha
irs.
Ana
l com
b ha
s 5–
6 da
rk b
row
n te
eth
Slim
with
the
cran
ial a
nd
the
caud
al e
nd
roun
ded.
Cre
mas
ter
with
8 h
ooke
d br
istle
s.
Fem
ale
pupa
e ge
nera
lly m
ore
stoc
ky
and
larg
er th
an m
ales
Eup
oeci
lia
ambi
guel
la
Tort
rici
dae,
To
rtri
cina
e W
ings
pan
12–1
5 m
m
0.6
× 0
.8 m
m
1 m
m
12 m
m
5–7
mm
Fo
rew
ings
yel
low
-bro
wn
with
da
rk b
row
n ba
nd. H
ind
win
gs
slat
e gr
ay f
ring
ed
Lig
ht y
ello
w w
hen
fi rst
laid
late
r be
com
ing
spot
ted
with
bri
ght o
rang
e
Cre
amy
whi
te
with
bro
wn
head
Red
dish
to b
row
nish
yel
low
w
ith b
rist
les
over
its
who
le b
ody.
Hea
d ca
psul
es a
nd p
roth
orac
ic
plat
e an
d th
orac
ic le
gs
are
blac
k. A
nal c
omb
with
6–7
teet
h
Red
dish
bro
wn.
C
rem
aste
r w
ith 1
6 ho
oked
bri
stle
s.
Fem
ale
pupa
e ge
nera
lly m
ore
stoc
ky
and
larg
er th
an m
ales
Cry
ptob
labe
s gn
idie
lla
Pyra
lidae
, Ph
yciti
nae
Win
gspa
n 15
mm
0.
7 ×
0.4
mm
1
mm
11
mm
7
mm
Fo
rew
ings
dar
k gr
ay, p
unct
uate
d by
tiny
bla
ck s
pots
, vei
led
in
whi
te a
nd d
otte
d w
ith r
eddi
sh
scal
es c
hara
cter
ized
by
indi
stin
ct li
ghte
r ba
nds.
Hin
d w
ings
shi
ny w
hite
and
st
reak
ed w
ith te
rmin
al g
ray
lines
. Hor
n-sh
aped
pro
cess
on
the
thir
d m
ale
ante
nnom
erou
s
Subc
irco
lar
shap
e w
ith o
ne s
light
ly
fl atte
r po
le. W
hite
at
layi
ng, i
t gr
adua
lly a
ssum
es
a ye
llow
ish
colo
r
Lig
ht y
ello
w
with
bro
wn
head
Yel
low
to li
ght b
row
n bo
dy
with
two
narr
ow
long
itudi
nal d
arke
r ba
nds
on th
e do
rsal
sid
e.
Bro
wn-
redd
ish
head
with
sm
all b
lack
are
as a
t the
ba
se o
f bl
acki
sh b
rist
les
Bro
wn-
redd
ish
whi
ch w
ill
turn
gra
dual
ly to
war
d th
e da
rk r
ed. V
ery
shar
p cr
eam
y ab
dom
en, d
ista
lly
disp
lays
obl
ique
ho
oks
Eph
esti
a pa
rasi
tell
a un
icol
orel
la
Pyra
lidae
, Ph
yciti
nae
Win
gspa
n 15
–18
mm
10
mm
7–
8 m
m
Fore
win
gs li
ght b
row
n w
ith
tran
sver
se r
eddi
sh b
row
n an
d ba
rely
per
cept
ible
str
ips
Subc
ircu
r sh
ape,
w
hite
at l
ayin
g W
hite
-pin
kish
w
ith d
ark
brow
n he
ad
Lig
ht y
ello
w b
ody
with
sh
ades
of
pink
and
nu
mer
ous
bris
tles
with
el
onga
ted
blac
kish
hai
r tu
berc
les.
Red
dish
hea
d
Red
dish
-bro
wn
Arg
yrot
aeni
a lj
ungi
ana
Tort
rici
dae,
To
rtri
cina
e W
ings
pan
12–1
6 m
m
18 m
m
Fore
win
g si
lver
-whi
te, s
trig
ulat
ed
with
gra
y; m
arki
ngs
dark
re
ddis
h br
own,
spr
inkl
ed w
ith
blac
k; m
argi
ns o
f ba
sal a
nd
med
ian
fasc
iae
irre
gula
r. H
indw
ing
gray
Bat
ches
of
40–5
0,
yello
w, t
urni
ng
brow
n du
ring
de
velo
pmen
t
Cle
ar g
reen
bod
y. L
ight
gr
een
or y
ello
wis
h gr
een
head
. Ana
l com
b w
ith
6–8
pron
gs
Pale
bro
wn;
