LITERATURE REVIEW

Biology and Management of Cuscuta in Crops

W.T. Lanini1 and M. Kogan2

1
Department of Plant Science
University of California, Davis
One Shields Ave, Davis, CA 95616
2
Facultad de Agronoma e Ingeniera Forestal
Pontificia Universidad Catlica de Chile
Casilla 306-22, Santiago, Chile

Abstract

W.T. Lanini and M. Kogan. Biology and management of Cuscuta in crops. Cuscuta
is a stem and leaf parasite that infects many broadleaf crops, ornamentals and weeds and
a few monocot crops. It lives entirely on the host plant, thus reducing the growth and yield
of the host. Preventing infestations by planting crop seed free of Cuscuta seed, rotating
to non-host crops, delaying crop planting until fall for sugarbeet (Beta vulgaris), use of
resistant varieties or large transplants, and preemergence herbicides have all been shown
to be successful in certain crops. Once Cuscuta attaches to a crop, some yield loss will
occur, regardless of the method of control, and selective control becomes very difficult.
Post attachment control often requires killing or severely injuring the host plant to avoid
spread of Cuscuta to surrounding plants. However, several herbicides have been shown
to selectively suppress attached Cuscuta, but complete control is rarely obtained. Cuscuta
attached to genetically modified, herbicide resistant crops, have not been successfully
killed by treatment with herbicide in all cases, indicating that these crops will only be a
partial solution to the problem. Cuscuta control will require an integrated approach
conducted over a period of many years.

Key Words: Cuscuta, dodder, parasitic plant, resistance, weed management.

Cien. Inv. Agr. 32(3): 127-141. 2005

INTRODUCCTION as important in Chilean crops. Dodder is a

The genera Cuscuta (known as dodder) are
obligate parasitic plants with approximately
170 different species distributed throughout
the world (Holm et al., 1997). Most of the
170 species are found primarily in the
Americas from Canada to Chile of which 7
or 8 are known in Chile (Navas, 1979).
Kogan (1992) recognized C. chilensis, C.
racemosa var. chiliana and C. campestris
nonspecific parasite that attacks, sometimes
simultaneously a wide range of host species
including many cultivated plant species and
dicotyledonous weeds, but not grasses or
monocotyledonous weeds (Table 1; Dawson
et al., 1994). Similarly, the same crop may
serve as a host of several dodder species
(Cudney and Lanini, 2000). The dodder
seedling coils around the host stem and
leaves, penetrates their tissue and vascular

Received on 16 March 2005; Accepted on 29 April 2005

1
Corresponding author: W.T. Lanini,wtlanini@ucdavis.edu
128 CIENCIA E INVESTIGACION AGRARIA

system via haustoria, and exploits the host
by withdrawing photosynthates and water
(Figure 1). Thus, the vigor of the host is
lowered and crop production is dramatically
reduced. Once a seedbank is established,
control is extremely difficult, as dodder seeds
can remain viable in soil for 20 years or
more, and continue to germinate and emerge
throughout the warm seasons. In addition,
the nature of attachment and association
between host and parasite requires a highly
selective herbicide to destroy the parasite
without crop damage (Fer, 1984). This article
reviews the life cycle, distribution and the
control measures currently used for control
of dodder in crops.

B C
Figure 1. Dodder, Cuscuta sp, common obligate parasitic plants. A. Onion field heavily
infested with dodder; B. Dodder haustoria (arrow) penetrating tomato stems; C. Dodder
wraping (arrow) arrow around tomato stems.
 VOL 32 N3 SEPTEMBER - DECEMBER 2005. 129

Table 1. Crops, ornamental plants, and common weeds susceptible to dodder (Cuscuta spp.) infestation.
Latin name Common name
Crops
Monocotyledonous Allium cepa Onion
Allium sativum Garlic
Dicotyledonous Asparagus officinalis Asparagus
Beta vulgaris Sugarbeet
Capsicum annuum Pepper
Carthamus tinctorius Safflower
Citrus spp. Citrus
Cucumis melo Melon
Cucumis sativus Cucumber
Daucus carota Carrot
Ipomoea batatas Sweet potato
Lycopersicon esculentum Tomato
Punica granatum Pomegranate
Solanum melongena Eggplant
Solanum tuberosum Potato
Ornamental
Monocotyledonous Impatiens spp. Impatiens
Chrysanthemum spp. Chrysanthemum
Ipomoea spp. Morningglory
Dicotyledonous Satureja hortensis Summer savory
Coleus blumei Coleus
Geranium spp. Geranium
Dahlia merckii Dahlia
Parthenocissus quinquefolia Virginia-creeper
Campsis radicans Trumpet-vine
Hedera helix English ivy
Petunia inflata Petunia
Foeniculum vulgare Fennel
Mentha spp. Mint
Origanum majorana Marjoram
Catharanthus roseus Periwinkle
Weeds
Monocotyledonous Solanum nigrum Black nightshade
Portulaca oleracea Common purslane
Amaranthus blitoides Prostrate pigweed
Dicotyledonous Bilderdykia convolvulus Wild buckwheat
Convolvulus arvensis Field bindweed
Salsola tragus Russian thistle
Chenopodium album Lambsquarters