in a
silk
en
coco
on in
spu
n le
aves
, or
ove
rwin
teri
ng in
a
coco
on in
deb
ris
on
the
grou
nd
342 C. Ioriatti et al.
14.2 Lobesia botrana and Eupoecilia ambiguella
14.2.1 Taxonomy and Occurrence
The European grapevine moth (EGVM) L. botrana was described in 1775 by Denis and Schiffermüller as Tortrix , and subsequently as Eudemis and Polychrosis . Currently, as Lobesia Guenée 1845, it is in the family Tortricidae, subfamily Olethreutinae, tribe Olethreutini (Razowski 1995 ) . Lobesia botrana is historically present in Europe, Asia, and Africa (CAB 1974 ) . Although widespread in all grapevine-growing areas, it is economically important mostly in southern Europe. In southern France, in central and southern Spain, Portugal, Greece, Italy, and the islands of the Mediterranean Basin, L. botrana is the only moth species to have an important impact on grapevine production (Thiéry 2005 ) . Recently, it has expanded its geographic range and was found in Chile in 2008, California in 2009, and Argentina in 2010 (Gonzales 2010 ; Varela et al. 2010 ) .
The European grape berry moth (EGBM), E. ambiguella , is considered a signifi -cant insect pest in many grapevine-growing areas where it can cause considerable damage to grapes. It is found from Britain to Japan, from the Mediterranean Basin to the Scandinavian countries (farther north than grapevine-growing regions) (CAB 1986 ) . First described by Hübner in 1796 as Tinea ambiguella , it was sub-sequently included in the genera Cochylis and Clysia and lastly, according to Razowski ( 1995 ) , in the genus Eupoecilia. Known as Einbindiger Traubenwickler, Polilla de la vid, Cochylis de la vigne and Tignola della vite, it was recognized as the major grape berry pest in Europe until the 1920s. More recently, and in many areas, it has been gradually replaced as a major pest by L. botrana. The shift started in the Mediterranean Basin and is now extending to Switzerland, Austria and southern Germany where populations of L. botrana and E. ambiguella overlap.
14.2.2 Host Plants
Lobesia botrana larvae feed on grapevine flowers and berries and on a number of other plants growing in warm-dry environments, such as one fi nds in most Mediterranean countries. Its host range includes about 40 species belonging to 27 different families (Coscollá 1997 ) . The spurge fl ax Daphne gnidium L. is hypoth-esized to be its original host plant (Bovey 1966 ; Lucchi and Santini 2011 ; Tasin et al. 2011 ) . However EGVM is frequently associated with other hosts in habitats where suitable host plants occur. These hosts include for example olive tree infl o-rescence, Virginia creeper, jujube, rosemary, evergreen clematis, dogwood, ivy, currant (Bovey 1966 ; Coscollá 1997 ; Stavridis and Savopoulou-Soultani 1998 ; Thiéry 2005 ) . Eupoecilia ambiguella is also polyphagous and shares several host plants with EGVM. Even though mugwort, Artemisia vulgaris L., is sometimes reported as its native host plant, grapevine is now accepted as its original host.
34314 Grape Berry Moths in European Vineyards
Other cultivated and wild host plants include black and red currant, lemon citrus, Virginia creeper, blackthorn, smooth bedstraw, wayfaring tree, highbush cranberry, laureltinus, privet, ash, maple, dogwood, and many others (Solinas 1962 ) . Since grape berry moths are mostly restricted to grapevines in grapevine-growing areas, the nutritional and ecological polyphagy of these two tortricid species needs to be researched.
14.2.3 Life History
Eupoecilia ambiguella and L. botrana have a similar biology, but slightly different climatic preferences: L. botrana prefers warm and dry conditions and E. ambiguella prefers cool and humid climates (Bournier 1977 ) . Lobesia botrana and E. ambiguella are typical multivoltine species with facultative diapause. In northern Europe and in the Mediterranean Basin, EGVM has 2–4 generations per year on Vitis vinifera L., depending on latitude and microclimates (Roditakis and Karandinos 2001 ; Harari et al. 2007 ) . Two to three generations per year are the rule in Germany, Switzerland, Austria, and northern France. It has three generations in the warmer climates of southern France, Spain, Portugal, Greece, and Italy, where the species can sometimes give rise to a fourth fl ight and a fourth generation, partial or complete. In Israel, Egypt, and Crete some EGVM populations do not undergo diapause and spend the winter in the larval stage, continuing to feed on unharvested clusters or on alternate hosts. EGBM usually has only two generations per year. A third generation has been frequently observed in France and Italy (Marcelin 1985 ; Varner and Mattedi 2004 ) .
Lobesia botrana moths are crepuscular insects fl ying at canopy levels at twilight (60–80 lux). Males are more active than females. The fl ight of virgin females is interrupted by several stationary phases, in which they release a pheromone, the main component of which is ( E,Z )-7,9-dodecadienyl acetate (Roelofs et al. 1973 ) . Eupoecilia ambiguella moths are active from twilight to dawn, resting motionless during daylight. They feed, call, and mate during the night and early morning. Females lay eggs in the afternoon and evening (Bovey 1966 ) . The main component of the pheromone is ( Z )-9-dodecenyl acetate (Arn et al. 1976 ; Saglio et al. 1977 ) .
Adults of L. botrana (Fig. 14.1a ) feed from emergence to sexual maturity. Mating occurs about 24 h after emergence and oviposition starts 3 days thereafter. Mating lasts from a few minutes to 2 h. Males mate several times with different females, and females have a rare tendency toward polyandry. Males are much less attracted to mated females than to virgins. On average a female lays from 50 to 80 eggs (Fig. 14.1b , c), most of which are laid in the fi rst week of life. The average life span of the adult moth is approximately 15–20 days, usually shorter for males than females. The fi rst generation of both species develops on fl owers (anthophagous) and the other generations on berries (carpophagous) (Figs. 14.1d and 14.2b–d ).
Single eggs are laid on bracts, caps and stems of the fl ower clusters in spring and on the berries in summer. First generation larvae feed on several pre-bloom fl owers
344 C. Ioriatti et al.
and web them together with silk. Webbing gradually thickens to form the so-called glomerulus or nest (Fig. 14.2a ).
In hot weather, E. ambiguella larvae sometimes bore into the rachis and peduncle, seeking moisture. Within the glomerulus, EGBM larvae construct a silk case to which they retreat during the hottest hours of the day. In these nests, the hidden larvae feed on fl owers whose remnants are used to increase the size of the case. First generation grape berry moths eventually pupate either inside the glomerulus or on the underside of a leaf (Figs. 14.1e and 14.3d ). Their pupal stage lasts about 2 weeks.
The emerging moths (Fig. 14.3a ) lay single eggs on the berries (Fig. 14.3b ). EGBM hardly ever lays eggs on the rachis or on peduncles. After hatching, the
Fig. 14.1 Lobesia botrana ( a ) adult, ( b ) egg, ( c ) egg in the black-head stage, ( d ) larva, ( e ) pupa
Fig. 14.2 Lobesia botrana ( a ) larval nest, ( b, c ) injured berries, ( d ) grape cluster infected by gray mold and sour rot
Fig. 14.3 Eupoecilia ambiguella ( a ) adult, ( b ) egg, ( c ) larva, ( d ) pupa
346 C. Ioriatti et al.
larvae (Fig. 14.3c ) feed on the berries and may cause severe damage. When dis-turbed they drop down on a silk thread. European grape berry moth larvae are slug-gish while EGVM larvae are highly mobile. Feeding on berries promotes infection by gray mold ( Botrytis cinerea Persoon ex Fries), which leads to even greater injury than that caused by the insect itself. Non-diapausing larvae pupate preferably within the cluster or on the leaves. Photoperiod induces diapause that lasts for months, but mature EGBM larvae stay for several months in a prepupal stage and low tempera-tures are necessary to advance to the pupal stage (Fig. 14.3d ). The diapausing larvae build their cocoons mainly under exfoliating bark, in crevices and cracks of the trunk and cordons.
14.2.4 Economic Importance and Control
14.2.4.1 Grape Berry Moth Damage
The two species are quite similar in behavior and damage. Sometimes they are pres-ent at the same time in the same vineyard but with a different degree of population density. Until the end of the nineteenth century, E. ambiguella was widespread in all the southern European regions. From the beginning of the twentieth century, L. botrana gradually became the predominant species south of the Alps.
Grape berry moth infestation levels depend on the growth characteristics of the cultivars, the agronomic practices, the climatic conditions, and the number of genera-tions per year. The anthophagous generation does not generally cause yield reduction. The following carpophagous generations are the most destructive due to larval feeding on green and ripe berries, which results in yield reduction. The presence of larvae, web-bing and rotten berries, leads to downgrading of table grapes. Moreover, secondary infections of gray mold, B. cinerea , develop rapidly on damaged berries causing bunch rot which substantially degrades wine quality. Black aspergilli’s rot, Aspergillus niger and A. carbonarius , producers of ochratoxin A (Cozzi et al. 2006 ) , are also often related to larval feeding activity. Because of these undesirable economic effects, L. botrana and E. ambiguella must be managed to keep their damage at an acceptable level.