DODDER LIFE CYCLE 1987). Favorable soil temperatures for

dodder germination and emergence are in
Cuscuta seeds germinate independently the range of 15 to 38C, with an optimum
of the presence of host plants (Dawson, around 30C (Hutchison and Ashton,
130 CIENCIA E INVESTIGACION AGRARIA
1979), which corresponds with the
prevailing temperatures during the spring
and summer. Because of the seed size (1
to 2 mm in diameter), emergence is limited
to the upper 1 to 1.5 cm of soil. Germi-
nating seeds will emerge as rootless and
long yellow-orange thread-like leafless
stems, which can grow to 2.5 to 7 cm in
height. After germination, the dodder
seedling circumnutates in a counter
clockwise direction in search of a host
stem or other objects to wind around. It is
possible that light or humidity may
influence growth toward a host plant.
These rootless seedlings will generally
only attach to hosts that are within a
distance of 2.5 to 5 cm. Cuscuta seedlings
possess a rudimentary autotrophic system,
containing only a small amount of
chlorophyll (Table 2) and other accessory
pigments (Dinelli et al., 1993). However,
the autotropic system is insufficient to
support growth. Thus, if no suitable host
is found within 3 to 5 days, the seedling
will die. Soon after the connection between
the parasite and host is established, the
dodder seedling loses its soil connection
and lives entirely from photosynthates and
water extracted from the host plant (Parker
and Riches, 1993). Dodder adheres to the
host with a cementing layer of pectin and
develops haustoria within a few days due
to thigmotropic responses and chemical
recognition of the host plants (Press et al.,
1990). The haustoria, single-cell hyphae,
elongates within the host tissue and meets
the vascular bundles, where they
differentiate into xylem and phloem
elements. This highly efficient absorption
system allows the parasite to divert
resources - water, amino acids and
assimilates - from the host into the parasite
(Dorr, 1987). It has been shown that when
Cuscuta and developing seeds from the
host plant fruit compete for assimilates,
the sink activity of dodder is much stronger
while the sink activity of the fruit can
disappear completely (Wolswinkel, 1984).
As the dodder plants grow, support is
maintained by continually reattaching to
the host. Dodder plants grow about 7 cm
per day and one plant can cover 3 m2 in a
growing season. When other suitable hosts
are nearby, dodder extends and attaches
to their stems, spreading from one host to
another, often forming a dense vegetative
mat of intertwined stems. Flowering can
occur from late spring through fall,
depending on the species and date of
emergence, however seed set is highest in
the late summer and fall. In a field study
of dodder (C. pentagona) developing on
processing tomato, the first dodder flowers
were observed at 51 days after initial
attachment and the first viable seeds were
observed at 60 days after attachment. First
flowering of dodder was observed to
initiate near the point of initial attachment
and later flowering progressed out away
from this point. Once seeds are shed to
the soil they can remain dormant, yet
viable, in the soil for 10 to 30 or more
years, depending on the species and
environmental conditions.
Table 2. Content of chlorophyll a and b in
Cuscuta campestris, Convolvulus arvensis and
Beta vulgaris.
Species Organ Chlorophyll a + b
(mgg-1)1
B. vulgaris Leaves 3.91 0.31
C. arvensis Leaves 2.54 0.21
C. campestris Stems 0.28 0.02
From: Dinelli et al., 1993.
1
Based on dry matter measurements. Standard
deviation
Economical Importance and geographical
distribution
The wide geographical distribution of
dodders (Table 3) and their wide range of
hosts, make them amongst the most
damaging parasites worldwide (Parker and
Riches, 1993). Studies have shown that
 VOL 32 N3 SEPTEMBER - DECEMBER 2005.
field dodder (C. pentagona) infestation
r e d u c e d t o m a t o ( Ly c o p e r s i c u m
esculentum) yield by 50 to 75% (Table 4;
Lanini, 2004). It was shown that field
dodder infestation reduced carrot (Daucus
carota) yield by 70 to 90% (Bewick et al.,
1988). Dodder is also considered a
troublesome weed in onion (Allium cepa).
Rubin (1990) stated that onion fields
heavily infested with dodder should be
destroyed, as there is no selective herbicide
to control it available for this crop. In
carrots or onions infested with dodder, the
roots or bulbs, respectively, fail to reach
a marketable size. When multiple dodder
plants attack the same host (tomato) plant,
death of the host often occurs. Alfalfa
(Medicago sativa) and clover (Trifolium
pratense) are the most common crops
infested by dodder (Dawson et al., 1994).
Although dodder does not generally kill
alfalfa, it weakens the crop, reduces stand
and can reduce yield of forage and seed
Table 3. The main Cuscuta species invading crops and their geographical distribution.
Species Distribution
C. pentagona Worldwide
(C. campestris)