14.2.4.2 Population Density and Risk Assessment
Time of the fi rst appearance of adults and hatching of the fi rst eggs can be forecasted by predictive models based on temperature requirements of individual instars and critical conditions for oviposition (Moravie et al. 2006 ) . Unfortunately, forecast models based on DD are empirical and their robustness is strongly dependent on the environment in which they have been validated. Alternative forecasting techniques are currently under development, such as the evaluation of larval age distribution during the previous generation in order to predict the distribution of female emer-gence (Delbac et al. 2010 ) .
34714 Grape Berry Moths in European Vineyards
Trapping females with food-baited traps is a valuable tool to predict the onset of oviposition, an event used to properly time insecticide treatments (Thiéry et al. 2006 ) . However, baited-trap maintenance and monitoring are very time-consuming chores for growers and consultants. Pheromone traps are easier to use. They are a sensitive tool to monitor fl ight of males. They can be useful to time an ovicidal treat-ment, and to properly schedule scouting activities in the vineyard.
Forecasting models and moth trapping alone do not provide suffi cient population density information and need to be supplemented with appropriate fi eld scouting of eggs and young larvae (Shahini et al. 2010 ) . Based on the resulting infestation assessment (% of injured clusters, number of nests per infl orescence, number of eggs and larvae per cluster, number of injured berries per cluster), insecticides are applied according to action thresholds (AT). The action thresholds vary widely depending on the generation, susceptibility of the cultivar to the subsequent infec-tion by B. cinerea , as well as whether berries are produced for table grape, raisins or wine production.
First generation larvae may not necessarily need to be controlled by chemical treatments, especially if they develop on cultivars with abundant blossom not subject to intense shedding. Between fl owering and harvest, damage to infl orescences is compensated by the increase in weight of uninjured berries in the majority of cultivars. That explains the lack of a defi ned injury threshold for the anthophagous generation (Moschos 2005 ) .
Chemical control for the fi rst generation is exclusively applied when the pest pop-ulation density is particularly high or if it exceeds an AT of more than 50% infested infl orescences (Bagnoli et al. 2009 ) . For the following generations, the suggested AT ranges from 5% to 15% of infested (eggs or young larvae) clusters respectively for compact and loosely-bunched cultivars, according to their susceptibility to rot.
Knowledge of the spatial distribution of the population is important for the devel-opment of effi cient sampling programs, that allow a more accurate estimate of the damage and AT.
14.2.4.3 Natural Enemies and Biological Control
The cohort of L. botrana and E. ambiguella natural enemies varies considerably in time and space due to insect physiology, activity and ecological niche of individual species. Fungi of the genera Spicaria , Beauveria , Paecilomyces , Aspergillus , Cephalosporium , Cladosporium , Penicillium , Citromyces , Verticillium and Stem-phylium can infect a large percentage of overwintering pupae. The bacteria Bacillus thuringiensis Berliner var. kurstaki (Btk) and var. aizawai are effectively and exten-sively used against EGVM and EGBM, both in conventional and organic vineyards (Scalco et al. 1997 ; Vidal 1997 ; Keil and Schruft 1998 ; Shahini et al. 2010 ) . Arthropods associated with grape berry moths include predators such as spiders (Clubionidae, Theridiidae, Tomisidae, Linyphiidae, Salticidae), mites (Thrombididae) and insects belonging to Dermaptera, Hemiptera, Neuroptera, Diptera and Coleoptera (Solinas 1962 ; Coscollá 1997 ) . Among insect parasitoids, species associated with EGVM in
348 C. Ioriatti et al.
Europe belong to the Hymenoptera (Ichneumonidae, Braconidae, Chalcididae, Pteromalidae, Eulophidae, Elasmidae, Trichogrammatidae) and Diptera (Tachinidae). As with most natural enemies including parasitoids, the natural con-trol achieved by each species varies greatly in time and space. Typically, the fre-quency of egg and larval parasitism is high in the fi rst two generations and decreases drastically in the overwintering generation, which is mainly affected by larval-pupal and pupal parasitoids.
Extensive scientifi c efforts to develop biological control as an effective solu-tion for practical use in the fi eld are still needed. Egg parasitoids of the genus Trichogramma have been mass-released in an inundative strategy with mixed results (Castaneda-Samayoa et al. 1993 ; Hommay et al. 2002 ; Ibrahim 2004 ) . The ptero-malids Dibrachys affi nis Masi and D. cavus (Walker) are gregarious generalist larval-pupal parasitoids of Lepidoptera, Diptera and Hymenoptera that can be readily reared in the laboratory. However, due to lack of host specifi city and because they are also hyperparasites, they are not good candidates for release. The most frequent and effi cient species in European vineyards is the larval parasitoid Campoplex capitator Aubert (Ichneumonidae). It is regarded as the best candidate for EGVM biological control, but to date, releases have not taken place because of the diffi culties associated with artifi cially mass-rearing the species (Thiéry and Xuéreb 2004 ) .
14.2.4.4 Chemical Control
Most insecticides applied in the past against grape berry moths have been gradually replaced by more selective and less toxic products. New neurotoxic insecticides (spinosyns and oxadiazines), chitin synthesis inhibitors, compounds accelerating molting, microbial insecticides, and more recently some avermectins and anthra-nilic diamides, have been introduced in current integrated control strategies. Nevertheless, the organophosphates chlorpyriphos and methyl chlorpyriphos are still largely used in European vineyards. Control with insecticides that are larvicidal with some ovicidal activity gives remarkable fl exibility on application timing. The effi cacy of these products depends on the optimal treatment of the most susceptible stages, so prediction of life cycle events is critical for each moth species. Because of increasing accuracy of the forecasting tools, a single insecticide application to control the second generation of either species is usually effective in most grapevine districts in Europe. More treatments are needed in the southern regions to control L. botrana . In terms of selectivity, B. thuringiensis has undoubtedly the highest ecological value, but its use is still limited due to its short persistence. Successful application timing can be achieved with adequate population monitoring with pher-omone traps and egg fi eld scouting.
14.2.4.5 Pheromone-Mediated Control Strategies
The use of pheromones for control of grape berry moths has increased in vine-yards due to the high selectivity and low environmental impact. Mating disruption
34914 Grape Berry Moths in European Vineyards
(MD) with hand-applied dispensers is the most well-studied and widely used pheromone-mediated control technique against grape berry moths in the European grapevine-growing regions (Stockel et al. 1992 ; Neumann et al. 1993 ; Charmillot and Pasquier 2000 ) . Currently it is applied on approximately 140,000 ha in European vineyards, i.e. about 3–4% of the total grapevine-growing area in the European Union (Table 14.2 ). Recent MD area-wide applications have also been conducted in Chile and California where EGVM was accidentally introduced (Witzgall et al. 2010 ; Ioriatti et al. 2011 ) .
The most common hand-applied dispensers available on the market for grape berry moths are Shin-Etsu twist-ties ropes (Isonet L ® , Lplus ® , LE ® in Europe; Isomate ® in the US), the BASF twin ampoules (RAK1+2 ® , RAK 2 ® ) and, for EGVM only, the Suterra ® membranes. The active ingredients in these dispensers are the main pheromone components, ( E,Z )-7,9-dodecenyl acetate and ( Z )-9-dodecenyl acetate for EGVM and EGBM, respectively.
Five hundred dispensers per hectare (the number of dispensers may vary depend-ing on manufacturer) must be deployed in the vineyards before the onset of the fi rst seasonal fl ight, because late deployment will likely cause control failures. Dispensers must be evenly distributed in the vineyard, and should be attached to vine shoots to ensure protection by foliage from direct exposure to sun and high temperatures. Twice as many dispensers must be hung along the vineyard edges. Border effects are obviously much reduced when MD is applied in area wide proj-ects as in certain growing regions of Germany, France, Switzerland, northern Italy, and Spain (Kast 2001 ; Ioriatti et al. 2008 ) .
Depending on the vineyard layout, the time to attach the dispensers on the vines may vary between 1.5 and 3 h/ha. The surface area of vineyards in Europe under pheromone-mediated (MD) control of grape berry moths is still limited, despite intensive research and substantial experience with practical applications during the last two decades. This is because of socio-cultural and economical conditions existing in the different vine growing areas where interest in innovative
Table 14.2 European vineyards treated with pheromone mating disruption for management of grape berry moth pests during 2010 (IBMA 2011 ) in relation to the total vineyard surface of each country (OIV 2007 )
Country Total vineyard surface (ha) Vineyard treated with MD (ha) % treated
Germany 102,000 70,000 68.6 France 867,000 20,000 2.3 Italy 847,000 16,500 1.9 Spain 1,169,000 14,500 1.2 Switzerland 14,800 7,000 47.3 Austria 49,900 2,400 4.8 Czech Republic 17,700 1,300 7.3 Portugal 248,000 1,200 0.5 Hungary 75,000 300 0.4 Slovakia 17,600 100 0.6 Cyprus 15,300 100 0.7 Total 3,423,300 133,400 3.9
350 C. Ioriatti et al.
methods is often low. Increasing quality standards for wine and table grapes, with respect to pesticide residues, are creating new opportunities for extensive adop-tion of MD in IPM programs. However, high costs of MD (about 110 €/ha for EGVM and 150 €/ha for both insects) have hampered the diffusion of this method to date. Cost reduction must be considered for a wider adoption of MD in European vineyards.