C. epithymum Worldwide
C. europaea Europe and North America
C. gronovii North America
C. indecora North and South America
C. planiflora Asia, Europe, and
North America
C. reflexa Asia
C. suaveolens South America,
Europe and Africa
131
production by more that 50% (Cudney et
al., 1992). Where present in harvested
alfalfa, its moist stems will not allow the
hay to cure properly resulting in moldy,
unmarketable hay. Other legume crops
vary in their sensitivity to dodders. Bean
(Phaseolus spp.) was reported as resistant
to China dodder (C. chinensis) (Rao and
Reddy, 1987) and to field dodder in India
(Nemli, 1987), but sensitive to C.
lupuliformis in France (Liu et al., 1991).
On the other hand, China dodder is a
noxious weed in soybean (Glycine max)
in China (Li, 1987). Ornamental shrubs,
trees and groundcovers are often infested
with dodder (Table 1). Dodder infestations
are readily apparent along roadsides in
weeds, shrubs and trees. Host plants are
rarely killed by dodder infestation. The
weakened state of the infected plants does
predispose them to loss from other
maladies (disease, insect, and nematode
invasions).
Comments
The most important Cuscuta species, attacking a wide
range of species, including vegetables, fruits,
ornamentals and woody plants. It is reported as a weed
in 25 crops in 55 countries.
This is a serious weed problem in Europe and Asia,
on clovers, other forage legumes and carrot.
A serious weed in Europe, but also found in USA.
This species typically is found in wet places and along
water courses. It has a wide host range, attacking
cranberry, other crops and shrubs.
This is an important species primarily found attacking
alfalfa.
A widespread species having a wide host range,
including alfalfa and clovers.
A serious weed problem in woody perennials.
Native to South America, but now found worldwide,
primarily attacking alfalfa
132 CIENCIA E INVESTIGACION AGRARIA
Table 4. Tomato fruit yield relative to the level of dodder infestation (% cover at harvest) at
Davis, California.
Dodder cover 1991
at harvest reds greens
(%) tonha-1
80 to 100 36.3 7.0
60 to 80 60.7 5.3
40 to 60 66.7 10.5
20 to 40 58.0 10.0
1 to 20 68.7 13.2
0 73.3 14.3
DODDER CONTROL
Dodder prevention can be achieved by
avoidance, and control by mechanical means
or hand removal, resistant varieties, and
herbicides. None of these methods work
100% of the time, but all offer some tools
for managing this parasitic weed. Another
complicating factor is that several dodder
species are often involved. Dodder
populations have also been observed to vary
in virulence and susceptibility to herbicide
treatment. Thus, an integrated approach to
managing dodder will be needed.
SANITATION
Avoidance and prevention are the most
effective and most economical methods to
reduce dodder infestations (Parker and
Riches, 1993). Dodder seeds are likely spread
by man, through seed international
commerce, movement of equipment, and in
the mud on tires and shoes (Cudney and
Lanini, 2000). Planting contaminated seed
can lead to severe infestations (Parker and
Riches, 1993). C. pentagona has been
distributed worldwide as a contaminant of
alfalfa seed, as both dodder and alfalfa seeds
are very similar in appearance. Dodder seed
has also been spread as a contaminant of
flax, linseed, and niger seed (Parker and
Riches, 1993). Thus, avoid using seed from
fields where dodder is present or use seed
1992
rots reds greens rots
tonha-1
0.0 22.9 1.1 0.7
0.0 39.5 1.6 1.8
0.0 73.4 3.1 2.0
0.3 93.6 3.4 3.6
0.1 95.4 5.2 2.7
0.2 93.4 7.4 1.6
certified free of dodder seed. To prevent
new dodder infestations, clean equipment
after working in a dodder infested field, prior
to moving to another field. Similarly, do not
allow animals feeding in a dodder infested
field to move to a dodder free field, as dodder
seed are known to survive passage through
the digestive system of many domestic
animals (Gaertner, 1950). Many broadleaf
weeds have been observed to host dodder
(Table 1; Ashton and Santana, 1976). Hence,
proper weed control in and around cultivated
areas will prevent dodder from parasitizing
weeds and would reduce the potential for
dodder seed production and dispersal.
Fluctuations in soil moisture and
temperature near the surface may initiate
dormancy breaking of superficially buried
dodder seed (Hutchison and Ashton 1979).
These authors observed dodder emergence
to begin in March and to cease in early to
mid-May (early spring to late spring) in the
central valley of California (N 3833, W
12147), despite temperatures in the
optimum range for germination and
emergence (Hutchison and Ashton 1980).
Parker and Riches (1993) observed a flush
of dodder seed germination often occurs
early in the spring season and they attributed
this to dodder seed losing their dormancy
during the cold winter months. Additionally,
the major flush of dodder emergence often
occurs close to the time of crop emergence,
 VOL 32 N3 SEPTEMBER - DECEMBER 2005. 133