Novel pheromone application systems to control Lepidoptera pests such as auto-confusion, lure and kill, aerosol puffers, microencapsulated sprayables, and nano-fi bers may represent future opportunity for grape berry moth control (Underwood et al. 2002 ; Charmillot et al. 2005 ; Nansen et al. 2007 ; Anfora et al. 2008 ; Hein et al. 2011 ) . New investments in fundamental research are critical for an effective improvement in semiochemical applications. The research should address the repro-ductive, physiological, and behavioral mechanisms by which the pheromone affects the target insects, as well as explain how volatile compounds are involved in tritrophic interactions.
14.2.4.6 Lobesia botrana as an Invasive Species in the Americas
It was fi rst detected in Chile in April of 2008. Grapes are grown from the region of Atacama in the north to the region of Araucanía in the south. In surveys conducted in the growing season of 2008–2009, moths were detected in all grapevine-growing regions. Low levels of catches were detected in the 2010–2011 season in grapevine-growing areas from Atacama to Araucanía. However, large urban areas remain as moth reservoirs. The current control strategy is to attempt eradication of the pest and major efforts are concentrated towards vineyards and urban areas surrounding grapevine-growing areas. After three seasons of control with insecticide and MD, moth catches in monitoring traps have decreased signifi cantly and the number of foci per region has also decreased considerably.
In the United States, the fi rst report of this pest was in September 2009 in the Napa Valley, California. Surveys conducted in 2010 show that the highest infesta-tion is in Napa County, with a few moths detected, and a few foci in nine other counties of California. No detection has been made in any other US state, despite trap-based surveillance in many of the primary grape production regions. The strat-egy in California is also eradication and in the fi rst year of control populations decreased dramatically from the fi rst to the third generation.
It is unclear how EGVM was fi rst introduced into Chile or California. At low popu-lation levels the damage caused by the larvae is inconspicuous. By the time the fi rst infestations are detected, the spread may be extensive due primarily to movement of grapevines with undetected infestations and movement of unsanitized machinery.
In both Chile and California the primary host for EGVM is grapevine fl ower clusters and berries. In extensive surveys in Chile it has only been detected in plums next to an infested vineyard. In California, it has been detected in low numbers only in olive fl owers adjacent to vineyards. To date, surveys conducted in riparian vege-tation in infested areas of California have not detected larvae in wild grapes.
35114 Grape Berry Moths in European Vineyards
Chile began eradication programs 2 years before California. In the fi rst year, control measures were applied to vineyards to a radius of 5,000 m from a detection site. This was reduced to 1,000 m the following year. The recommendations are to make two insecticide applications for the fi rst generation, one for the second and one for the third generation, plus the application of pheromone dispensers for mat-ing disruption. In urban districts surrounding grapevine-growing areas, homeown-ers have a choice of destroying the fruit or accepting insecticide treatments.
In California, EGVM has three generations a year. In the fi rst year control mea-sures were applied to a radius of 1,000 m from a detection site and it was reduced to 500 m the following year. With the goal of eradicating the pest, one insecticide application is targeted for each of the fi rst and second generations, when the larvae are most exposed. To further suppress populations the use of MD dispensers (the product registered as Isomate®-EGVM in the US) is highly encouraged. Control in urban areas has been limited to those counties with low trap catches. Homeowners mostly adopted fruit removal as a management technique.
Given that the fi rst fl ight and egg laying period is very extended, if the application for this generation is done before egg laying or too early in the egg laying period, a second application may be needed to cover the prolonged egg hatch. Furthermore, at this time, the fl ower cluster is rapidly expanding, decreasing the surface covered by an insecticide. Thus, it is best to wait and time the control of the fi rst generation when the highest proportion of larvae is about to emerge from eggs. The timing for this event can be determined by following the male moth fl ight with pheromone traps and monitoring egg development. If the insecticide used has some ovicidal properties the recommendation is to make the application when the heads of the larvae are visible in 20% of the eggs. When eggs are too few to monitor, treatment is applied shortly after peak fl ight. Insecticides registered for organic production are strictly larvicidal and are applied at egg hatch. Due to the short residue of organic materials, two or more applications are recommended starting at egg hatch and weekly for as long as larvae are detected forming glomeruli in the fl ower cluster.
The insecticide timing for the second generation depends on whether the insec-ticide has some ovicidal properties or if it is strictly larvicidal. If it is ovicidal, the applications can start a few days after the fi rst males of the second fl ight are caught in a trap. For larvicidal insecticides (conventional or organic), the applications can start 10–14 days after the fi rst moths of the second fl ight are caught, if eggs are too few to monitor. The second generation is substantially shorter than the fi rst, lasting approximately 4 weeks. This makes timing for control of the second generation easier to predict. If treatments are timed appropriately for the fi rst and second gen-erations, treatment of the third generation should not be necessary. Treatments for the third generation are limited in their effi cacy in California because of overlap in generations, the diffi culty in penetrating a closed cluster and the short period between egg hatch and the larvae penetrating the berry.