indicating that field preparation, crop Table 5. Number of dodder attachments to

seeding and irrigation or rainfall may also selected tomato varieties when scarified dodder
be involved in germination, perhaps by was seeded into pots when tomato reached the
scarification of the seed coat or leaching 1 leaf stage or 3 leaf stage.
of chemicals out of the seed coat. Delayed Tomato variety Dodder attachments
planting can reduce dodder infestation, but 1 leaf 3 leaf
that is not always practical due to marketing Campbell CXD 233 3 2
considerations for certain crops. In direct Heinz 1100 0 0
seeded crops, the use of transplants may Heinz 9492 0 2
allow a later planting without sacrificing Halley 3155 4 5
early market delivery. Larger transplanted SVR 024 1 0312 2 3
tomatoes have also been observed to be Campbell CXD 234 5 0
more resistant to dodder attachment (Table Heinz 1400 4 2
5), as older stems may become more SVR 024 2 0662 3 0
lignified (Lanini, 2004). Early seeding of SVR 024 2 0664 1 0
the crop in spring might delay infestation SVR 024 2 0665 3 2
due to low temperatures, which together 4863N 4 0
AB 2 3 2
with a delayed irrigation can prolong the
APT 410 4 1
period before dodder emerges (Dawson,
Campbell CXD 179 4 0
1987). These means will allow the Campbell CXD 222 4 0
development of dense crop foliage that will ENP 113 5 0
suppress dodder's initial light dependent Heinz 0830 3 0
growth. Heinz 2501 3 0
Heinz 2601 2 2
Crop rotation can be an effective method Heinz 8892 4 1
to reduce dodder infestations. However, the HMX 3859 5 0
large weed and crop host range makes it Heinz 9663 0 0
difficult to avoid this pest by crop rotation Heinz 9665 0 0
and thus, careful crop selection is essential Heinz 9780 2 0
(Parker, 1991). Dodder is often observed Heinz 9888 0 0
coiling around grasses, but cannot form Heinz 9997 2 0
haustoria connections and will die unless Average 2.69 0.85
a suitable host is found. Thus, growing
cereals or other grass crops (which are not Due to the absence of roots, dodder
parasitized by most dodder species) seedlings (before attachment to the crop)
continuously for several years, may facilitate are easy to control by shallow cultivation
the exhaustion of dodder seed bank in the (Parker and Riches, 1993). In addition,
soil (Dawson, 1987). Lanini (2004) found tillage may hasten drying the soil surface,
that growing wheat followed by corn in a thus preventing further dodder germination
field heavily infested with C. pentagona and emergence. Tillage can be employed
reduced the number of dodder plants in crops grown in rows, which allow
infesting tomato by 90%. Thus, two years cultivation such as sugar beet and alfalfa
of growing a non-host crop was effective grown for seed production. However, the
in reducing the population. Of course these tillage only controls the dodder growing
grass crops must be well weeded, as dodder between the rows and not in the rows. Deep
can easily develop and set seed on broadleaf tillage with implements that invert the soil
weeds in these crops. can greatly reduce emergence of recently
134 CIENCIA E INVESTIGACION AGRARIA
shed seed (Parker and Riches, 1993), but
may bring previously deep buried seed to
the soil surface. In cranberry, burying C.
gronovii seed with 2.5 cm of sand reduced
infestation, indicating emergence was
primarily in the superficial soil layer
(Sandler et al., 1997). Late season tillage
may facilitate the distribution of vegetative
twigs of dodder from an infested row to
other plants, and should be avoided. Dodder
often spreads beyond the typical crop row
by late season and tillage can break off
vegetative pieces of dodder which can attach
to non-infected plants if they end up in
close proximity to these plants.
Removal by hand crews, of the crop with
dodder attached, remains a viable but
expensive option, when infestations are
small patches. When infestations are
extensive, hand removal is prohibitively
expensive, not to mention the loss in the
crop stand. If done, hand removal should
be done when dodder is first detected to
prevent spread and seed production. In a
trial area, farm workers were observed to
remove approximately 90% of the attached
dodder (Lanini, 2004). The remaining 10%
was generally missed, because it was too
small to be easily detected. If a hand crew
can be sent back through a field about 15
to 21 days after the first hand weeding, the
remaining dodder plants can be removed
prior to any dodder seed production. Dodder
can reattach to a new host if left in close
proximity to living crop plants, but if the
plants that are removed are moved six or
more inches from the remaining crop plants,
the dodder will not be able to reach a new
host. Dodder seedlings can only survive a
few days without a host, however dodder
stems which are attached to a host plant,
have been known to survive for several
weeks after being removed from the host,
before drying. These twigs are able to attach
to a new host plant and rapidly produce
haustoria (Dawson, 1984).
RESISTANCE
Although not much effort has been made
to assess variety differences to dodder
attachment or damage, some resistance has
been observed among sensitive crops.
Different levels of resistance to C.
pentagona were observed within four wild
Lycopersicon species (Al-Menoufi and
Ashton, 1991). Incompatible interactions
between Cuscuta species and
resistant/tolerant tomato plants have been
reported by Loffler et al., (1995). They
described incompatible interactions between
30 tomato varieties and C. reflexa. This
resistance was characterized by enlargement
of epidermis, hypodermis and collenchyma
of tomato cells, necrotic tissue developing
around the pre-haustoria and hardening of
cell walls adjacent to the necrotic tissue,
preventing haustoria formation. Ihl and
Miersch (1996) observed a similar
resistance phenomenon in 22 tomato
varieties and four wild tomato species to
four Cuscuta species: C. reflexa, C.
japonica, C. odorata and C. europaea. In
all cases, no functional haustoria were
formed due to hypersensitive response of
external cell layers of the tomato stems or
petioles after pre-haustoria contact. The
layer of dead tomato cells prevented
intrusion of initial haustoria, followed by
an increase in peroxidase activity.
Nevertheless, there are contradicting reports
of resistance of tomato to field dodder.
Nemli (1987) found resistance to C.
pentagona (C. campestris) in all five tomato
varieties tested, however, Ashton and
Santana (1976), Hutchison and Ashton
(1980) and Nir et al., (1996) reported that
all commercial tomato varieties were
seriously attacked by C. pentagona. These
contradictions may be due to variation in
the tomato varieties used in the studies or
variation in C. pentagona populations.
Recently, Miranda-Sazo (2003) observed
different levels of virulence among several
 VOL 32 N3 SEPTEMBER - DECEMBER 2005.
populations of C. pentagona, perhaps
partially explaining the contradictions in
the reports on resistant tomato varieties. In
an examination of 33 commercial tomato
varieties, sensitivity to a highly virulent C.
pentagona population varied considerably
(Goldwasser et al., 2001; Lanini, 2004).
Thus far, six commercially available tomato
varieties have been identified as dodder
resistant Heinz 9492, Heinz 9553, Heinz
9992, Heinz 9888, Heinz 1100, and
Campbells CXD 233. Some dodder plants
are able to attach and survive on these
varieties, but generally, tomato yields are
not reduced and dodder seed production is
very low or non-existent. The Heinz
varieties were originally developed to be
bacterial canker (Clavibacter michiganensis
subs. michiganesis) resistant, and thus, the
mechanism of disease resistance could be
what prevents dodder growth.
BIOLOGICAL CONTROL
The potential of using insects and pathogens
for dodder control has been reviewed
(Parker, 1991). In spite of the optimism,
there is still much research to be done before
any commercial biological agent for dodder
control can be released (Bewick et al.,
1987). Several fungi have been found that
damage dodder, including Fusarium
tricinctum and Alternaria spp. which attack
swamp dodder (C. gronovii) and A.
alternata and Geotrichum candidum which
both attack field dodder (C. pentagona).
Additionally, researchers in China have
found that a conidia suspension of
Colletotrichum gloeosporioides has
provided selective control of C. chinensis
and C. australis in soybeans. In field tests
with Alternaria destruens, Bewick et al.,
(2000) reduced C. gronovii attack by 90%
in cranberry (Vaccinium macrocarpon) and
carrot (Daucus carota), but it was not