The strategy of targeting control measures towards the fi rst and second generation supplemented with mating disruption has proven extremely successful at drastically reducing populations. This is probably aided by the fact that all control measures are taking place in an area wide manner since all growers are strongly encouraged
352 C. Ioriatti et al.
to participate. So far, alternate hosts do not appear to contribute signifi cantly to population levels. In California, chlorpyriphos is not registered for seasonal use in vineyards, and the insecticides most used to control L. botrana are insect growth regulators, diamides and Btk, and to a lesser extent, spinosyns and avermectin. A major concern of the program was to avoid disruption of natural control of native pseudococcids pests. This was achieved by using selective insecticides.
A major challenge is to achieve complete control in urban areas. Another chal-lenge is to determine when a population has truly been eradicated. Delimitation of infestations is done using pheromone traps. Pheromone traps are effective; however the male moth does not fl y more than 100 m, with an average distance <50 m (Roehrich and Carles 1981 ) . This entails having a very high density of traps with no catches during several generations. The risk of a false negative is signifi cant since it will be tempting to declare the pest eradicated when in reality populations are breed-ing at undetectable levels.
In February 2010, EGVM was also detected in the major grape production region of the Province of Mendoza in Argentina. As of 2011, the fi rst year of control is underway.
14.3 Cryptoblabes gnidiella
14.3.1 Taxonomy and Occurrence
Among Pyralidae Phycitinae, the honeydew moth (HM), C. gnidiella (Fig. 14.4a–e ), is the most frequent and harmful species on grapes in the Mediterranean Basin. Described for the fi rst time by Millière in 1867 as Ephestia gnidiella , it was then reported by Briosi as Albinia wockiana in 1877. The current classifi cation is due to Hartig ( 1939 ) , who redescribed the species from specimens collected in central Italy. Widespread throughout the Mediterranean region, HM is reported from Malaysia, New Zealand, Hawaii, some African and Asian countries, and many trop-ical and subtropical regions of North and South America.
14.3.2 Host Plants
Cryptoblabes gnidiella is a polyphagous species associated with about 60 different host plants belonging to 30 families. These include Actinidia deliciosa (Chevalier), Citrus spp., Daphne spp., Daucus carota L., Diospyros kaki L., Eriobotrya japonica (Thunberg) Lindley, Gossypium herbaceum L., Malus spp., Persea Americana Miller, Prunus spp., Pyrus spp., Ricinus communis L., Tamarix spp., and Vitis spp. (Zocchi 1971 ; Yehuda et al. 1991–1992 ; Sing and Sing 1997 ) . Very frequently HM shares the host plant with other insects, either Lepidoptera (e.g., the European grapevine moth L. botrana ) or Hemiptera (aphids and pseudococcids) which
35314 Grape Berry Moths in European Vineyards
produce honeydew of which HM larvae are active consumers, hence the common name “Honeydew moth”.
14.3.3 Life History
In the Mediterranean Basin, C. gnidiella has 3–4 generations per year depending on latitude, with a fi rst fl ight in May–June, a second in July, a third in August–September and a fourth in October–November, with possible overlapping generations on late harvest grape varieties (Bagnoli and Lucchi 2001 ) . In Israel the species can have up to seven generations on grapes and citrus (Avidov and Harpaz 1969 ) . In the grapevine-growing areas of northeast Brazil, where climatic conditions allow two annual crops, HM can have as many as nine generations per year (Bisotto-de-Oliveira et al. 2007 ) . It overwinters as a larva and pupation takes place inside a silken cocoon. The sex pheromone of Cryptoblabes gnidiella is a mixture of quaternary aldehydes (Bjostad et al. 1981 ) .
14.3.4 Economic Importance and Control
The economic importance of C. gnidiella varies greatly according to geographical areas. Though it mainly occurs in coastal areas characterized by heavy infestation of
Fig. 14.4 Cryptoblabes gnidiella ( a ) adult, ( b ) young larva, ( c ) mature larva, ( d ) pupa, ( e ) infested clusters
354 C. Ioriatti et al.
L. botrana and Planococcus spp., it is also able to infest healthy pre-veraison grapes, feeding on cluster stems. During ripening, larvae may feed superfi cially on berry skins. Regardless of the feeding damage, since it is highly gregarious the number of larvae determines the level of damage caused (Lucchi et al. 2011 ). Because of the highly aggregated larval distribution, affected clusters are fully compromised by the inevitable and rapid development of rots enhanced by the presence of drosophilids and by nitidulids. In Israel and Brazil, the species is considered a key pest of vine-yards (Harari et al. 2007 ; Bisotto-De-Oliveira et al. 2007 ) . Protection of grapes from C. gnidiella infestation is achieved with effective control of L. botrana . If well timed, it eliminates the need for a specifi c spray against phycitin larvae. Moreover, Btk can be usefully employed in case of asynchronous outbreaks.