effective against C. pentagona under
California conditions, possibly due to the
135
dry climate and the different dodder species
(Lanini, 2004). Difficulty in culturing and
efficacy under field conditions has limited
the use and commercialization of fungi.
Solarization has also been tested and found
to not be effective, as dodder has a hard
seed coat.
CHEMICAL CONTROL
The use of herbicides to control dodder has
been intensively studied worldwide.
Treatments may be divided according to
the timing of application; applications made
prior to attachment to the host or
applications made after attachment.
Dodder control prior to attachment
This approach attempts to control dodder
before it attaches to the host and eliminate
any possible damage (Parker, 1991).
Fumigants, such as methyl bromide, or
metam sodium controls many weeds prior
to vegetable planting, but species with a
hard seed coat, such as C. pentagona, are
not affected (Lanini, 2004). However,
Ashton and Santana (1976) reported some
success in controlling dodder ahead of
transplanting tobacco. However, since
transplants were used, it is difficult to
determine if the fumigant was effective or
if the transplants were too large for dodder
to successfully attach.
There are many preemergence (PRE)
herbicides that have been shown to affect
germinating dodder seedlings. These
herbicides are applied to the soil either
before seeding or before emergence of the
crop. When these herbicides are used,
dodder seeds germinate but initial growth
is reduced and attachment to the host fails.
Chloropropham (CIPC), a carbamate
herbicide, was the first soil-applied
herbicide used for dodder control in various
crops (Lee and Timmons, 1956).
Chloropropham is no longer used, however,
136 CIENCIA E INVESTIGACION AGRARIA
pronamide, a benzamide herbicide, is now
used to prevent dodder attachment in alfalfa,
sugarbeet, cranberry (Bewick et al., 1989)
and onion (Rubin, 1990). It is typically
used at 1-2 kgha-1, of active ingredient (a.i.)
and its persistence is longer than
chloropropham. Pronamide is also used in
sugarbeets in Chile as a sequential
application for control of dodder, with 1
kgha-1 applied at cotyledon to first true leaf
sugarbeets and a second and possibly a
third application at the same rate at 20 day
intervals. DCPA applied at 7 to 14 kgha-1
(a.i.), provides selective dodder control in
carrot (Shlevin and Golan, 1982),
ornamentals, and onion. Unlike
chloropropham and pronamide, DCPA is
non-volatile and presumably acts on dodder
by direct contact (Bayer et al., 1965).
Dinitroaniline herbicides have been
examined for their effectiveness to control
dodder in alfalfa (Orloff and Cudney, 1987).
Although they differ in their persistence of
dodder control, all dinitroanilines reduced
dodder infestation. Pendimethalin and
prodiamine were more effective than
trifluralin and controlled dodder for a longer
period of time. Prodiamine, applied at high
rate (2.6 kgha-1 (a.i.)), is the longest lasting
of the dinitroaniline herbicides, but its
persistence restricts its use only to
ornamental and non-crop areas. Trifluralin,
at fairly high rates (22.5 kgha-1 of the TR-
10 granular formulation), is also used for
selective dodder (C. indecora and C.
pentagona) control in alfalfa. Trifluralin is
surface applied in a granular formulation
in alfalfa, which concentrates the herbicide
near the surface where dodder seed
germinates. Using the liquid formulation
of trifluralin has not been successful in
controlling field dodder in tomatoes and
other crops, as trifluralin on or near the soil
surface degrades rapidly, and thus the
shallow germination depth and lack of roots
permits dodder to escape control.
Pendimethalin is also recommended for
selective dodder control in carrot, onions,
and alfalfa (Orloff and Cudney, 1987).
Pendimethalin is less volatile than trifluralin
and thus incorporation is not necessary, but
overhead irrigation or rainfall improves
activity (Parker and Riches, 1993).
Various herbicides including ethofumesate
at 2.2 kgha-1 (a.i.), were also reported to
control dodder selectively in sugarbeet
(Orloff and Cudney, 1987). Ethofumesate
can be used both pre and postemergence,
and is selective in sugarbeet at rates of 0.25
to 2.0 kgha-1 (a.i.), (Foschi and Rapparini,
1977). Dichlobenil is used to prevent dodder
growth in ornamentals and blueberries.
Thiazopyr (thiazole herbicide), which is
currently recomended in tree and vine crops,
has been shown to stunt dodder seedlings
and prevent attachment to host crops, such
as alfalfa. Following thiazopyr treatment,
field dodder seedling emergence is delayed,
and seedling length is reduced, while width
is increased. The seedlings are not able to
circumnutate normally and attachment to
a host does not occur.
P o s t - a t t a c h m e n t d o d d e r c o n t ro l
Parker (1991) stated that post-attachment
control of a parasite is very important in
order to reduce its seed production and further
spread, although irreversible damage was
already done. Rapid eradication of dodder
patches should be done when first detected
in the field before they have a chance to
produce seeds. In alfalfa, Cudney et al.
(1992) suggested spraying both host and
parasite, with a contact herbicide, such as
paraquat or by searing with a flame-throwing
torch or hand burner. In infected alfalfa for
hay, flail mowing or burning can be effective
(Orloff and Cudney, 1987). In tomatoes or
other row crops with new or small
infestations, hand roguing host plants with
attached dodder while they are small, before
many plants are infected can be a reasonable
 VOL 32 N3 SEPTEMBER - DECEMBER 2005.
practice. Patches should be marked with a
flag or other method, and examine the fields
again two or three weeks later to insure all
dodder was removed. Diquat, a contact
herbicide, has been used for dodder control
in alfalfa and clovers (Gimesi, 1966). Diquat
application following forage harvest limits
the crop foliage that is damaged, since both
crop and dodder are desiccated by this
treatment. Paraquat is used in a similar way
to diquat, but generally alfalfa is less tolerant,
and thus its use is typically limited to spot
treatment of small patches (Cudney and
Lanini, 2000).
The nature of attachment and association
between host and parasite requires a highly
selective herbicide to destroy the attached
dodder without crop damage.
Postemergence applications of herbicides
such as ethofumesate, pronamide, and
pendimethalin can suppress the parasite,
but dodder generally recovers (Orloff and
Cudney, 1987). It was suggested that the
low transpiration rate of dodder may limit
movement of xylem mobile, soil applied
herbicides into the parasite, and thus, should
not be used for dodder control (Fer, 1984).
The author argued that xylem-mobile
compounds move and accumulate mostly
in the host organs with high transpiration
rate and may damage the host more than
the parasite. As expected, Liu and Fer (1990)
found that pendimethalin, an 'ambimobile'
herbicide, applied to mung beans
(Phaseolus aureus) roots does not move to
the attached dodder via the transpiration
stream. However, when pendimethalin was
applied to one leaf, 56% of the herbicide
was accumulated in the parasite.
A phloem-mobile herbicide applied
broadcast to the host plant should
accumulate selectively in the parasite
because of its stronger sink (Nir et al.,
1996). Bewick et al., (1991) reported that
14
C-labeled glyphosate foliar applied to
137
carrot plants infected with swamp dodder
accumulated in the parasite tissue more than
in any part of the host plant. Along this line,
studies have been conducted using phloem-
mobile herbicides, such as glyphosate, for
selective post-attachment control of dodder
in alfalfa (Liu and Fer, 1990; Dawson,
1989a). No damage to alfalfa was observed
following treatment with low doses of
glyphosate (75 to 150 gha-1(a.i.) when both,
parasite and host, were still vegetative and
vigorously growing (Dawson, 1989b).
Similarly, glyphosate at 400 gha-1(a.i.)
applied late in the life cycle of carrot when
swamp dodder was in full flower,
satisfactorily controlled the parasite and
increased carrot root yield (Bewick et al.,
1988). Unfortunately, host crops can be
injured by glyphosate treatment (Orloff and
Cudney, 1987) and the parasite may not be
adequately controlled (Frolisek, 1987).
Several acetolactate synthase (ALS)
inhibiting herbicides have been shown to
control or suppress dodder. When imazaquin
is applied to dodder-infected soybean and
mung beans plants, the herbicide
accumulates in the apical part of the dodder
stem (Liu et al., 1991). It has also been