14.4 Ephestia parasitella unicolorella
Larvae of E. parasitella unicolorella (Pyralidae: Phycitinae) (Fig. 14.5a–f ) are found within the cluster after veraison as a secondary pest on wilted or dried berries and very often hidden within them. Winter is spent in the larval stage, in a thin cocoon spun by the mature larva on woody structures of the vine or on support poles. It is not yet clear where this insect resides outside the vineyard, especially during the spring, nor the number of generations that the species has in Europe. The economic importance of E. parasitella unicolorella is negligible. Deseo ( 1980 ) reports that the young larva feeds on the rachis and petiole of the bunch, whereas the older larva can penetrate and develop on a single berry, feeding on the pulp. At harvest mature larvae are frequently found in the most internal parts of the bunch associated with or inside rotten or dried berries, almost motionless and folded in a C shape. Xuéreb et al. ( 2003 ) advised to destroy the unharvested clusters to avoid the further development of the species in the vineyard. In a recent review the name of Ephestia unicolorella woodiella Richard & Thomson has been proposed for this species (Huertas Dionisio 2007 ) .
14.5 Argyrotaenia ljungiana
Argyrotaenia ljungiana (Fig. 14.6a–d ) (syn. A. pulchellana Haworth) is present throughout the Palaearctic region with the exception of Japan. Females deposit their eggs in batches of 40–50 eggs, usually on the upper surface of the leaves. Larvae feed primarily on leaves of host plants. Pupation occurs in a silken cocoon inside webbed leaves. In the Mediterranean region, the species has three generations per year, the fi rst generation occurring in April and May, the second from the end of June to July and the third in August–September. Argyrotaenia ljungiana overwinters in the pupal stage, inside a cocoon in debris on the ground. A highly polyphagous species, it feeds on many wild and cultivated plants, including grapevine and apple.
35514 Grape Berry Moths in European Vineyards
Fig. 14.5 Ephestia parasitella unicolorella ( a ) adult, ( b, c, d ) larva, ( e ) overwintering larva, ( f ) pupa
Occasionally, A. ljungiana can give rise to important outbreaks in vineyards, where it feeds on infl orescences and berry clusters. The harmfulness of this tortricid on grapes has been described in Italy (Varner et al. 2001 ) , France (Marcelin 1985 ) , Bulgaria (Kharizanov 1976 ) , and Hungary (Voigt 1972 ) . On the berries it may cause superfi cial but extensive excavations, which are different from those caused by the other two tortricids, but it can also provide entry points to rots when feeding on ripening grapes. Sometimes the larvae can deeply abrade the cluster rachis causing desiccation of the grapes.
356 C. Ioriatti et al.
14.6 Conclusion
Of the fi ve Lepidoptera species feeding on clusters in European vineyards, EGVM and EGBM have the highest adverse economic impact. The other three species described here, two Pyralidae and one Tortricidae, are occasional or secondary pests. EGVM has increased its geographic range in the twentieth century throughout Europe and the Middle East, invading the Americas early in the twenty-fi rst century. In the Mediterranean region south of the Alps, EGBM is being replaced by EGVM as the major lepidopteran pest. Where EGVM is established, insecticide control for the fi rst generation is not practiced given that, in most varieties, damage to the infl o-rescence has no impact on yield. Insecticidal control is targeted primarily at the second generation larvae. In recent years, more selective insecticides have been introduced, with some having ovicidal activity. In Europe, the area-wide approach based on the use of pheromones to control EGVM and EGBM represents an impor-tant development in a more environmentally acceptable control of these insects. In regions were eradication is being pursued a program combining mating disruption
Fig. 14.6 Argyrotaenia ljungiana ( a ) adult, ( b ) larva, ( c, d ) damage on grapes
35714 Grape Berry Moths in European Vineyards
and insecticides targeted for fi rst and second generation larvae has achieved a drastic reduction in population levels.
Acknowledgements We are much indebted to Bruno Bagnoli and Vittorio Veronelli for productive discussions on grape moths. Photo acknowledgments: Bruno Bagnoli: Figs. 14.1b , c; 14.3c ; 14.4b–d ; 14.5b , c. Paolo Giannotti: Figs. 14.1a , e; 14.3a , d. Mauro Varner: Fig. 14.6b , c. Elena Pozzolini: Fig. 14.6d .
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