shown that imazethapyr and thiazopyr
(thiazole herbicide) applied to carrot plants
infested with dodder were less damaging
than when applied to non-infested carrot,
indicating the potential of using low rates
of non-selective herbicides for selective
control of parasitic weeds (Nir et al., 1996).
Possibly the herbicide accumulates
selectively in the dodder due to it strong sink
activity. Imazethapyr at 100-150 gha-1 (a.i.)
applied preemergence reduces dodder
infestations in seedling alfalfa and
suppression of more established dodder
stems (Cudney and Lanini, 2000).
Rimsulfuron applied at 35 gha -1 (a.i.)
suppresses dodder in tomatoes, particularly
when split applications are used (Mullen
et al., 1998). Further studies have shown
138 CIENCIA E INVESTIGACION AGRARIA
that treatments made soon after dodder
attachment were more effective than
applications made after dodder became well
established. However, the season-long
control achieved by the best rimsulfuron
treatments was only about 50% and differed
substantially in different fields, possibly
indicating dodder population differences
(Lanini, 2004). Sulfosulfuron applied at 20
to 50 gha-1 (a.i.) was shown to be very
effective at controlling attached dodder in
direct seeded tomatoes, with excellent
selectivity (Eizenberg et al., 2003), however,
it is only registered for use in wheat in the
USA.
Introduction of transgenic crops resistant
to phloem-mobile broad-spectrum
herbicides to selectively control parasitic
weeds was suggested for dodder control
(Rubin, 1991). However, Nadler-Hasser
and Rubin (2003) found that dodder
growing on transgenic glyphosate resistant
sugarbeet was not controlled when treated
with glyphosate (1.08 kgha-1(a.i)), nor when
sulfometuron resistant tomatoes were
treated with sulfometuron (22.5 gha-1 (a.i.)).
Dodder may be obtaining amino acids from
the resistant host plant and thus are able to
survive these treatments. Since dodder, in
a sense, is just another stem on the host
plant, we believe that some crop injury may
be necessary to achieve dodder control with
herbicides. Thus, rates higher than are
currently registered for use may be needed,
regardless of the herbicide used.
CONCLUSIONS
The most successful control of dodder
involves a systematic approach whereby
several methods of control are used together.
Effective management requires control of
any new populations, prevention of seed
production and monitoring in subsequent
years to insure complete control. Where
more extensive infestations exist, rotation
to non-host crops for several years may be
required. Once a host crop is planted again,
treatments must be made to contain escaped
plants.
RESUMEN
La cscuta (Cuscuta spp.) es un parsito
foliar y de los tallos, que infecta muchos
cultivos de hoja ancha, ornamentales y
malezas y slo algunos cultivos
monocotiledneas. Vive completamente en
la planta hospedera, reduciendo el
crecimiento y los rendimientos. La
prevencin por medio de rotacin de
cultivos con especies no hospederas,
retrasando la fecha de siembra hasta el
otoo en el caso de la remolacha azucarera
(Beta vulgaris), uso de cultivares resistentes
y el uso de herbicidas de pre emergencia,
han demostrado ser tiles en ciertos cultivos.
Una vez que la cscuta ataca el cultivos,
se registrarn prdidas de rendimiento,
independientemente del mtodo de control
aplicado, siendo en estas circunstancias,
muy difcil disponer de mtodos selectivos.
El control luego de la fijacin de la cscuta
a la planta a menudo requiere de la
destruccin o de daos muy severos en la
planta hospedera para evitar la dispersin
de la cscuta a las plantas vecinas. Sin
embargo, existen varios herbicidas que han
demostrado selectividad permitiendo
detener el ataque de cscuta, pero rara vez
se obtienen un total control. La cscuta
adherida a plantas modificadas
genticamente, en cultivos resistentes a
herbicidas, no ha sido exitosamente
controlada con tratamientos herbicidas,
indicando que estos cultivos permiten una
solucin parcial al problema. El control de
la cscuta requiere de una estrategia de
control integrado aplicada por varios aos.
Palabras clave: Cabello de ngel, Cuscuta,
plantas parsitas, resistencia, manejo
de maleza.
 VOL 32 N3 SEPTEMBER - DECEMBER 2005.
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